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SMITH, Eric Howard, 194-3- SYSTEMATIC REVISION OF THE MACULATE SPECIES OF THE GENUS PHYLLOTRETA CHEV. OF AMERICA NORTH OF MEXICO (COLEOPTERA: CHRYSOMELIDAE, ALTICINAE).

The Ohio State University, Ph.D., 1973 Entomology

University Microfilms, A XEROX Company , Ann Arbor, Michigan SYSTEMATIC REVISION OF THE MACULATE SPECIES OF THE

GENUS PHYLLOTRETA CHEV. OF AMERICA NORTH OF MEXICO

(COLEOPTERA: CHRYSOMELIDAE, ALTICINAE)

DISSERTATION

Presented in Partial Fulfillment of the Requirements for the Degree Doctor of Philosophy in the Graduate School of The Ohio State University

By

Eric Howard Smith, B.A., M.S.

*****

The Ohio State University

1973

Reading Committee: Approved by

Dr. Donald J. Borror

Dr. Charles A. Triplehorn Adviser Dr, Barry D. Valentine Department of Zoology tr

TABLE OF CONTENTS

Page

ACKNOWLEDGEMENTS ...... iv

VITA ...... vi

LIST OF F I G U R E S ...... viii

LIST OF ABBREVIATIONS ...... xv

INTRODUCTION ...... 1

METHODS ...... 3

CLASSIFICATION ...... 8

Subfamily Alticinae ...... 8

Genus Phyllotreta ...... 8

Key to the males of the maculate sp ec i e s...... 18

Key to the females of the maculate sp e cies...... 27

Phyllotreta arcuata n.sp...... 36

Phyllotreta armoraciae (Koch) ...... 39

Phyllotreta attenuate n.sp...... 46

Phyllotreta bipustulata (F.) 51

Phyllotreta bisinuata n . s p ...... 61

Phyllotreta constricta n .sp...... 64

Phyllotreta decipiens Horn ...... 69

Phyllotreta denticomis H o r n ...... 74

Phyllotreta dolichophalla n.sp...... 78

Phyllotreta emarginata n.sp...... 81

ii TABLE OF CONTENTS ( C o n t ,)

i Page

Phyllotreta lepidula (LeC.) ...... 86

Phyllotreta liebecki Schffr...... 90

Phyllotreta oblonga Chttn...... 95

i ' Phyllotreta Oregonensis (Crotch) ...... 98

Phyllotreta ramosa (Crotch) ...... 104 " T_ I ...... i | Phyllotreta ramosoides n.sp...... 107

| Phyllotreta rohusta (LeC.) ...... 113

| ...... Phyllotreta spatulata n.sp...... 117 ! j Phyllotreta striolata (F.) 121

I ! Phyllotreta utana Chttn...... 135 | ! Phyllotreta utanula n.sp...... 139 i Phyllotreta zimmermanni (Crotch) ...... 143

Species considered nomina dubia ...... 153 i FIGURES ...... 155

LITERATURE CITED ...... 21S

iii ACKNOWLEDGEMENTS

Assistance in the form of the opportunity to examine and the loan of specimens has been extended by many individuals and institutions. I would like to particularly thank the following who have permitted examination of Phyllotreta under their care: Lee H.

Herman, Jr. (AMNH), W. Wayne Moss and David C. Rentz (ANSP), Melville

Hatch (BMUW), Hugh B. Leech (CASC), Terry N. Seeno (CDAE), Edward C.

Becker (CNCI), L. L. Pechpman (CUIC), Phillip J. Clausen (DEFW),

Brett C. Ratcliffe (DEUN), W. J. Hanson (EMUS), Edward U. Balsbaugh,

Jr. (EUBC and SDSU), Henry Dybas (FMNH), George C. Wallace (ICCM), the late Jean L. Laffoon (ISUI), Charles L. Hogue and Roy R. Snelling

(LACM), Larry Burgess (LBCC), Richard E. Morel and Jan White (MCZC),

Charles A. Triplehorn (OSUC), Paul Oman (OSUO), Edward E. Simons

(PADA), Ke Chung Kim (PSUC), E. Phil Rouse (UADE), Saul Frommer (UCRC)

F. W. Wood (UMDC), Eben A. Osgood, Jr. (UMDE), Richard E. White (USNM)

Lutz J. Bayer (UWEM), Vernon M. Kirk (VMKC), Michael Kosztarab (VPIC),

William J. Turner (WSUC), Jan Edelmann (University Zoological Museum,

Copenhagen), and F. Hieke (Zoological Museum of Humboldt University,

Berlin, DDR).

I would like to thank the members of my reading committee,

Donald J. Borror and Charles A. Triplehom, and my major professor,

Barry D. Valentine, for their assistance, encouragement, and personal friendship extended to me throughout my stay at Ohio State. Special appreciation is extended to Dr. Valentine for his guidance, help in solving many nomenclatorial problems, his patience, and his personal faith in me and support throughout our association.

My sincere thanks are extended to Dr. John Briggs, Dr. Frank

Fisk, and especially to Dr. Rodger Mitchell for th£ use of equipment under their care, to Dr. Triplehom for my constant use of the Ohio

State Collection and museum facilities, and to Dr. Donald E. Johnston for help with the variation analysis work.

And lastly, I would like to thank my wife, Cheryl, and my sister,

Margaret, for their help in recording label data and preparation of the distribution maps.

v VITA

July 4, 1943 . . . B o m - Cincinnati, Ohio

1966 ...... B.A., Miami University, Oxford, Ohio

1966-196 9 ...... Research Assistant. Department 'of Entomology, Purdue University, Lafayette, Indiana

1967-196 9 ...... Member of the Publication Board of the "Coleopterists* Bulletin"

1968-1969. .... Business Manager of the "Coleopterists* Bulletin"

1969-197 2 ...... Teaching Associate. Department of Entomology, The Ohio State University, Columbus, Ohio

1970 ...... M.S., Purdue University, Lafayette, Indiana

1970-197 3...... Graduate Student Member of the Graduate Committee. Department of Entomology, The Ohio State University, Columbus, Ohio

1971 ...... Member of the Council, Coleopterists Society

1971 ...... NSF Summer Traineeship for Graduate Teaching Associates. The Ohio State University, Columbus, Ohio

1972 ...... Summer Research Associate. Department of Entomology, The Ohio State University, Columbus, Ohio

1972-1973...... University Dissertation Year Fellowship. The Ohio State University, Columbus, Ohio

1973 ...... Contributing Author, North American Fauna Project

PUBLICATIONS

1968. Young, R. M . , E. H. Smith, E. C. Mignot, et al. A partial list of active Coleopterists, by specialty. Coleopterists* Bull. Suppl., 28 pp. 1969. Arnett, R, H., Jr., E. C. Mignot, and E. H. Smith. North American Coleoptera fauna: notes on Pyrophorinae, Elateridae. Coleopterists* Bull. 23(1):9-15.

1969. Arnett, R. H., Jr. and G. A. Samuelson (eds.), G. F. Flory, E. C, Mignot, C. D. Schaaf, and E. H. Smith. A directory of Coleoptera collections of North America. Purdue Univ. (C. S. C.) 123 pp.

1969. Smith, E. H. Book review: World Crop Production, volume I 6 II.' Coleopterists* Bull. 23(3):88.

MAJOR FIELD

Entomology

vii LIST OF FIGURES

Figure Page

1. Male 5th abdominal sternum and pygidium, ventral v i e w ...... 156

2. Female 5th abdominal sternum, ventral vi e w ...... 156

3. Label of lectotype of Phyllotreta armoraciae ...... 156

4. Male genitalia with structures labeled, dorsal v i e w ...... 156

5. Male genitalia with structures labeled, lateral view ...... 156

6. Female spermatheca with structures labeled, lateral v i e w ...... 156

7. Phyllotreta arcuata, holotype, dorsal view ...... 158

8. Phyllotreta armoraciae, dorsal view ...... 158

9. Phyllotreta attenuata, holotype, dorsal view ...... 158

10. Phyllotreta bipustulata, dorsal v i e w ...... 158

11. Phyllotreta bipustulata, right elytron showing color pattern variation, dorsal v i e w ...... 158

12. Phyllotreta bisinuata, holotype, dorsal view ...... 160

13. Phyllotreta constricta, holotype, dorsal view ...... 160

14. Phyllotreta decipiens, dorsal view ...... 160

15. Phyllotreta denticomis, dorsal v i e w ...... 160

16. Phyllotreta denticornis, right elytron showing color pattern variation, dorsal v i e w ...... 160

17. Phyllotreta dolichophalla, holotype, dorsal view ...... 162

viii LIST OF FIGURES CCont.)

Figure Page

18. Phyllotreta emarginata, holotype, dorsal view ...... 162

19. Phyllotreta' lepidula, dorsal view ...... 162

20. Phyllotreta liebecki, holotype, dorsal view ...... 162

21. Phyllotreta oblonga, holotype, dorsal view ...... 164

22. Phyllotreta oregonensis, lectotype, dorsal view ...... 164

23. Phyllotreta oregonensis, right elytron showing color pattern variation, dorsal ...... 164

24. Phyllotreta ramosa, dorsal v i e w ...... 164

25. Phyllotreta ramosoides, holotype, dorsal view ...... 164

26. Phyllotreta robusta, dorsal view ...... 166

27. Phyllotreta robusta, right elytron showing color pattern variation, dorsal vi e w ...... 166

28. Phyllotreta spatulata, holotype, dorsal view ...... 166

29. Phyllotreta striolata, dorsal view ...... 166

30. Phyllotreta striolata, right elytron showing color pattern variation, dorsal vi e w ...... 166

31. Phyllotreta striolata, right elytron showing color pattern variation, dorsal vi e w ...... 166

32. Phyllotreta striolata, right elytron showing color pattern variation, dorsal view ...... 166

33. Phyllotreta striolata, right elytron showing color pattern variation, dorsal view ...... 168

34. Phyllotreta striolata, right elytron showing color pattern variation, dorsalvi e w ...... 168

35. Phyllotreta striolata, right elytron showing color pattern variation, dorsal view ..... 168 LIST OF FIGURES CCont.)

Figure Page

36. Phyllotreta utana, dorsal v i e w ...... 168

37. Phyllotreta utamila, holotype, dorsal view ...... 168

38. Phyllotreta zimmermanni, dorsal vi e w ...... 168

39. Phyllotreta zimmermanni, right elytron showing color pattern variation, dorsal view ...... 168

40. Male antenna of Phyllotreta arcuata, holotype, anterodorsal view of right ...... 170

41. Female antenna of Phyllotreta arcuata, allotype, anterodorsal view of right ...... 170

42. Male antenna of Phyllotreta armoraciae, anterodorsal view of right ...... 170

43. Female antenna of Phyllotreta armoraciae, anterodorsal view of right ...... 170

44. Male antenna of Phyllotreta attenuata, holotype, anterodorsal view of right ...... 170

45. Female antenna of Phyllotreta attenuata, allotype, anterodorsal view of.right ...... 170

46. Male antenna of Phyllotreta bipustulata, anterodorsal view of right ...... 170

47. Female antenna of Phyllotreta bipustulata-, anterodorsal view of right ...... 170

48. Male antenna of Phyllotreta bisinuata, holotype, anterodorsal view of.right ...... 170

49. Female antenna of Phyllotreta bisinuata, allotype, anterodorsal view of right ...... 170

50. Male antenna of Phyllotreta constricta, holotype, anterodorsal view of left...... 170

51. Female antenna of Phyllotreta constricta, allotype, anterodorsal view of right ...... 172

x LIST OF FIGURES (Cont.'j

Figure Page

52. Male antenna of Phyllotreta decipiens, anterodorsal view of right ...... 172

53. Female antenna of Phyllotreta decipiens, anterodorsal view of right ...... 172

54. Male antenna of Phyllotreta denticomis, anterodorsal view of right ...... 172

55. Male antennal segments 5-8 of Phyllotreta denticomis, anterior view of right ...... 172

56. Female antenna of Phyllotreta denticomis, anterodorsal view of r i g h t ...... 172

57. Male antenna of Phyllotreta dolichophalla, holotype, anterodorsal view of right ...... 172

58. Female antenna of Phyllotreta dolichophalla, allotype, anterodorsal view of l e f t ...... 172

59. Male antenna of Phyllotreta emarginata, holotype, anterodorsal view of right’"." ...... 172

60. Female antenna of Phyllotreta emarginata, allotype, anterodorsal view of r i g h t ...... 172

61. Male antenna of Phyllotreta lepidula, anterodorsal view of right ...... 172

62. Female antenna of Phyllotreta lepidula, anterodorsal view of right ...... 172

63. Male antenna of Phyllotreta liebecki, holotype, anterodorsal view of right ...... 174

64. Male antennal segments 4-6 of Phyllotreta liebecki, holotype, anterior view of right ...... 174

65. Female antenna of Phyllotreta liebecki, anterodorsal view of right ...... 174

66. Male ant’eiiina of Phyllotreta oblonga, anterodoisal. view of right ...... 174 LIST OF FIGURES (Cont.)

Figure Page

67. Female antenna of Phyllotreta obloriga, holotype, anterodorsal vievr of ri g h t ...... 174

68. Male antenna of Phyllotreta oregonensis, anterodorsal view of ri g h t ...... 174

69. Female antenna of Phyllotreta oregonensis, anterodorsal view of ri g h t ...... 174

70. Male antenna of Phyllotreta raniosa, anterodorsal view of right ...... 174

71. Female antenna of Phyllotreta ramosa, anterodorsal view of right ...... 174

72. Male antenna of Phyllotreta ramosoides, holotype, anterodorsal view of right ...... 174

73. Female antenna of Phyllotreta ramosoides, allotype, anterodorsal view of right ...... 174

74. Male antenna of Phyllotreta robusta, anterodorsal view of right ...... 176

75. Female antenna of Phyllotreta robusta, anterodorsal view of right ...... 176

76. Male antenna of Phyllotreta spatulata, holotype, anterodorsal view of right ...... 176

77. Female antenna of Phyllotreta spatulata, allotype, anterodorsal view of right ...... 176

78. Male antenna of Phyllotreta striolata, anterodorsal view of right ...... 176

79. Female antenna of Phyllotreta striolata, anterodorsal view of right ...... 176

80. Male antenna of Phyllotreta utana, paratype, anterodorsal view of right ...... 176

81. Female antenna of Phyllotreta utana, paratype, anterodorsal view of right ...... 176

m * X ll LIST OF FIGURES CCont.)

Figure Page

82. Male antenna of Phyllotreta utqjjula, holotype, anterodorsal view of right ...... 176

83. Male antenna o f 'Phyl16treta ziriunermanni, anterodorsal view of right ...... 176

84. Female antenna of Phyllotreta zimmermanni, anterodorsal view of right ...... 176

85. Male genitalia of Phyllotreta arcuata, paratype, dorsal v i e w ...... 178

86. Male genitalia of Phyllotreta arcuata, paratype, lateral view . ^ I ...... 178

87. Male genitalia of Phyllotreta armoraciae, dorsal v i e w ...... 178

88. Male genitalia of Phyllotreta armoraciae, lateral v i e w ...... 178

89. Male genitalia of Phyllotreta attenuata, holotype, dorsal v i e w ...... 178

90. Male genitalia of Phyllotreta attenuata, holotype, lateral view...... 178

91. Male genitalia of Phyllotreta bipustulata, dorsal v i e w ...... 178

92. Male genitalia of Phyllotreta bipustulata, lateral v iew ...... 178

93. Male genitalia of Phyllotreta bisinuata, paratype, dorsal v i e w ...... 178

94. Male genitalia of Phyllotreta bisinuata, paratype, lateral view ...... 178

95. Male genitalia of Phyllotreta constricta, paratype, dorsal view I I I ...... 178

96. Male genitalia of Phyllotreta constricta, paratype, lateral view ...... 178

xiii LIST OF FIGURES CCont.)

Figure Page

97. Male genitalia of Phyllotreta decipiens, dorsal view ...... 180

98. Male genitalia of Phyllotreta decipiens, lateral v i e w ...... 180

99. Male genitalia of Phyllotreta denticomis, dorsal v i e w ...... 180

100. Male genitalia of Phyllotreta denticomis, lateral v i e w ...... 180

101. Male genitalia of Phyllotreta dolichophalla, holotype, dorsal v i e w ...... 180

102. Male genitalia of Phyllotreta dolichophalla, holotype, lateral view ...... 180

103. Male genitalia of Phyllotreta emarginata, paratype, dorsal view ...... 180

104. Male genitalia of Phyllotreta emarginata, paratype, lateral v i e w ...... 180

105. Male genitalia of Phyllotreta lepidula, dorsal v i e w ...... 180

106. Male genitalia of Phyllotreta lepidula, lateral v i e w ...... 180

107. Male genitalia of Phyllotreta liebecki, dorsal v i e w ...... 180

108. Male genitalia of Phyllotreta liebecki, lateral v i e w ...... 180

109. Male genitalia of Phyllotreta oblonga, dorsal v i e w ...... 182

110. Male genitalia of Phyllotreta oblonga, lateral v i e w ...... 182

111. Male genitalia of Phyllotreta oregonensis, dorsal v i e w ...... 182

xiv LIST OF FIGURES CCont.)

Figure Page

112. Male genitalia of Phyllotreta. oregonensis. lateral v i e w ...... 182

113. Male genitalia of Phyllotreta ramosa, dorsal v i e w ...... 182

114. Male genitalia of Phyllotreta ramosa, lateral v i e w ...... 182

115. Male genitalia of Phyllotreta ramosoides, holotype, dorsal v i e w ...... 182

116. Male genitalia of Phyllotreta ramosoides, holotype, lateral view . I ...... 182

117. Male genitalia of Phyllotreta robusta, dorsal v i e w ...... 182

118. Male genitalia of Phyllotreta robusta, lateral v i e w ...... 182

119. Male genitalia of Phyllotreta spatulata, holotype, dorsal view I I I ...... 182

120. Male genitalia of Phyllotreta spatulata, holotype, lateral view i I ...... 182

121. Male genitalia of Phyllotreta striolata, dorsal v i e w ...... 184

122. Male genitalia of Phyllotreta striolata, lateral v i e w ...... 184

123. Male genitalia of Phyllotreta utana, dorsal v i e w ...... 184

124. Male genitalia of Phyllotreta utana, lateral v i e w ...... 184

125. Male genitalia of Phyllotreta utanula, holotype, dorsal view I ^ I ...... 184

126. Male genitalia of Phyllotreta utanula, holotype, lateral view . II ...... 184

xv LIST OF FIGURES CCont.)

Figure Page

127. Male genitalia of Phyllotreta zirirniermarini. dorsal view ...... 184

128. Male genitalia of Phyllotreta zimmermarini, lateral v i e w ...... 184

129. Female spermatheca of Phyllotreta arcuata, paratype, lateral v i e w ...... 186

130. Female spermatheca of Phyllotreta armoraciae, lateral v i e w ...... 186

131. Female spermatheca of Phyllotreta attenuata, allotype, lateral view ...... 186

132. Female spermatheca of Phyllotreta bipustulata, lateral v i e w ...... 186

133. Female spermatheca of Phyllotreta bisinuata, allotype, lateral v i e w ...... 186

134. Female spermatheca of Phyllotreta constricta, paratype, lateral v i e w ...... 186 • • 135. Female spermatheca of Phyllotreta decipiens, lateral v i e w ...... 186

136. Female spermatheca of Phyllotreta denticomis, lateral v i e w ...... 186

137. Female spermatheca of Phyllotreta do1ichophalla, allotype, lateral view ...... 186

138. Female spermatheca of Phyllotreta emarginata, paratype, lateral v i e w ...... 186

139. Female spermatheca of Phyllotreta lepidula, lateral view ...... 188

140. Female spermatheca of Phyllotreta liebecki, lateral v i e w ...... 188

141. Female spermatheca of Phyllotreta oblonga, holotype, lateral view ...... 188 LIST OF FIGURES CCont.)

Figure Page

142. Female spermatheca of Phyllotreta oregonensis. lateral v i e w ...... 188

143. Female spermatheca of Phyllotretaramosa, lateral v i e w ...... I88

144. Female spermatheca of Phyllotreta ramosoides, paratype, lateral view ...... I88

145. Female spermatheca of Phyllotreta robusta, lateral v i e w ...... 188

146. Female spermatheca of Phyllotreta spatulata, paratype, lateral view ...... I88

147. Female spermatheca of Phyllotreta striolata, lateral v i e w ...... 1®®

148. Female spermatheca of Phyllotreta utana, lateral v i e w ...... I®8

149. Female spermatheca of Phyllotreta zimmermanni, lateral v i e w ......

150. Distribution of Phyllotreta ar c u a t a ...... 1®9

151. Distribution of Phyllotreta armoraciae ...... 190

152. Distribution of Phyllotreta attenuata ...... 191

153. Distribution of Phyllotreta bipustulata ...... 192

154. Distribution of Phyllotreta bisinuata ...... 193

155. Distribution of Phyllotreta constricta ...... 194

156. Distribution of Phyllotreta decipiens ...... 195

157. Distribution of Phyllotreta denticomis ...... 196

158. Distribution of Phyllotreta dolichophalla ...... 197

159. Distribution of Phyllotreta emarginata ...... 198

xvii LIST OF FIGURES CCont.)

Figure Page

160. Distribution of Phyllotreta lepidula ...... 199

161. Distribution of'Phyllotreta ~liebecki...... 200

162. Distribution of Phyllotreta oblonga ...... 201

163. Distribution of Phyllotreta oregonensis ...... 202

164. Distribution of Phyllotreta ramosa ...... 203

165. Distribution of Phyllotreta ramosoides ...... 204

166. Distribution of Phyllotreta robusta ...... 205

167. Distribution of Phyllotreta spatulata ...... 206

168. Distribution of Phyllotreta striolata...... 207

169. Distribution of Phyllotreta utana ...... 208

170. Distribution of Phyllotreta utanula ...... 209

171. Distribution of Phyllotreta zimmermanni ...... 210

172. Phyllotreta alberta, holotype, dorsal view ...... 212

173. Phyllotreta obtusa, holotype, dorsal view ...... 212

174. Phyllotreta perspicua, holotype, dorsal view ...... 212

175. Female antenna of Phyllotreta alberta, holotype, anterodorsal view of left I ...... 212

176. Female antenna of Phyllotreta obtusa, holotype, anterodorsal view of left I ...... 212

177. Female antenna of Phyllotreta perspicua, holotype, anterodorsal view of right ...... 212

178. Female spermatheca of Phyllotreta alberta, holotype, lateral vi e w ...... 212

179. Female spermatheca of Phyllotreta obtusa, holotype, lateral view ...... 212

xviii LIST OF FIGURES CCont.)

Figure Page

180. Female spermatheca of Phyllotreta perspicua. holotype, lateral view ...... 212 181. Phenograms representing color pattern variation of 14 samples of Phyllotreta striolata; information given: state or province, n - sample size, X = mean, V = coefficient of variation...... 214

xix LIST OF ABBREVIATIONS1

AMNH American Museum of Natural History

ANSP Academy of Natural Sciences at Philadelphia

BMUW University of Washington '

CASC California Academy of Science

CDAE California Department of Agriculture

CNCI Canadian National Collection of

CUIC Cornell University

DEFW University of Minnesota

DEUN University of Nebraska

EHSC Eric H. Smith Collection

EMUS Utah State University

EUBC Edward U. Balsbaugh, Jr. Collection

FMNH Field Museum of Natural History

ICCM Carnegie Museum

ISIU Iowa State University

LACM Los Angeles County Museum of Natural History

LBCC Larry Burgess Collection

MCZC Museum of Comparative

MSUC Michigan State University

*From Arnett and Samuelson (1969), except EUBC and LBCC were coined for collections not listed.

xx NCSU North Carolina State University

NMDC N. M. Downie Collection

OSUC Ohio State University

OSUO Oregon State University

PADA Pennsylvania Department of Agriculture

PSUC Pennsylvania State University

SDSU South Dakota State University

UADH University Arkansas

UCRC University of California, Riverside

UMDC University of Maryland

UMDE University of Maine

USNM United States National Museum

UWEM University of Wisconsin

VMKC Vernon M. Kirk Collection

VPIC Virginia Polytechnic Institute

WSUC Washington State University INTRODUCTION

The Alticinae (flea ) is the largest subfamily of the

Chrysomelidae, consisting of about 6,000 species; about 400 species occur in America north of Mexico. Except for the European and some

/ North American species, primarily those of economic importance, this group is very poorly known. In America north of Mexico 42 genera occur, of which only 7 (3 as recent unpublished theses) have been completely reviewed since 1889. The biology is unknown for the great majority of species. Thus, a great deal of basic systematic work remains to be done on North American Alticinae before more advanced studies can be undertaken.

The genus Phyllotreta was erected by Chevrolat in the 3rd edition of the Dejean Catalogue of Coleoptera (1837) and included 14 species,

2 occurring north of Mexico. Crotch (1873), the first American work of any note o i North America flea beetles, described all the species at the Academy of Natural Sciences of Philadelphia which included the

LeConte and Horn Collections. Chapuis (1875) was the first to present a complete classification of the Alticinae. He placed Phyllotreta in the group "Aphthonites” which was later considered of tribal rank. Horn

(1889) monographed the Alticinae of America north of Mexico, the only time this has been done. He slightly altered Chapuis* classification and placed Phyllotreta along with Longitarsus, Glyptina, and Aphthona in the group ”Aphthonae,r which is now the tribe Aphthonini, with the addition of Palaeothona.

Horn (1 8 8 9 ) presented the first really comprehensive revision of

Phyllotreta. The most recent revision was by Chittenden (1927) in which 17 new species and 5 new variations or subspecies were described.

Unfortunately, his key is very poor and his descriptions are somewhat inaccurate and inadequate. He studied neither the male nor female

/ genitalia; the former provide the critical characters for separation of many species.

This genus contains many species of economic importance (see the biology section for the genus and for each species). Biological work has recently been done and is being continued on several species by researchers at Cornell University [see Root (1969), Feeny, Paauwe, and

Demong (1970), Tahvanvainen (1972), and Hicks (1972)]..

The present work is a systematic revision of the genus Phyllotreta for all maculate species known to occur north of Mexico. A key to these species is presented. The biology for each species has been summarized and the present distribution is given. This revision is based on a study of about 8,000 specimens; in addition, representatives of most of the immaculate species, including their male genitalia, were studied. METHODS

Measurement s

For each species, the series of specimens used for measurements included those representing both the minimum and maximum length. The measurements were made in the following manner using a Leitz model

RS binocular dissecting scope with ocular grid:

1. Beetle length: sum of elytral length, see 6 below, and distance measured along midline from pronotal base to most anterior part of frontal carina.

2. Beetle width: measured on a line perpendicular to elytral suture at greatest breadth.

3. Interocular Distance/Maximum Diameter of eye: interocular distance measured on dorsal aspect of head between most median portions of each compound eye, and maximum diameter of eye measured on lateral aspect of head across maximum dimension of compound eye.

4. Antennal segment Clength/width}: measurements are given in ocular micrometer units and represent maximum length or width; only half units are estimated, if dimension was only slightly more or less than a unit, this is indicated by a plus or minus sign.

5. Pronotal length and width:length measured on dorsal midline from basal to anterior edge, and width measured on a line perpendicular to pronotal midline at greatest breadth. 4

6. Elytral length and width: length measured along dorsal line from base of scutellum along suture to apex of elytra; for width see

2 above.

7. Length of male genitalia; measured dorsally or laterally from base of basal piece to apex.

8. Length of female genitalia: measured in lateral view from most anterior part of sclerotized portion of spermathecal duct to posterior most extent of pump.

Genitalia Preparation

The beetles were first relaxed by placing them in a beaker of water just below the boiling point for about 2 minutes. Each was then transferred to a drop of water in an elliptically-grooved depression slide. The beetle was held with a pair of jeweler*s forceps, and a minuten-pin hooked probe was inserted between the metacoxae and the first visible abdominal segment and the abdomen was gently separated. The abdomen was placed in a beaker of concentrated KOH

(5 pellets/15 ml distilled water in 30 ml. beaker, using 2 glass boiling beads, and the liquid was not allowed to evaporate below 7.5 ml ) for about 3 minutes. The abdomen was then transferred back to the water in the grooved slide dorsum up and while holding it along the right margin with a pair of jeweler*s forceps, the left tergal margin was slit from base to apex with a minuten-pin scalpel (if upon opening up the abdomen the tissue was not fairly well dissolved the abdomen was briefly returned to the KOH for another minute or two).

The abdomen was then placed in acid alcohol for 1-2 minutes, then into a 95% ETOH (males) or distilled water (females) rinse for another

1-2 minutes. While in this rinse, the male genitalia or female sper- mathecae were separated from the tissue residue and dissected out using a pair of jeweler*s forceps and minuten-pin hooked probe. The abdomen was placed ventral surface up in a drop of LePage’s water- soluble glue on the point next to the corresponding specimen. / The male genitalia were transferred to glycerine for study and later to glycerine in a small size polyethylene genitalia vial which was attached to the pin below the corresponding remounted specimen.

All transfers of the genitalia alone were done using a minuten probe.

The female spermatheca was transferred to a diluted Hoyer*s mounting medium solution (2 drops Hoyer*s/4 drops distilled water in a

Plant Quarantine watch glass). This solution was allowed to evaporate at room temperature, while shielded from dust, until the consistency was that of the Hoyer's used for slide mounting (4-6 hours, or less if a slide drying tray is used). The spermatheca was then mounted in

Hoyer*s and allowed to dry for 5 days on a slide drying tray (about 45°C). All transfers of spermathecae alone were made using a dis­ posable pipette, except a minuten probe was used for the final transfer to the slide.* After study and/or illustration, the slide was soaked in distilled water to dissolve the Hoyer's. The spermatheca was transferred to a drop of LePage's water-soluble glue on a 3 x 7 mm card and covered with a square of cover glass (easily cut with a diamond pencil).

This card was then pinned below the corresponding specimen.

The procedure for male genitalia is a greatly modified version of

the method used by Arnett (1947) and that for the female genitalia is a greatly modified version of the technique suggested by Samuelson

C1972).

Illustration Preparation

Male genitalia were placed in a drop of glycerine in a shallow

circular depression slide necessary, 1 or 2 grains of white sand

were used for positioning). The general features were drawn using a

■Ken-A-Vision technical model microprojector and the details were added

using a Leitz RS scope. Female spermathecae were drawn with the aid

of a camera lucida equipped Wild M20 compound binocular scope. The

beetle drawings as well as those of the antennae and sterna were made

with the aid of a camera lucida equipped Wild M5 binocular dissecting

scope. The final illustrations of the above were made by transferring

the original drawing to #3 bristle board.

Distribution Maps

The maps represent the distribution of specimens personally

examined. The localities are recorded in 2 ways. When a locality

within a state is known, it is represented by a solid dot at this

locality. If only a state locality is known (and no other locality

within the state is known), it is represented by a star placed in

approximately the center of that state.

Systematic Formats

Key Format

The key to species is artificial and dichotomous, but a word is needed about the order of characters within the couplet. If color will separate, it is used first because it is easiest to use. Color

is followed by morphological characters, principally of the antennae

and genitalia. The male genitalia provide the most reliable characters.

Species Format

r The descriptions are based on holotypes, lectotypes, or neotypes.

All descriptions of variation are placed in parentheses. In the

biology section tinder host plants of adults, records which are

probably potential hosts are marked with an asterisk; the habits

section includes dates of collection.

The specimens examined section is organized in the following

manner: total number examined; state (in capitals, listed alphabeti­

cally; Canada last with provinces in capitals, listed alphabetically),

county or parish (underlined, arranged alphabetically), other locality

details; number of specimens, colon, location of specimens (all in

parentheses). If only a county record is given, number of specimens

and location are recorded in a separate set of parentheses after all

other listings for that particular county. If only a state record is

given, number of specimens and location appears in brackets after all

listings for counties of that particular state. The four-letter

abbreviations denoting the location of the specimens are listed above

under LIST OF ABBREVIATIONS .(p. xx). The complete label data were

recorded and remain in my files; they are available upon request. CLASSIFICATION

Subfamily Alticinae Illiger

Head rounded, more or less inserted into prothorax; antennae

short, often dilated towards tip, rarely long and filiform. Pronotum

.transverse, almost as wide as elytra, less often square or subrec-

tangular; mostly punctate. Elytra usually oblong oval, always

covering tergites. Prosternum more or less convex between coxae, very

rarely narrow with coxae contiguous. Legs of medium length, rather

robust; metafemora always more or less swollen, grooved beneath;

tibiae most often grooved on outer face; tarsi short, dilated, last

segment sometimes globose or swollen; claws appendiculate, rarely

bifid or simple.

For a key to the genera of the Alticinae, see Arnett (1963).

Genus Phyllotreta Chevrolat

Phyllotreta Chevrolat, 1837. In Pejean, Cat. Coleoptera, 3rd ed.:391.

■ Type-species: Chrysomela brassicae F. (Designated by Chevrolat,

in d'Orbigny, 1845:6.)

Orchestris Crotch, 1873 (nec. Kirby, 1837). Proc. Acad. Nat. Sci.

Philadelphia. 25:65. (Synonymy by LeConte, 1878:615.)

MAJOR REFERENCES: Crotch (1873), Horn (1889), Blatchley (1910), Duckett (1920}, Chittenden (1927), Beller and Hatch (1932),

Heikertinger and Ciski (1939), Wilcox (1954), Hatch (1971), Balsbaugh

(1972) .

DIAGNOSIS: Small size (less than 3.7 mm), elongate oblong shape, glabrous, completely dark or elytra each with a median pale stripe or

1 or 2 pale marks. In addition, antennae 11-segmented, slender (males

/ of some species with 5 and often 4 dilated); postantennal tubercles obsolete; pronotal width about 2x length; elytra confusedly punctate; procoxal cavities open; metatibial apical spur short, simple; 1st metatarsal segment much less than 1/2 tibial length, 5th segment not swollen apically; claws simple. DESCRIPTION: Small, 1.80-3.62 mm long, 0.90-1.85 mm wide; elongate oval to oblong oval; moderately convex to subdepressed; usually glabrous above but lightly pubescent ventrally, moderately pubescent on apical antennal segments, with vesture short, fine, and pale; dark, elytra may have pale pattern.

Head: Dark; covered by prothorax to just behind eyes; eyes prominent, not emarginate; postantennal tubercles obsolete, reduced to flat spots without sharp superior delimination; frons usually evenly punctate. Antennae 11-segmented, about half body length, slender with

1st and apical segments slightly wider (males of some species with one or more of segments 4-7 dilated, usually 5 and often 4). Maxillary palpi with last segment about as long as preceding, elongate conical.

Prothorax: Width about 2x length, somewhat narrower apically; moderately convex, without distinct antebasal or lateral impressions; 10

basal margin adjacent to hind angles transverse, not oblique;

procoxal cavities open.

Elytra: Oblong oval, moderately convex to subdepressed; entirely

dark or each elytron with a median pale stripe or 1 or 2 pale marks;

confusedly punctate; humeri never prominent.

Wings: Normally with metathoracic wings well developed, but in

both males and females of £. arcuata the wings are usually vestigial

(sometimes well developed)« in P. bisinuata wings are reduced in males

and vestigial in female.

Legs: Metafemora greatly enlarged, saltatorial; tibiae normal, without a tooth above followed by a ciliate emargination; apical spurs

inserted medially on apex, short, with a single point, 1st metatarsal

segments much less than 1/2 tibial length, 5th metatarsi simple, not

swollen apically; claws simple with, at most, a very small basal

swelling. Normal color sequence is trochanters pale, femora darkest,

tibiae intermediate, tarsi pale (often with apical segments darker),

all articulations paler than ajoining segments; rarely a different

sequence. When "Normal color sequence" appears in the species

description, it means the sequence described here.

■Abdomen: 5th sternum sexually dimorphic, simple in outline in

females (Fig. 2), but males (Fig. 1) with an apical median lobe which is

median 1/3 of apical margin set off by 2 narrow, moderately deep

emarginations (pygidiura fitting this 3-part configuration), median lobe

usually concave with concavity extending toward but ending before

sternal base, male 5th sternum usually with a median linear impression 11 extending part of length; females sometimes with a median linear or oblong impression in apical 1/3 of 5th sternum; both sexes usually with shallow lateral subbasal impressions. Dark, often with apical margins of sterna 1-4 paler or darker, male usually with apical 1/2 to 1/3 of 5th paler.

Male genitalia (Figs, 4, 5; terminology modified from that used by / Sharp and Muir (1912) and Powell (1941)) . Genitalia a single slender tube arched dorsoventrally; rather uniform in width. Basal piece narrower in basal half, almost entirely open ventrally. Tegmen ventral, attached medially about at midpoint of basal piece; slender, usually about 1/3 to 1/4 total length and bifurcate about 1/3 to 1/2 from its base, the branches wrapped around median lobe but not meeting dorsally.

Median lobe usually slightly tapering to apex which, in dorsal view, is usually rounded or acute, but may be emarginate or truncate; orifice dorsal, located subapically (in lateral view, on a plane oriented about 30° from longitudinal axis of median lobe), oblong in outline, occluded by a transparent area reinforced by 3 longitudinal sclerotized strips; sometimes, dorsally and/or ventrally with a

"washboard" which consists of a median series of short transverse parallel heavily sclerotized evenly spaced ridges, which usually start

subbasal to basal piece and extend onto orifice base, a ventral wash­ board may be interrupted medially by a longitudinal concavity resulting

in 2 parallel rows of teeth; usually with a ventral concavity or groove

at least in apical 1/4.

Female genitalia (Fig. 6; terminology modified from that used by Samuel son (1972) ) : Spermatheca, in lateral view, with pump of uniform width, about 1/3 to 1/2 length of receptacle, usually more heavily sclerotized in apical 1/4^Tapex bluntly rounded. Receptacle (lateral view) usually cylindrical to elongate oval, slightly constricted sub- medially, but may be elongate pear-shaped or oblong; sometimes with a ring collar at junction with pump; uniformly sclerotized; in / dorsal view, laterally compressed, shallowly "C" shaped. Spermathecal duct (lateral view) usually attached to anterior end of receptacle submedially (in dorsal 1/2), rarely on dorsum (Fig.132), usually parallelling receptacle outline anteriorly and ventrally to just beyond gland valve attachment; usually constricted at anterior end of gland valve attachment, sometimes swollen just before constriction; usually gradually tapering posteriorly beyond gland valve attachment, uniformly sclerotized to where this tapering ceases, then not visibly sclerotized (most of unsclerotized portion of duct usually broken off during dissection). Gland valve (lateral view) globose, moderately developed, prominent; spermathecal gland attached on ventral posterior surface (often broken off during dissection).

Sexual dimorphism; This is exhibited in overall size (females usually slightly larger), 5th abdominal sterna (see section on abdomen above), and in middle antennal segments of male in several species

(see section on head above).

IMMATURE STAGES: The egg, larva, and pupa have been described

for only 3 species considered in this revision; these are £. armoraciae by Chittenden (1917), P. striolata by Shimer (1869) and Riley (1885),

and P. zimmermanni by Riley (1885). All 3 species have the larva and 13 pupa illustrated but only P_. armoraciae has the egg drawn. The descriptions of Riley and Chittenden are fairly detailed, but Shimer's is sketchy. Other species have had the adults reared from larvae but no description has been published of the immature stages.

BIOLOGY: The members of this genus are for the most part confined to the Cruciferae as host plants, but a few feed on the closely related Capparidaceae, Limnanthaceae, and Tropaeolaceae. It has been demonstrated that these beetles key on the mustard oils and mustard oil glucosides found in these plants (Gomitz, 1953; Feeny,

Paauwe, and Demong, 1970; Matsumoto, 1970).

In the temporate zone, a generalized life cycle would be as follows: adults overwinter; mating and egg laying occur in early spring, the eggs are laid on or close to the part of the plant upon which the larvae feed (some are leaf miners, few are stem miners, and others are root feeders); the prepupal and pupal stages are spent in an earthen cell in the ground; and the adults emerge in late summer and are active until late fall.

The larvae are leaf miners in P. constricta, liebecki, P. oregonensis, and P^. zimmermanni and '’stem miners" in P. armoraciae

(restricted to the petiole and midrib), whereas the larvae of £. striolata are root feeders. The transition from leaf to stem or stem to root feeding has been observed in 2 species. In one experiment, a larva of P. zimmermanni after having consumed the leaves moved into the stem and did quite well (Chittenden, 1927:21). Larvae of P^. armoraciae have been known to complete their development in the roots of horse­ radish after the leaves have died, and later emerge as adults 14

(Chittenden, 1917:10). Contrary to the impression given here that most larvae are leaf miners, in considering the genus worldwide, the majority of known larvae are stem and root borers or root feeders

(Frost, 1924).

Among the North American species considered here, P^. armoraciae,

P. bipustulata, P. decipiens, P. oregonensis, P. ramosa, and P. 1 / zimmermanni have been more or less periodically of economic importance.

Recently several researchers at Cornell University have been working on such biological aspects as phenology and microhabitat selection (Tahvanainen, 1972), the influence of vegetation diversity on population ecology (Tahvanainen, 1972), host plant specificity

(Feeny, Paauwe, and Demong, 1970), and the role of temporal hosts in seasonal development of the crucifer fauna (Root, 1969).

DISCUSSION: Phyllotreta is worldwide in distribution, excluding the Arctic and Antarctic regions. Two of the maculate species found north of Mexico are considered introductions, P_. armoraciae first found at Chicago in 1893 and P_. striolata apparently introduced some time before 1801 from Eurasia. Of the species considered here, 10 are restricted to the Pacific Coast and adjacent states, 1 to northwestern

Canada, 3 to the Rocky Mountains and westward, 1 to east of the Rocky

Mountains, 2 to the Great Plains, 2 to the northern 1/2 of the U.S. and

Canada, and 3 are found mostly east of the Mississippi River.

The genus Phyllotreta is most closely related to Longitarsus,

Glyptina, Apthona, and Palaeothona, the other members of the Apthonini.

This tribe is characterized by the following: small size, oval form,

antennae 11-segmented, procoxal cavities open, pronotum without any 15 antebasal impression, mesostemum always visible, metatibiae not sulcate followed by a sinuation on outer edge, 1st metatarsal segments long and slender, 5th metatarsal segments slender, and claws simple.

Phyllotreta can be separated from these 4 genera on the basis of it having the metatibiae not grooved on the outer edge but slightly excavated near the tip and the spur inserted medially, the 1st metatarsal segments not much more than 1/3 tibial length at most, the elytra confusedly punctate, each often with a pale stripe or marks, and the postantennal tubercles obsolete.

Superficially, Phyllotreta might be confused with Systena

(Chevrolat, 1837), Epitrix (Foundras, 1860), and Chaetocnema (Stephens,

1831), but these 3 genera have the procoxal cavities closed instead of open. In addition, Epitrix is usually pubescent instead of glabrous,

Chaetocnema is more robust and its metatibiae have a preapical tooth followed by a ciliate emargination, and Systena is twice as long for most species and if it has pale stripes, these are simple and straight.

It should be pointed out that most of the external morphological characters used by previous workers are highly variable, with the exception of those of the antennae. In particular this includes the texture, punctation, and luster of the head, pronotum, and elytra, the color of the antennae and legs, and the elytral pattern for most species. Contrary to Chittenden (1927), the sexes may be easily distinguished by the apical margin of the 5th abdominal sternum (see section on abdomen, above). The male genitalia have the most reliable characters for species separation, whereas, the female spermathecae are usually reliable only for separation to groups of closely related 16 species. The male antennae provide very reliable characters for most of the species, particularly those which exhibit sexual dimorphism.

The authorship of the genus Phyllotreta has been credited to several people but White (1970) has clarified this situation,

Chevrolat (1837) being the correct author. Also, several species have been cited as the type-species but Chevrolat (in d'Orbigny, 1845) / satisfactorily designated Chrysomela brassicae Fabricius (1789) as the type-species (White, 1970); it is now a synonym of P. exclamationis

(Thunberg, 1874). Crotch*s unfortunate adoption of Kirby*s name

Orchestris for this genus has been adequately refuted by LeConte

(1878:615).

Chittenden's 1927 revision of Phyllotreta deserves a few comments, at least concerning his treatment of the maculate species. Excluding

armoraciae, his key to species is almost completely dependent on male characters, yet 4 new species (P. alberta, P. perspicua, P^. oblonga,

P. obtusa), which are separated by male characteristics were known only from females (incorrectly sexed). Of his 5 new variations, 3

(P. armoraciae biplagiata, P^. vittata vemicosa, P^. vittata artivitta) were not included in his key nor was P^. vittata monticola Weise which he recognized as being present in the U.S. Also, the unique type of his new species P. amplicomis is a perfectly good male of P^. denti- comis with its very distinctive male antennal characteristics. If

Chittenden had been careful in determining sex and had studied the male genitalia, many of his errors could have been avoided.

Hatch's 1971 review of Phyllotreta for the Pacific Northwest deserves a note of caution to its users. Much of the material bearing his determination label that I have reworked is misidentified.

Therefore, his listed localities should be accepted provisionally.

Also, the type of his new species aequalis is a male of P^. denticomis with its very distinctive male antennal characteristics.

The name Phyllotreta comes from the Greek, phyllo- meaning leaf, and treta meaning to perforate; leaf perforation is the characteristic

/ damage. The accent is on the penult syllable (based on the Greek vowel n , which is long): Phyllotreta. 18

Key to Males (Fig, 1) of Maculate Species of Phyllotreta

1, Elytra each appearing pale yellow, outlined in

brown or black with sutural margin slightly

wider (Fig* 8); northern 1/2 U.S. and adjacent

Canada east of Rocky Mts., I d a h o armoraciae (Koch)

1*. Elytra brown to black, with pale stripes or marks ...... 2

2 (1 *) . Elytra each with 1 or 2 marks ...... 3

2 *. Elytra each with a median stripe ...... 6

3(2). Antennal segment 6 with a sharp anteroventrally

directed process (Fig. 55); elytra each with a

subbasal and/or subapical somewhat indistinct

reddish mark; California, Oregon......

...... (in part) denticornis Horn

3*. Antennal segment 6 simple, no process; elytra each

. with 1 somewhat indistinct small subapical reddish

mark or each with 2 yellowish marks ...... 4

4(3?). Elytra each with 1 small subapical reddish brown

to yellow mark; Rocky Mts. and

westward, Texas ...... decipiens Horn

4*. Elytra each with 2 yellowish marks ...... 5 19

5(4'). Antennal segments 5 and 6 subequal in length, 5th

simple; elytral subapical mark with sutural margin

straight, not incurved toward suture (Fig. 10);

east of Rocky Mts...... (in part) bipustulata (F.)

S'. Antennal segment 5 about 2x length of 6, 5th

enlarged (Fig. 78); elytral subapical mark

incurved apically toward suture (Fig. 35);

widespread ...... (in part) striolata (F.)

6(2') Elytral stripe simple, of uniform width and without

lateral or terminal dilations, slightly incurved

toward suture apically (Fig. 7, 12, 15, 19, 28);

5th antennal segment expanded bilaterally,

appearing dorsoventrally flattened (Fig. 40) ...... 7

6*. Elytral stripe with 1 or 2 lateral dilations

and/or terminal dilations; 5th antennal segment

may be enlarged or expanded ...... 11

7C6). Antennal segment 6 with a sharp anteroventrally

directed process (Fig. 55); California, Oregon ..

...... (in part) denticomis Horn

7*. Antennal segment 6 simple, without a process ...... 8

8(7'). Antennal segments 7-11 darkest, in contrast to

paler basal segments; antennal segment 5 with

venter longitudinally concave on inner 1/2 only;

southern California ...... arcuata n.sp. 20

Antennal segments 5-11 darkest at least dorsally;

antennal segment 5 with venter longitudinally

concave medially or flat ...... 9

9(8.'), Antennal segments with at least 1-5 paler on

venter, 2 may be paler; antennal segment 5 with

venter flat; northern California ...... spatulata n.sp.

9 ** Antennal segments 2-3 pale; antennal segment 5 with venter longitudinally concave medially ...... 10

10(9'). Metathoracic wings normal, well developed; male

genitalia in lateral view slightly sigmoid

(Fig. 106); California, New York ...... lepidula (LeC.)

10*. Metathoracic wings reduced to less than elytron

length; male genitalia in lateral view with a

very abrupt, strong bend submedially (Fig. 94);

northern California ...... bisinuata n.sp.

Elytral median stripe greatly expanded at least in

apical 1/3 over most of elytral width (expended

portion rarely only slightly paler than background

color) reaching lateral, posterior, and suture

margins apically (Figs. 20, 26) ...... 12

11*. Elytral median stripe at most moderately expanded subapically, never covering most of subapical

area, stripes never reaching apical margins ...... 13

12(11). Antennal segment 5 with a long pointed apical 21

prolongation (Fig. 74); east o£ Rocky Mts. in

northern 1/2 U.S. and adjacent Canada, Colorado,

Utah, N e v a d a ...... robusta LeC.

12*. Antennal segment 5 with apical 1/2 at most

moderately expanded laterally, to one side

(Fig. 64); Great Plains and eastward ... liebecki Schffr.

. 13(11*). Antennal segment 4 moderately and 5 strongly

expanded bilaterally, appearing dorsoventrally

flattened (Figs. 80, 83) ...... 14

13*. Antennal segment 4 simple and 5 simple or only

slightly enlarged, essentially cylindrical

(Figs. 46, 70, 78) ...... 19

14(13). Antennal segments 2-5 pale, in distinct contrast

to darker segments 7-11; segment 6 may be pale

or dark ...... 15

14*. Antennal segments 2-4 pale and/or dark, segments

5-11 d a r k ...... 16

15(14). Longer, greater than 2.8 mm (about 3 mm); male

genitalia in lateral view slightly sigmoid with

apex almost straight dorsally, and strongly

swollen medially on venter (Fig. 124); Oregon ....

...... utana Chttn. 22

15*. Shorter, less than 2.8 mm (about 2.6 nun); male

genitalia in lateral view with dorsum gradually

arched, apex curving slightly ventrad, and

venter only very slightly swollen medially

(Fig. 126); Pacific Northwest ...... utanula n.sp.

t 16(14f) Antennal segment 5 with a distinct concavity in

basal 1/2 anteriorly or to outside, 5th much

wider than 4 (Fig. 83); apex of male genitalia

with a narrow median emargination (Fig. 127),

moderately curved in lateral view with a

ventral subapical concavity (Fig. 128);

widespread ...... zimmermanni (Crotch)

16*. Antennal segment 5 evenly flattened ventrally

although slightly concave, 5 subequal in width

to 4'(Figs. 50, 59, 68); apex of male genitalia

entire (Figs. 95, 111) or if emarginate, then

ventrally straight or flat in lateral view, no

subapical concavity ventrally (Fig. 103) ...... 17

17(16*) Elytral stripe width medially distinctly less,

usually much less, than distance from stripe to

suture (Fig. 18); apex of male genitalia with a

narrow median emargination (Fig. 103), straight

or flat ventrally in lateral view with no

subapical concavity (Fig. 104); Oregon, northern

California...... emarginata n.sp. 23

17*. Elytral stripe width medially about equal or

usually greater than distance £rom stripe to

suture (Figs. 13, 22), rarely less, if so then

apex of male genitalia entire, with no median

emargination (Figs. 95, 111) ...... 18

/ 18(17*) Pronotal punctures usually more widely separated

than elytral punctures; male genitalia

abruptly narrowed in apical 1/2 and apex

distinctly directed ventrad in lateral view

(Fig. 96); east of Rocky Mts. and west of

Mississippi River ...... constricta n.sp.

18*. Pronotal and elytral punctures usually separated

by about same distance; male genitalia gradually

narrowed toward apex which is essentially

straight in lateral view (Fig. 112); Rocky Mts.

and westward ...... oregonensis (Crotch)

19(13') Elytral stripe apical dilation with inner margin

straight, not incurved toward suture (Fig. 11);

east of Rocky Mts...... (in part) bipustulata (F.)

19*. Elytral stripe apex strongly incurved toward

suture (Figs. 17, 24) ...... 20

20(19*). Elytral stripe width medially almost equal to

distance from stripe to suture, basal and

subbasal lateral dilations fused and usually 24

with a shallow lateral subbasal emargination

Figs. 9, 17); antennal segment 5 simple,

segments 4-5-6 subequal in length (Figs. 44,

57) ...... 21

2 0 *. Elytral stripe width medially much less than

interval, basal and subbasal lateral dilations

appearing as 2 distinct dilations (Figs. 21,

24, 25, 29); antennal segment 5 enlarged in

diameter and/or longer than 4 or 6 (Figs. 66,

72, 78) OR antennal segment 5 simple, segments

4-5-6 subequal in length (Fig. 70) ...... 22

21C20). Elytral stripe basally somewhat and apically

abruptly incurving toward suture (Fig. 17);

antennal segments 2-5 pale, 6-7 transitional,

8-11 dark; male genitalia longer, about 1.4 mm,

in lateral view apex distinctly directed

ventrad (Fig. 102); northern California ......

...... dolichophalla n.sp.

2 1'. Elytral stripe basally parallelling suture,

gradually incurving toward suture in apical

1/3 to 1/2 (Fig. 9); antennal segments 2-4 pale,

5 transitional, 6-11 dark; male genitalia

shorter, about 1.2 mm, in lateral view almost

straight (Fig. 90); Colorado, South Dakota ....

...... attenuata n.sp. 25

22(20*). Elytral stripe with median 1/3 very narrow, of

uniform width, subbasal lateral dilations with

posterior margin almost at 90° angle to stripe,

apical 1/3 of stripe abruptly widened,

appearing notched basally on outside (Figs. 24,

25) ...... 23

22*. Elytral stripe with median 1/3 gradually

widening toward ends, basally into subbasal

lateral dilation, apically into wider apical

1/3, sometimes with a lateral notch 1/3 from

apex (Figs. 21, 29) ...... 24

23(22). Antennal segment 5 longer than 4 or 6, 4 and 6

subequal (Fig. 72); longer, about 2.5 mm;

male genitalia in lateral view moderately

and evenly arched dorsoventrally with apex

gradually curved ventrad (Fig. 116);

Pacific Northwest ...... ramosoides n .sp.

23*. Antennal segments 4-5-6 subequal in length

(Fig. 70); shorter, about 2.2 ram; male

genitalia in lateral view only very slightly

arched dorsoventrally with apex bent ventrad

at about 45° angle (Fig. 114); Pacific Coast

states, Nevada, New York ...... ramosa (Crotch)

24(22') Antennal segment 5 more than 2x length of 6, 4 26

longer than 6 (Fig. 78); elytral stripe strongly

incurved toward suture basally and apically

(Fig. 29); widespread...... (in part) striolata (F.)

24*. Antennal segment 5 about 1.25x length of 6, 4 and

6 subequal in length (Fig. 66); elytral stripe

inner margin almost parallelling suture, except

strongly incurved toward suture apically

(Fig. 21); Alberta, Northwest Territory ....

...... oblonga Chttn. 27

Ke y to Females (Fig. 2) of Maculate Species of Phyllotreta

1. Elytra each appearing pale yellow, outlined in

brown or black with sutural margin slightly

wider (Fig. 8); northern 1/2 U.S. and adjacent

Canada east of Rocky Mts...... armoraciae (Koch)

1* . Elytra brown to black with pale stripes or marks ...... 2

2(1*). Elytra each with 1 or 2 marks ...... 3

2*. Elytra each with a median stripe ...... 8

3(2). Elytra each with 1 mark ...... 4

3'. Elytra each with 2 marks ...... 6

4(3). Elytra each with a subbasal mark; California,

Oregon ...... (in part) denticomis Horn

4'. Elytra each with a subapical mark ...... 5

5(4*). Shorter than 2.5 mm (about 2-2.4 mm); spermathecal

receptacle shorter, distinctly tapered in apical

1/3 to a moderately developed apical ring collar

(Fig. 135); Rocky Mts. and westward decipiens Horn

5 *. Longer than 2.5 mm (about 2.6 mm); spermathecal

Teceptacle longer, slightly narrowed apically

with an indistinct ring collar, receptacle very

slightly narrowed submedially (Fig. 136);

California, O r e g o n (in part) denticomis Horn 28

6(3'). Antennal segments 5 and 6 equal or subequal in

length (Fig. 47); east of Rocky Mts......

...... (in part) bipustulata (F.)

6*. Antennal segment 5 about 1.5x length of 6

(Figs. 56, 79) ...... 7

7C6'). Shorter than 2.5 mm (about 2-2.3 mm); spermathecal receptacle shorter, distinctly tapered in apical

1/3 to a moderately developed apical ring collar,

pump much longer (Fig. 147); widespread ......

...... (in part) striolata (F.)

7*. Longer than 2.5 mm (about 2.6 mm); spermathecal

receptacle longer, slightly narrowed apically

. with an indistinct ring collar, pump shorter

(Fig. 136); California, Oregon ......

...... (in part) denticornis Horn

8 (2') . Elytral stripe simple, of uniform width and

without lateral or terminal dilations, slightly

incurved toward suture apically (Figs. 7, 12,

15, 19, 28); antennal segment 5 about 1.5x

length of 6 ...... 9

Elytral stripe with 1 or 2 lateral dilations

and/or terminal dilations; antennal segment 5

from subequal to l.Sx length of 6 ...... 14

9(8). Metathoracic wings vestigial 10 29

9*. Metathoracic wings normal, well developed ...... 11

10(9). Spermathecal receptacle unique (Fig. 146), basal

end tapering to a blunt predorsal point, rather

abruptly and strongly swollen ventrally in

basal 1/2 and elongate, almost parallel-sided

f in apical 1/2, ring collar moderately developed;

northern California ...... spatulata n.sp.

10*. Spermathecal receptacle with basal end evenly

rounded, slightly narrower in apical 1/2, ring

collar slightly developed (Fig. 129); southern

California ...... (in part) arcuata n.sp.

11C9*). Spermathecal receptacle of rather uniform width,

with a distinct dorsoventral bend, ring collar

moderately developed (Fig. 133); northern

California bisinuata n.sp.

11'. Spermathecal receptacle slightly narrower in

apical 1/2, not bent, ring collar at most

slightly developed (Figs. 129, 136, 139) ...... 12

12(11') Spermathecal receptacle (Fig. 129) much shorter,

about 0.22 mm; southern California ......

...... (in part) arcuata n.sp.

12*. Spermathecal receptacle (Figs. 136, 139) longer,

about 0.26-0.27 m m ...... 13 30

13(12'). Elytral stripes usually somewhat indistinct with

borders blending into background color* somewhat

pale reddish brown; spermathecal receptacle less

robust (Fig. 136); California, Oregon ......

...... (in part) denticomis Horn

13*. Elytral stripe usually with borders clearly / delimited, straw-yellow; spermathecal

receptacle more robust (Fig. 139); California,

New York ...... lepidula (LeC.)

14(8*). Elytral median.stripe greatly expanded laterally

at least in apical 1/4 over most of elytral

width, reaching lateral, posterior, and

sutural margins apically (Figs. 20, 26) ...... 15

14*. Elytral median stripe at most moderately expanded

subapically, never convering most of subapical

area, stripes never reaching apical margins ...... 16

15(14). Elytral median stripe with a moderate subbasal

lateral dilation and usually greatly expanded

laterally only in apical 1/4, leaving median

1/3 of stripe narrow (Fig. 20); Great Plains

and eastward ...... liebecki Schffr.

IS*. Elytral stripe with a strong subbasal lateral

dilation and greatly expanded in apical 1/3

to 1/2, leaving usually much less than median

1/3 of stripe narrow (Figs. 26, 27); east of 31

Rocky Mts. in northern 1/2 of U.S. and adjacent

Canada ...... Tobusta LeC.

16 C14 *). Elytral stripe with median 1/3 very narrow, of

uniform width, subbasal lateral dilations with

posterior margin almost at 90° angle to stripe,

apical 1/3 of stripe abruptly widened,'

appearing notched basally on outside (Figs.

24, 25) ...... 17

16*. Elytral stripe with median 1/3 gradually widening

toward ends, basally into subbasal lateral

dilation, apically into wider apical 1/3,

sometimes with a lateral notch 1/3 from apex

CFigs. 11, 21, 22, 36, 38) ...... 18

17(16). Antennal segment 5 longer than 4 or 6, 4 and 6

subequal (Fig. 73); longer, about 2.5 mm;

Pacific Northwest ...... ramosoides n.sp.

17*. Antennal segments 4-5-6 subequal in length (Fig.

71); shorter, about 2.2 mm; Pacific Coast states,

Nevada, New Y o r k ...... ramosa (Crotch)

18(16') Elytral stripe width medially almost equal to

or usually greater than distance from stripe

to suture (Figs. 9, 13, 17, 22), basal and

subbasal lateral dilations fused and usually

with a shallow lateral subbasal emargination ...... 19 18*. Elytral stripe width medially much less than

distance from stripe to suture, basal and

subbasal lateral dilations appearing as 2

distinct dilations (Tigs* 18» 21, 23, 29,

36, 38, 39) ...... 22

t 19(18). Antennal segment 5 about l.Sx length of 6

(Figs. 51, 69) ...... 20

19*. Antennal segments 5 and 6 subequal in length

Figs. 45, 58) ...... 21

20(19). Spermathecal receptacle strongly constricted

submedially on venter (Fig. 134); east of Rocky

Mts. and west of Mississippi River .... constricta n.sp.

20*. Spermathecal receptacle only slightly narrowed

submedially on venter (Fig. 142); Rocky Mts. and

westward ...... (in part) oregonensis (Crotch)

21(19*). Elytral stripe basally somewhat and apically

abruptly incurving toward suture (Fig. 17);

antennal segments 2-5 pale, 6-7 transitional,

8-11 dark; spermathecal receptacle unique

(Fig. 137), basal 1/2 strongly swollen

ventrally, apical 1/2 of rather uniform width,

ring collar very wide, strongly developed;

northern California ...... dolichophalla n.sp. 33

21*. Elytral stripe basally parallelling suture,

gradually incurving toward suture in apical

1/3 to 1/2 (Fig. 9); antennal segments 1-3

paler, 4 and 5 each transitional or dark (if

4 dark then 5 also dark), 6-11 dark;

spermathecal receptacle without ventral con­

striction, ring collar at most slightly

developed (Fig. 131); Colorado, South Dakota

...... attenuata n.sp.

22(18') . Antennal segments 2-S pale (5 may be slightly

darker) in distinct contrast to darker segments

6-11 ...... 23 2 2 *. Antennal segments 2-4 pale and/or dark, segments

5-11 da r k ...... 24

23(22). Antennal segments 5 and 6 subequal in length

(Fig. 47); elytral stripe very strongly dilated

subbasally and subapically (Fig. 11); spermathecal

receptacle oval, spermathecal duct attached on

dorsum of receptacle (Fig. 132); east of Rocky

Mts...... (in part) bipustulata (F.)

23'. Antennal segment 5 about 1.5x length of 6; elytral

stripe of rather uniform width, with moderate

subbasal and slight subapical dilations (Fig. 81);

spermatheca as in Fig. 148; Oregon Cfemale of 34

utanula n.sp. is unknown but would most probably

key out here, differing only in spermatheca and

being shorter; Pacific Northwest I! utana Chttn.

24 (221) Elytral stripe inner margin almost parallelling

suture except strongly incurving toward suture

/ apically (Fig. 21); antennal segment 5 about

1.25x length of 6 (Fig. 67); spermathecal

receptacle distinctly narrowed and almost

parallel-sided in apical 1/3 (Fig. 141):

Alberta, Northwest Territory ...... oblonga Chttn.

24'. Elytral stripe inner margin sinuous (if almost

parallelling suture, then basally strongly

incurved toward suture), strongly incruved toward

suture apically (Figs. 18, 23, 29, 38, 39);

antennal segment 5 at least 1.5x length of 6

(Figs. 60, 69, 79, 84); spermathecal receptacle

elongate or oblong ...... 25

25(24*) Shorter, less than 2.6 mm (about 2.2-2.4 mm);

spermathecal receptacle much shorter, evenly

tapering apically to moderately developed ring

collar, pump longer than 1/2 receptacle length

(Fig. 147); widespread...... (in part) striolata (F.)

25*. Longer, more than 2.5 mm (2.8-3.4 mm); spermathecal

receptacle much longer, with a slight submedial 35

dorsoventral bend, pump much less than 1/2

receptacle length ...... 26

26(25') Spermathecal receptacle shorter, more robust

(Fig. 138), ring collar slightly developed;

Oregon, northern California ...... emarginata n.sp.

26'. Spermathecal receptacle longer, more elongate

CFigs. 142, 149); ring collar strongly

developed or absent ...... 27

27(26') Spermathecal receptacle with ring collar strongly

developed (Fig. 149); widespread ......

...... zimmermanni (Crotch)

27*. Spermathecal receptacle with no ring collar

(Fig. 142); Rocky Mts. and westward ....

...... (in part) oregonensis (Crotch) 36

Phyilotreta arcuata NEW SPECIES

(Figs. 7, 40-41, 85-86, 129, 150)

Holotype: Male, deposited at LACM.

Type locality: Griffith Park, Los Angeles, California.

DIAGNOSIS: Elytra each with the median stripe simple, of uniform t width and without lateral or terminal dilations, slightly incurved toward suture apically; male 4th and 5th antennal segments expanded bilaterally, dorsoventrally flattened, 5th with venter longitudinally

concave on inner 1/2 only; antennal segments 2-6 pale, in contrast

to darker segments 7-11.

DESCRIPTION: Holotype (variation, excluding punctation, in paren­

theses) : Fig. 7; elongate oval, length 2.50 mm (male 2.28-2.50, female

2.18-2.58), width 1.05 mm (male 1.05-1.12, female 1.02-1.20); head and pronotum black with slight metallic luster, elytra brownish black, each

with a median straw-yellow stripe. Head: Black; texture finely

granulate basally to slightly roughened anteriorly (entirely finely

granulate); moderately punctate, punctures separated by less than to 2x

their diameter, mostly by about one diameter; Interocular Distance/

Maximum Diameter of eye, 1.63 (male 1.50-1.63, female 1.50-1.63).

Antennae: Figs. 40, 41; segments 4 and 7 about 2x length of 6, 5 about

2x length of 4; segments 4 and 5 expanded bilaterally and ventrally

flattened; segment 5 distinctly longest, with venter longitudinally

concave on inner 1/2 only; segment (length/width]: 1(9,5/3], 2(4/2+],

3(4,5/3], 4(4.5/S], 5(8.5/6], 6(2+/3], 7(5/3.5], 8(5/3], 9(5/3.5],

10(5/3.5], 11(7/3], total length 1.50 mm (Allotype: 1(8.5/3], 2(4/2+], 37

3[4/2j, 4(4.5/2,5], 5(6/2.5], 6(4/2.5], 7(5/3], 8(5/3,5], 9(5/3.5],

10(5/3.5], 11(7/3+], total length 1.45 mm; simple); antennae various shades of brown, basal 6(5) segments paler with dorsum of 1 darker,

6 Cor none) intermediate, 7-11 darkest. Pronotum: Length 0.45 mm

(male 0.42-0.45, female 0.38-0.48), width 0.75 mm (male 0.72-0.78, female 0.72-0.78); black; texture slightly roughened (finely granulate);

t moderately to coarsely punctate, punctures separated by less than to equal their diameter, mostly by about one diameter. Elytra: Length

1.80 mm (male 1.60-1.85, female 1.62-1.88), width 1.05 mm (male 1.05-

1.12, female 1.02-1.20); brownish black (dark brown), median stripe as in Fig. 7; texture slightly roughened (usually smooth); coarsely punctate in basal 1/2, moderately punctate in apical 1/2, punctures

separated by less than 1/2 to equal their diameter, mostly by less than one diameter. Legs: Normal color sequence. Abdomen: 5th sternum with an apical median lobe moderately concave, concavity oval in its

extension to within 1/3 of sternal base, with a median linear impression

extending from sternal base and ending subapically (male median linear

impression ending 1/4 from base; female: simple in outline, usually

with apical 1/2 shallowly concave, usually with median linear impression

in apical 1/2); black with apical 1/3 paler except along midline (dark

brown to brownish black).

Male genitalia: Figs. 85, 86; length approximately 0.82 mm; mod­

erately arched dorsoventrally, of rather uniform width; apex in dorsal

view very bluntly rounded with shallow, moderately wide median emar-

gination, in lateral view gradually but strongly curved ventrad, end

bluntly rounded. Female genitalia: Fig. 129; length approximately 0.35 mm; recep­ tacle elongate, ring collar slightly developed; pump long.

BIOLOGY: Host plants and Immature stages are unknown.

Habits; Adults have been collected in southern California during

May.

DISCUSSION: Distribution: Fig. 150; £. arcuata has been found

/ only in Los Angeles and Santa Marguerita, California. New name: The

specific epithet arcuata comes from the Latin arcus- meaning an arch,

and refers to the male genitalia in lateral view being moderately and

evenly arched dorsoventrally.

Relationships: £. arcuata is most similar to _P. lepidula and P^.

bisinuata, and less similar to _P. utanula and P_. utana. Phyllotreta

arcuata shares having a simple median elytral stripe with £. lepidula,

p< bisinuata, P_. spatulata, and the vittate specimens of P. denticornis.

The antennal character of having the 5th segment appearing dorso­

ventrally flattened is shared with £. utana, utanula, £. oregonensis,

P. emarginata, P^. constricta, and P^. zimmermanni, but arcuata shares

the additional feature of having antennal segments 2-5 pale in con­

trast to 7-11 dark, with P. utana and _P. utanula. The apex of the male

genitalia in dorsal view is almost identical in P^. arculata, P. lepidula,

and P. bisinuata; however, in lateral view, arcuata has the apex

strongly curved ventrally and with no median subapical lobe, whereas

P. lepidula and P. constricta both have the apex essentially straight

and have a median subapical lobe.

TYPE MATERIAL: The holotype, allotype, and 3 paratypes (Id, 2?) 39

aTe at the LACM; ’2 paratypes (1(5, 1?) are at the USNM; and 3 paratypes

(Id, 2?) are at the CASC.. The holotype is mounted on a point and the pin bears 2 labels (decending order): "Griffith Pk LA 4*4*25"

"Holotype Phyllotreta arcuata"; and a polyethylene genitalia vial. I added the type label and vial. The allotype bears the same locality label; an allotype label and card bearing the spermatheca were added by me. The 3 paratypes at the LACM and the 2 at the USNM have the same locality label as the holotype; all 3 paratypes at the CASC have the same 2 labels (decending): "Santa Marguerita [sic.!] Calif. IV-26-

1933" and "R. HOPPING COLLECTION".

SPECIMENS EXAMINED: Total 10: CALIFORNIA: Los Angeles Co., Griffith

Park (Holotype, Allotype, LACM:3, USNM;2), San Luis Obispo Co., Santa

Margarita (CASC:3).

Phyllotreta armoraciae (Koch)

(Figs. 3, 8, 42-43, 87-88, 130, 151)

Haltica armoraciae Koch, 1803. Entomologische Helft, 2:75-76; table 3,

fig. 6.

Lectotype (here designated): Male, Zoological Museum of Humboldt

Univ., Berlin, DDR.

Type locality: ?

Phyllotreta armoraciae, Foudras, 1860. Ann. Soc. Linn. Lyon, 6(ser.2):

344.

Haltica Vittata Stephens, 1831 (nec. Fabricius, 1801). Illus. British

ent., Mandibulata, 4:297. (Synonymy by Weise, 1888:865) 40

Phyllotreta armoraciae (var.) biplagiata Chittenden, 1927. Ent.

Americana, 8(n.s.,no.l):16.

DIAGNOSIS: Elytra each appearing pale, outlined in brown or black with sutural margin slightly wider; antennae simple.

DESCRIPTION: Lectotype (variation, excluding punctation, in paren­ theses): Fig. 8; elongate oval, length 2.82 mm (mal'e 2.48-3.38, female

.3.08-3.62), width 1.35 mm (male 1.30-1.62, female 1.60-1.85); head and pronotum black with slight metallic luster (no luster), elytra dark brown, each with a very wide median straw-yellow stripe covering most of elytron. Head: Black (brownish black); texture finely granulate basally to roughened anteriorly (finely granulate, finely alutaceous basally to roughened anteriorly, finely alutaceous); moderately punctate, punctures separated by l-2x their diameter, mostly by about one diameter; Interocular Distance/Maximum Diameter of eye, 1.30 (male

1.30-1.50, female 1.36-1.67). Antennae: Figs. 42, 43; segments 4-6 subequal in length, simple; segment length/width : 1(9/3.53, 2(4.5/3-D,

3(5/2.53, 4C7+/3-3, 5(7/33, 6(6/3-3, 7[6+/33, 8(6+/33, 9(6+/3+3,

10(6/3+3, 11(8/3.53, total length 1.76 mm (female Paralectotype:

1(11/43, 2(4.5/33, 3(5/33, 4(8/2.53, 5(7..5/33, 6(6+/3-], 7(7/33,

8(7/3.53, 9(7/43, 10(6.5/4-3, 11(8.5/43, total length 1.95 mm; simple); antennae various shades of brown, basal 3 paler, (4 intermediate),

4-11(5-11) darker. Pronotum: Length 0.50 mm (male 0.42-0.60, female

0.55-0.62), width 0.92 mm (male 0.85-1.05, female 1.05-1.15) black

(brownish black); texture finely granulate to slightly roughened

(finely granulate, slightly roughened); moderately punctate, punctures 41

separated by less than 1/2 to equal their diameter, mostly by about

one diameter. Elytra: Length 2.10 mm (male 1.90-2.50, female 2.38-

2.78), width 0.92 mm (male.1.30-1.62, female 1.60-1.85); dark brown

Cbrown, usually brownish black), median stripe pattern as in Fig. 8;

texture slightly roughened (usually smooth); moderately to coarsely punctate, punctures separated by less than 1/2 to equal their / diameter, mostly by less than one diameter. Legs: Normal color

sequence except tibiae and tarsi same color (normal color sequence).

Abdomen: 5th sternum with a shallowly concave apical median lobe,

concavity extending to within 1/3 of base; a median linear impression

extending subbasally to base of median lobe, impression somewhat

indistinct or interrupted (male concavity moderate, linear impression

distinct, extending to sternal base; female: simple in outline, usually with a shallow narrow median elliptical impression extending

from apex to within 1/3 of base or to base); brownish black, median

lobe area paler (5th sternum with apical 1/2 paler).

Male genitalia: Figs. 87, 88; length approximately 1.08 mm;

slightly arched dorsoventrally, almost uniform in width, with a dorsal

concavity extending from about midpoint and ending subapically; apex

in dorsal view with a very wide, moderately deep median emargination,

in lateral view with a subapical dorsal swelling and a rounded end.

Female genitalia: Fig. 130; length approximately 0.36 mm; recep­

tacle oblong, ring collar slightly developed; pump long.

BIOLOGY: Host plants, adult:*horse-radish (Armoracia rusticana

Gaertn., Mey. § Scherb., =Radicula armoracia) (Koch, 1803:75); *marsh 42 cress (Rorippa islandica var. Ferrialdiana Butters 6 Abbe, -Radicula palustris (Chittenden, 1895:405); *cabbage, not observed feeding

(Chittenden, 1917:8). New Collection Records: alfalfa: "Sauk Co.,

Wis., 5 mi. N. LaValle, My 3, 1948, J.T. Medler Coll." (UWEN-.l);

DuPuits alfalfa: "Savage Farm, Ithaca, N.Y., 9 June 1967, C.U. Lot

946" (CUIC:1): *Raphanus: "Ithaca, N.Y., 30 July '30 (CUIC:3).

Host plants, larva: horse-radish (Chittenden, 1917:2).

Immature stages: Egg, larva, and pupa described by Chittenden

(1917).

Habits: Larvae are stem miners on basis of plant anatomy, being restricted to the petiole and midrib of horse-radish leaves; rarely may they mine the roots (Chittenden, 1917). Adults cause either the typical shot-hole leaf damage or gouge pits in the midrib and petiole

(Chittenden, 1917). Adults of .P. armoraciae have been collected in northern half of the U.S. and adjacent Canada east of the Rocky Moun­

tains from mid-April (as early as mid-February in the midwest) until mid-November, and in Idaho in mid-August. The adults are long-lived,

they overwinter and then the females, with periodical mating, lay eggs

until early August; in Wisconsin, larvae from eggs laid in late May

emerged as adults in late July or early August (Chittenden, 1917). The

most complete biological account of this species is by Chittenden, 1917.

DISCUSSION: Distribution: Fig. 151; P. armoraciae has been found

mainly in the northern half of the U.S. and adjacent Canada east of the

Rocky Mountains, but also in southern Missouri and northern Idaho. It

is an introduced species and was first collected in 1893 at Chicago, 43 Illinois, by Chittenden (Chittenden, 1895).

Nomenclature: Koch (1803) apparently described this species from

a series of 8 specimens, placing it in Haltica. Foudras (1860)

transferred it to Phyllotreta. In 1831, Stephens described Haltica

vittata which was synonymized by Weise (1888); this is not the

Fabriciah vittata described in Crioceris in 1801. The variety P.

a. biplagiata, described by Chittenden (1927) is only a specimen which has the median pale stripe somewhat irregularly interrupted by

various shades of brown; otherwise, it is a fairly typical P^.

armoraciae except that overall, it is somewhat darker. According to

the Rules of Zoological Nomenclature, this variety holds no taxonomic

rank and is included only for completeness and clarity.

Relationships: P. armoraciae shows no close similarities with any

other species in this genus found north of Mexico.

TYPE MATERIAL: The lectotype and 7 apparent paralectotypes (Id1,

6?) are in the Zoological Museum of Humboldt University, Berlin, DDR.

This series is accepted as the type series on the authority of Dr. F.

Hieke of this museum. It should be pointed out that within this series

only two specimens bear labels and 5 different methods of mounting are

used. The syntype selected as lectotype was chosen because it was the

first in the series, has 2 more detailed labels associated with it,

and its mount is unique in the series. The lectotype is pinned through

the right elytron with a short (23-24 ram long) silvery pin of about #5

thickness and the pin bears 3 labels (decending order): ”55898"

(Armoraciae EH.* Crioc. Nasturtii Fab? Hung. Jr. Rhen. Koy. Koch."

(see Fig. 3D and "LECTOTYPE Phyllotreta armoraciae Koch BY Eric H. Smith 44

'73"; the lectotype label was added by me. The paralectotypes are as follows: 2?, each mounted on a short white point supported by a thin silvery pin; Id, pinned with a very thin silvery pin bearing a very small cream colored to white square of paper below; 1?, pinned with a thin silvery pin bearing a very small cream colored to white square of paper below; 1?, mounted on a short gray point supported by a moderate size silvery pin and bearing the label "Halae. Erichs.”

[this could be Crisks.]; 2$, each mounted identically to the previous female but without the label.

SPECIMENS EXAMINED: Total: 763: CONNECTICUT: Litchfield Co.,

Cornwall (CUIC:1); IOWA: Clayton Co., Guttenburg (ANSP:1, USNM:2),

Henry Co., Rome (ISUI:1), Johnson Co., Iowa City (LACM:1, USNM:2),

Muscatine Co., Muscatine (OSUO:4); Scott Co., Pleasant Valley (ISUI:5),

Story Co., Ames (ISUI:7), Woodbury Co., Sioux City (DEFW:1); IDAHO:

Bonner Co., Sandpoint (JSCC:1, NMDC:5); ILLINOIS: Champaign Co., Urbana

(DEFW:1, WSUC:1), Cook Co., Chicago (CASC:3, CUIC:1, DEFW:1, MCZC:3,

OSUC:1, UWEM:4), Willow Springs (CASC:2, CUIC:2, FMNH:2), (MCZC:2),

(AMNH:2, CASC:43; INDIANA: Lake Co., Pine (FMNH:1, CASC:4), Tippecanoe

Co., Lafayette (CNCI:1); MASSACHUSETTS: Middlesex Co., Framingham

(MCZC:1), Stoneham (CASC:10; CUIC:10); MAINE: Oxford Co., Paris

(MCZC:2); MICHIGAN: Allegan Co., Fennville (CASC:5, LACM:2, MSUC:4),

Ingham Co., Ag. Coll. Mich. =Michigan St. Univ. (CASC:5, MSUC:23),

Ionia Co., Belding (MSUC:1), Lapeer Co. (MSUC:1), Oakland Co., Pontiac

(MSUC:1), (MSUC:1), St. Clair Co., Port Huron (MSUC:2); MINNESOTA:

Fairbault Co., Winnebago (CNCI:2, DEFW:7), Hennepin Co., Minneapolis 45

(DEFW:37), Houston Co. {'DEFW:21. Olmstead Co. (DEFW:4), Polk C o ..

E. Grand Forks (UCRC:2), Ramsey Co., St. Paul, University Farm

(DEFW:20), Rice Co., Fairbault (CNCI:1), Wabasha Co., Lake City

(CNCI:3, DEFW: 6), Winona Co., Dresbach (DEFW;15); MISSOURI: Pike Co.,

Louisana (ISUI:4), Mississippi Co., Charleston (USNM:1), Montgomery

Co., Montgomery City (CNCI:3); NEBRASKA: Butler Co., Bellwood (DEUN:

6); NEW HAMPSHIRE: Coos Co ., Mt. Washington (CNCI:1); NEW JERSEY:

Bergen Co., Hillsdale (CASC:1, NMDC:1), Morris Co., Chester (USNM:3);

NEW YORK: Broome Co., Binghamton (CUIC:1), Erie Co., E. Aurora (CASC:2),

Genesee Co., Batavia (ISUI:2), Lawrence Co., Potsdam (CASC:1, MCZC:4),

Madison Co., Bridgeport (JSCC:1, OSUO:3, USNM:5), Niagara Co., Olcott

(CUIC:14), Onondaga Co. (NMDC:4), Putnam Co., West Point (USNM:1),

Suffolk Co., Orient Point (USNM:3), Tompkins Co., Ithaca (CUIC:8),

(NMDC:2), (CUIC:223; OHIO: Ashtabula Co., Jefferson (OSUC:8), Crawford

Co., Plankton (CASC:2), Franklin Co., Columbus (WSUC:1), Wood Co.,

Bowling Green (0SUC:6), ECNCI:43 ; PENNSYLVANIA: Allegheny Co., Aspin­ wall (ICCM:4, MCZC:2), Pittsburgh (ICCM:20), Wilmerding (ICCM:4),

Centre Co., State College (PSUC:5), Dauphin Co., Hummelstown (PADA:1),

Forest Co. (PSUC:3), Monroe Co., Tobyhanna (PADA:19), Northhampton Co.,

Easton (CASC:2), Philadelphia Co., Bustleton, Pa. Sta. Col. Lab

(PSUC:2), Germantown (ANSP:5, CASC:6), Philadelphia (USNM:5), SOUTH

DAKOTA: Brookings Co., Brookings (MSUC:2, SDSU:16), Minnehaha Co.,

Colton (SDSU:2); WISCONSIN: Adams Co., Big Hats (UWEM:2), Brown Co.,

Green Bay (CNCI:1, DEFW:2, FMNH:1, MCZC:6, OSUC:7, PSUC:3, USNM:1,

UWEM:17), Dane Co., Madison (CNCI:1, USNM:43, UWEM:57), Door Co., 46

Sturgeon Bay (JSCC: 1, OSUO:1, UWEM:4), Oneida Co., Minocqua (UWEM:2) ,

Sauk Co., LaVille (UWEN:1), Walworth Co., Lake Geneva (CUIC:1, USNM:2,

UWEM:9), Waukesha Co., Oconomowoc (USNM:2), Okauchee (AMNH:2, CASC:6,

CUIC:4, MCZC:7, 0SUC:7), Waupaca C o .. Waupaca (UWEM:5), CAMNH:4l.

CANADA: MANITOBA: Brandon (CNCI:4), Rock Lake (CNCI:7); NEW BRUNSWICK:

Fredrickton (CNCI:1), Millville (CNCI:9); ONTARIO: Britannia (CNCI:1),

Burlington (CNCI:1), Chatham (CNCI:2), Dresden (CNCI:1), Leamington

(CNCI:1), Manotick (CNCI:1), Port Rowan (CNCI:4), Prince Edward Co.

(CNCI:6), Tilbury (CNCI:23), Welland (PSUC:1); QUEBEC: Duparquet

(CASC:1), Laprairie (CNCI:1), Montreal (CNCI:2), Outermont (CASC:4,

CNCI:8, LACM:10, MCZC:1, UCRC:3); SASKATCHEWAN: Blaine Lake (EHSC:1,

LBCC:1), Donovan (EHSC:1, LBCC:3), Duck Lake (LBCC:1), D u ndum

(LBCC:1), Forestry Farm (LBCC:1), Moose Jaw (CNCI:1), Saskatoon

(CNCI:2), Strasbourg (CNCI:1). NO DATA: [DEFW:2, MCZC:1, WSUC:l3.

Phyllotreta attenuata NEW SPECIES

(Figs. 9, 44-45, 89-90, 131, 152)

Holotype: Male, deposited at USNM.

Type locality: Cottonwood, South Dakota.

DIAGNOSIS: Elytra each with a median pale stripe with dilations which basally parallel suture and gradually incurve toward suture in apical 1/3 to 1/2 but never meet, medially stripe usually wider than distance from stripe to suture; antennal segments 4-5 simple; male antennal segments 6-11 darkest in contrast to paler basal segments; 47 male genitalia in lateral view with apex very slender, almost straight.

DESCRIPTION: Holotype (variation, excluding punctation, in paren­ theses): Fig. 9; elongate oblong, length 2.58 mm (male 2.20-2.62, female 2.30-2.65), width 1.18 mm (male 1.05-1.20, female 1.02-1.30); head and pronotum black with slight metallic luster, elytra brownish black (rarely with slight metallic luster), each with a median straw- r yellow stripe. Head: Black (rarely dark reddish brown or brownish black); texture finely granulate basally to smooth anteriorly (usually

finely granulate basally to slightly roughened anteriorly); moderately punctate, punctures separated by less than to 2x their diameter, mostly by less than one diameter; Interocular Distance/Maximum Diameter of eye, 1.33 (male 1.30-1.44, female 1.33-1.56). Antennae: Figs. 44,

45; segments 5 and 6 equal in length, slightly longer than 4 and

slightly shorter than segment 7; segments 4 and 5 simple, cylindrical;

segment Clength/width): 1(8.5/33, 2[4/2+3» 3(4/23, 4(4+/23, 5(5/2+3,

6(5/2.53, 7(5+/3-3, 8(5.5/33, 9(5.5/3+3, 10(5/3+3, 11(7/4-3, total

length 1.46 mm (Allotype: 1(8.5/33, 2(4+/23, 3(4/23, 4(4.5/23,

5(5.5/2.53, 6(5.5/3-3, 7(6/33, 8(6/3+3, 9(6-/3+3, 10(5/3+3, 11(7/3+3,

total length 1.55 mm; simple); antennae various shades of brown,

basal 4 (3) segments paler, 5 (4-5 or none) transitional, 6-11 (4-11 or

5-11 darkest). Pronotum: Length 0.50 mm (male 0.40-0.50, female 0.45-

0.50), width 0.82 mm (male 0.75-0.85, female 0.75-0.85); black; texture

finely granulate; coarsely punctate with moderate punctures interspersed,

punctures separated by less than 1/2 to equal their diameter, mostly

by less than one diameter. Elytra: Length 1.88 ram (male 1.60-1.95, 48 female 1.62-2.00), width 1.18 mm (male 1.05-1.20, female 1.02-1.30); brownish black (dark brown or black), median stripe as in Fig. 9; texture smooth ( u s u a l l y smooth to slightly roughened or entirely slightly roughened); coarsely punctate with very coarse punctures in­ terspersed in basal 1/2, punctures separated by less than 1/2 their diameter. Legs: Normal color sequence. Abdomen: 5th sternum with an apical median deeply concave lobe, concavity oblong and extending to within 1/3 of sternal base, with a median linear impression extending from sternal base to base of concavity (usually extending to base of median lobe, sometimes interrupted; female: simple in outline, usually with a shallow longitudinal median impression in apical 1/3); black

(brownish black) with median concavity paler (5th sternum unicolorous; females usually with apical 1/3 paler).

Male genitalia: Figs. 89, 90; length approximately 1.20 mm, mod­ erately arched dorsoventrally, of rather uniform width except slightly narrower in apical 1/3 in dorsal view and very slender in apical 1/4 in lateral view; with a dorsal washboard; apex in dorsal view acute, gradually tapering to a blunt point, in lateral view very narrow, ending in a sharp point.

Female genitalia: Fig. 131; length approximately 0.38 mm; recep­ tacle elongate, moderately swollen basally and tapering posteriorly to a slightly developed ring collar.

BIOLOGY: Host plants, adult: *marsh cress (Radicula palustris)

(Chittenden, 1923:135; recorded as £. oregonensis). New Collection

Record: sugar beets, "Rocky Ford, Col. 29 Aug 11, H O Marsh Collector, 49

Phyllotreta oregonensis Chttn. Ent. Am. 1927” CUSNM:1).

Immature stages: Unknown.

Habits: Adults have been collected in South Dakota from late May until mid-June and in Colorado in late August.

DISCUSSION: Distribution: Fig. 152; P_.’ attenuata has been collected in Colorado and South Dakota. New name: The specific epithet attenuata comes from the Latin attenuate meaning thin, and refers to the apical 1/4 of the male genitalia which is very narrow in lateral view.

Relationships: _P. attenuata is most similar to P. dolichophalla and the specimens of 1?. bipustulata which have the prebasal and pre- apical marks connected. These 3 species share having antennal segments

4 and 5 simple, segments 5 and 6 subequal in length, and the male genitalia very similar. The male genitalia differ mainly in that those of P. bipustulata are longest, those of £. dolichophalla are intermediate in length, and those of P. attenuata are shortest, and that the apex in lateral view is almost straight in P^. attenuata, whereas in bipustulata and P. dolichophalla it is distinctly directed ventrad.

However, the female spermathecae of these 3 species are very different: the receptacle of P. bipustulata is oval and robust with the ring collar only slightly developed and the spermathecal duct is attached dorsally, in P. dolichophalla the receptacle is elongate, strongly swollen in the basal 2/3 and rather uniformly narrow in the apical 1/3 with the ring collar strongly developed and the spermathecal duct attached pre- dorsally on the base, and in P; attenuata the receptacle is moderately 50 swollen basally and tapering posteriorly with the ring collar at most slightly developed and the spermathecal duct attached predorsally on the base. attenuata and JP. dolichophalla share the additional character of having the elytral stripe very wide medially, whereas in

P. bipustulata the stripe is narrow medially.

Phyllotreta attenuata might be confused with P. oregonensis on f the basis of having the elytral stripe wide medially, but P^. attenuata has antennal segments 5 and 6 subequal in length, 5 simple and cylin­ drical, whereas P. oregonensis has antennal segment 5 about 1.5x length of 6 and in the male, segment 5 is dorsoventrally flattened.

TYPE MATERIAL: The holotype, allotype, and 5 paratypes ([2d, 3?) are at the SDSU, 3 paratypes (2d, l?) are at the USNM, and 2 paratypes

(Id, 19) are in my collection, EHSC. The holotype is mounted on a point and the pin bears 2 labels (decending order): "Cottonwood S.D.

June 9, 1954 H.C. Severin Coll." and "HOLOTYPE Phyllotreta attenuata

By Eric H. Smith '73": also I added a polyethylene genitalia vial.

The allotype bears the same locality-date label and an allotype label.

The 5 paratypes at the SDSU are labeled with a paratype label and the following locality-date labels: same as holotype (Id, 1?); same as holotype but date is July 9, 1954 (15); "Midland S.D. June 9, 1954

H.C. Severin Coll." (2?). The 3 paratypes at the USNM are labeled with a paratype label and the following additional labels (decending order):

"Rocky Ford Col 26 Aug 15" "Radicula palustris" "H 0 MaTsh Collector"

"Chittenden No 1578" "oregonensis Phyllotreta Chttn. Ent Am 1927" (id, l9); "Rocky Ford Col 29 Aug 11" "Sugar Beets" "H 0 Marsh Collector" 51

"oregonensis Phyllotreta Chttn. Ent Am 1927" (1(J). The 2 paratypes

C1

SPECIMENS EXAMINED: Total 12: COLORADO: Crowley Co., Rocky Ford

(USNM: 3); SOUTH DAKOTA: Union Co., Elk Point (SDSU:2), Jackson Co.,

Cottonwood (EHSC:2; Holotype, Allotype, SDSU:3). /

Phyllotreta bipustulata (F.)

(Figs. 10-11, 46-47, 91-92, 132, 153)

Crioceris bipustulata Fabricius, 1801. Systema Eleutheratorum, 1:464.

Neotype (here designated): Male, deposited at USNM.

Type locality: "Carolinae"; neotype, Myrtle Beach, South Carolina,

Orchestris bipustulata, Crotch, 1873. Proc. Acad. Nat. Sci.

Philadelphia, 25:66.

Phyllotreta bipustulata, Horn, 1889. Trans. American Ent. Soc., 16:300.

DIAGNOSIS: Elytra each with a subbasal and subapical pale mark or

with these marks connected, sutural margin of subapical mark straight,

not incurved toward suture; antennal segments 4 and 5 simple and cylin­

drical, segments 5 and 6 subequal in length.

DESCRIPTION: Neotype (variation, excluding punctation, in paren­

theses): Fig. 10; elongate oblong, length 2.68 mm (male 2.0-2.72,

female 2.18-3.02), width 1.22 mm (male 0.95-1.32, female 1.18-1.40);

head and pronotum black with slight metallic luster, elytra black

(rarely with slight metallic luster), each elytron with a subbasal and subapical straw-yellow mark. Head: Black ( r a r e l y dark brown or brownish black); texture finely granulate basally to slightly roughened anteriorly (entirely finely granulate); moderately punctate, punctures separated by less than 1/2.to equal their diameter, mostly by less than

1/2 a diameter; Interocular Distance/Maximum Diameter of eye, 1.40

(male 1.22-1.56, female 1.33-1.50). Antennae: Figs. 46, 47; segment 4 — / shorter than 5, segments 5 and 6 subequal (equal) in length, segments

6 and 7 subequal (equal) in length; segments 4 and 5 simple and cylindrical; segment (length/width]: 1(8.5/3), 2(4-/2.5], 3(4/2],

4(4.5/2.53, 5(5.5/2+3, 6(5+/2+3, 7(5.5/3-3, 8(6-/3+3, 9(5.5/3.5],

10(5+/4-3, 11(6/43, total length 1.49 mm (female: 1(9/33, 2(4/2+3,

3(4-/23, 4(4+/23, 5(5-/2+3, 6(5+/2.53, 7(5/33, 8(5/3+3, 9(5/3.5],

10(5/33, 11(6.5/3], total length 1.44 mm; simple); antennae various shades of brown, basal 4 (5 or 6; 1st rarely darker) paler, 5 (5-6 or

5-7) transitional, 6-11 (5-11 or 7-11 or 8-11) darkest. Pronotum:

Length 0.60 mm (male 0.38-0.60, female 0.42-0.65), width 0.85 mm (male

0.68-0.95, female 0.70-0.95); black (rarely dark brown or brownish black); texture finely granulate (rarely finely alutaceous); coarsely punctate, punctures separated by less than 1/2 to equal their diameter, mostly by less than one diameter. Elytra: Length 1.88 mm (male 1.45-

2.02, female 1.55-2.20), width 1.22 mm (male 0.95-1.32, female 1.18-

1.40); black (often brownish black), color pattern as in Fig. 10

(sometimes as in Fig. 11); texture slightly roughened (smooth to slightly roughened); coarsely punctate, punctures separated by less than 1/2 to equal their diameter, mostly by less than one diameter. Legs: 53

Normal color sequence (often with tibiae and taTsi same color).

Abdomen: 5th sternum with an apical median deeply concave lobe, con­ cavity oblong and extending to within 1/3 of sternal base (oval in its extension to midpoint), with a median linear impression from sternal base to base of concavity (female: simple in outline, usually with a shallow median oblong impression in apical 1/3); black

(brownish black, rarely dark brown) with apical 1/2 of 5th sternum paler (entirely dark or with median lobe to apical 1/3 of 5th sternum paler).

Male genitalia: Figs. 91, 92; length approximately 1.32 mm; mod­ erately arched dorsoventrally, of rather uniform width except slightly narrowed preapically in dorsal view, slender and tapering toward apex in apical 1/3 in lateral view; with a dorsal washboard; apex in dorsal view acute, bluntly rounded, in lateral view very narrow and distinctly directed ventrad, ending in a sharp point.

Female genitalia: Fig. 132; length approximately 0.26 mm; recep­ tacle oval, very robust with ring collar slightly developed; speimathe- cal duct attached dorsally with distance to gland valve attachment very short.

BIOLOGY: Host plants, adult: *cabbage, *turnip, *hedge-mustard

(Sisymbrium officinale), *charlock (Brassica arvensis), *shephard's purse (Bursa bursapastdris) (Chittenden, 1902:77); *yellow rocket

(Barbarea barbarea), *marsh cress (Radicula palustris), *horse-radish,

*wild cress(Chittenden, 1923:136); *mustard, *radish, occasionally rose

(Duckett, 1920:137); *Lepidiuro virginicum (Chittenden, 1927:34); *Dentaria diphylla Michx. (Tahvanainen, 1972:125). New Collection

Records: *Barbarea verna, College Park Md May 19, 1920 (USNM:1);

♦Barbarea vul(garis), College Park Md May 19, 1920 (USNM:1); *Brassica kaber (feeding on), Delaware Co., 0. 13?VIII*72 E.H. Smith Lot No.

367 (EHSC:1); *Cardamine douglassi, Cedar Springs Ont. May 2 1945 A.A.

Wood (CNCI: 2); lettuce, W. Lafayette, Ind. 5-VIII-71 E$C Smith Lot No.

377 (EHSC:2); *mustard (feeding on), College Park Md 7-9-11 A B

Duckett Coll (UMDCrl), and Bladnsbg Md 8-4-12 Duckett Collection

(UMDC:1), and Bladnsbg Md 8-12-12 Duckett Collection (UMDC:4); *pepper- grass, LaFayette (sic.), Ind., Aug 3 1922 G.M. Stirrett (CNCI:1);

Prunis virginana flowers, Sherbom, Mass. V*30*38 C.A. Frost (MCZC:1); red clover, N. Bend Jackson Co., Wis My 3 1948 J.T. Medler Col.

*UWEM:1).

Immature stages: Unknown.

Habits: The larva is with little doubt, a root feeder (Chittenden,

1923:136). The adults overwinter in the top soil close to the host plants (Hicks, 1972) and become active in early spring (early May).

Eggs are laid in the soil around the host plant (Hicks, 1972) and the

adults decline thereafter and are rarely encountered after June. The

larvae probably spend most of the summer feeding and adults appear again

in the fall.

This species has been a pest in past years but it has apparently

been displaced for the most part by the more competitive £. cruciferae

(Goeze) (Hicks, 1972), which was apparently introduced from Europe in

the early 1920*s on the West Coast and probably again around 1940 on the 55

East Coast (Milliron, 1953). In the Ithaca, New York, area P^. bipustulata is restricted to woodland inhabiting Dentaria diphyila with the adults common on it until the leaves die off in late June; it has been collected via pitfall traps in rich woods in Franklin Co., Ohio.

In central Ohio I have collected it feeding on Brassica kaber along the roadside In mid-summer' and in northwestern Indiana feeding on turnips and on lettuce having flea beetle damage in early August; the collections consisted of predominately JP. cruciferae and P^. striolata, and a few £. zimmermanni.

Adults have been collected throughout their range from late March- early April to late June and again from mid-July until late August or until late October in northeastern and southeastern U.S. and Canada.

DISCUSSION: Distribution: Fig. 153; P_. bipustulata has been collected east of the Rocky Mountains but mostly east of the Mississippi

River in the U.S. and in Manitoba and Ontario in Canada.

Nomenclature: Fabricius (1801 a) described P^. bipustulata in the genus Crioceris. In 1873, Crotch wrongly placed bipustulata in

Orchestris (see LeConte, 1878:16S) and Horn (1889) placed this species

in Phyllotreta.

'The "type" of Crioceris bipustulata F., given as coleopteran number 1604 by Zimsen (1964:105), was borrowed from the University

Zoological Museum, Copenhagen, but the specimen proved to be a species of the genus Zeugophora Kunze and is very near Z. scutellaris Suffrian.

Fabricius* original description agrees with what we presently

call P. bipustulata and so does Olivier's redescription of the Fabrician 56 type (1808a:723-724) with the exception that both describe the elytral pale subapical mark as "lunatis" or crescent-shaped. This verbal description best fits JP. striolata discendens Wise, but

01ivier*s illustration of P. bipustulata (1808b:pl.5, fig.97) shows that the apex of this mark is not incurved toward the suture and it therefore matches bipustulata as presently understood.

Clavareau (in the Junk Catalogue, 1913:52) indicates that

Fabricius also described (1801b:40) another species, using the same specific epithet, in the genus Clytra. Clavareau lists this species as being in doubt. The Fabrician original description of

Clytra bipustulata does not fit either P^. bipustulata or the '‘type" sent. Unfortunately, Olivier does not treat £. bipustulata in his

Entomologie. Although there is no check on the identity of this species, it does not appear to be involved in the present problem.

The type locality is given as "Carolinaeu and the location of the type as in the Bose Collection. This suggests that the type locality be restricted to near Charleston, South Carolina (Blake, 1952). Blake

(1952) did not find C . bipustulata in the Bose Collection but she did

comment that all specimens were in good condition because "probably all

specimens that were not had long since been discarded." The specimen

sent from Copenhagen was accompanied by Fabricius* determination label which consists of a torn piece of brown paper with "2 pustulata"

written in black ink. This specimen is from the Kiel Collection which

was Fabricius* personal collection and was therefore probably not the

type specimen he referred to as in the Bose Collection. 57

Based on the "type” sent, the original description, and Blake's observations, it is most probable that the type of Crioceris bipustulata F. (1801:464, vol.l) has been lost and that the Fabrician determination label has been erroneously associated with another species of chrysomelid. Therefore, a neotype has been designated which represents the closest collection to the type locality known to me and which agrees with Fabricius* original description.

Relationships: £. bipustulata is most similar to P^. dolichophalla and P^. attenuata; for a discussion of these relationships see this

section under JP. attenuata.

Phyllotreta bipustulata has often been confused with the specimens of P. striolata which have the elytral stripe reduced to a subbasal

and subapical pale mark but in P^. bipustulata the elytral subapical mark has the sutural margin straight, not incurved toward the suture,

whereas in P. striolata the subapical mark is incurved apically toward

the suture. Also, in P. bipustulata antennal segment 5 is simple and

segments 5 and 6 are sub equal in length, whereas in P. striolata

antennal segment 5 in the male is enlarged, and in both sexes segment

5 is about 2x longer than 6.

TYPE MATERIAL: The neotype is at the USNM. It is mounted on a

point and the pin bears 3 labels (decending order): "Myrtle Beach, S.C.

21 March 1963 V.M. Kirk" "Washed up on beach" and "NEOTYPE Phyllotreta

bipustulata By Eric H. Smith ’73".

SPECIMENS EXAMINED: Total 613: ARKANSAS: Washington Co. (UADE:1);

CONNECTICUT: Litchfield Co., Cornwall (CUIC:1); DISTRICT OF COLUMBIA:

Washington (ANSP:5, CASC:1, DEFW:3, USNM:6); FLORIDA: Highlands Co., 58

Archbold Biol. Sta. (PSUC:1); ILLINOIS; Champaign Co., Urbana (MCZC:1);

CCNCI;1), Cook Co., Chicago (FMNHil, UWEM:1), Dekalb Co., DeKalb

(UWEM:1), McLean Co., Heyworth (CASC:8, CNCI:1), Piatt Co.„ White

(JSCC:2), St. ClaiT Co. (MCZC:2), Will Co., Joliet (FMNH:1), [DEFW:7,

FMNH;2, ICCMil, JSCC:8, MCZC:2, MSUC:1, WSUC;3]; INDIANA: Boone Co..

Ogden Dunes (FMNH:1 )f Crawford Co. (UWEM:4), Howard Co.. Kokomo f (CNCI:1), Knox Co. (CUIC:1), Lake Co ., Long Lake (FMNH:2), Marion Co .

(UWEM:2), Monroe Co. (ISUI:1), Perry Co . (UWEM:1), Starke Co.. Knox

(USNM:1), Tippecanoe Co., Lafayette (CNCI:8), W. Lafayette (EHSC:6),

(NMDC:8), CFMNH:33; IOWA: Boone Co., Ledges State Park (ISUI:5, JSCC:1),

Clayton Co., Guttenburg (USNM:3), Iowa Co.. Williamsburgh (ISUI:2),

Johnson Co., Iowa City (LACM:1, MCZC:1), Kossuth Co.. 2 mi. S. Ledyard

(ISUI:1), Lee Co., Keokuk (MCZC:1), Muscatine Co.. Muscatine (USNM:2),

Sac Co., Little Wall Lake, 2 mi. S. Jewell (ISUI:1), Scott Co ..

Davenport (ISUI:1), Pleasant Valley (CNCI:1), Story Co .. Ames (ISUI:21,

JSCC:2, LACM:3, UCRC:3), (ISUI:1), CCASC:1, ISUI:2, USNM:73; KANSAS:

Douglas Co.» Lawrence (CNCI:20), Pottawatomie Co.. Onago (CASC:2,

MSUC:1), ClCCM:1, USNM:1]; KENTUCKY: Christian C o .. 5 mi. W. Hopkins­

ville (CNCI:1), Henderson Co.. Henderson (LACM:1), Todd Co.. Trenton

(LACM:1), CUSNM:23; LOUISIANA: Lafavette Par.. Lafayette (SDSU:1);

MARYLAND: Baltimore Co.. Baltimore (CASC:1), Prince George1s Co..

Bladensburg (UMDC:5, UCRC:1), College Park (UMDC:3, USNM:2), [DEFW:3,

FMNH:1, USNM:3]; MASSACHUSETTS: Bristole Co., Fall River (MCZC:1),

Hampshire Co., Amherst (UCRC:1), Middlesex Co., Framingham (MCZC:2,

UCRC:1), Sherbom (MCZC:2), MICHIGAN: Eaton Co., Grand Ledge (USNM:1), 59

Gratiot Co. (MSUC:1). Ingham Co., Ag College (=Michigan State Univ.)

(CASC:3, MSUC:8), Lapeer Co. (MSUC:2), MaComb Co., Mt. Clemens,

Selfridge field (FMNH:5), Midland Co. (>1SUC:3); MINNESOTA: Crowing Co.,

Mille Lacs (DEFW:1), Goodhue Co. (DEFW:1), Kittson Co. (DEFW:1), Lake

Co., Two Harbors (DEFW:1), LeSueur Co., Mim. River (DEFW:1), ^Lincoln

Co., Hendricks (VMKC:1), Ramsey Co., near Gray Cloud Is. (CNCI:2,

DEFW:1), Scott C o ., Blakely (DEFW:1), Dunes at Jordan (DEFW:1), Sibley

Co., River near Blakely (DEFW:16), Traverse Co. (DEFW:1), Wabasha Co.,

Lake City (DEFW:1), [DEFW:6, WSUC:13; MISSISSIPPI: [CUIC:l]; MISSOURI:

Boone Co., Columbia (LACM:48), St. Louis C o., St. Louis (CNCI:3),

[DEFW:1, USNM:lH; NEBRASKA: Antelope Co ., Neligh (DEUN:3), Lancaster

Co., Lincoln (DEUN:4); NEW JERSEY: Burlington Co., Columbus (MCZC:20),

Riverton (MCZC:2), Camden Co ., Camden (MCZC:1, OSUC:6), Ocean Co.,

Lakehurst (AMNH:3), Warren Co., Phillipsburg (CASC:3), (CASC:1, SDSU:3j;

NEW YORK: Erie Co., Lancaster (USNM:1), Herkimer Co., Newport (MCZC:2),

Monroe Co., Rochester (LACM:6), Nassau Co., Flushing L.I. (CNCI:1),

Onondaga Co., Canton (NMDC:2), Elbridge (NMDC:1, USRC:2), Orange Co .,

Long Beach, L.I. (CUIC:1), Suffolk Co., Riverhead L.I. (VMKC:1), Tomp­ kins Co., Ithaca (USNM:1), Westchester Co., Yonkers (FMNH:1), (CASC:2,

DEFW:1, USNM:13; NORTH CAROLINA: Union Co. (VMKC:1), Wake Co., Raleigh

(NCSU:1); OHIO: Deleware Co. (EHSC:1, OSUC:2), Fairfield Co. (EHSC:1),

Franklin Co., Columbus (PADA:1, OSUC:2), (EHSC:1, Univ. Zool. Mus.,

Copenhagen), Green Co. (0SUC:1), Guernsey Co. (EHSC:1), Hamilton Co., Cin­ cinnati (MCZC:1), Hocking Co. (EHSC:1, 0SUC:1), Pickaway Co. (0 SUC:1 ),

Washington Co., Marietta (USNM:1), Wood Co., Bowling Green (0SUC:1), CANSP:1, USNM:l]; PENNSYLVANIA: Allegheny Co., Pittsburg (ICCM:4),

Cumberland Co., N. Cumberland (PADA:2, VMKC:1), Berks Co., Reading (FMNH:

2), Dauphin Co., Hummelstn (ANSP:1, 0SUC:1), Delaware Co., Glenolden

(OSUC:4), Lancaster Co., Lancaster (PSUC:2), Northampton Co., Easton

(CASC:4), Monroe Co., Water Ga (MCZC:1), Perry Co., Blain (PADA:1),

Philadelphia Co., Angora (MCZC:2), Germantown (ANSP:1), Philadelphia

(CASC:2, FMNH:1), Westmoreland Co., Jeanette (ICCM:7, PADA:3), St.

Vine. (ICCM:4), CANSP:1, MCZC:2, OSUC:21; SOUTH CAROLINA: Horry Co.,

Myrtle Beach (USNM:Neotype, VMKC:1); SOUTH DAKOTA: Brookings Co.,

Brookings (SDSU:11, VMKC:2), Volga (VMKC:2), Brule Co., Chamberlain

(SDSU:2), Butte Co., Newell (SDSU:1), Union Co., Canton (SDSU:1), Elk

Point (SDSU;6, VMKC:1), Yankton Co., Yankton (SDSU:1, VMKC:1) [VMKC:l3;

TENNESSEE: Smith Co., Elmwood (CASC:1), [AMNH:l]; VIRGINIA: Arlington

Co., Arlington (USNM:1), Augusta Co., Staunton (VPIC:1), (USNM:l3;

WEST VIRGINIA: Berkley Co., Berkley (USNM:1), CCASC:l]; WISCONSIN:

Dane Co., Madison (CASC:2, CUIC:1, JSCC:1, 0SU0:3, PADA:1, UWEM:14),

(UWEM:1), Jackson Co., N. Bend (UWEM:1), Monroe Co., Sparta (UWEM:1),

St. Croix Co., St. Croix Falls (UWEM:1), Sauk Co. CJSCC:1), (ANSP:l3.

CANADA: MANITOBA: Aweme (CNCI:2), Strathem (CNCI:3); ONTARIO: Black­ burn (CNCI:1), Cedar Springs (CNCI:6), Chatham (CNCI:1), Leamington

(CNCI:2), Ottawa (CNCI:1), Pelee Island (CNCI:1), Port Credit (CUIC:3),

Ridgeway (CASC:1), Toronto (CUIC:1); QUEBEC: Brome (CNCI:1), Knowlton

(CNCI:1). NO DATA: (AMNH:1, ANSP:4, CUIC:5, DEFW:2, FMNH:1, 0SUC:1,

USNM:4, WSUC:2 ]. 61

Phyllotreta bisinuata NEW SPECIES

(Figs. 12, 48-49, 93-94, 133, 154)

Holotype: Male, deposited at CASC.

Type locality: Green Point,:Humboldt Co., California.

DIAGNOSIS: Elytra each with median pale stripe ^simple, of uniform width and without dilations; male antennal segments 4 and 5 moderately expanded bilaterally, dorsoventrally flattened, 5 moderately concave ventrally; antennal segments 5-11 darkest in contrast to paler basal segments; metathoracic wings reduced to less than elytron length; male genitalia in lateral view with a very abrupt, strong bend sub- medial ly.

DESCRIPTION: Holotype (variation, excluding punctation, in paren­ theses): Fig. 12; elongate oblong, length 2.65 mm (male 2.65-2.75, female 2.68), width 1.28 mm (male 1.28-1.30, female 1.42); head and pronotum black with slight metallic luster, elytra dark brown, each with a median straw-yellow stripe. Head: Black; texture finely granulate basally to slightly roughened anteriorly (finely alutaceous basally); moderately punctate, punctures separated by less than to equal their diameter, mostly by less than one diameter; Interocular Distance/Maximum

Diameter of eye, 1.44 (male 1.44-1.53, female 1.56). Antennae: Figs.

48, 49; segment 4 longer than 6 and equal in length to 7, segment 5 more

than 2x length of 6 and almost 2x length of 4; segments 4 and 5 moderately expanded bilaterally, dorsoventrally flattened, 5 with venter deeply concave in basal 1/2 to outside; segment (length/width]: 62

1(8.5/4-3, 2(5-/3-3, 3(S/33, 4(5/4.53, 5(9/5.53, 6(3.5/3+3, 7(5/3.53,

8(5.5/3.53, 9(5/3.53, 10(5/4-3, 11(7/3+3, total length 1.59 mm

(Allotype: 1(8.5/43, 2(4.5/33, 3(4+/33, 4(5-/33, 5(7/3+3, 6(4.5/33,

7(5+/3+3, 8(5.5/43, 9(5.5/43, 10(5+/43, 11(6/43, total length 1.51 mm; simple); antennae various shades of brown, basal 4 (female 5) segments paler. 5 darkest, 6-11 darker than basal segments. Pfonotum: Length / 0.50 mm (male 0.50, female 0.52), width 0.88 mm (male 0.85-0.88, female

0.88); black; texture finely granulate to slightly roughened (entirely slightly roughened); coarsely punctate, punctures separated by less than 1/2 to equal their diameter, mostly by less than one diameter.

Elytra: Length 1.95 mm (male 1.95-2.00, female 1.95), width 1.28 mm

(male 1.28-1.30, female 1.42); dark brown, brownish black in basal 1/8, (usually entirely brownish black), median stripe pattern as in Fig. 12; texture smooth (smooth to slightly roughened); moderately punctate with coarse punctures interspersed basally, punctures separated by less than

1/2 to equal their diameter, mostly by about one diameter. Legs: Normal color sequence. Abdomen: 5th sternum with an apical median deeply concave lobe, concavity tapering as it extends almost to midpoint, with a median linear impression extending subbasally to just inside concavity

(extending from sternal base to base of median lobe, sometimes interrupted; female: simple in outline, with a very shallow oval median impression in apical 1/3); brownish black (black) with apical 1/3 paler (with only median lobe paler; female entirely dark).

Male genitalia: Figs. 93, 94; length approximately 1.32 mm; strongly sigmoid or twice bent, of rather uniform width except median lobe 63 strongly constricted just anterior to junction with basal piece in dorsal view, swollen medially in lateral view; apex in dorsal view ! broadly rounded, with a moderately deep and moderately wide emargxnation, in lateral view gradually tapering to a wide bluntly rounded end which is directed ventrad, with a small median subapijcal lobe on venter.

Female genitalia: Fig. 133; length approximately 0.38 mm; recep­ tacle elongate, of rather uniform width, moderately bent dorsoventrally, ring collar moderately developed.

BIOLOGY: Host plants and Immature stages are unknown.

Habits: Adults have been collected in northern California in mid-

June.

DISCUSSION: Distribution: Fig. 154; P. bisinuata is known only from

Humboldt Co., California. New name: The specific epithet bisinuata comes from the Latin bi- meaning two and the Latin sinua- meaning bend, that is, twice bent which refers to the male genitalia in lateral view being strongly sigmoid.

Relationships: P^. bisinuata is most similar to P_. lepidula and less similar to P. arcuata, P. spatulata, and the striped specimens of P. denticomis; for a discussion of these relationships see this section under P^. lepidula.

TYPE MATERIAL: The holotype and allotype are at the CASC and one paratype (3) is in my collection (EHSC). The aolotype is mounted on a point and the pin bears 4 labels (decending order): "Green Point

Humboldt Co., Cal. VI-9-16" "F.E. Blaisdell Collector" "Blaisdell

Collection" and "HOLOTYPE Phyllotreta bisinuata By Eric H. Smith *73"; 64

I added a polyethylene genitalia vial. The allotype and paratype bear the same first 3 labels, and then an allotype and paratype label respectively.

SPECIMENS EXAMINED: Total 3: CALIFORNIA: Humboldt Co., Green

Point (CASC:Holotype, Allotype, EHSC:1).

Phyllotreta constricta NEW SPECIES

(Figs. 13, 50-51, 95-96, 134, 155)

Holotype: Male, deposited at USNM.

Type locality: Rocky Ford, Colorado.

DIAGNOSIS: Elytra each with a median pale stripe with dilations which never meet at suture, stripe medially wider than distance from stripe to suture; male antennal segment 5 dark, expanded bilaterally and dorsoventrally flattened, subequal to 4 in width, with venter flat; male genitalia in lateral view abruptly narrowed in apical 1/2 and apex distinctly directed ventrad.

DESCRIPTION: Holotype (variation, excluding punctation, in paren­ theses) : Fig. 13; elongate oblong, length 2.62 mm (male 2.55-2.85, female 2.80-3.10), width 1.22 mm (male 1.15-1.30, female 1.32-1.48); head and pronotum black with distinct metallic luster, elytra brownish black with slight metallic luster (no luster), each with a median straw- yellow stripe. Head: Black; texture finely granulate basally to slightly roughened anteriorly (entirely finely granulate); moderately punctate, punctures separated by less than 1/2 to 2x their diameter, 65 mostly by less than one diameter; Interocular Distance/Maximum Diameter of eye, 1.56 (male 1.40-1.56, female 1.25-1.44). Antennae: Figs. 50,

51; segment 4 longer than 3,‘ 6, or 7, segment 5 about 1.4x ( u s u a l l y

1.5x) length of 4 and more than 2x length of 6, 6 distinctly shortest; segments 4 and 5 expanded bilaterally, dorsoventrally flattened; segment

5 distinctly longest, subequal to 4 in width, with venter flat; segment / (length/width]: 1(10/4], 2[4.5/2.5], 3(5/3+], 4(6/5], 5(8/5+], 6(3+/3],

7(4.5/3], 8(5/3], 9(5.5/3+], 10(5/3+], 11(7/3+], total length 1.56 mm

(allotype: 1(10/4-], 2(4.5/2.5], 3(5/3-], 4(5/2.5], 5(7-/3], 6(4.5/2.5],

7C5+/3.5], 8(5.5/3.5], 9(6-/3+], 10(5+/3+], 11(7/3.5], total length

1.61 mm; simple); antennae various shades of brown, basal 3 segments pale with dorsum of 1 and 3 darker (3 usually entirely pale), 4-5 darkest (4 sometimes only dark), segments 6-11 dark (female 4-11 dark).

Pronotum: Length 0.45 mm (male 0.42-0.50, female 0.48-0.95), width 0.82 mm (male 0.78-0.90, female 0.90-1.02); black; texture finely granulate to slightly roughened (usually entirely finely granulate); moderately punctate with fine and coarse punctures interspersed, punctures

separated by less than 1/2 to 3x their diameter, mostly by about one diameter. Elytra: Length 1.88 mm (male 1.80-2.02, female 1.90-2.32), width 1.22 mm (male 1.15-1.30, female 1.32-1.48); brownish black (dark brown laterally), median stripe pattern as in Fig. 13; texture slightly roughened (smooth to slightly roughened, or smooth); coarsely punctate with several moderate punctures interspersed, punctures separated by

less than 1/2 their diameter. Legs: Normal color sequence. Abdomen:

5th sternum with an apical median moderately concave lobe, concavity 66 tapering (moderately rounded) posteriorly and extending to midpoint, a median linear impression extending from sternal base to median lobe

(ending subbasally and/or extending to subapex, sometimes interrupted; female simple in outline, usually with a median longitudinal im­ pression in apical 2/5); black (brownish black) with median lobe paler.

Male genitalia: Figs. 95, 96; length approximately 0.95 mm; / slightly arched dorsoventrally in dorsal view* of rather uniform width, in lateral view abruptly narrowed in apical 1/2; apex in dorsal view bluntly rounded, in lateral view abruptly tapered to a ventrally directed, rounded end.

Female genitalia: Fig. 134; length approximately 0.42 mm; recep­ tacle elongate, strongly constricted submedially on venter.

BIOLOGY: Host plants, adult: *Cleome serrulatum, *marsh cress

(Radicula palustris), and *turnip (Chittenden, 1923:135; recorded as

P. oregonensis). Host plants, larva: Cleome serrulatum (Chittenden,

1923:135; recorded as P. oregonensis). These host records represent

Chittenden's Fowler, Colo, record of Cleome serrulatum, his Childress,

Tex. and Garden City, Kans. records of turnip, and part of his Rocky

Ford, Colo, record of Radicula palustris.

Immature stages: Undescribed.

Habits: The larvae are leaf miners of Cleome serrulatum (Chittenden,

1923:135). Adults have been collected in southern Saskatchewan and

Alberta in late April and late May, in the central Great Plains from mid-June until late August, and in New Mexico and Texas in early July.

DISCUSSION: Distribution: Fig. 155; £. constriCta has been collected in southcentral Canada, the central part of the Great Plains, southern 67

New Mexico, and northern Texas. New name: The specific epithet

constricta comes from the Latin constrictus meaning drawn together or

contracted, and refers to the male genitalia in lateral view being

abruptly narrowed in the apical 1/2.

Relationships: _P. constricta is most similar to JP. oregonensis and

is less similar to emarginata, zimmermarini, £. utanula, and _P.

/ utana. For a discussion of these relationships see this section under

P. oregonensis.

Phyllotreta constricta might be confused with £. attenuata and

P. dolichophalla on the basis of having the elytral stripe wide medially, but, P^. constricta has the male 5th antennal segment appearing

dorsoventrally flattened, whereas in _P. attenuata and P^. dolichophalla

it is simple.

TYPE MATERIAL: The holotype, allotype, and 20 paratypes (17d, 4$)

are at the USNM; 5 paratypes (d) are at CASC; 3 paratypes (6) are at

CNCI; 4 paratypes (2d, 2?) are at EHSC; 1 paratype (d) at JSCC; 4

paratypes (3d, 1$) are at MCZC; and 5 paratypes (4o, 1?) are at SDSU.

The holotype is mounted on a point and the pin bears 6 labels (decending

order): "Rocky Ford Col 26 Aug 15" "Radicula palustris" "H 0 Marsh

Collector" " d " "oregonensis Phyllotreta Chttn. Ent Am 1927" and

"HOLOTYPE Phyllotreta constricta By Eric H. Smith '73"; I also added

a polyethylene genitalia vial. The allotype bears the same labels as

the holotype except in the place of the sex label is "Chittenden No

1578" and an allotype label was added.

The paratypes all have a paratype label added. The 5 d1 paratypes

at CASC are as follows: 2, "Eldorado Springs, Colo VII-12-1939 J.W. 68

Green"; 2, "Porvenir, N. Mex. Dr. A. Fenyes" "A. Fenyes Collection"; and 1, "Show Low, Ariz.. IX-11-1941 0. Bryant." The 3 d paratypes at CNCI: 1, Elbow, Sask. 24-VII 1954 Brooks-Wallis"; 1, "Ft. Collins

Colo." TESTE 16067 Chittenden"; and 1, "Ft. Collins Colo 6-13-22"

"Colo 28" "TESTE 16067 Chittenden". The 4 paratypes in EHSC are: Id and 1$, "Childress Tex. July 8, 09" On turnip" "H 0 Marsh Collector"

/ and female has in addition "oregonensis Phyllotreta Chttn. Ent. Am

1927"; 1<5 and 1$, "Ft. Collins Colo. 7-11.22" sex symbol label

"oregonensis Phyllotreta Chttn. Ent Am 1927" and female has in addition

"Colo 25". The d paratype at JSCC is labeled the same first 4 labels as the holotype. The 4 paratypes at MCZC are all labeled "F.H. Snow" and "Frederick Blanchard Collection". The 5 paratypes at SDSU are as follows: Id and 1?, "Elk Point, S.D. June 27, H.C. Severin Coll.";

2d, "Elk Point, S.D. June 27, 1946 J.A. Lofgren, Coll."; and Id,

"Freeman, S.D. June 27, 1946 J.A. Lofgren, Coll." The 21 paratypes at the USNM are as follows: Id and 19, same first 4 labels of allotype;

Id and 19, same 5 labels as allotype; o, Medicine Hat Alta 31.V.25 F.S.

Carr" " d " "oregonensis Phyllotreta Chttn. Ent Am 1927"; Id, "3 mi. N

Tularosa N.M. 7/2/29 L. alyssoides" "from VE Romney"; Id, "Fowler

Col 9 June 04" "ESG Titus Collector" "Cleome" " d " ; Id and 1?, LARAMIE,

WYO. 6-14-93" "WICKHAM Collection 1933"; Id, "Ft. Collins Colo.

7*3*32" "Colo 26" "oregonensis Phyllotreta Chttn. Ent Am 1927"; 19,

"Ft. Collins Colo. 7-11*22" "9" "oregonensis Phyllotreta Chttn. Ent Am

1927"; d, "Garden City Kans"; 2d1, l9, "Childress Tex. July 8, 09"

"On turnip" "H 0 Marsh Collector" sex symbol label "oregonensis

Phyllotreta Chttn. Ent Am 1927"; 2d, "Childress Tex. July 8, 09" "On 69 turnip” "H 0 Marsh Collector; lo, "Lincoln Neb June".

SPECIMENS EXAMINED: Total 33: COLORADO: Crowley Co., Fowler

(USNM:1), Rocky Ford (JSCC:1, Holotype, Allotype, USNM:4), Larimer Co.,

Ft. Collins CCNCI:2, EHSC:2, USNM:2); KANSAS: Finney Co., Garden City

(USNM:1); NEBRASKA: Lancaster Co., Lincoln (USNM:1); NEW MEXICO:

Otero Co., 3 mi. N. Tularosa (USNM:1); SOUTH DAKOTA: Union C o ., Elk

Point (SDSU:5); TEXAS: Childress Co., Childress (EHSC:2, USNM:5);

WYOMING: Albany Co., Laramie (USNM:2). CANADA: ALBERTA: Medicine Hat

(USNM:1); SASKATCHEWAN: Elbow (CNCI:1).

Phyllotreta decipiens Horn

(Figs. 14, 52-S3, 97-98, 135, 156)

Phyllotreta decipiens Horn, 1889. Trans. American Ent. Soc., 16:298-299.

Lectotype (here designated): Male, at ANSP.

Type locality: Oregon and Washington Territory.

Phyllotreta decipiens (var.) ordinata Chittenden, 1927. Entomologica

Americana, 8(1):38. Type locality: Elko, Nevada.

DIAGNOSIS: Elytra each with 1 small subapical reddish brown to

yellow mark; male antennal segment 5 dark, simple but slightly enlarged,

about 2x length of 6.

DESCRIPTION: Lectotype (variation, excluding punctation, in paren­

theses): Fig. 14; oblong, length 2.35 mm (male 1.85-2.40, female 2.38-

2.55), width 1.15 mm (male 0.92-1.15, female 1.20-1.30); head and

pronotum black with slight metallic luster (head no luster), elytra

brownish black with slight metallic luster (no luster), each with a subapical yellowish brown mark. Head: Black; texture finely granulate

(to slightly roughened anteriorly); moderately punctate, punctures separated by less than to equal their diameter, mostly by less than one diameter; Interocular Distance/Maximum Diameter of eye, 1.22

(male 1.00-1.28, female 1.22-1.38). Antennae [segments 3-11 of left missing]: Figs. 52, 53; segments 4 and 7 equal (subequal) in length,

/ longer than 6, segment 5 about 2x length of 6; segment 5 simple, slightly enlarged in diameter; segment (length/width]: 1(9/3],

2C5-/2.5], 3(4+/2+], 4(4.5/2+], 5(7/3], 6(3.5/2+], 7(4.5/2.5],

8(5-/2.5], 9(5/3], 10(5/3+], 11(7/3+], total length 1.49 mm (female:

1(9/3], 2(4/2], 3(4/1.5],' 4(4/2-], 5(5.5/2], 6[3.5/2], 7(4.5/2+],

8(4.5/3], 9C4.5/3+], 10(4/3+], 11(5.5/3+], total length 1.32 mm; simple); antennae various shades of brown, basal 3 segments paler,

4-6 darkest (4 intermediate or usually 4-6 only dark), 7-11 dark

(4-11 or usually 5-ir darkest) (female: basal 2 or 3 segments paler,

3 or 4 intermediate, 4-11 or 5-11 darkest). Pronotum: Length 0.45 mm

(male 0.35-0.45, female 0.42-0.48), width 0.75 mm (male 0.60-0.75, female 0.60-0.80); black; texture finely granulate to slight roughened

(usually entirely finely granulate); coarsely punctate, punctures

separated by less than 1/2 to equal their diameter, mostly by less than

1/2 a diameter. Elytra: Length 1.68 mm (male 1.40-1.80, female 1.75-

1.90), width 1.15 mm (male 0.92-1.15, female 1.20-1.30); brownish black

(often black, rarely dark brown), color pattern as in Fig. 14; texture

slightly roughened (often smooth); coarsely punctate, punctures

separated by less than 1/2 a diameter, strong (to slight) serial 71 tendency. Legs: Normal color sequence. Abdomen: 5th sternum with an apical median shallowly concave, concavity oval and extending to apical 1/3 (oblong or tapering, extending to midpoint), with a median linear impression extending from midpoint to subapex (extending from sternal base to subapex, sometimes interrupted; female: simple in outline, usually with a shallow median oval impression in apical 1/4); / black (often brownish black.) with median lobe dark brown except median linear impression black (apical margin to apical 1/3 paler or entirely dark).

Male genitalia: Figs. 97, 98; length approximately 0.79 mm; mod­ erately arched dorsoventrally, of rather uniform width except slightly narrowed preapically in dorsal view, with a dorsal washboard; apex in dorsal view broadly acute (about 45°), lateral edges almost straight, in lateral view gradually tapering in apical 1/4 to very narrow apex, abruptly directed ventrad with tip pointed.

Female genitalia: Fig. 135; length approximately 0.30 mm; recep­ tacle elongate oblong, wider basally and gradually tapering to posterior moderately developed ring collar.

BIOLOGY: Host plants, adult: *radish» *turnip, sugar beet, and potato (Chittenden, 1927); the records for turnip and potato have been verified by me. New Collection Record: Solidago, Creston, B.C. 2*VI

1950 G. Stace Smith, C.A. Frost Collection 1962 (MCZC:1).

Immature stages: Unknown.

Habits: Adults have been collected in the Pacific Northwest and

Montana and Utah from late April until early August and again in late 72

August and mid-October; no dates are associated with other collection

records.

DISCUSSION: Distribution: Fig. 156; P; decipiens has been

collected mainly in the Pacific Northwest. The Texas record may be a

case of mislabeling.

Nomenclature: Horn (188®) described P^. decipiens from at least 3 f specimens and noted that 2 of them had a subapical pale mark on each

elytron, whereas the third was entirely black (it actually has a

slightly paler, indistinct subapical mark). Chittenden (1927) named

the color variation with the subapical pale marks £. decipiens ordinata.

However, the male genitalia are the same, and the other characters all

fall within the observed variation for this species. According to the

Rules of Zoological Nomenclature, this variety holds no taxonomic rank

and is included here only for completeness and clarity.

Relationships: P. decipiens is a rather distinctive species. The

male genitalia of I?, decipiens are most similar to those of P^. striolata,

but the apex in dorsal view in P^. decipiens is broadly acute whereas,

in P. striolata it is broadly rounded with an acute blunt tip. The

female spermathecae of P_. decipiens are most similar to those of

striolata, and differ mainly in that those of P. decipiens are longer,

more robust, and the pump is less than 1/2 the receptacle length.

TYPE MATERIAL: The lectotype and 2 paralectotypes (Id, 1$) are

at the ANSP. The lectotype is mounted on a point and the pin bears

3 labels (decending order): "Or.” "PARA-TYPE 3852.3" and "LECTOTYPE

Phyllotreta decipiens By Eric H. Smith *73". The $ paralectotype is 73 labeled "W.T." "PARA-TYPE 3852.2" and "PARALECTOTYPE Phyllotreta decipiens By Eric H. Smith '73" and the male paralectotype is labeled

"W.T." "LectoTYPE 3852" and "PARALECTOTYPE Phyllotreta decipiens By

Eric H. Smith ’73". This lectotype label on the male designated para­ lectotype by me was added by a subsequent curator and is not a valid designation. The male selected as lectotype was chosen because it has one complete antenna, whereas the other male lacks/antennae.

SPECIMENS EXAMINED: Total 148: CALIFORNIA: [ANSP:1, USNM:l3;

IDAHO: Booner Co., Sandpoint (NMDC:4); MONTANA: Gallatin Co., Mont.

Exp. Stat, LoLo (USNM:2); NEVADA: Elko Co., Elko (ANSP:1, CNCI:1,

USNM:1); OREGON: Clatsop C o ., Astoria (USNM:1), Columbia Co., St.

Helens (JSCC:1), Scappose (JSCC:24, 0SU0:1, NMDC:3), Vemo n i a (JSCC:1),

Crook Co ., 10 mi. SE of Prineville, Crooked River (JSCC:1), Deschutes

Co., F.edmond (JSCC:1), Harney Co. (0SU0:1), Jackson Co., Butte Falls

(NMDC:2), Talent (JSCC:2), Klamath Co., 9 mi. N.E. Bly (JSCC:1),

Crescent Lake (JSCC:1, NMDC:1), Crooked Cr. Sprg., Hwy. 62 Bridge

(JSCC:2), Crystal Cr., Upper Klamath Lk. (JSCC:1), Ft. Klamath, Crooked

Creek (JSCC:1), Klamath Falls, Poe Valley (NMDC:1), Klamath Fall's

above Ceary Ranch (JSCC:1), Klamath Falls, Ceary Ranch (JSCC:1), Klamath

Falls (JSCC:2), Lake of the Woods (CUIC:4, MC2C:4), Odessa (JSCC.-2,

NMDC:1), Spregus Riv., 5 mi. E. Bly (JSCC:4), Williamson Riv. (JSCC:2),

Upper Klamath Lk., 3 mi. Creek (JSCC:1), Upper Klamath Marsh, Military

Crossing (JSCC:1), 2.5 mi. N. Hwy. 66, Buck Lake Road (JSCC:1), Lake Co.,

Chewavcan R, near Paisley (JSCC:1), Chewavcan River near Valley Falls

(JSCC:1), Lane Co., Beaver Marsh (JSCC:1), Multnomah Co., Portland

(AMNH:2, ANSP:1, MC2C:1, USNM:4), Tillamook Co., Tillamook Naval Base 74

(JSCC:1), Umatilla Co,, Weston (JSCC:1), Yamhill Co., McMinnville

CUCRC:1), CANSP:1, DEFW:1, 0SU0:1, USNM:23; TEXAS: [MCZC:ll; UTAH:

Wasatch Co., Heber (MSUC:1); WASHINGTON: Benton Co., Kennewick

(JSCC:1), King Co., Auburn (USNM:1), Skagit Co., Mt. Vernon (WSUC:3),

Thurston Co., Olympia (MCZC:2), Tenino (USNM:4), Whatcom Co., Femdale

(WSUC:6), Whitman Co., Pullman (WSUC:2) [WSUC:6]. [Washington t Territory, (ANSP:2)3. CANADA: ALBERTA: Watertown (CNCI:1); BRITISH

COLUMBIA: Creston (MCZC:1), Terrace (AMNH:6, CUIC:9, MCZC:2), Victoria

(USNM:1).

Phyllotreta denticomis Horn

(Figs. 15-16, 54-56, 99-100, 136, 157)

Phyllotreta denticornis Horn, 1889. Trans. American Ent. Soc., 16:

297-298, fig. 19.

Lectotype (here designated): male, at ANSP.

Type locality: California.

Phyllotreta amphicomis Chittenden, 1927. Entomologica Americana,

8(n.s.,1):35-36. Type locality: Wawawai, Washington. NEW SYNONYMY-

Phyllotreta aequalis Hatch, 1971. The beetles of the Pacific Northwest.

Part V:210. Type locality: Bear Springs, Oregon. NEW SYNONYMY.

DIAGNOSIS: Elytra each with median pale stripe simple, of uniform width and without dilations, or reduced to a subbasal and/or subapical mark, OR elytron entirely dark; male antennal segment 5 expanded bilaterally, appearing dorsoventrally flattened, segment 6 with a sharp anteroventrally directed process. 75

DESCRIPTION: Lectotype (variation, excluding punctation, in paren­ theses): Fig. 15; elongate oblong, length 2.65 mm (male 2.45-2.80, female 2,55-2.80), width 1.20 mm (male 1.00-1.20, female 1.20-1.28); head and pronotum black with slight metallic luster, elytra black with slight metallic luster (no luster), each with a somewhat indistinct, slightly paler median stripe. Head: Black; texture finely granulate basally to slightly roughened anteriorly (entirely slightly roughened); moderately punctate, punctures separated by less than 1/2 to equal their diameter, mostly by less than 1/2 a diameter; Interocular

Distance/Maximum Diameter of eye, 1.62 (male 1.44-1.74, female 1.56-

1.75). Antennae: Figs. 54, 55, 56; segment 4 longer than 3 or 7, shorter than 5, segment 5 more than 2x length of 6; segment 5 bilater­ ally expanded, appearing dorsoventrally flattened, venter concave in basal 2/3; segment 6 with a sharp anteroventral process, segments 7 and 8 with a short blunt apical process; segment (length/width]:

1C8+/43, 2(5/33, 3(5/43, 4(6/5.53, 5(7.5/6-3, 6(2.5/3, 4.5 with process3, 7(S-/4], 8(5/4+3, 9(5/4-3, 10(4.5/33, 11(6.5/33, total length

1.50 mm (female: l(8/3+3, 2(5-/3-3, 3(5-/2.53, 4(5.5/3-3, 5[6-/3-3,

6(4/3-3, 7(5/33, 8(6-/3.53, 9(4.5/4-3, 10(5-/4-3, 11(6.5/3.53, total length 1.51 mm, simple); antennae various shades of brown, basal 2

(3) segments paler with dorsum of segment 1 dark, 3-6 (4-5 or none) darkest, 7-11 (6-11 or 4-11) darker (female: basal 3 or 4 segments paler, 4-11 or 5-11 darker). Pronotum: Length 0.45 mm (male 0.42-0.48, female 0.42-0.48), width 0.82 mm (male 0.75-0.85, female 0.80-0.82); black (rarely brownish black); texture finely granulate (finely 76

granulate to slightly roughened, rarely finely alutaceous to finely

granulate); coarsely punctate, punctures separated by less than 1/2

their diameter. Elytra: Length 1.92 mm (male 1.75-2.02, female 1.88-

2.02), width 1.20 mm (male 1.00-1.20, female 1.20-1.28); black (dark brown, usually brownish black), median stripe indistinct, slightly paler .than background color (stripe distinct as in Fig. 15, or

/ reduced to a subbasal and/or subapical mark as in Fig. 16, OR elytron

entirely dark); texture smooth (smooth to slightly roughened); moderately and coarsely punctate, punctures separated by less than 1/2

to equal their diameter, mostly by less than 1/2 a diameter. Legs:

Normal color sequence. Abdomen: 5th sternum with an apical median moderately concave lobe, concavity tapering and extending to midpoint

(extension ending subbasally, oblong and extending to midpoint), with

a median linear impression extending prebasally to apex (female: simple

in outline, usually with a shallow to moderately deep oval impression

in apical 1/3); black (usually brownish black) with concavity and apical margin paler (apical 1/3 paler or entirely dark).

Male genitalia: Figs. 99, 100; length approximately 0.99 mm;

essentially straight except for basal piece slightly and apex strongly

curved ventrad, of rather uniform width except apical 1/4 slightly

wider; apex in dorsal view very bluntly rounded with shallow, moder­

ately wide median emargination, in lateral view strongly and rather

abruptly curved ventrad with a small ventral median subapical lobe, of

rather uniform width with end bluntly rounded.

Female genitalia: Fig. 136; length approximately 0.34 mm; recep­

tacle elongate slightly swollen in basal 1/3, gradually tapering to a 77 slightly developed ring collar posteriorly.

BIOLOGY: Host plants and Immature stages are unknown.

Habits: Adults have been collected in California, Oregon, and

Washington from late April until early July.

DISCUSSION: Distribution: Fig. 157; P. denticomis is known only from California, Oregon, and Washington.

Nomenclature: Horn (1889) described JP. denticomis from male specimens. In 1927, Chittenden described JP. amphicomis remarking that its antennae were similar to those of £. denticomis but that the 6th antennal segment lacked the acute process; the acute process is present on the 6th antennal segment of his unique type and the specimen is otherwise within the normal variation of P_. denticomis. Hatch (1971) described P. aequalis from a striped specimen of JP. denticomis and not noting the acute process on the 6th antennal segment, placed it close to P. lepidula. The type of PJ. aequalis has a well developed acute process on the 6th antennal segment, and is a normal, striped _P. denticomis.

Relationships: P. denticomis is most similar to P. spatulata, and less similar to P. arcuata, P^. bisinuata, and P^. lepidula with which it shares having a simple elytral stripe and having the male 5th antennal segment dorsoventrally flattened. Phyllotreta denticomis and P. spatulata have very similar male genitalia, differing mainly in the ventral aspect of apex in lateral view which in _P. denticomis has a small subapical lobe, whereas P. spatulata there is a subapical notch; in dorsal view of the apex, JP. denticomis is shallowly emarginate, whereas P. spatulata is faintly truncate. Phyllotreta denticomis has 78 a sharp process on the male 6th antennal segment which is unique.

TYPE MATERIAL: The lectotype and paralectotype (5) are at the

ANSP. The lectotype is mounted on a point and the pin bears 4 labels

(decending order); "Cal" "LectoTYPE 3850" "P. denticomis Horn" and

"LECTOTYPE Phyllotreta denticomis By Eric H. Smith '73". The para­

lectotype pin bears 3 labels (decending order): "Cal" "PARA-TYPE

3850" and "PARALECTOTYPE Phyllotreta denticomis By Eric H. Smith *73".

The curatorially added lectotype 3850 label is not a valid designation.

SPECIMENS EXAMINED: Total 23: CALIFORNIA: Los Angeles Co., Los

Angeles (CASC:1), San Luis Obispo C o ., Santa Margarita (CASC:1),

Shasta Co ., Castle Crag. (CASC:2), ANSP:1 ; OREGON: Deschutes Co.,

Redmond (JSCC:1), Hood River Co., Hood River (USNM:2), Josephine C o .,

Cave Junction (JSCC:l)f Wasco Co., Bear Springs (BMUW:1), The Dalles

(JSCC:1, USNM:8), 5 mi. S. The Dalles, 8 Mile Creek (JSCC:2), Rowena

Loops Viewpoint (JSCC:1); WASHINGTON: Whitman Co ., Wawawai (USNM:1).

Phyllotreta dolichophalla NEW SPECIES

(Figs. 17, 57-58, 101-102, 137, 158)

Holotype: Male, deposited at CASC.

Type'locality: Stinson's Beach, Marin Co., California.

DIAGNOSIS: Elytra each with a median pale stripe with dilations

basally somewhat and apically abruptly incurved toward suture, but never

meeting at suture; antennal segments 4-5 simple, male antennal segments

6-11 darkest in contrast to paler basal segments; male genitalia in

lateral view with apex very narrow, directed ventrad. 79

DESCRIPTION: Holotype (variation, excluding punctation, in paren­ theses) : Fig. 17; oblong, length 2.78 mm (female 3.12), width 1.30 mm

(female 1.45); head and pronotum black with slight metallic luster, elytra brownish black, each with a median straw-yellow stripe. Head:

Black; texture finely granulate basally to slightly roughened anteriorly; finely to moderately punctate, punctures separated by / less than 1/2 to 1.5x their diameter, mostly by about one diameter;

Interocular Distance/Maximum Diameter of eye, 1.67 (female 1.50).

Antennae: Figs. 57, 58; segments 4-5-6 equal (subequal) in length, segment 7 longer than (equal to) 6; segments 4 and 5 simple, cylindri­ cal; segment (length/width]: 1(10/4], 2(5-/3], 3(5-/3-], 4(5-/3],

5(5.5/3], 6(6/3.5], 7(6/4-], 8(6/4-], 9(6/4-], 10(5.5/4], 11(7/4], total length 1.68 mm (Allotype: 1(10/4], 2(5/3+], 3(6/3], 4(6/3],

5(6/3], 6(6/3], 7(7.5/4-3, 8(7+/4], 9(7+/4.5], 10(6/4.5], 11(9.5/4+3, total length 1.90 mm; simple); antennae various shades of brown, basal

5 segments paler, 6-7 intermediate, 8-11 darkest. Pronotum: Length

0.58 mm (female 0.60), width 0.92 mm (female 1.02); black; texture slightly roughened to finely alutaceous (finely granulate to slightly roughened); finely to moderately punctate, punctures separated by less than 1/2 to 2x their diameter, mostly by about two diameters. Elytra:

Length 1.98 mm (female 2.28), width 1.30 mm (female 1.45); brownish black, median stripe as in Fig. 17; texture slightly roughened (smooth to slightly roughened); coarsely punctate, punctures separated by less than 1/2 to equal their diameter, mostly by less than 1/2 a diameter.

Legs: Normal color sequence except tibiae and tarsi same color. 80

Abdomen: 5th sternum with an apical median deeply concave lobe, concavity tapering and extending to within 1/3 of sternal base, with a median linear impression extending subbasally to base of median lobe, interrupted at base of concavity (female: simple in outline, with a median longitudinal impression in apical 1/2); black (brownish black with apical 1/3 of 5th sternum paler).

Male genitalia: Fig. 101, 102; length approximately 1.42 mm; mod­ erately arched dorsoventrally, of rather uniform width except gradually tapering toward apex in apical 1/3 in lateral view; apex in dorsal view acute with tip bluntly rounded, in lateral view very narrow, directed ventrad, ending in a sharp point.

Female genitalia: Fig. 137; length approximately 0.38 mm; recep­ tacle unique, basal 1/2 strongly swollen ventrally, apical 1/2 of rather uniform width, ring collar very wide, strongly developed.

BIOLOGY: Host plants and Immature stages are unknown.

Habits: Adults have been collected in Marin Co., California in mid-April.

DISCUSSION: Distribution: Fig. 158; P. dolichophalla has been collected only in Marin Co., California. New name: The specific epithet dolichophalla comes from the Greek dolicho- meaning long and the Greek phallo meaning penis, which refers to the male genitalia being much

longer than the average length for the genus.

Relationships: P. dolichophalla is most similar to P^. attenuata and the specimens of bipustualta which have the prebasal and preapical marks connected; for a discussion of these relationships, see this • 81 section under P. attenuata.

Phyllotreta do 1 ichophalla might be confused with P^. oregoriensis on the basis of having theelytral stripe wide medially, but P^. dolichophalla has antennal segments 5 and 6 subequal in length, 5 simple and cylindrical, whereas P^. oregonensis has antennal segment

5 about 1.5x length of 6 and in the male, segment 5 is dorsoventrally t flattened.

TYPE MATERIAL: The holotype and allotype are at the CASC. The holotype is mounted on a point and the pin bears 4 labels (decending order): "Stinson's B Cal 4*11*33" "COL BY E J BLUM" "J.E. Blum Coll­ ection, gift of W.H. Nutting. Calif. Acad. Sci. Accession 1968" and

"HOLOTYPE Phyllotreta dolichophalla By Eric H. Smith *73"; I added a polyethylene genitalia vial. The allotype bears the same first 3 labels and the 4th label reads "ALLOTYPE Phyllotreta dolichophalla By

Eric H. Smith '73".

SPECIMENS EXAMINED: Total 2: CALIFORNIA: Marin Co., Stinson's

Beach (CASC:Holotype, Allotype).

Phyllotreta emarginata NEW SPECIES

(Figs. 18, 59-60, 103-104, 138, 159)

Holotype: Male, deposited at CASC.

Type locality: Williamson River Ranch, Klamath Co., Oregon.

DIAGNOSIS: Elytra each with a median pale stripe with dilations which never meet at suture, medially stripe width distinctly less than 82 distance from stripe to suture; male antennal segment 5 dark, expanded bilaterally and dorsoventrally flattened, subequal to 4.in width, with venter flat; apex of male genitalia in dorsal view with a narrow deep median emargination, straight ventrally in lateral view with no subapical concavity.

DESCRIPTION: Holotype (variation, excluding pqnctation, in paren­ theses): Fig. 18; elongate oblong, length 2.78 mm (male 2.25-2.78, female 2.50-2.80), width 1.32 mm (male 0.95-1.32, female 1.22-1.50); head and pronotum black with slight metallic luster, elytra brownish black, each with a median straw-yellow stripe. Head: Black; texture finely granulate (finely granulate basally to slightly roughened anteriorly); moderately punctate, punctures separated by less than 1/2 to equal their diameter, mostly by less than one diameter; Interocular

Distance/Maximum Diameter of eye, 1.40 (male 1.30-1.50, female 1.44-

1.76). Antennae: Figs. 59, 60; segments 4 and 7 equal (subequal) in length, 6 about 1/2 length of 4, segment 5 about 2x length of 4 and subequal to 4 in width; segments 4 and 5 expanded bilaterally, dorso­ ventrally flattened; segment 5 distinctly longest, with venter flat, not concave; segment Clength/width3: lCll/4.53, 2t4/33, 3C4.5/43,

4C5.5/S], 5C10/63, 6C3.5/3+3, 7C5.5/3.53, 8C5+/4-3, 9[5+/4-3, 10C5/4-3,

11[6.5/3+3, total length 1.64 mm (female paratype: 1C9/3.S3, 2C4/2.53,

3C4/2.53, 4C5/2.53, 5[6/3-3, 6[4/3-3, 7[S3/33, 8E5/33, 9[5/33,

10C5-/33, 11E6/33, total length 1.45 mm; simple); antennae various shades of brown, basal 2 (females 3) segments pale with dorsum of 1 dark, segments 3 and 8-11 (usually 7-11; females 4-11) dark, 4-7 83

(usually 4-6) darkest. Pronotum: Length 0.50 mm (male 0.40-0.50, female 0.48), width 0.92 mm (male 0.72-0.92, female 0.82-0.92); black; texture finely granulate; coarsely punctate with moderate punctures interspersed, punctures separated by less than 1/2 to equal their diameter, mostly by less than one diameter. Elytra: Length

2.02 iran (male 1.42-2.02,- female 1.80-2.05), width 1.32 mm (male 0.95- / 1.32, female 1.22-1.50); brownish black, median stripe pattern as in

Fig. 18; texture slightly roughened (smooth); coarsely punctate with moderate punctures interspersed, punctures separated by less than 1/2 to equal their diameter, mostly by less than 1/2 a diameter. Legs:

Normal color sequence. Abdomen: 5th sternum with an apical median lobe moderately concave, concavity tapering as it extends to within

1/3 of sternal base, a median linear impression in middle 1/3 only

(female simple in outline, no median impression); black.

Male genitalia: Figs. 103, 104; length approximately 0.94 mm; slightly sigmoid, of rather uniform width; apex in dorsal view abruptly triangular with tip broadly rounded each side of a median narrow moderately deep emargination, in lateral view almost straight ventrally, not directed ventrad, with no ventral subapical concavity, tip bluntly rounded.

Female genitalia: Fig. 138; length approximately 0.36 mm; recep­ tacle elongate oblong with a slight dorsoventral bend, basal 1/2 slightly swollen, with a slightly developed ring collar at junction with pump.

BIOLOGY: Host plants and Immature stages are unknown.

Habits: Adults have been collected in California in mid-June and 8 4 in Oregon from late May until late July.

DISCUSSION: Distribution: Fig. 159; £. emarginata has been collected in California and Oregon. New name: The specific epithet emarginata comes from the Latin emarginatus meaning notched at the apex, and refers to the apex of the male genitalia in dorsal view which has a narrow and moderately deep emargination.

/ Relationships: P. emarginata is most similar to £. zimmermanni and is less similar to P. utana, P. utanula, £. oregonensis, and P. constricta with which it shares elytral color pattern and having the male 5th antennal segment appearing dorsoventrally flattened. However, the elytral stripe of P. oregonensis and P. constricta is usually much wider medially than that of the other 4 species. The male 5th antennal

segment of P. zimmermanni has a basal concavity ventrally, whereas

that of the other 5 species is evenly flattened ventrally; also antennal

segments 2-5 are pale in P. utanula and £. utana, whereas in the

other 4 species antennal segments 2-3 are pale and/or dark and segments

5-11 are dark. Phyllotreta emarginata has the male 5th antennal segment

subequal to the 4th, whereas zimmermanni has the 5th distinctly wider

than the 4th. The male genitalia of P. emarginata, zimmermanni, and

P. liebecki form a distinct group which in dorsal view have the apex

narrowly and shallowly or moderately emarginate. However, those of P.

zimmermanni and P. liebecki in dorsal view have the apex broadly

rounded, whereas those of P. emarginata are abruptly triangular, and

those of P. emarginata and P. zimmermanni have the median emargination

moderately deep, whereas those of P. liebecki have the median emargina­

tion shallow; in lateral view P. emarginata and P. zimmermanni are 85 slightly sigmoid, whereas P. liebecki is evenly and moderately arched dorsoventrally. The female spermathecae of P. emarginata and _P. zimmermanni are distinctive from the other 3 species (females of .P. utanula are unknown) because the receptacle is slightly bent dorso- ventrally and the ring collar is well developed. However, P_. emarginata have the receptacle more robust than those of IV. zimmermanni.

TYPE MATERIAL: The holotype, allotype, and 1 paratype (3) are at the CASC. Two paratypes (3, 9) are at the USNM, 2 paratypes (3, 9) are in my collection (EHSC), 1 paratype (3) at CNCI, 1 paratype (3) at MCZC, and 5 paratypes (3) are in the JSCC collection. The holotype is mounted on a point and the pin bears 2 labels (decending order):

"Ore.; Klamath Co. Williamson Riv. Rch.; alfalfa 8 cruciferous weeds;

5-28-70 Joe Schuh, Coll." and "HOLOTYPE Phyllotreta emarginata By

Eric H. Smith '73"; also a polyethylene genitalia vial was added. The allotype bears 2 labels; "Ore.; W. side of Upper Klamath La. July 30,

1969 Joe Schuh, Coll." and "ALLOTYPE Phyllotreta emarginata By Eric H.

Smith *73".

The paratypes all have a paratype label added. The male at the

CASC is labeled: "Orinda, Cal Con Cos Co VI-10-1939" "B.E. White

Collector" and "BURDETTE E. WHITE Collection Calif. Acad. Sci.

Accession 1967". The 2 paratypes at the USNM and the 2 in my collection

(EHSC) are all labeled: "Mare's Egg Spring, Ore.; Klamath County May

30, 1962 Joe Schuh, Collector". The paratype at the CNCI and the one at

MCZC are both labeled with the same locality-date label as the holotype.

The paratypes in the JSCC are as follows: 3, same as allotype, 6 86

"Sprague Riv., Ore. 5 mi.:E. Bly June 16, 1957 Joe Schuh, Coll.";

<$ and $, "Crystal Cr., Ore. Upper Klamath Lk. May 30, 1960 Joe

Schuh, Coll."

SPECIMENS EXAMINED: Total 13: CALIFORNIA: Contra Costa Co.,

Orinda (CASC:1); OREGON: Klamath Co., Crystal Creek, Upper Klamath

Lake (JSCC:2), Mare's Egg Spring (EHSC:2, USNM:2), ^Sprague River, 3 mi. E. Bly (JSCC:1), W. side of Upper Klamath Lake (CASC:Allotype,

JSCC:1), Williamson River Ranch (CASC:Holotype, CNCI:1, MCZC:1).

Phyllotreta lepidula (LeConte)

(Figs. 19, 61-62, 105-106, 139, 160)

Haltica lepidula LeConte, 1857. Report of insects collected on the

survey. Washington, p.68.

Lectotype (here designated): Male, at MCZC.

Type locality: San Jose and San Diego, California.

Orchestris lepidula, Crotch, 1873. Proc. Acad. Nat. Sci. Philadelphia,

25:65-66.

Phyllotreta lepidula, Horn, 1889. Trans. American Ent. Soc., 19:294-295.

DIAGNOSIS: Elytra each with its median pale stripe simple, of

uniform width and without dilations; male antennal segment 5 moderately

expanded bilaterally, dorsoventrally flattened, moderately

concave ventrally; antennal segments 2-3 pale in contrast to darker

5-11; male genitalia in lateral view slightly sigmoid.

DESCRIPTION: Lectotype (variation, excluding punctation, in paren­

theses) : Fig. 19; elongate oval, length 2.60 mm (male 2.60-2.75, female 2.55-2.72), width 1.22 mm (male 1.18-1.28; female 1.18-1.32); head and pronotum black with slight metallic luster, elytra brownish black

(with slight metallic luster), each with a median straw-yellow stripe.

Head: Black (brownish black), texture finely roughened, alutaceous basally (finely granulate; smooth but finely granulate or roughened basally); moderately punctate, punctures separated by less than 1/2 their diameter; Interocular Distance/Maximum Diameter of eye, 1.33

(male 1.33-1.50, female 1.33-1.38). Antennae: Figs. 61, 62; segment 4

slightly longer than (subequal to) 7, 5 almost 2x length of 4, 6 about

1/2 length of 4; segment 5 moderately expanded bilaterally, appearing dorsoventrally flattened, venter moderately concave entire length;

segment (length/width): 1(8/3], 2(4/2+], 3(4/2), 4(4.5/3], 5(8/4.53,

6(2.5/23, 7(4/3-3, 8(4.5/3], 9(4/3], 10(4/3], 11(6/3], total length

1.34 mm (female: l(8/2.5], 2(4/2], 3(3.5/3-3, 4(4/2], 5(6/2+], 6(4/2+],

7(4/3-], 8(4.5/3], 9(4.5/3-3, 10(4/2.5], 11(5.5/3-], total length 1.30

mm; simple); antennae various shades of brown, basal 4 (3) segments

paler with segments 1 and 4 dorsally darker (4 sometimes completely

darker), apical 7 (8) segments darker. Pronotum: Length 0.45 mm

(male 0.38-0.45, female 0.42-0.42), width 0.79 mm (male 0.75-0.85,

female 0.78-0.85); black; texture slightly roughened with wrinkles

radiating from most punctures (smooth but with wrinkles); coarse to

very coarsely punctate, punctures separated by less than 1/2 their

diameter. Elytra: Length 1.98 mm (male 1.92-2.10, female 1.90-2.08),

width 1.22 mm (male 1.18-1.28, female 1.18-1.32); brownish black (dark

brown), median stripe pattern as in Fig. 19; texture slightly

roughened (usually smooth); coarsely punctate, punctures separated by 88 less than 1/2 their diameter. Legs: Normal color sequence. Abdomen:

5th sternum with an apical median deeply concave lobe, its concavity extending and broadly tapering basad to within 1/3 of base, trans­ versed by a median linear impression extending subbasally to sub- apical ly (female simple in outline, with a shallow lanceolate median impression starting at midpoint and ending subapically); sterna 1-4 black, 5 brownish black to dark brown in apical 1/2 with median linear impression brownish black (entirely black).

Male genitalia: Figs. 105, 106; length approximately 0.90 mm; slightly sigmoid dorsoventrally, rather uniform in width but constricted just posterior to basal piece and preapically; apex in dorsal view shallowly and moderately emarginate, almost truncate, in lateral view abruptly tapering to a very blunt, somewhat ventrally directed point, with a short preapical median ventrally directed rounded projection.

Female genitalia: Fig. 139; length approximately 0.35 mm; recep­ tacle oblong, of rather uniform width, no ring collar.

BIOLOGY: Host plants and Immature stages are unknown.

Habits: Adults have been collected in California from March until early September; no dates are associated with the questionable New York record, see below.

DISCUSSION: Distribution: Fig. 160; JP. lepidula is known only from

California, except for 2 specimens in the MSUC that are labeled "N.Y.", which may be a case of midlabeling.

Nomenclature: LeConte (1857) apparently described I\ lepidula from

5 specimens and placed it in Haltica. In 1873, Crotch wrongly placed 89

P^. lepidula in Orchestris (see LeConte, 1878:615) and Horn (1889) placed this species in Phyllotreta.

Relationships: _P. lepidula is most similar to P^. bisinuata and less similar to P. arcuata, P. spatulata, and the striped specimens of £. denticomis with which it shares having a simply elytral stripe and having the male 5th antennal segment dorsoventrally flattened. / Phyllotreta denticomis has a sharp process on the male 6th antennal segment which is absent in the other 4 species, P. arcuata has the male antennal segments 2-6 pale which separates it, P. spatulata has the male antennae dark but with the venter of 1-5 paler and has the venter of the male 5th antennal segment flat which separates it from the other 4 species, and P^. bisinuata is unique in that it has no metathoracic wings. Phyllotreta lepidula and j^. bisinuata are almost iden­ tical externally and the male genitalia are more similar to each other than either is to the other 3 species. The male genitalia in lateral view of P. lepidula are only very slightly sigmoid, whereas those of P^. bisinuata are very abruptly and strongly bent submediallyj the apices are almost identical in both species and P^. lepidula in dorsal view have the median lobe not as strongly constricted before the junction with the basal piece. The female spermathecae of P. lepidula are most similar to those of P_. arcuata but are more robust, P. spatulata and

£. bisinuata are very similar to each other and distinct from the other

3 species, and those of P. denticomis are distinct but are more similar to P. lepidula than to any of the others.

TYPE MATERIAL: The lectotype and 4 paralectotypes (2(J, 2?) are at the MCZC. The lectotype is mounted on a point and the pin bears only a 90

gold circle (^California) and my lectotype label, "LECTOTYPE

Phyllotreta lepidula LeConte By Eric H. Smith '73". The 4 paralecto-

types have the same gold circle but one female has the additional

2 labels (decending order): "Type 4428" and "Phyllotreta lepidula Lee.

S. Jose, S.D." The type label was added by a subsequent curator and

is not a valid designation. The determination-locality label does

/ not appear to be in LeConte*s writing.

SPECIMENS EXAMINED: Total 60: CALIFORNIA: Humboldt Co. (CASC:5),

Los Angeles Co., Los Angeles (CASC:4, USNM:1), (USNM:4), San Diego Co.

Poway (CASC:1), (CASC:1), San Luis Obispo Co., Cholame (CASC:2), San

Margarita (CASC:3), Solano Co., Birds Landing (USNM:5), Yolo Co.,

Rumsey (CASC: 14), (ANSP:2, CNCI:2, DEFW:3, MCZC:8, MSUC:2, USNM.-lJ;

NEW YORK: [MSUC:2]. NO DATA: (ANSP:!).

Phyllotreta liebecki Schaeffer

(Figs. 20, 63-65, 107-108, 140, 161)

Phyllotreta liebecki Schaeffer, 1919. J. New York Ent. Soc., 27:339-

340.

Holotype: Male, USNM type #42423.

Type locality: Enterprise, Florida.

DIAGNOSIS: Elytra each with a median pale stripe greatly expanded

in apical 1/3, reaching margin and suture apically; male antennal

segment 5 moderately enlarged in diameter, with apical 1/2 at most

moderately expanded ventrally, and venter deeply concave.

DESCRIPTION: Holotype (variation, excluding punctation, in paren­ 91 theses): Fig. 20; elongate oval, length 2.18 ram (male 2.02-2.28, female

2.22-2.32), width 1.01 ram (male 0.95-1.15, female 0.98-1.10); head and pronotum black with slight metallic luster, elytra dark brown, each with a median straw-yellow stripe. Head: Black; texture finely gran­ ulate (also finely alutaceous basally); moderately punctate, punctures separated by less than their diameter; Interocular Distance/Maximum

Diameter of eye, 1.43 (male 1.38-1.43, female 1.38-1.47). Antennae:

Figs. 63, 64, 65; segments 4 and 6 equal (subequal) in length, 5 more than 2x length of 4, 6 shorter than 7; segment 5 distinctly longest moderately enlarged in diameter, with apical 1/2 moderately (slightly) expanded ventrally, venter deeply concave entire length; segment

[length/width]: 1(7/3], 2(4/2], 3(3/2], 4(2.5/2.53, 5(7/4), 6(2.5/2),

7(4/3), 8(4/3), 9(4.5/3], 10(4.5/3], 11(6/3), total length 1.22 mm

(Allotype: 1(7/3], 2(4/2+], 3(3.5/2-], 4(3/2], 5[5+/2+], 6(4/2],

7(4.5/3], 8(4+/3], 9(4/3-], 10(4+/3+], 11(6/3+], total length 1.24 mm; simple); antennae various shades of brown, basal 2 (4) segments paler,

3-4 (4 or none) intermediate, 5-11 darkest. Pronotum: Length 0.40 mm

(male 0.38-0.42, female 0.40-0.42), width 0.68 mm (male 0.62-0.72, female 0.70-0.75); black; texture finely granulate with fine wrinkles radiating from punctures (only finely granulate); coarsely punctate, punctures separated by less than to 2x their diameter, mostly by about one diameter. Elytra: Length 1.55 mm (male 1.50-1.68, female 1.68-

2.20), width 1.01 mm (male 0.95-1.15, female 0.98-1.10); dark brown changing to brown at lateral margins, median stripe pattern as in Fig.

20; texture slightly roughened including extremely fine punctures 92

(smooth* slightly roughened only); coarsely punctate, punctures separated by less than to 2x their diameter, mostly by about one diameter. Legs: Normal color sequence. Abdomen: 5th sternum with an apical median lobe and a very shallow, narrow transverse median impression (female simple in outline, no median impression); dark brown (brown, brownish black; 5th often paler, especially apically).

/ Male genitalia: Figs. 107, 108; length approximately 0.68 mm; moderately arched dorsoventrally, broadest subapically; apex in dorsal view very bluntly rounded with a median shallow notch, in lateral view somewhat abruptly tapering to a point.

Female genitalia: Fig. 140; length approximately 0.32 mm; recep­ tacle elongate, slightly constricted submedially, no ring collar.

BIOLOGY: Host plants, adult: *mustard, *radish, Chinese cabbage or pe-tsai (Chittenden, 1923:135). New Collection Record: *tumip,"Baton

Rouge, La., 27-1-19, T.H. Jones"(USNM:2). Host plants, larva: Rorippa obtuse (Nutt.), Arabis virginica (L.), Radicula walteri (Ell.) (Smith,

1921:78); Lepidium virginicum L. (Chittenden, 1927:28).

Immature stages: Undescribed.

Habits: The larvae are leaf miners. Smith (1921) found £. liebecki breeding extensively as a miner in the foliage of the 3 host plants credited above. Chittenden (1923) reported that the larva had been reared from mines in the leaves of Lepidium virginicum. Blatchley

(1914) reports collecting this species [as P. robusta] in large numbers in the herbage along borders of cypress swamps at Sanford, Florida.

Adults of P. liebecki have been collected in northeastern U.S. in mid-

May, in northcentral U.S. and Ontario in May and June, in the midwest 93 during May, and in southeastern U.S. in January and from March until late June. The distribution (see below) falls within that of the known larval host plants and the collection dates within their growing season. Lawrence (1951) states that Lepidium is a weed of economic importance and that Arabis or Rock Cress is an ornamental. Thus, this species may be of beneficial as well as injurious potential.

DISCUSSION: Distribution: Fig. 161; liebecki has been found mainly in the Gulf Coast states. Records not varified by me include

Dunford and Sarasota, Florida, and Liberty, Texas (Gentner, 1926).

Nomenclature: Schaeffer (1929) described this species from specimens sent to him by Charles Liebeck who also mentioned the possible wrong identification of this species as P^. robusta by Blatchley (1914:142).

Blatchley (1920:263) corrected this error.

Relationships: P. liebecki is very similar to P^. robusta with which only it shares the distinctive elytral color pattern of the median pale stripe greatly expanded in apical 1/3, meeting the margin and suture apical ly; but in P^. liebecki the stripe tends to be narrower and less expanded laterally. These species both have the male 5th antennal segment enlarged, in P_. liebecki there is no prolongation and the segment is deeply concave on the venter, whereas in P. robusta there is a bluntly pointed apical prolongation with the segment otherwise simple.

The male genitalia differ, in £. liebecki the apex is bluntly rounded and with a shallow median notch, wheTeas in £. robusta the apex is

sharply tapered to a narrow point. The spermathecae are almost identi­ cal, but those of P. liebecki tend to be much less heavily sclerotized 94

and the gland valve tends to be slightly less rounded.

TYPE MATERIAL: The holotype and allotype are at the USNM and one paratype (9) is at the MCZC. The holotype is mounted on a point and

the pin bears 5 labels (decending order); "Type if" "Enterprise Fla v*14" "BROOKLYN MUSEUM COLLN 1929" "U.S.N.M. Type No. 42423" [orange]

and "Phyllotreta liebecki Schffr." The allotype has the 1stlabel

"Alotype [sic.] the 2nd and 3rd labels the same, the 4th

"liebecki U.S.N.M. Allotype No. 42423" and no other labels. The MCZC paratype is labeled: "Enterprise Fla V-8" "Paratype" and "Liebeck

Collection".

SPECIMENS EXAMINED: Total 60: CONNECTICUT: Litchfield Co., Corn­

wall (CUIC:1); FLORIDA: Seminole Co., Sanford (AMNH:4, CUIC:3, UWEM:4),

Volusia Co., Enterprise (CASC:13, MCZC:18; USNM:Holotype, Allotype),

[CNCI:lD; GEORGIA: [MSUC:4]; IOWA: Story Co., Ames (ISUI:2); KENTUCKY:

[AMNH:2]; LOUISIANA: East Baton Rouge Par., Baton Rouge (CASC:2, USNM:8),

Madison Par., Tallulah (USNM:2), Tangipahoa Par., Ponchatula (USNM:1);

MASSACHUSETTS: Middlesex Co., Tyngsboro (MCZC:1); OHIO: Fairfield Co.,

E.H. Smith Lot No. 302 (EHSC:1), Fairfield or Jefferson or Scioto Co.,

Sugar Grove (0SUC:1), Hocking Co., E.H. Smith Lot No. 346 (EHSC:1),

E.H. Smith Lot No. 300 (EHSC:14),(EHSC:2); PENNSYLVANIA: Dauphin Co.,

Hummelston (ANSP:1); SOUTH DAKOTA: Deuel Co., Gary (SDSU:1); TEXAS:

Colorado Co., Columbus (CNCI:1, USNM:3). CANADA: ONTARIO: Normandale

(CNCI:1). NO DATA: [ANSP:3]. 95

Phyllotreta oblonga Chttn.

(Figs. 21, 66-67, 109-110, 141, 162)

Phyllotreta oblonga Chittenden, 1927. Entomologica Americana, 8(1):

31-32.

Holotype: Female, USNM type #28,840.

Type locality: Edmonton, Alberta, Canada. ,

DIAGNOSIS: Elytra each with a median pale stripe whose inner margin almost parallels suture except strongly incurved toward suture apically but never meeting at suture; male antennal segment 5 simple, about 1.25x length of 6, segments 4 and 6 subequal in length; male genitalia in lateral view with apex abruptly and strongly directed ventrad.

DESCRIPTION: Holotype (variation, excluding punctation, in paren­ theses): Fig. 21; oblong, length 2.72 mm (male 2.62-2.68, female 2.58-

2.82), width 1.35 mm (male 1.22-1.25, female 1.32-1.40); head and pronotum black with slight metallic luster (no luster), elytra black

(with slight metallic luster), each with a median straw-yellow stripe.

Head: Black; texture finely granulate (to slightly roughened anteriorly in addition also finely alutaceous basally); moderately punctate, punctures separated by less than to equal their diameter; Interocular

Distance/Maximum Diameter of eye, 1.44 (male 1.33-1.63, female 1.33-

1.53). Antennae: Figs. 66, 67; segments 4 and 6 equal (subequal) in

length, segments 5 and 7 equal (subequal) in length, longer than 4 or

6 (male segment 5 about 1.25x length of 6); segments simple; segment

[length/width!]: l[9/3+], 2t4.5/2.5], 3[4/2], 4C4/2], 5C5/2], 6C4/2.5], 7(5/33, 8(5/33, 9(5.5/3+3, 10(5/33, 11(7/3+3, total length, 1.45 ram

(male: 1(8.5/33, 2(4.5/2+3, 3(5-/2+3, 4(4+/2+3, 5(5/2.53, 6(4.5/2.53,

7(5.5/33, 8(5/33, 9(5/33, 10(5/3+3, 11(7/3.53, total length 1.48 mm;

simple); antennae various shades of brown, basal 3 (2) segments paler,

segment 4 (3 or none) intermediate, segments 5-11 (4-11) darkest

(male: same sequence as female or segment 1 paler, 2-3 and 8-11 dark, / 4-7 darkest). Pronotum: Length 0.52 mm (male 0.50-0.52, female 0.52-0.55

0.55), width 0.82 mm (male 0.85, female 0.82-0.88), black: texture

finely granulate (slightly roughened or smooth to slightly roughened); moderately punctate, punctures separated by less than to equal their

diameter, mostly by about one diameter. Elytra: Length 2.02 mm (male

1.85-2.00, female 1.85-2.08), width 1.35 mm (male 1.22-1.25, female

1.32-1.40); black (usually brownish black), median stripe pattern as

in Fig. 21; texture finely roughened (rarely smooth); coarsely

punctate, punctures separated by less than to equal their diameter,

mostly by about one diameter. Legs: Normal color sequence. Abdomen:

5th sternum simple in outline (with a shallow median oval impression

in apical 1/3) (male: 5th sternum with an apical median deeply concave

lobe, concavity tapering posteriorly and extending to apical 1/3 or > extending to within 1/3 of sternal base, with a median linear

impression extending from sternal base to base of concavity or ending

subapically, sometimes interrupted); brownish black (black) with apex

of 5th paler (apical margin to apical 1/3 paler; median lobe paler but

median linear impression dark; or entirely dark). 97

Male genitalia: Figs. 109, 110; length approximately 1.05 mm; slightly arched dorsoventrally, of rather uniform width except slightly narrower preapically in dorsal view, in lateral view tapering on dorsum from base of median lobe to apex with venter essentially

straight; with dorsal washboard; apex in dorsal view rounded with acute

tip, in lateral view moderately narrow, rather abruptly and strongly / directed ventrad, with a subapical notch on venter.

Female genitalia: Fig. 141; length approximately 0.40 mm; recep­

tacle elongate, tapering in median 1/3 to a narrow, almost parallel-

sided apical 1/3; ring collar slightly developed.

BIOLOGY: Host plants, adult: *Lepidium virginicum (Chittenden,

1927:31).

Immature stages: Unknown.

Habits: Adults have been collected in the Northwest Territory and

Alberta during June.

DISCUSSION: Distribution: Fig. 162; £. oblonga has been collected

in the Northwest Territory and Alberta.

Nomenclature: Chittenden (1927) described this species from a

series of females (about half the series including the holotype were

erroneously sexed) and the male is described here for the first time,

from 2 specimens.

Relationships: P. oblonga is a rather distinctive species; having

the antennae simple with segment 5 longer than 6, and 4 and 6 subequal,

along with the distinctive elytral stripe pattern of having the inner

margin of the stripe essentially parallelling the suture except strongly

incurved apically easily separate this species. 98

TYPE MATERIAL: The holotype and 6 paratypes (?) are at the USNM.

The holotype is mounted on a point and the pin bears 4 labels

(decending order): "Edmonton Alta 1»VI*18 F.S. Carr" "Type No.

28840 U.S.N.M." and a determination label. The female spermatheca is mounted on a card immediately under the point.

SPECIMENS EXAMINED: Total 16; OREGON: Benton Co., Mary S Pk near / Corvallis (CASC:1). CANADA: ALBERTA: Edmonton (CASC:4, Holotype,

USNM:6); NORTHWEST TERRITORY: Aklavik (CASC:2, CNCI:2).

Phyllotreta oregonensis (Crotch)

(Figs. 22-23, 68-69, 111-112, 142, 163)

Orchestris oregonensis Crotch, 1873. Proc. Acad. Nat. Sci. Philadel­

phia, 25:66..

Lectotype (here designated): Male, at ANSP.

Type locality: Oregon.

Phyllotreta oregonensis, Horn, 1889. Trans. American Ent. Soc., 16:

296-297.

DIAGNOSIS: Elytra each with a median pale stripe with dilations which never meet at suture, medially stripe usually wider than distance

from stripe to suture; male antennal segment 5 dark, expanded bilater­

ally and dorsoventrally flattened, subequal to 4 in width, with venter

flat; apex of male genitalia in dorsal view entire, in lateral view

almost straight, not directed ventrad.

DESCRIPTION: Lectotype (variation, excluding punctation, in paren­

theses): Fig. 22; oblong oval, length 2.80 mm (male 2.28-3.10, female 2.28-3.35), width 1.38 mm (male 1.22-1.55, female 1.18-1.50); head and pronotum black with slight metallic luster, elytra dark brown

(with slight metallic luster), each with a median straw-yellow stripe.

Head: Black; texture finely granulate (finely granulate basally to slightly roughened anteriorly); finely to moderately punctate, punctures separated by less than 1/2 to equal their diameter, mostly by less than / 1/2 a diameter; Interocular Distance/Maximum Diameter of eye, 1.56

(male 1.30-1.56, female 1.44-1.56). Antennae: Figs. 68, 69; segment 4 longer than 3, 6, or 7, segment 5 about l.Sx length of 4, 6 distinctly shortest; segments 4 and 5 expanded bilaterally, dorsoventrally flattened; segment 5 distinctly longest, subequal to 4 in width, with venter flat; segment (length/width): 1(10/4-], 2(4.5/2.5], 3(4/3],

4(5/4), 5(7.5/4.5), 6(3/3), 7(4.5/3), 8(5/3+), 9(5/3+], 10(4.5/3+),

11(8/3), total length 1.52 mm (female paralectotype: 1(10/3.5),

2(4.5/2.5), 3(4.5/3-], 4(5.5/2.5], 5(7/3), 6(4.5/3), 7(6/3.5), 8(6/3+],

9(5.5/3+), 10(5+/3+], ll(7+/3.5], total length 1.64 mm; simple); antennae various shades of brown, basal 2 (3) segments pale with dorsum of 1 dark, 3 (3-4 or none) intermediate, 4-11 (5-11) darkest. Pronotum:

Length 0.50 mm (male 0.48-0.55, female 0.45-0.58), width 0.90 mm

(male 0.72-1.02, female 0.82-1.02); black; texture slightly roughened

(in addition with wrinkles radiating from some punctures; finely granulate); moderately to coarsely punctate, punctures separated by

less than 1/2 to equal their diameter, mostly coarsely punctate with punctures separated by less than 1/2 a diameter. Elytra: Length 2.02 mm male 2.00-2.25, female 1.55-2.42), width 1.38 mm (male 1.22-1.55, female

1.18-1.50); dark brown (usually brownish black, sometimes black) median 100

stripe pattern as in Fig. 22 (sometimes as in Fig. 23); texture smooth

(to slightly roughened or entirely slightly roughened); coarsely punctate with moderate punctures interspersed, punctures separated by

less than 1/2 to equal their diameter, mostly by less than 1/2 a

diameter. Legs: Normal color sequence. Abdomen: 5th sternum with an

apical median moderately concave lobe, concavity tapering posteriorly t and extending to midpoint (extending to within 1/3 o£ sternal base),

a median linear impression extending subbasally to subapically

(extending from sternal base to base of concavity; female simple in

outline, sometimes with a median linear impression in apical 1/3 to

1/2); brownish black (usually black) with median lobe paler except

median linear impression black.

Male genitalia: Figs. Ill, 112; length approximately 1.00 mm;

slightly arched dorsoventrally, of rather uniform width except median

lobe slightly narrowed submedially; apex in dorsal view narrowly

truncate, in lateral view tapering to a moderately wide, rounded tip,

venter almost straight.

Female genitalia: Fig. 142; length approximately 0.40 mm; recep­

tacle elongate, slightly narrowed medially, ring collar slightly

developed.

BIOLOGY: Host plants, adult: leaves, seed-heads, and flowers of

sugar beet (Chittenden, 1923:135). New Collection Records: on potatoes,

from "Tulelake Cal. July 20, 1946" (JSCC:2) and *Brassica, from

"Tulelake, Siskiyou Co., Cal., Joe Schuh, Coll." (JSCC:2). 101

The other host records cited by Chittenden (1923:135) belong to other species (see I\ attenusta and P. constricta) except the following two which have not been verified by me because of lack of specimens: Fowler and Paonia, Colo., by Titus, on foliage and flowers of peppergrass,

Lepidium (sopulorum) spathalatum, and Rocky Ford, Colo., by Marsh, on radish.and turnip. The peppergrass citation also represents the larval

/ leaf mining record.

Immature stages: Undescribed or unknown (see above).

Habits: The larvae may be leaf miners (see above). Adults have been collected in Colorado in mid-June, in Oregon in mid-May, and in

California in mid-July; no date is associated with the specimen from

Utah.

DISCUSSION: Distribution: Fig. 163; P. oregonensis has been collected in the Rocky Mountains and westward, specifically in Colorado,

Utah, Oregon, and California.

Nomenclature: Crotch (1873) apparently described £. oregonensis from 7 specimens and placed it in the genus Orchestris (Crotch, not

Kirby). LeConte (1878:615) pointed out Crotch*s error in using this generic name and Horn (1889) transferred this species to Phyllotreta.

Relationships: £. oregonensis is most similar to P_. constricta and is less similar to P_. emarginata, P^. zimmermanni, £. utanula, and 1?. utana with which it shares elytral color pattern and having the male 5th antennal segment appearing dorsoventrally flattened. However, the elytral stripe of P^. oregonensis and constricta is usually much wider medially than that of the other 4 species. The male 5th antennal segment of P. zimmermanni has a basal concavity ventrally, whereas that of the 102 other 5 species is evenly flattened ventrally; also antennal segments

2-5 are pale in P_. utana and P. utanula, whereas in the other 4 species

antennal segments 2-3 are pale and/or dark and segments 5-11 are dark.

The male genitalia of these 6 species separate into 3 distinct groups

(based on dorsal view of apex) as follows: £. utana and P^. utanula

which have the appex very broad and its margin moderately and broadly

/ emarginate; J?. zimmermanni and emarginata with the apex not broadened and with its margin moderately but narrowly emarginate medially; and £. oregonensis and P_. constricta with the apex not

broadened and with its margin entire. However, P^. oregonensis has

its apex in dorsal view narrowly truncate and in lateral view almost

straight, whereas P^. constricta has its apex evenly rounded and

directed ventrally. The female spermathecae of these 5 species

(female of P^. utanula is unknown) are fairly distinct with those of P^.

oregonensis and P_. constricta most similar. Phyllotreta utana have the

receptacle oblong, not narrowed, and a slight posterior collar, whereas

P. emarginata have the receptacle elongate oblong with a slight bend

and a moderately developed ring collar. Those of P^. oregonensis, P^.

constricta, and zimmermanni have the receptacle elongate, but P.

oregonensis and £. constricta have the receptacle slightly narrowed

submedially and a slightly developed collar or no collar, whereas ]?.

zimmermanni have the receptacle slightly bent and a prominent collar.

The spermathecae of P^. oregonensis tend to have the receptacle less

strongly swollen in the basal 1/2 and have a slight collar, whereas those

of P. constricta tend to be more strongly swollen and have no collar. 103

Phyllotreta oregonensis might be confused with P. attenuata and

£. dolichophalla on the basis of medially wide elytral stripe, but _P.

oregonensis has the male 5th antennal segment appearing dorsoventrally

flattened, whereas in iP . attenuata and P^. dolichophalla it is simple.

TYPE MATERIAL: The lectotype and 5 paralectotypes (1

the ANSP and 1 possible paralectotype (9) is at the MCZC. The lecto- / type is mounted on a point and the pin bears 4 labels (decending order):

"Or." "5" "Horn Coll H 7025" and "LECTOTYPE Phyllotreta oregonensis

(Crotch) By Eric H. Smith *73". Four of the ANSP paralectotypes are

labeled: "Or." (l£ has the period partially cut off, 1? has the period

completely cut off) and "Horn Coll H 7025" and the fifth (?) is

labeled as follows: "Or." "TYPE No. 2711" and "P. oregonensis Crotch".

The specimen at the MCZC is labeled: "Or." "J.L. LeConte Coll." Type

5015" and "oregonensis Cr."

The series at the ANSP is accepted as Crotch's type series based

on his statement (1873:19) that "All the species described are from the

cabinets of Drs. Leconte (sic.] and Horn," the type locality is given

as "Oregon(Horn)," and the sexually dimorphic male antennae is men­

tioned in the description. Crotch did not designate a type as was the

custom of the day. The type label on the one specimen was added by a

subsequent curator and is not a valid designation; the Horn Collection

label was either never added to this specimen or was accidently not

returned to the specimen, possibly at this time. Dr. Rentz, Curator

of the Entomology Collection, could add nothing to my conclusions. A

male was selected because of the diagnostic male genitalia and antennae.

The specimen labeled as type at the MCZC may have been examined by 104

Crotch because he studied LeConte's Collection and it was customary for Horn and LeConte to exchange material, but it is possible that it was added after Crotch’s study (1873) because LeConte was still active and Crotch specifically indicated that the specimens were from Horn's

Collection. Therefore, it was not labeled a paralectotype. The type label was added by a subsequent curator and is not a valid designation.

SPECIMENS EXAMINED: Total 26: CALIFORNIA: Siskiyou Co., Tule

Lake (JSCC:3); COLORADO: Delta Co., Paonia (USHN:12), Montrose C o .,

Montrose (USNM:2); OREGON: Crook Co ., Prineville (JSCC:1), [Lectotype,

ANSP:S; MCZC:l3 ; UTAH: Cache C o ., Logan (USNM:1).

Phyllotreta ramosa (Crotch)

(Figs. 24, 70-71, 113-114, 143, 164)

Orchestris ramosa Crotch, 1874. Trans. American Ent. Soc., 5:80.

Holotype: Female, MCZC type #5022.

Type locality: Lake Port, North California.

Phyllotreta ramosa, Horn, 1889. Trans. American Ent. Soc., 16:299.

DIAGNOSIS: Elytra each with a median pale stripe with abrupt lateral dilations, apically curved towards but never meeting suture, median 1/2 uniformly very narrow; antennal segment 5 simple, almost same

length as 4 or 6, 4 and 6 subequal.

DESCRIPTION: Holotype (variation, excluding punctation, in paren­

theses): Fig. 24; oblong oval, length 2.20 mm (male 1.92-2.35, female

1.98-2.25), width 1.05 mm (male 0.88-1.12, female 0.92-1.08); head and pronotum black with a slight metallic luster, elytra dark brown with a slight metallic luster (no luster), each elytron with a median straw-yellow stripe. Head: Black (brownish black); texture finely granulate (in addition finely alutaceous basally or slightly roughened anteriorly); moderately punctate, punctures separated by less than

1/2 to equal their diameter, mostly by less than 1/2 one diameter; r Interocular Distance/Maximum Diameter of eye, 1.3. (male 1.25-.133, female 1.25-1.38). Antennae Cleft missing): Figs. 70, 71; segments 4 and 6 subequal (equal) in length; segment 5 simple, slightly longer than (subequal to) 4, 6 or 7, 7 slightly longer than (subequal to) 6; segment (length/width) : l(7/2+3, 2(4/23, 3(3.5/2.5+3, 4(3.5/3.5+3,

5(4/23, 6C3.5+/2), 7(4/2.53, 8(4/2.53, 9(4.5/2.53, 10(3.5/2.53,

11(5-/2.53, total length 1.16 mm (male: 1(7/2.53, 2(3+/2), 3(3/1.53,

4(3/23, 5(3.5/2+3, 6(3/23, 7(4/2.53, 8(4/2.53, 9(4/2.53, 10(3.5/2.53,

11(5/33, total length 1.08 mm); antennae various shades of brown, basal

2 (3) segments paler with segment 1 darker medially (not darker to basal 1/2 darker), 3 (4 or none) intermediate, 4-11 (5-11) darkest.

Pronotum: Length 0.40 mm (male 0.35-0.42, female 0.35-0.42), width

0,70 mm (male 0.60-0.78, female 0.65-0.72); black (rarely brownish black); texture finely granulate; very coarsely punctate, coarsely punctate medially in apical 1/2, punctures separated by less than 1/2 their diameter. Elytra: Length 1.65 mm (male 1.40-1.72, female 1.45-

1.75), width 1.05 mm (male 0.88-1.12, female 0.92-1.07); dark brown but basally brownish black between stripes (entirely dark brown; with suture and margins darker), median stripe pattern as in Fig. 24; texture smooth; very coarsely punctate in basal 1/2, coarsely punctate 106 in apical 1/2, punctures separated by less than 1/2 their diameter.

Legs: Normal color sequence. Abdomen: 5th sternum with outline simple, with a shallow median impression in apical 1/3 (female: in addition apical 1/2 shallowly concave; impression in apical 2/3 but very narrow or no median impression but this area flat; male: 5th sternum with an apical- median, moderately to deeply concave lobe and a shallow median / linear impression; median concavity sometimes extended and tapering toward base but ending before reaching basal 1/3, median linear impression can extend complete length of sternum, but usually ends 1/3 of way into median concavity, or rarely is expanded in basal 1/3); dark brown (to black), (male usually with apical 1/3 of 5th paler).

Male genitalia: Figs. 113, 114; length approximately 0.85 mm; very slightly arched dorsoventrally, width rather uniform but slightly tapering toward base in basal 1/2; with a dorsal washboard; apex in dorsal view moderately acute, tip slightly rounded, in lateral view extreme apex very slender, abruptly bent ventrally at about 45°, tip pointed.

Female genitalia: Fig. 143; length approximately 0.31 mm; recep­ tacle elongate pear-shaped, tapering towards pump, with slightly developed ring collar at junction with pump; pump uniformly sclerotized.

BIOLOGY: Host plants, adult: *cabbage, *cauliflower, *brussels sprouts, *radish, *rape, *mustard, *stocks, *tumips, *wallflower,

♦water cress (Essig, 1926:481). New Collection Records: *Brassica nigra

(L.), "Guadalupe, Cal., July 29, 1908, I.J. Condit Collector" (USNM:4);

Asclepias sp., "mouth Williams R., Klamath County, Ore., June 17, 1958, 107

Joe Schuh Coll." (JSCC:1); and Algoma Sp., "Klamath Falls, Ore.,

July 19, 1946, Joe Schuh, Coll." (JSCC:1).

Immature stages: Unknown.

Habits: Essig (1926:280-81) remarks that this species is often very abundant and destructive to the above recorded host plants.

Adults have been collected in the Pacific Northwest from mid-March until mid-June and in late January (perhaps overwintering), in Cali­ fornia from March until early August and in early December; no dates are associated with the 2 Nevada specimens; the questionable New York record was collected in mid-June (see below).

DISCUSSION: Distribution: Fig. 164; P. ramosa has been collected in the Pacific Coast states, 2 specimens are recorded from Nevada, and

1 specimen is recorded from New York which is either mislabeled or is an important new record for this western economic species.

Nomenclature: There are no synonyms and Crotch (1874) described

P. ramosa from a single specimen.

Relationships: ramosa is most similar to P^ ramosoides and less

similar to bipustulata, IP. s trio lata and P^. decipiens. Of the maculate species, these 5 have the male genitalia of the same general

shape, have the dorsal washboard, and an acute apex which in lateral

view is angled ventrally. Only P^. ramosa and P. ramosoides share the

distinctive elytral color pattern having the stripe with abrupt lateral

dilations, apically curved towards but never reaching the suture, and

with the median 1/2 uniformly very narrow; but occasionally, these two might be confused with a P_. striolata having the median part of the

stripe narrow, but the antennae will easily separate the three species. 108

Phyllotreta ramosa and P. bipustulata share the additional character of having the 5th antennal segment in both sexes simple and not elongate, whereas in P^. ramosoides and £. decipiens it is simple but elongate in both sexes and in P^. striolata it is simple and elongate in the female and enlarged and elongate in the male. The female spermathecae of these five species are each distinctive, but those of P. ramosa and / P. ramosoides are somewhat similar. However, the spermathecae of ramosa are less robust with a longer pump and have the spermatheceal duct arising closer to the receptacle dorsum in lateral view than in

P. ramosoides.

TYPE MATERIAL: The holotype is at the MCZ. It is mounted on a point and the pin bears 4 labels (decending order): "Cala.r" "J.L.

Leconte Coll.” "Type 5022" and "ramosa sp.n."

SPECIMENS EXAMINED: Total 106: CALIFORNIA: Alameda Co., Berkeley

(CASC:5), Niles (CADE:1), Oakland (CASC:2), (CASC:S), Contra Costa Co.,

Antioch (CASC:1), Brentwood (CNCI:1), Lafayette (MCZC:1), Lake C o .,

Lucerne (CASC:1), Los Angeles Co., Los Angeles (CASC:2), Pasadena

(CASCil), Orange Co ., Los Alomitos (USNM:1), Sacramento C o., Sacramento

(CADE:1, NMDC:1, UCRC:3), (JSCC:1), San Bernardino C o ., Chino (USNM:3),

San Francisco Co. (MCZC:1), San Mateo C o ., Crystal Lakes (CASC:5), E.

Palo Alto (CASC:1), (CADE:2), Santa Barbara Co., Guadalupe (USNM:4),

Santa Clara Co., Milpitas (USNM:1), San Jose (CASC:1), Santa Clara

(CASC:1), Searsvilie L. Stanford U. (CASC:1), Stanford U. (LACM:1),

Siskiyou C o ., Grass Lake (CASC:4, CADE:4, NMDC:2), Tulelake (JSCC:4),

Sonoma Co., Rio Nido (CASC:1), (LACM:3), Yolo Co., Clarksburg (CASC:2), 109

Davis (CASC: 2 , USNM:1), [AMNH:4, ANSP:2, CASC:4, MCZC:1, WSUC:2H;

NEVADA: Elko Co., Elko (ANSP:1, USNM:1), (CASC:ID, Washoe Co., Reno

Hot Spring (CASC:1); NEW YORK: Monroe Co., Rochester (USNM:1); OREGON:

Benton Co., Corvallis (MCZC:1), Jackson C o., Phoenix (JSCC:1), Talent

(MCZC:3), (ICCM:1), Klamath Co., Bly Mt. (JSCC:1), Klamath Falls

(JSCC:1, 0SU0:3), Mouth Williamson R. (JSCC:1), Lower Klamath Lk.

(JSCC:1), Worden (JSCC:1), Union Co ., 4 mi. S. LaGrande (CADE:1),

Union (JSCC:1), Yamhill Co ., McMinnville (UCRC:1).

Phyllotreta ramosoides NEW SPECIES

(Figs. 25, 72-73, 115-116, 144, 165)

Holotype: Male, deposited at CNCI.

Type locality: Pender Harbor, British Columbia, Canada.

DIAGNOSIS: Elytra each with a median pale stripe with abrupt lateral dilations, apically curved toward but never meeting suture, median 1/2 uniformly very narrow; antennal segment 5 simple, longer than 6, segments 4 and 6 subequal; male genitalia in lateral view moderately and evenly arched dorsoventrally.

DESCRIPTION: Holotype (variation, excluding punctation, in paren­ theses): Fig. 25; oblong, length 2.45 mm (male 2.38-2.65, female 2.48-

2.78), width 1.20 mm (male 1.18-1.25, female 1.22-1.30); head brownish black (with slight metallic luster), pronotum black with slight metallic luster (no luster), elytra dark brown (with slight metallic

1luster), each with a median straw-yellow stripe. Head: Brownish black 110

(usually black); texture finely granulate basally to smooth anteriorly (basally, finely alutaceous or alutaceous and granulate; anteriorly, slightly roughened or smooth to roughened; entirely slightly roughened); moderately punctate with coarse and very fine punctures interspersed, punctures separated by less than 1/2 to equal their diameter, mostly by less than 1/2 a diameter; Interocular Distance/

/ Maximum Diameter of eye, 1.33 (male 1.33-1.39, female 1.26-1.39).

Antennae: Figs. 72, 73; segments 4 and 6 subequal, shorter than segment

7, segment 5 about 1.5x length of 6; segment 5 simple; segment

(length/width]: 1(9/3+], 2(4.5/2.5], 3(4.5/2+], 4(4+/2+3, 5[6-/3],

6(4/2.53, 7(5/2.53, 8(5/33, 9(5/3], 10(5-/3], 11(6.5/3], total length

1.46 mm (Allotype: 1(8/3], 2(5/2+], 3(4/2], 4(4+/2], 5(5+/2+], 6(4/2+],

7(4.5/3-], 8(4.5/3-3, 9(5-/3-3, 10(4.5/3-], 11(7/3+], total length

1.39 mm; simple); antennae various shades of brown, basal 3 segments paler with dorsum of 1 darker (females with 1 entirely paler), segment

4 intermediate, 5-11 darkest. Pronotum: Length 0.48 mm (male 0.45-

0.50, female 0.45-0.52), width 0.80 (male 0.80-0.88, female 0.80-0.90); black; texture smooth (slightly roughened, finely granulate, smooth to roughened, alutaceous to roughened); coarsely punctate, punctures

separated by less than 1/2 to equal their diameter, mostly by less than

1/2 a diameter. Elytra: Length 1.80 mm (male 1.78-1.92, female 1.80-

2.08), width 1.20 mm (male 1.18-1.25, female 1.22-1.30); dark brown

(black, sometimes brownish black between stripes basally), median

stripe pattern as in Fig. 25; texture smooth (slightly roughened or

smooth to slightly roughened); coarsely punctate, punctures separated Ill by less than 1/2 their diameter. Legs: Normal color sequence.

Abdomen: 5th sternum with an apical median moderately (deeply) con­ cave lobe, concavity oval and extending to apical 1/3 (oblong or tapering extending to within 1/3 of sternal base), with a median linear impression extending from sternal base to apex (ending at base of median lobe; female: simple in outline; with no impression, with a

/ median short narrow shallow transverse subapical impression, or with a median shallow oval impression in apical 1/3); brownish black

(black or dark brown) with median lobe paler (entirely dark).

Male genitalia: Figs. 115, 116; length approximately 1.10 mm; moderately and evenly arched dorsoventrally, of rather uniform width, with a dorsal washboard; apex in dorsal view roundly acute with tip rounded, in lateral view narrow, gradually curved ventrad with tip pointed.

Female genitalia: Fig. 144; length approximately 0.35 mm; recep­ tacle elongate pear-shaped, tapering toward pump, ring collar slightly developed.

BIOLOGY: Host plants and Immature stages are unknown.

Habits: Adults occur in the Pacific Northwest from late May until

late June.

DISCUSSION: Distribution: Fig. 165; jP. ramosoides has been

collected in the Pacific Northwest, specifically the Northwest Terri­

tory, southern British Columbia, northern Idaho, and Oregon. New name:

The specific epithet ramosoides comes from the Latin ramo- meaning

branch and the Latin suffic -oides meaning like; this refers to the

elytral stripe pattern which this species shares with P_. ramosa and the 112 name implies this similarity.

Relationships: P. ramosoides is most similar to _P. ramosa and less similar to P. bipustulata, P. striolata, and P. decipiens; for a discussion of these relationships, see this section under P. ramosa.

TYPE MATERIAL: The holotype, allotype, and 8 paratypes (3#, 3$) are at the CNCI; 21 paratypes (lid, 10?) are at CASC; 4 paratypes / (2d, 29) are in my collection, EHSC; 4 paratypes (3d, 1?) are at

JSCC; 1 paratype (d) at the NMDC; and 2 paratypes (Id, 1?) are at the

USNM. The type is mounted on a point and the pin bears 2 labels

(decending order): "Pender Harbor B.C. V-27-28 G.R. Hopping" and

"HOLOTYPE Phyllotreta ramosoides By Eric H. Smith ’73"; I also added a polyethylene genitalia vial. The allotype has the same locality- date label as the holotype and "ALLOTYPE Phyllotreta ramosoides By

Eric IJ. Smith '73".

The paratypes all have a paratype label added. Those at CNCI are as follows: 3d and 2?, same locality-date label as holotype, 1?

"Aklavik, N.W.T. 25-VI-1956 E.F. Cashman"; the 21 paratypes at the CASC all have the same locality-date label as the holotype and the additional label "R. HOPPING COLLECTION"; the 4 paratypes in my collection are labeled identically to those of CASC; those in the JSCC are as follows: lcf, "Scappoose, Ore. VI-8-1937 R.E. Reider, col.", Id1, "Scappoose, Ore.

VI-12-1937 R.E. Reider, col.", lo, "Scoppoose, Ore. VI-25-1937 R.E.

Reider, col.", and 1$, "Sprague, Riv., Ore 5 mi. E. Bly June 16, 1957

Joe Schuh, Coll."; the male paratype in NMDC is labeled "Sandpoint,

Ida VI-11 1951 N.M. Downie" "may be oblonga & Chitt" "also as 16073 113 I as Det. L.G.G. but too large"; the 2 paratypes at the USNM are labeled identically to those o£ CASC.

SPECIMENS EXAMINED: Total 41: IDAHO: Bonner Co., Sandpoint

(NMDC:1); OREGON: Columbia Co., Scappoose (JSCC:3), Klamath Co .,

Sprague Riv., 5 mi. E. Bly (JSCC:1). CANADA: BRITISH COLUMBIA: Pender

Harbor (CASC:21, Holotype, Allotype, CNCI:6, EHSC:4, USNM:2);

NORTHWEST TERRITORY: Aklavik (CNCI:1).

Phyllotreta robusta LeConte

(Figs. 26-27, 74-75, 117-118, 145, 166)

Phyllotreta robusta LeConte, 1878. Proc. American Phil. Soc., 17:614-

615.

Holotype: Female, MCZ type #4429.

Type locality: Detroit, Michigan.

DIAGNOSIS: Elytra each with a median pale stripe greatly expanded

in apical 1/3, reaching margin and suture apically, and basally,

usually laterally expanded; male antennal segment 5 moderately ex­

panded, with a ventrally directed bluntly pointed apical prolongation.

DESCRIPTION: Holotype (variation, excluding punctation, in paren­

theses): Fig. 26; elongate oval, length 2.38 mm (male 1.98-2.30,

female 2.08-2.38), width 1.25 mm (male 1.00-1.12, female 1.10-1.18);

head and pronotum black with slight metallic luster (no luster),

elytra brownish black (rarely with faint metallic luster), each with

a median straw-yellow stripe. Head: Black; texture finely granulate (sometimes in addition, finely alutaceous basally); moderately punctate, punctures separated by less than 1/2 to equal their diameter, mostly by less than 1/2 diameter, many punctures confluent; Inter­ ocular Distance/Maximum Diameter of eye, 1.22 (male 1.22-1.38, female

1.22-1.38). Antennae: Figs. 74, 75; segments 4 and 6 subequal in length, 6 shorter than 7, 5 distinctly longer than 4, 6 or 7 and

/ simple (male segment 5 moderately expanded, with a long ventrally directed bluntly pointed apical prolongation; segment 6 shorter than

4 or 7);.segment (length/width): 1(8/3-), 2(4.5/2+), 3(4/2), 4(3.5/2],

5(5.5/2+3, 6(4-/2], 7E5/3], 8(5+/3+], 9t5/3+], 10(4/33, ll(6/3], total length 1.36 mm (male: lC8/3], 2(4/2+], 3(3.5/2], 4(3.5-/2.5],

5(6/5.5], 6(2.5/2+], 7(4/3-], 8(4.5/3], 9(4.5/3], 10(4.5/3], 11(5/3], total length 1.25 mm); antennae various shades of brown, basal 3 (4) segments paler, 5 (4-7) intermediate, 6-11 darkest (5-11), (dorsum of

1-5 sometimes darker in any combination, usually 1 and 4-5). Pronotum:

Length 0.45 mm (male 0.38-0.42, female 0.40-0.45), width 0.75 mm

(male 0.65-0.75, female 0.72-0.78); black; texture finely granulate; coarsely punctate, punctures separated by less than 1/2 to equal their diameter, mostly by less than 1/2 diameter. Elytra: Length 1.75 mm

(male 1.48-1.75, female 1.52-1.75), width 1.25 nun (male 1.00-1.12, female 1.10-1.18); brownish black between stripes to dark brown (brown) laterally, median stripe pattern as in Fig. 26 (sometimes as in Fig.

27); texture slightly roughened (sometimes smooth); coarsely punctate in basal 1/2 with punctures separated by less than 1/2 to less than their diameter, mostly by less than one diameter, moderately punctate 115 in apical 1/2 with punctures separated by less than to equal their diameter, mostly by about one diameter. Legs: Normal color sequence.

Abdomen: 5th sternum simple in outline (female rarely with median shallow linear impression in apical 1/3; male with an apical median deeply concave lobe, a deep, trough-like wide median impression extending subbasally to base of median lobe concavity); brownish black | r (black, male median lobe usually paler).

Male genitalia: Fig. 117, 118; length approximately 0.80 mm; slightly.arched dorsoventrally, rather uniform in width but slightly wider just basad of orifice in dorsal view and strongly tapered dorsally from midpoint of median lobe to apex in lateral view; apex in dorsal view strongly, somewhat abruptly acute, in lateral view tapering to a narrow rounded tip but shallowly interrupted subapically on dorsum.

Female genitalia: Fig. 14S; length approximately 0.325 mm; recep­ tacle elongate, slightly constricted submedially, ring collar slightly developed.

BIOLOGY: Host plants, adults: *radish and *turnip (Chittenden,

1927:19). New Collection Records: *Brassica arvensis (=Sinapis arvensis

L.), "Waupaca, Wis., 6-21-20 L.G. Gentner Collector" (CNCI:1, UWEM:1);

*Lepidium sp., "'Madison, Wis. 6-3-19, L.G. Gentner Collector" (UWEM:1);

Poa pratensis L , "Whitewater L. 4 mi. N. Whitewater, Man. 22*VI*1958,

R.D. Bird" (CNCI!:1); Elaegnus commutata, "5 mi. S.W. Shile, Man., Flood- plain Community nr. Tamarack Bog at margin of Beaver Pond, 28-V*1958,

R.B. Madge" (CNCI:1). Immature stages: Unknown.

Habits: Adults have been collected in northeastern U.S. and adjacent Canada from late May until late June and in mid-January

(probably overwintering), in the midwest and adjacent Canada from early

May until late July, in northcentral U.S. and adjacent Canada from

April -until late September, and in western U.S. and adjacent Canada / from early May until early October.

DISCUSSION: Distribution: Fig. 166; P. robusta has been collected east of the Rocky Mountains in the northern half of the U.S. and adjacent Canada, and in Colorado, Utah, and Nevada.

Nomenclature: There are no synonyms and LeConte (1878) described

P. robusta from a single specimen.

Relationships: P_. robusta is most similar to P_. liebecki, see this section under P^. liebecki.

TYPE MATERIAL: The holotype is at the MCZC. It is mounted on a point and the pin bears 4 labels (decending order): "Detroit Mic 26/5"

"34" "Type 4429" (red) and "0. robusta Lee."

SPECIMENS EXAMINED: Total 96: COLORADO: Costilla Co., Garland

(ANSP: 1, USNM.-l); CONNECTICUT: Litchfield C o ., Cornwall (CUIC:1);

INDIANA: Lake Co., Long Lake (FMNH:1); MAINE: Oxford Co., Paris (MCZC:1)

MASSACHUSETTS: Middlesex Co., Farmingham (CASC:3), S h e rbom (MCZC:1),

CMCZC:1]; MICHIGAN: Ingham Co., Ag College =Michigan St. Univ. (MSUC:2),

Marquette Co., Marquette (USNM:1); MINNESOTA: Anoka Co., Bunker Pr.

Dunes (DEFW: 1), Mille Lacs Co., Mille Lacs (DEFW.-2), Nicollet Co., St.

Peter, Fish Hatch, pond (DEFW:1), Ramsey C o ., near Gray Cloud Is. 117

(CNCI:1), (DEFW:1), Red Lake Co ., Plummer (DEFW:1), Sibley Co., Riv. near Blakeley (DEFW:1); NEVADA: Elko Co ., Elko (MSUC:1, USNM:3);

NEW HAMPSHIRE: Hillsborough Co., Antrim (MCZC:1); NEW YORK: CMCZC:2];

SOUTH DAKOTA: Brookings Co., Brookings (SDSU:2), Lawrence C o .,

Spearfish Creek 3 mi. N. Spearfish (SDSU:2); UTAH: Salt Lake C o .,

Salt Lake (USNM:1), Utah Co., Provo CCASC:5, MCZC:10, USNM:4); ' / WISCONSIN: Dane Co., Madison (PADA:1), Waupaca Co ., Waupaca (CNCI:1,

UWEM:1). CANADA: ALBERTA: Hussar (CASC:1), Bilby (CASC:1); MANITOBA:

Awene (CNCI:4, USNM:2), Morgate (LBCC:1), Onah (CNCI:1), Teulon

(USNM:1); ONTARIO: Green1s Creek (CNC1:1), Toronto (CUIC: 1); QUEBEC:

Ste Foy (NMDC:1); SASKATCHEWAN: Alvena (EHSC:2, LBCC:2), Baine Lake

(LBCC:1), Buffer Lake (LBCC:1), C.B.C. Tower (LBCC:1), Elbow (CNCI:1),

Forestry Farm (LBCC:2), Ogema (CNCI:1), Perdue (LBCC:1), Saskatoon

(CNCI:14, USNM:1).

Phyllotreta spatulata NEW SPECIES

(Figs. 28, 76-77, 119-120, 146, 167)

Holotype: Male, deposited at CASC.

Type-locality: Humboldt Co., California.

DIAGNOSIS: Elytra each with the median stripe simple, of uniform width and without dilations, slightly incurved toward suture apically; male Sth antennal segment expanded bilaterally, dorsoventrally

flattened, with venter flat, not concave; antennal segments dark

except at least 1-5 paler on venter. DESCRIPTION: Holotype (variation, excluding punctation, in paren­ theses): Fig. 28; elongate, length 2.52 mm (male 2.45-2.52, female

2.55-2.62), width 1.15 nun (male 1.12-1.15, female 1.25-1.28); head and pronotum black with slight metallic luster, elytra brownish black with slight metallic luster (no luster), each with a median straw- yellow stripe. Head: Black; texture finely granulate basally to slightly roughened anteriorly (entirely finely granulate); moderately punctate, punctures separated by less than to 2x their diameter, mostly by about one diameter; Interocular Distance/Maximum Diameter of eye, 1.88 (male 1.88, female 1.81). Antennae: Figs. 76, 77; segments

3-5 progressively longer, 6 less than 1/2 length of 4, segment 7

slightly longer than 6; segments 4 and 5 expanded bilaterally

dorsoventrally flattened; segment 5 distinctly longest, with venter

flat, not concave; segment(length/width3 : l(9/4-3, 2(5.5/2.53,

3(5.5/3.53, 4(7/4.53, 5(8.5/5.53, 6(2.5/33, 7(4/33, 8(4+/3+3, 9(4/33,

10(4+/33, 11(6/3.53, total length 1.50 mm (Allotype: 1(8.5/3+3,

2(5/2+3, 3(4.5/23, 4(5-/2.53, 5(6.5/3-3, 6(4+/33, 7(5-/33, 8(S/3+3,

9[5/3+3, 10(4.5/3+3, 11(6/3+3, total length 1.48 mm; simple); antennae

dark brown with segments 1-7 (at least 1-5) paler ventrally (female

with basal 2 or 3 segments or with segments 2 or 2-3 paler, ventrally

segments 3-5 or 4-5 somewhat darker than basal segments but paler than

dorsum). Pronotum: Length 0.45 mm (male 0.42-0.45, female 0.45), width

0.80 mm (male 0.80, female 0.82); black; texture finely granulate;

moderately punctate, punctures separated by less than 1/2 to equal

their diameter, mostly by about one diameter. Elytra: Length 1.82 mm 119

(male 1.75-1.82, female 1.90-1.92), width 1.15 mm (male 1.12-1.15, female 1.25-1.28); brownish black (dark brown), median stripe as in

Fig. 28; texture smooth (slightly roughened); coarsely punctate with moderate punctures interspersed, punctures separated by less than

1/2 their diameter. Legs: Normal color sequence. Abdomen: 5th sternum with an apical median lobe moderately concave, concavity / tapering (oval) in its extension to midpoint, with a median linear impression from sternal base to subapex, frequently interrupted

(female simple in outline, with an oblong median impression in apical

1/2 or with apical 1/2 flat medially); brownish black (dark brown) with median lobe paler.

Male genitalia: Figs. 119, 120; length approximately 0.95 mm; essentially straight except for basal piece slightly and apex strongly curved ventrad, of rather uniform width except gradually broadening in apical 1/3, broadest subapically; apex in dorsal view broadly

rounded, faintly truncate medially, in lateral view strongly but

gradually curved ventrad with a ventral subapical notch, of rather uniform width with tip bluntly rounded.

Female genitalia: Fig. 146; length approximately 0.35 mm; recep­

tacle elongate, with basal margin tapering to a blunt preapical point,

rather abruptly and strongly swollen ventrally in basal 1/2 and

elongate, almost parallel-sided in apical 1/2, with a moderately

developed ring collar at junction with pump; pump long; spermathecal

duct attached on dorsum of receptacle.

BIOLOGY: Host plants, Immature stages, and Habits are unknown. 120

DISCUSSION: Distribution: Fig. 167; P. spatulata has been collected only in Humboldt Co., California. New name: The specific epithet spatulata comes from the Latin spatul- meaning a spoon and refers to the male genitalia in lateral view resembling a scoop or spoon.

Relationships: P^. spatulata is most similar to the striped specimens of JP. denticomis, and is less similar to P_. lepidula, P. arcuata, and P. constrieta with which it shares having a simple elytral stripe and the male 5th antennal segment dorsoventrally flattened. Phyllotreta spatulata also shares the flattened male 5th antennal segment with P^. utana, P_. utanula, P_. oregonensis, P^. emarginata, and P_. zimmermanni, but these 6 species have the elytral stripe dilated, not simple. Phyllotreta denticomis is unique in having the male 6th antennal segment with a sharp process. The male genitalia of P. spatulata and denticomis are very similar and distinct as a group, based particularly on the lateral view. However,

in dorsal view P. spatulata is more abruptly broader subapically

and the apex is broadly rounded and faintly truncate medially, whereas

in P. denticomis the median lobe is somewhat broadened in the apical

1/4 and the apex is broadly rounded but moderately and shallowly

emarginate; in lateral view on the venter P^. spatulata has a shallow

subapical notch, whereas P^. denticomis has a small ventrally directed

subapical lobe. The female spermathecae of P^. spatulata are unique,

but are most similar to those of £. dolichophalla. The spermathecae

of P_. spatulata have the dorsum straight and the ring collar is not

as wide or strongly developed, whereas in dolichophalla the 121 receptacle dorsum is slightly swollen in the basal 1/2 and the ring collar is much more strongly developed.

TYPE MATERIAL: The holotype, allotype, and one paratype (9) are at the CASC, and one paratype (d) is at the USNM. The holotype is mounted on a point and the pin bears 3 labels (decending order):

"Humboldt Co. Cal." "Van Dyke Collection" and "HOLOTYPE Phyllotreta t spatulata By Eric H. Smith '73"; also there is a polyethylene genitalia vial added by me. The allotype bears the same first two labels and a third label "ALLOTYPE Phyllotreta spatulata By Eric H.

Smith *73" and the paratype at the CASC also bears the same first two labels and a paratype label; the paratype at the USNM bears the following: "Humboldt Co. Cal." "WICKHAM Collection 1933" "Phyllotreta lepidula Lec." and a paratype label.

. SPECIMENS EXAMINED: Total 4: CALIFORNIA: Humboldt Co., CCASC:1,

Holotype, Allotype, USNM:1).

Phyllotreta striolata (F.)

(Figs. 29-35, 78-79, 121-122, 147, 168)

Crioceris vittata Fabricius, 1801. Systema Eleutheratorum, 1:469.

(not £. vittata Fabricius, 1775:122)

Neotype (here designated): Male, at MCZC.

Type locality: "Carolinae"; neotype, South Carolina.

Crioceris striolata Fabricius, 1803. Index alphabeticus in J.C.

Fabricii Systema Eleutheratorum, genera et species continens,

p. 38. (Substitute name for C. vittata F., 1801.) 122

Haltica striolata Illiger, 1807. Mag. Insektenk., 6:148.

Phyllotreta striolata, Barber, 1947. Proc. Bnt. Soc. Washington,

49(6):156-157.

Haltica sinuata Redtenbacher, 1849. Fauna Austriaca, p. 532. (not

Stephens, 1831; not Horn, 1889)

Phyllotreta sinuata, Foudras, 1860. Ann. Sco. Linn. Lyon, 6(ser.2):345.

/ Orchestris vittata, Crotch, 1873. Proc. Acad. Nat. Sci. Philadelphia,

25:66.

Phyllotreta vittata, Horn, 1889. Trans. American Ent. Soc., 16:294.

Phyllotreta sinuata (Var.) discedens Weise, 1888. Naturgeschichte der

Insecten Deutschlands . . ., Abt. 1, Coleoptera, 6 (Lief.5):871.

Type locality: "Nordafrika", Syria, Siberia to Wladiwostok.

Phyllotreta vittata (variety) discedens, Chittenden, 1927.

Entomologica Americana, 8(n.s.,1):25.

Phyllotreta vittata (fa.) maculipennis (ab.) discedens, Heikertinger

and Csiki, 1939. Coleopterorum catalogus, pars 166, p. 55.

Phyllotreta maculipennis (abber.) discedens, Balsbaugh, 1972. The

leaf beetles of Alabama, p. 172.

Phyllotreta sinuata (Var.) monticola Weise, 1888. Naturgeschichte der

Insecten Deutschlands . . ., Abt. 1, Coleoptera, 6 (Lief.5):871.

Type locality: "Kamthen" and Krain” .

Phyllotreta vittata (variety) monticola, Chittenden, 1927.

Entomologica Americana, 8(n.s.,1):24.

Phyllotreta vittata (fa.) tenuelimbata (ab.) monticola, Heikertinger

and Csiki, 1939. Coleopterorum catalogus, pars 166, p. 55. 123

Phyllotreta tenuelimbata (abber.) monticola, Balsbaugh, 1972. The

leaf beetles of Alabama, p. 172.

Phyllotreta vittata (variety) artivitta Chittenden, 1927.

Entomologica Americana, 8(n.s.,1):26. Type locality: Muscatine,

Iowa.

Phyllotreta vittata (ab.) artivitta, Heikertinger and Csiki, 1939.

Coleopterorum catalogus, pars 166, p. 56.

Phyllotreta vittata (variety) lineolata Chittenden, 1927. Entomologica

Americana, 8(n.s.,1):25. Type locality: Corvallis, Texas.

Phyllotreta vittata (ab.) lineolata, Heikertinger and Csiki, 1939.

Coleopterorum catalogus, pars 166, p. 56.

Phyllotreta vittata (variety) vernicosa Chittenden, 1927. Entomologica

Americana, 8(n.s.,1):25. Type locality: Fieldbrook, California.

Phyllotreta vittata (ab.) vernicosa, Heikertinger and Csiki, 1939.

Coleopterorum catalogus, pars 166, p. 56.

DIAGNOSIS: Elytra each with a median pale stripe with dilations,

which may be reduced to a subbasal and subapical pale mark, subapical

part of stripe or mark incurved apically but never meeting at suture;

male antennal segment 5 enlarged, cylindrical, about 2x length of

segment 6.

DESCRIPTION: Neotype (variation, excluding punctation, in paren­

theses): Fig. 29; elongate oblong, length 2.12 mm (male 1.80-2.42,

female 2.12-2.60), width 1.02 mm (male 0.90-1.22, female 1.02-1.35);

head and pronotum black with slight metallic luster, elytra brownish

black (often with slight metallic luster), each with a median 124 straw-yellow stripe. Head; Black (rarely brownish black); texture finely granulate (rarely finely granulate to finely alutaceous or slightly roughened); moderately punctate, punctures separated by less than 1/2 to equal their diameter, mostly by less than

1/2 a diameter; Interocular Distance/Maximum Diameter of eye, 1.25 (male 1.11-1.28, female 1.11-1.28). Antennae Cleft

/ with segments 8-11 missing]: Figs. 78, 79; segment 4 longer than

6, subequal to 7, segment 5 about 2x length of 6; segment 5 moderately (often only slightly) enlarged, cylindrical (shape may vary slightly); segment [length/width]: 1(8/3], 2(4/2.5], 3(4+/2],

4(4/3-], 5(6.5/3.5], 6(3/2], 7(4.5/2+], 8[4+/3], 9(4.5/3], 10(4/3],

11(6.5/3], total length 1.32 mm (female: 1(8/3-], 2(4/2+], 3(3.5/2-],

4(4/2-], 5(5+/2], 6(3/2], 7(4.5/2+], 8(4.5/2.5], 9(4.5/2.5], 10(4/2.5],

11(5.5/3], total length 1.26 mm; simple); antennae various shades of brown, basal 2 segments paler with dorsum of 1 dark, 3 (or none) intermediate, 4-11 (3-11) darkest (females: basal 3 segments paler, dorsum of 1 may be dark, segment 4 or 4-5 intermediate, segments 5-11 or 6-11 darkest). Pronotum: Length 0.40 ram (male 0.32-0.48, female

0.40-0.48), width 0.68 mm (male 0.60-0.78, female 0.65-0.90); black; texture finely granulate (rarely finely alutaceous, or finely alutaceous to slightly roughened, or smooth to slightly roughened to finely granulate); coarsely punctate, punctures separated by less than

1/2 their diameter. Elytra: Length 1.58 mm (male 1.38-1.80, female

1.25-2.60), width 1.02 mm (male 0.90-1.22, female 1.02-1.35); brownish black (black), median stripe pattern as in Fig. 29 (pattern varies as in

Fig. 30 to Fig. 35); texture smooth to slightly roughened (often 125 entirely slightly roughened); coarsely punctate, punctures separated by less than 1/2 to equal their diameter, mostly by less than 1/2 a diameter. Legs Cleft metatarsus with segments 2-5 missing]: Normal color sequence. Abdomen: 5th sternum with an apical median shallowly

(to deeply) concave lobe, concavity tapering (often oblong) extending to within 1/3 of sternal base (extending only to midpoint), with a somewhat indistinct median linear impression extending from sternal base to base of concavity (distinct; extending from sternal base or subbasally to subapically, sometimes interrupted; female: simple in outline, often with a median longitudinal to oval shallow impression in apical 1/3); black (dark brown, usually brownish black) with apical

1/3 (to 1/4) paler.

Male genitalia: Figs. 121, 122; length approximately 0.80 mm; mod­ erately arched dorsoventrally, of rather uniform width except slightly narrowed preapically in dorsal view, with a dorsal washboard; apex in dorsal view broadly rounded with an acute blunt tip, in lateral view gradually tapering in apical 1/4 to a very narrow, ventrally directed apex, tip pointed.

Female genitalia: Fig. 147; length approximately 0.26 mm; recep­ tacle elongate, wider basally and gradually tapering to posterior moderately developed ring collar; pump long, more than 1/2 receptacle length.

EI0L0GY: Host plants, adult (the following represent those records verified by me via label data): Alfalfa (Kansas, Vernal), *Brassica, bluegrass, *cabbage, *cauliflower, clover, c o m , *horse-radish, *kale, lettuce, *mustard, *Texas mustard green, *Nasturtium palustre, *pepper- 126

grass, potato vines, Pelfer grass, *Radicula palustris (=Rorippa

islandica Feraaldiana Butters $ Abbe), *radish, *rape, ♦rutabaga,

♦turnip. Riley (1885a) records in addition: *charlock (Sinapis),

♦shepherd's purse (Capsella), *stock (Matthiola), and *rocket

(Hesperis). Duckett (1920) lists in addition: ♦brussels sprouts,

gallberry (Ilex glabra), and sugar and garden beets. Chittenden (1927) records in addition: ♦mustaird (Brassica nigra and JS. arvensis), *stock

(Matthiola incana et al.), *hoary alyssum (Berteroa incana), ♦sweet

alyssum (Alyssum maritimus et al.), ♦wallflower (Chieranthus cheiri),

♦candy tuft (Iberis), ♦yellow cress (Radicula silvestris), sea lavender

(Static** s *.nuata), and ♦Erysimum spp. Feeny, Paauwe, and Demong (1972)

list collections off 24 plants of which the following are probable hosts not listed above: ♦broccoli (Brassica oleraces botrytis L.) and

♦spider plant (Cleome pungens Willd.). Balsbaugh (1972) adds ♦Brassica

napobrassica.

Host plants, larva: Cabbage (Shimer, 1869).

Immature stages: Descriptions include egg by Riley (1885a), larvae

by Shimer (1869) and Riley (1885a), and pupa by Shimer (1869).

Habits: The larvae are root feeders (Shimer, 1869). The adults

feed mainly on the Cruciferae, including most of the genera (Tahvanainen,

1972). However, they probably feed on some members of the closely

related plant families Capparidaceae, Tropaeolaceae, and Limnanthaceae

(Feeny, Paauwe, and Demong, 1970). If populations become particularly

large, they will feed on plants of other families, for example, I have

collected them feeding on lettuce (Lactuca sativa) belonging to the 127

Compositae which was located at the end of turnip rows. This species is an important economic pest and insecticide treatment is usually required if cruciferous vegetables are grown.

In the midwest and northeastern U.S. the adults overwinter (in the top soil in New York; Hicks, 1972) and become active in early spring.. Eggs are laid on or near the host (Riley, 1885a; Hicks, 1972).

/ A few adults are present throughout the summer but beginning in late

July the populations start to rapidly increase in size due to the emergence of the new adults from the spring-deposited eggs.

This species was introduced from Eurasia sometime before 1801 and is now widespread. Adults have been collected starting in mid-June in the Northwest Territory, in early April in Iowa and Wisconsin, in late March in North Carolina and early March in Louisiana. They have been collected as late as mid-September in Canada to late October in

Louisiana; occasionally, they have been collected in December and

January which probably represent overwintering adults.

An analysis of elytral color pattern variation was done for the available material. Seven categories of patterns were selected and are represented in Figs. 29-35; these are in order of category 1 (very wide stripe) to category 7 (stripe represented by two marks) with the exception that Fig. 29 fits between Fig. 31 and Fig. 32, representing category 3. For the population samples, the individuals were placed in one of these seven categories and the results are represented by the phenograms of Fig. 181, which are arranged (left to right) geograph­ ically north to south with the disjunct California population represented last. In general, the sample size was relatively small 128 for variation analysis work and only a coefficient of variance has been run on the data. However, a general trend can be seen which is that progressing north to south, there is an increase in dark versus light area, or the elytral stripe tends to be more interrupted

(represented by two marks). Until more and larger samples from a wider geographical range can be obtained, little more can be said.

DISCUSSION: Distribution: Fig. 168; P_. striolata is widespread but has been more frequently collected in northeastern and midwestem

U.S. and central Canada.

Nomenclature: Fabricius (1801a) described this species in the genus Crioceris using the already occupied specific epithet, vittata.

In 1803, Fabricius published an index to his 1801 work and proposed the specific epithet striolata to replace this 1801 vittata. Illiger

(1807) transferred £. striolata to Haltica. The Fabrician replacement name and Illiger*s transfer were ignored until Barber (1947) made the transfer from Haltica to Phyllotreta and clarified the previous confusion. In 1849, Redtenbacher described this species as Haltica sinuata (not Stephens, 1831; not Horn, 1889, as Phyllotreta sinuata) which was synonymized by Foudras (1860). Crotch (1873) wrongly placed P. vittata in Orchestris (see LeConte, 1878:165) and Horn

(1889) transferred it to Phyllotreta.

Weise (1888) described two varieties of P. striolata as P^. sinuata discedens and IP. sinuata monticola; Chittenden (1927) transferred them to P. vittata and retained their varietal status. Heikertinger and

Csiki (1939) listed these as aberrations of two "fa.*1 (? form), 129 maculipennis and tenuelimbata; that is, P^. vittata maculipennis discedens and £. vittata tenuelimbata monticola. Balsbaugh (1972) forgot to include the specific epithet vittata in his citation of the

Heikertinger and Csiki (1939) listing and miscited the year and page references. Chittenden (1927) described 3 color varieties of P. vittata (artivitta, lineolata, and vemieosa) which were listed by

Heikertinger and Csiki (1939) as aberrations. According to article 1 and article 45 a, b, c, e(i) of the Rules of Zoological Nomenclature, these 5 varieties or aberrations hold no taxonomic rank and are listed here only for completeness and clarity. In each of several long series of over 100-150 specimens which I have collected, all 3 of Chittenden*s varieties are present. Also the European varieties discedens and monticola of Weise, which Chittenden recognized as being present in the U.S., are both represented in the same long series as Chittenden*s

3 varieties (see Figs. 29-35 and Fig. 181).

According to the Rules of Zoological Nomenclature, the type specimen of Crioceris vittata F. is also the type specimen of the replacement name Crioceris striolata F.; for additional discussion, see

Barber (1947:156-157).

The "type" of Crioceris vittata Fabricius, given as coleopteran number 1618 by Zimsen (1964:105), was borrowed from the University

Zoological Museum, Copenhagen. However, the "type" sent is not a

Phyllotreta but, based on my experience (1970), is most probably

Systena elongata (F.). The specimen has the head and prothorax missing

and is a female which means that until the female genitalia are studied, 130 further confirmation of identity is not possible. The differences between the Fabrician description and elongata are: 1) the size is given as somewhat small, whereas jS. elongata is of moderate size for a flea beetle, and 2) the elytral stripe is given as incurved toward the suture basally and apically, but not touching apically, whereas in S. elongata the stripe is straight basally and only very slightly incurved apically at most. Fabricius1 original description agrees with what we presently call P^. striolata but unfortunately Olivier

(1808a, 1808b) didn't review and/or illustrate this species when he reworked the Fabrician collection so there is no check.

The type locality is given as "Carolinae" and the location as being

1 in the Bose Collection, which suggests that the type locality be restricted to the vicinity of Charleston, South Carolina (Blake, 1952).

Blake (1952) did not find C. vittata in the Bose Collection but she did comment that all specimens present were in good condition because

"probably all specimens that were not had long since been discarded".

The specimen sent from Copenhagen was accompanied by Fabricius' deter­ mination label which consists of a t o m piece of brown paper with

"vittata" written in Black ink. This specimen is from the Kiel

Collection which was Fabricius* personal collection and is probably not the type specimen he referred to as in the Bose Collection.

Based on the "type" sent, the original description, and Blake's observations, it is most probable that the type of Crioceris vittata

F. (1801a:469) has been lost and that the Fabrician determination label has been erroneously associated with a different species of striped flea beetle. Therefore, a neotype has been designated which was collected as near to the type locality as available and which agrees with Fabricius' original description.

Relationshipst P^. striolata is a rather distinctive species.

Specimens with the typical elytral stripe pattern have been confused with P_. zimmermanni which are much longer and more robust and have the male 5th antennal segment dorsoventrally flattened instead of cylin-

/ drical. Specimens of P_. striolata which have the elytral stripe reduced to a subbasal and subapical pale mark have often been confused with P. bipustulata but in P. striolata the elytral subapical mark has incurved toward the suture instead of having its sutural margin straight; also male antennal segment 5 is enlarged and about 2x length of 6 in P. striolata,’ whereas in P. bipustulata segment 5 is simple and segments 5 and 6 are subequal. The male genitalia of P_. striolata is most similar to those of P^. decipiens, 1?. bipustulata, P. dolicho- phalla, £. ramosa, and P_. ramosoides with which it shares general shape and having a dorsal washboard, but differs in the shape of the apex.

The female spermathecae of P. striolata are most similar to those of

decipiens, but those of P_, striolata are shorter and have the pump much longer.

TYPE MATERIAL: The neotype is at the MCZC. It is mounted on a point and the pin bears 2 labels (decending order): "S.C." and "NEOTYPE

Phyllotreta striolata By Eric H. Smith '73".

SPECIMENS EXAMINED: Total 1536: ALABAMA: Jefferson Co., Birmingham

(USNM:4), Mobile Co., Mobile (MCZC:1); ALASKA: CUSNM:4); ARKANSAS:

Crawford Co. (UADE:5), Hempstead Co., Hope CCUIC:8, LACM:22); CALIFORNIA

Inyo Co., Bishop (LACMrl), Los Angeles Co., N. Long Beach (UCRC:6), San Marino (LACM:4, UCRC:3), Orange Co., Huntington Beach (USNM:1),

Santa Ana CLACM:3), San Diego Co., Descanso (FMNH:1), [ANSP:3, MCZC:9,

USNM:l3; COLORADO: Arapahoe Co., Denver (ICCM:1); FLORIDA: Highlands

Co., Archbold Bio. Stat. (PSUC:7), Orange Co., Orlando (CNCI:1, USNM:2),

[ICCM:l3; GEORGIA: Bryon Co., Richmond Hill (USNM:19); IOWA: Adair Co.

(USNM:1), Lee Co., Ft. Madison (ICCM:4), Scott Co., Pleasant Valley

(CNCI:1), Story Co., Ames (ISUI:16, 0SU0:5), Taylor Co. (USNM:1),

Woodbury Co., Sioux City (DEFW:1), CDEFW:1, USNM:l3; ILLINOIS: Cook Co.,

Chicago (UWEM:1), Palos Park (FMNH:2), DeKalb Co. (UWEM:4),CDEFW:16,

WSUC:43; INDIANA: LaPorte Co., LaPorte (ICCM:1), Tippecanoe Co.,

Lafayette (CNCI:10), West Lafayette, ESC Smith Lot No. 375 (EHSC:40),

ESC Smith Lot No. 376 (EHSC:153), ESC Smith Lot No. 377 (EHSC:130);

KANSAS: Douglar Co., Lawrence (CNCI:4), CMSUC:l3;LOUISIANA: East Baton

Rouge Par., Baton Rouge (CNCI:2 , MCZC:1, USNM:1), Evangeline Par.,

Mamou (SDSU:2), Lafayette Par., southwestern Lafayette (SDSU:1), Rapides

Par., Ball (USNM:2), St. Bernard Par., Haraham (MCZC:1),EMCZC:23;

MASSACHUSETTS: Middlesex Co., Arlington (MSUC.-l), N. Cambridge (MCZC:1),

Sherborn (MCZC:2), CMCZC:153; MARYLAND: Anne Arundel or Howard Co.,

Jessup (UCRC:1), Baltimore Co., Baltimore (MCZC:6), Frederick Co.

(MCZC:1), Garrett Co., Oakland (UMDC:1), Prince GeorgeTs Co., Blandens-

burg (UMDC:3), College Park (UMDC:22, USNM:10), Riverdale (UMDC:3),

Somerset Co., Westover (CUIC:2), EMCZC:1, UMDC:2]; MAINE: Hancock Co.,

Isle-au-Haut (MCZC:3), Penobscot Co., Orono (CNCI:1, UMDE:4), C0SUC:13;

MICHIGAN: Allegan Co., Fennville (MSUC:3), Ingham Co., Ag College

(MSUC:14), CMSUC:1] ; MISSOURI: Boone Co., Columbia (LACM:24), [DEFW:23; 133

MINNESOTA: Anoka Co. (DEFW:1), Cook Co ., Grand Marais (DEFW:2), poplar

Lake (DEFW:1), (DEFW:81, Dodge Co., Mantorville (CNCIsl, DEFW:3,

FMNH:1), Fairbault C o., Winnebago (CNCI:2, DEFW:2), Mille Lacs Co.

(DEFW:3), Olmsted Co. (DEFW:6), Polk Co., Crookston, Red Lake River

(DEFW:5), Crookston (DEFW:6), Ramsey Co,, St. Anthony Pk. (DEFW:4),

St. Paul (DEFW:2), St. Paul, Univ. Farm (DEFW:13), (DEFW:8), Sibley

Co., Blakely (DEFW:17), Steele C o., Owatonna (CUIC:2), Winona Co.,

Dresbach (DEFW:6), [CNCI:1, DEFW:2, MCZC:lD; MISSISSIPPI: Copiah Co.,

Crystal Sprs. (USNM:1), George Co., Lucedale (CUIC:18); NEBRASKA:

Lancaster Co., Lincoln (DEUN:5); NORTH CAROLINA: Ashe Co., Jefferson

(NCSU:1), Pender Co., St. Helena (NCSU:9), Wake Co ., Raleigh (UCRC:2),

[MCZC:3]; NEW HAMPSHIRE: [MCZC:5, MSUC:1); NEW JERSEY: Ocean Co.,

Seaside Park (FMNH:4), [MCZC:53; NEW YORK: Chautaugua Co., Westfield

(MCZC:9), Monroe Co., Rochester (LACM:7), Madison Co., Hubbardsville

(CUIC:2), Nassau Co. (CUIC:4), Queens Co., Flushing L.I. (CUIC:2),

St. Lawrence Co., Canton (NMDC:1), Onondaga Co. (NMDC:2), Tompkins Co.,

Ithaca, Savage Farm (CUIC:2), Ithaca (CUIC:8), (NMDC:4), Wyoming Co.,

Pike (MCZC:2), CAMNH:3, ANSP:1, CUIC:9, DEFW:1, MSUC:1, USNM:1]; OHIO:

Delaware Co., E.H. Smith Lot No. 310 (EHSC:63), E.H. Smith Lot No.

367 (EHSC:119), E.H. Smith Lot No. 371 (EHSC:65), Franklin Co., Columbus

(LACM:1, OSUC:5, SDSU:2), Wyandot Co ., ESC Smith Lot No. 391 (EHSC:52);

OREGON: Multnomah Co., Portland (USNM:1), [0SU0:1); PENNSYLVANIA:

Allegheny Co., Harmarville (ICCM:2), Pittsburg (ICCM:9), (ICCM:10),

Centre C o ., State College (PSUC:1), Dauphin Co., Harrisburg (PADA:5),

Hummelstown (PADA:9), Lancaster Co., Lancaster (PSUC:7), Northhampton Co., Bangor (PADA:5), Westmoreland Co.. Jeannette (ICCM:3, PADA:4),

CANSP:3, ICCM:5, MCZC:5, 0SUC:6, PADA:23; SOUTH CAROLINA: (ANSP:1,

Neotype, MCZC:1); SOUTH DAKOTA: Beale Co., Huron (SDSU:1), Brookings

Co., Brookings (SDSU:38), Brown Co., Hecla (SDSU:3), Clay Co .,

Vermilion (SDSU:1), CSDSU:4]; TEXAS: Bexar (UCRC:1), Colorado Co .,

Columbus (USNM:1); UTAH: Utah Co., Benjamin (JSCC:1), Provo (MCZC:1);

VIRGINIA: Fairfax Co ., Falls Church (MCZC:2), York C o., Norfork CCUIC:2,

UCRC:2), CANSP:1, VPIC:1]; WISCONSIN: Dane Co., Madison (JSCC:2,

0SU0:4, USNM:6, UWEM:46), Grant Co. (JSCC:2), Kenosha Co., Kenosha

CUWEM:3), Lafayette Co., Belmont (MSUC:1), Milwaukee Co., Milwaukee

(MSUC:1, SDSU:7), Oneida Co., Minosqua (UWEM:2), Racine Co., Racine

(UWEM:3), Wood Co ., Griffith CUWEM:1), [CUIC:3]. CANADA: ALBERTA:

Edmonton (ANSP:4, CNCI:1, CUIC:4, MCZC:4, USNM:1), McMurray (CNCI:1);

QUEBEC: Aylmer (CNCI:1), Hemningford (CNCI:1), Laniel (CNCI:1),

LaTrappe (MSUC:4), Montreal (MCZC:1), Outremont (LACM:3); ONTARIO:

Aldershot (CNCI:3), Burlington (CNCI:1), Constance Bay (CNCI:1),

Eberts (CNCI:1), Erican (CNCI:2), Gainston (CNCI:1), Leamington (CNCI:1),

Manotick (CNCI:1), Mer Bleve (CNCI:1), Ottawa (CNCI:18), Pelee Island

(CNCI:1), Toronto (CUIC:18)i MANITOBA: Aweme (CNCI:11, MCZC:6), Berens

River (CNCI:2), Brandon (LBCC:1), Grandview (LBCC:2), Lena (LBCC:1),

Morris (CNCI:1), Ninette (CNCI:1), Shoal Lake (LBCC:2), Westbaurne

(LBCC:1), Winnipeg (LBCC:1); NEWFOUNDLAND: Harmonfield (CNCI:4),

Stephenville Cross (CNCI:1); NEW BRUNSWICK: Fxederickton (CNCI:5),

Millville (CNCI:1); NORTHWEST TERRITORY: Aklarik (CNCI:2), Norman Wells

(CNCI:23), Yellow Knife (CNCI:1); NOVA SCOTIA: Truto (CNCI:1); 135

SASKATCHEWAN: Aberdean (LBCC:1), Alvena (LBCC:2), Aylsham (LBCC:1),

Buffer Lake (EHSC.'l, LBCC:1), Cochin (LBCC:2), Duck Lake (LBCC:1),

Edam (EHSC:1), Flin Flan (SDSU:1), Forestry Farm (LBCC:3), Goodsoil

(LBCC:1), MacDowall (LBCC:1), Marcelin (EHSC:1, LBCC:1), Meskavaw

(LBCC:1), Saskatoon (CNCI:1), Shell Lake (LBCC:1), Smeaton (LBCC:2),

Strawbourg (CNCI: 2) , Sutherland (EHSC:1, LBCC:1), Zenon Pk. (LBCC:1).

NO DATA: CCNCI:1, DEFW:2, MCZC:6, UNDC:4, UWEM:1, WSUC:2l3.

Phyllotreta utana Chittenden

(Figs. 36, 80-81, 123-124, 148, 169)

Phyllotreta utana Chittenden, 1920. J. Washington Acad. Sci., 10:

389-390, fig. 1.

Holotype: Male, USNM type #23114.

Type locality: Logan, Utah.

DIAGNOSIS: Elytra each with a median pale stripe with dilations

which never meet at suture; male antennal segment 5 expanded bilater­

ally, dorsoventrally flattened; antennal segments 2-5 pale, in distinct contrast to darker segments 7-11; greater than 2.8 mm long (about 3 mm); male genitalia in lateral view slightly sigmoid with apex almost straight dorsally, and strongly swollen medially on venter.

DESCRIPTION: Holotype (variation, excluding punctation, in paren­

theses): Fig. 36; elongate oval, length 3.25 mm (male 2.85-3.25, female

2.85-3.02), width 1.20 mm (male 1.20-1.50, female 1.42-1.52); head and

pronotum black with a slight metallic luster (no luster), elytra brown­

ish black with a slight metallic luster (no luster), each elytron with a median straw-yellow stripe. Head: Black; texture finely granulate

(finely alutaceous basally to slightly roughened anteriorly); moder­ ately punctate, punctures separated by less than to equal their diameter; Interocular Distance/Maximum Diameter of eye, 1.55 (male

1.47-1.70, female 1.50-1.78). Antennae: Figs. 80, 81; segments 4 and 6 in length (subequal), 5 about 1.5x longer than 4 (almost 2x 4),

/ 6 shorter than 7; segments 4 and 5 strongly expanded bilaterally, dorsoventrally flattened; segment 5 distinctly longest, venter essen­ tially flat; segment (length/width): 1(10/4], 2(4/3], 3(6/3], 4(6/5),

5(9/6), 6(4/3.5], 7(6/3.5], 8(6/4), 9(6.5/3.5), 10(6/3.5], 11(7/4), total length 1.76 mm (female paratype: 1(9/3+]* 2(4.5/3-), 3(4.5/2.5),

4(5/2.5], 5(6.5/2.5), 6(4.5/3-], 7(5.5/3+], 8(6/3.5+], 9(6/3.5],

10(5.5/3-], 11(7/3], total length 1.60 mm; simple); antennae various shades of brown, basal 5 (7) segments paler with dorsum of 1 dark,

8 (non) intermediate, 5-11 (6 or 7-11) darkest. Pronotum: Length

0.58 mm (male 0.50-0.58, female 0.52), width 1.05 mm (male 0.95-1.05, female 0.92-1.02); black; texture finely granulate and alutaceous

(slightly roughened, finely granulate only or with wrinkles radiating from punctures); coarsely punctate, punctures separated by less than to equal their diameter. Elytra: Length 2.42 mm (male 2.10-2.42, female 2.12-2.30), width 1.20 mm (male 1.20-1.50, female 1.42-1.52); brownish black (black), median stripe pattern as in Fig. 36; texture smooth; coarsely punctate, punctures separated by less than to equal their diameter, with several moderate punctures irregularly inter­

spersed. Legs: Normal color sequence except tibiae and tarsi same color. 137

Abdomen: 5th sternum with an apical median moderately concave lobe,

concavity tapering posteriorly and restricted to apical 1/3, a shallow

linear impression from base of segment to edge of median concavity

(median linear impression sometimes extending 1/3 into median con­ cavity or in basal 1/2 of sternum only, rarely recessed to median

concavity or only in basal 1/3; female with apical area broadly and r shallowly concave, a narrow median longitudinal impression in apical

1/3); brownish black (dark brown, black).

Male genitalia: Figs. 123, 124; length approximately 1.15 mm;

slightly sigmoid dorsoventrally, broadest apically in dorsal view,

strongly swollen medially on venter in lateral view; apex in dorsal

view moderately deep and broadly emarginate, in lateral view tapering

to a blunt posteriorly directed point with a short subapical median,

ventrally directed lobe.

Female genitalia: Fig. 148; length approximately 0.38 mm; recep­

tacle oblong with a slight ring collar at junction with pump.

BIOLOGY: Host plants, adult: Sugar beet, *hedge mustard (Chittenden,

1920:390). The hedge mustard record has not been verified by me

because no material with such labels was sent from the USNM.

Immature stages:• Unknown.

Habits: Adults of utana have been collected in Oregon from late

May until late July and in Utah from mid-April until mid-July.

DISCUSSION: Pistribution: Fig. 169; P. utana has been collected

in Oregon and Utah.

Nomenclature: There are no synonyms and Chittenden (1920)

described P. utana from 11 specimens. 138

Relationships: P. utana is most similar to P.. utanula and less similar to JP. emargiriata, zimmermanni, P^. constricta, and P^. oregonensis which all share the elytral color pattern and having the male 5th antennal segment appearing dorsoventrally flattened.

However, the elytral stripe of £. oregonensis and constricta is usually much wider medially than that of the other 4 species. The male 5th antennal segment of jP. zimmermanni has a basal concavity ventrally, whereas that of the other 5 is evenly flattened ventrally; also antennal segments 2-5 are pale in £. utana and £. utanula, whereas in the other 4 species antennal segment 2-3 are pale and/or dark and segments 5-11 are dark. The male genitalia of these 6 species separate into 3 distinct groups.(based on dorsal view of apex) as follows: P^. zimmermanni and P. emarg5.nata have the apex not broadened and its margin moderately but narrowly emarginate medially; oregonensis and P. constricta have the apex not broadened and its margin entire; and P. utana and P^. utanula which have the apex very broad and its margin moderately and broadly emarginate. However, the male genitalia in lateral view of utana are almost straight dorsally including the apex, and medially strongly swollen on the venter, whereas in P. utanula the dorsum is gradually arched, the apex curving slightly to the venter, and the venter is only very slightly swollen medially. The female spermathecae of P^. utana have the receptacle oblong or much more robust than those of the other 4 species (female of P. utanula is unknown), which are more elongate and also somewhat narrower medially. 139

Phyllotreta utana might be confused with P. arcuata on the basis of antennal structure and color, but P. arcuata has the male 5th antennal segment venter concave on the inner half instead of entirely flat and the elytral stripe is simple, without lateral dilations.

TYPE MATERIAL: The holotype and 10 paratypes are at the USNM. The holotype is mounted on a hair which is glued to a njale sex label and the pin bears 4 labels (descending order); "Logan UT" "6" "U.S.N.M.

Type No. 23114" (orange) and a determination label not in Chittenden’s writing.

SPECIMENS EXAMINED: Total 31: OREGON: Klamath Co., Crooked Cr.

Sprg. Hwy. 62 Bridge (JSCC:2), Crystal Cr. Upper Klamath Lk. (JSCC:1)

Klamath Falls (JSCC:2), 5 mi. E. Kirk (0SU0:1), 3 mi. S. Saddle Mt.

(JSCC:1), Williamson Riv. Rch. (JSCC:10), Wood River Valley (JSCC:1);

UTAH: Cache Co ., Logan (EMUS:1; holotype, USNM:1), Logan Canyon

(EMUS:1).

Phyllotreta utanula NEW SPECIES

(Figs. 37, 82, 125-126, 170)

Holotype: Male, deposited at USNM.

Type locality: Clatskanie, Oregon.

DIAGNOSIS: Elytra each with a median pale stripe with dilations

which never meet at suture; male antennal segment 5 expanded bilaterally,

dorsoventrally flattened; antennal segments 2-5 pale, in distinct con­

trast to darker segment 6-11; less than 2.8 mm long (about 2.6 mm); 140 male genitalia in lateral view with dorsum gradually arched, apex curving slightly ventrad, and venter only very slightly swollen medially; female unknown.

DESCRIPTION: Holotype (variation, excluding punctation, in paren­

theses): Fig. 37; elongate oblong, length 2.60 mm (2.48-2.75), width

1.18 mm (1.18-1,35); head and pronotum black with slight metallic

luster (luster pronounced), elytra dark brown, brownish black basally between stripes, with slight metallic luster, each elytron with a median straw-yellow stripe. Head: Black; texture slightly roughened

(finely alutaceous and roughened or granulate, finely granulate to roughened); moderately punctate, punctures separated by less than to

equal their diameter, mostly by about one diameter; Interocular

Distance/Maximum Diameter of eye, 1.67 (1.44-1.67). Antennae: Fig. 82;

segment 4 about 2x length of 6, longer than 7, segment 5 about 2x

length of 4; segments 4 and 5 strongly expanded bilaterally, dorso-

ventrally flattened; segment 5 distinctly longest, venter essentially

flat; segment (length/width]: 1(10/43, 2(4+/33, 3(4.5/33, 4(5.5/53,

5(11/73, 6(2.5/33, 7(4/33, 8(4.5/3.53, 9(5/3.53, 10(4.5/33, 11(6.5/33,

total length 1.55 mm; antennae various shades of brown, basal 6 (7)

segments pale with dorsum of 1 darker, apical 5 (4) segments dark.

Pronotum: Length 0.48 mm (0.42-0.50), width 0.82 mm (0.78-0.92); black;

texture slightly roughened (smooth, slightly roughened and/or finely

granulate); coarsely punctate, punctures separated by less than 1/2 to

equal their diameter, mostly by less than one diameter. Elytra: Dark

brown, brownish black basally between stripes (brownish black, black), 141 median stripe pattern as in Fig. 37; texture smooth to slightly roughened (smooth or slightly roughened); coarsely punctate, punctures separated by less than 1/2 a diameter. Legs: Normal color sequence.

Abdomen: 5th sternum with an apical median moderately concave lobe, concavity oval and extending to apical 2/5 (usually extending to midpoint), with a median linear impression extending from sternal base to apex (interrupted); black (brownish black).

Male genitalia: Figs. 125, 126; length approximately 0.89 mm; slightly arched dorsoventrally, gradually widening from midpoint to apex with apex boradest in dorsal view, widest medially in lateral view, not swollen on venter; apex in dorsal view moderately deep and broadly emarginate, in lateral view curving slightly ventrad, tapering to a blunt tip, with a short subapical median, ventrally directed lobe.

Female genitalia: Unknown.

BIOLOGY: Host plants and Immature stages are unknown.

Habits: Adults of P. utanula have been collected in Pacific North­ west from late May until early June and in early August and mid-October.

DISCUSSION: Distribution: Fig. 170; P. utanula has been collected in the Pacific Northwest east of the Rocky Mountains, specifically in southern British Columbia, eastern Washington, and eastern Oregon.

New name: The specific epithet utanula comes from a combination of the specific epithet of P. utana and the Latin diminutive suffix -ula referring to the fact that this species is very similar to P_. utana but smaller.

Relationships: P. utanula is most similar to P. utana, and less 142 similar to P_. emarginata, P. zimmermanni, P_. constricta, and P. oregonensis; for a discussion of these relationships, see this section under P^. utana.

Phyllotreta utanula might be confused with ]?. arcuata on the basis of antennal structure and color, but P. arcuata has the male 5th antennal segment venter concave on the inner half instead of entirely / flat, and the elytTal stripe is simple, without lateral dilations.

TYPE MATERIAL: The holotype is at the USNM. All paratypes are males and are distributed as follows: 1 at ANSP, 2 at CASC, 1 at CNCI,

2 at EHSC, 2 at JSCC, 2 at NMDC, and 1 at OSUO. The holotype is mounted on a point and the pin bears 3 labels (decending order):

"Clatskanie, Ore. Aug. 4, 1956 Cline of printing marked out with pencil! Coll. J. Schuh" "Phyllotreta utana

Gentner" and "HOLOTYPE Phyllotreta utanula By Eric H. Smith ?73"; I added a polyethylene genitalia vial.

The paratypes all have a paratype label added. The paratype at

ANSP is labeled "Portlant Ore.” "268" "Horn Coll H 7025"; those at CASC

are one labeled "Fort Lewis, Wn. V I *12*1945 P.H. Amaud", the other

"Corvallis Or." "Van Dyke" "Van Dyke Collection"; the paratype at

CNCI is labeled "B.C.; 12 mi. E. Hope, VI.2.1969 Cambell 8 Smetana"

"ex river debris"; one at EHSC has the same first two labels as the

holotype, the other "Crystal Cr., Ore. Upper Klamath Lk. May 30, 1960

Joe Schuh, Coll."; the two paratypes at JSCC are "Kelsey Valley, Ore.

Douglas County June 26, 1962 J.D. Vertress, Coll." and the other,

"Corvallis Ore." one paratype at NMDC is labeled "Montesano, Wn.

* 143

14 June 1928 Win. W. Baker" and the other "Puyallup, Wn. 10 Oct. 1927

Wn. W. Baker"; and the paratype at 0SU0 is labeled "Salem Ore. 5-15-25."

SPECIMENS EXAMINED: Total 12: OREGON: Benton Co., Corvallis

(JSCC:1), Columbia Co., Clatskanie (EHSC:1, Holotype:USNM), Douglas Co.,

Kelsey Valley (JSCC:1), Crystal Cr. Upper Klamath Lk. (EHSC:1),

Marion Co., Salem (OSUO:l), Multnomah Co., Portland, (ANSP:1);

WASHINGTON: Grays Harbor Co., Montesano (NMDC:1), Pierce Co., Puyallup

(NMDC:1). CANADA: BRITISH COLUMBIA: 12 mi E. Hope (CNCI:1).

Phyllotreta zimmermanni (Crotch)

(Figs. 38-39, 83-84, 127-128, 149, 171)

Orchestris zimmermanni Crotch, 1873. Proc. Acad. Nat. Sci. Philadelphia,

25:66.

Neotype (here designated): Male, deposited at USNM.

Type locality: Missouri; neotype, Devils Elbow, Phelps Co.,

Missouri.

Phyllotreta zimmermanni, Riley, 1885. Ann. Rep. Dept. Agr. for 1884,

Rep. Entomologist, p.304.

Phyllotreta sibirica Weise, 1887. Archiv fur Naturgeschichte, 53:200.

(Synonymy from Heikertinger and Csiki, 1939:59)

Phyllotreta sinuata Horn, 1889. Trans. American Ent. Soc., 16:295,

table VI, fig. 15. (not Stephens, not Redtenbacher) (Synonymy

by Heikertinger, 1911:12) 144

DIAGNOSIS: Elytra each with a median pale stripe with dilations which never meet at suture; male antennal segment S dark, expanded bilaterally and dorsoventrally flattened, much wider than 4, with a

distinct concavity in basal 1/2 anteriorly; apex of male genitalia

in dorsal view with a narrow deep median emargination, moderately

curved ventrad in lateral view with a ventral subapical concavity.

/ DESCRIPTION: Neotype (variation, excluding punctation, in paren­

theses): Fig. 38; elongate oval, length 2.75 mm (male 2.18-2.78,

female 2.52-3.10), width 1.28 mm (male 1.00-1.30, female 1.18-1.50);

head and pronotum black with slight metallic luster, elytra brownish

black to black with slight metallic luster, each with a median straw-

yellow stripe. Head: Black; texture finely granulate basally to

slightly roughened anteriorly (usually entirely finely granulate);

moderately punctate, punctures separated by less than 1/2 to equal

their diameter, mostly by less than one diameter; Interocular Distance/

Maximum Diameter of eye, 1.56 (male 1.56-2.00, female 1.50-1.62).

Antennae: Figs. 83, 84; segments 4 and 7 subequal in length, 4 longer

than 6, segment 5 more than 3x length of 6 and more than 2x length of

4, 5 distinctly wider than 4; segments 4 and 5 strongly expanded

bilaterally, dorsoventrally flattened; segment 5 distinctly longest,

venter with a distinct moderately deep concavity in basal 1/2 anteriorly

or to outside; segment Clength/width]: lClO.5/4], 2C5-/3-3, 3C5-/3.5),

4C4/5D, 5C10/7-D, 6C2.5/2.43, 7C4.S/3], 8C5/3.S], 9C5/43, 10C5/43,

11E6.5/4], total length 1.58 mm (female: 1C9.5/3], 2[4.5/2.5], 3C4/2+],

4C4/2+3, 5C6/2+), 6C3.5/2], 7[5/3], 8[5/3], 9C4.5/3], 10C4.S/3),

11C7/3+], total length 1.44 mm; simple); antennae various shades of 145 brown, basal 3 (4) segments paler with dorsum of 1 dark, (4 intermediate),

4-11 (5-11) darkest. Pronotum: Length 0.50 mm (male 0.40-0.50, female

0.40-0.58), width 0.90 mm (male 0.70-0.90, female 0.80-0.95); black;

texture finely granulate; coarsely punctate, punctures separated by

less than 1/2 to equal their diameter, mostly by less than 1/2 a

diameter. Elytra: Length 2.00 mm (male 1.55-2.05, female 1.85-2.22), width 1.28 mm (male 1.00-1.30, female 1.18-1.50); brownish black to black (brownish black or black, rarely dark brown), median stripe

pattern as in Fig. 38 (sometimes as in Fig. 39); texture slightly

roughened (often smooth)coarsely punctate with very coarse punctures

basally between stripes, punctures separated by less than 1/2 a diameter.

Legs: Normal color sequence. Abdomen: 5th sternum with an apical

median moderately concave lobe, concavity tapering in its extension to

midpoint (extending to within 1/3 of sternal base), with a median linear

impression extending from sternal base to base of median lobe (extending

to apex, sometimes interrupted) (female: simple in outline, sometimes

with a shallow oblong preapical impression); black (brownish black)

with median lobe paler (apical 1/2 of 5th paler or entirely dark).

Male genitalia: Figs. 127, 128; length approximately 0.88 mm;

moderately arched dorsoventrally with a slight submedian ventrally

directed bend in lateral view, of rather uniform width except slightly

narrower in apical 1/4; apex in dorsal view bluntly rounded with a short

median extension, with a median narrow moderately deep emargination, in

lateral view broadly tapering to an abruptly narrower, rounded tip which

is directed ventrad, with a moderately deep subapical ventral concavity. 146

Female genitalia: Fig. 149; length approximately 0.42 mm; recep­ tacle elongate, of rather uniform width, with a submedian dorsoventral bend, ring collar strongly developed.

BIOLOGY: Host plants, adults (the following represent those records verified by me via label data): *Arabis, alfalfa, *Brassica,

•cabbage, clover, Fragaria, hickory (feeding on), *horse-radish, / •Lepidium virginicum, *Lepidium, pepper (feeding on), *pepper-grass, wild plum, potato vine, *Rorippa palustris, *Radicola, *radish, straw­ berry, tobacco, *white turnip, and *water-cress. Duporte (1914; as

P. sinuata) adds *tumip; Duckett (1920; as P^. sinuata) adds *mustard;

Tahavanainen (1972) records in addition, *Barbarae vulgaris, *Brassica

oleracea acephala, and *Dentaria diphylla; Hicks (1972) adds *Brassica

nigra and Lepidium compastre; and Balsbaugh (1972) records Trifolium

inca matum.

Host plants, larva: Wild pepper-grass (Lepidium virginicum) and

Arabis ludoviciana (Riley, 1885b:304-305); cress (Duport, 1914:433;

as sinuata).

Immature stages: Egg, larva and pupa described by Riley (1885b:

305-306; pi. IV, fig. 1) with the larva and pupa illustrated; Duporte

(1914:433; as P. sinuata) also illustrates the larva.

Habits: The larvae are leaf miners, preferring Lepidium virginicum

and sparingly attacking Arabis ludoviciana, and are not confined to a

single mine or even one leaf (Riley, 1885b:304-305). The adults over­

winter (Hicks, 1972) and become active in early spring. Eggs are laid

on the upper leaf surface along the midvein (Riley, 1885); however, 147

Hicks (1972) says that "P^. zimmermanni lay their eggs in the leaf tissues." Pupation takes place in an earthen cell just beneath the soil surface (Riley, 1885b). According to Riley (1885b) in May and

June the entire life cycle may be completed in 20-23 days in Missouri.

This species has been a pest in past years but it has apparently been displaced for the most part by the more competitive introduced species P. cruciferae (Goeze) (Hicks, 1972) which was apparently introduced from Europe in the early 1920*s on the West Coast and probably again around 1940 on the East Coast (Milliron, 1953).

DISCUSSION: Distribution: Fig. 171; P. zimmermanni is widespread but has been more frequently collected in the northeastern and mid- western U.S.

Nomenclature: Crotch (1873) described P. zimmermanni in the genus

Orchestris; this unfortunate choice of generic name was corrected by

LeConte (1878:615). In 1885, Riley transferred this species to

Phyllotreta. Heikertinger and Csiki (1939) listed P^. sibirica Weise

as a synonym but I haven't seen the type to verify this. Horn (1889)

thought that Crotch had failed to realize that his new species was the

common European JP. sinuata Stephens, but Heikertinger (1911) recognized

that'P. zimmermanni was native to North America and a distinct species,

whereas Stephens' Haltica sinuata was native to Europe and Asia.

A search for the type of P. zimmermanni has proven unsuccessful.

Crotch gives the type locality and collector as "Missouri(Riley)'' and

it was apparently a male because the diagnostic male antennal character

is given. The specimen labeled as the type at the MCZC is a female 148 labeled "NX." and there is no indication that Riley was the collector; also a search turned up no other P_. zimmermarini material. Since it is well known that Crotch and Horn freely exchanged material, a search, was requested at the ANSP; no such specimen was found. Finally, since the Riley collection is at the USNM, a search was also requested there; again, no such specimen was found. These unsuccessful investigations lead to the conclusion that the type of P^. zimmermanni is most probably lost. Therefore, a neotype has been designated from a long series which

I collected at Devils Elbow, Missouri, a site which is part of the original type locality. This specimen also agrees with Crotch*s original description and is a male.

SPECIMENS EXAMINED: Total 1258: ALASKA: Matanuska (USNM:1);

ARKANSAS: Crawford C o . (UADE:3); ARIZONA CDEFW:l3; CALIFORNIA: Los

Angeles Co., Los Angeles (LACM:1), San Mateo Co . (LACM:2), Stanislaus

Co., Modesto (DEFW:1); COLORADO: Douglas Co., Larkspur (LACM:2),

Mineral Co., Creede (ISUI:3); CONNECTICUT: Fairfield Co., Norwalk

(USNM:2), Litchfield Co., Cornwall (CUIC:1), Kent (MCZC:4), New Haven

Co., New Haven (PADA:1), Windham Co., Nth Windham (MCZC:1); DISTRICT

OF COLUMBIA: Washington (CNCI:1, MCZC:2, USNM:50); GEORGIA: Rabun Co .,

Pine Mountain (CNCI:1), (ANSP:l3; ILLINOIS: Champaign Co., Urbana (ISUI:1),

Cook Co., Chicago (ANSP: 5, USNM:3, UWEM:2), Homewood (FMNH:4), Gallatin

Co., Bowmanville (FMNH:3), Lake Co., Ft. Sheridan (FMNH:2), McHenry Co.,

McHenry (WSUC:2), Union Co. (USNM:4), (DEFW:2, WSUC:33; INDIANA: Knox

Co., Vincennes (CNCI:1, USNM:1), Lake Co., Longlake (FNMH:10), Marion

Co. (CUIC:3), Pulaski Co. (NMDC:1), Tippecanoe Co., Lafayette (CNCI:9, 149

NMDC:1), (FMNH:4, MSUC:13]; IOWA: Appanoose Co. (USNM:1), Boone Co.,

Ledges State Park (ISUI:1), Davis_Co. (USNM:!), Hancock Co. (ISUI:1),

Jasper Co., Colfax (ISUI:1), Johnson Co., Iowa City (MCZC:2, LACM:1,

USNM:1), Howard Co., Cresco (ISUI:1), Lee Co., Montrose (ISUI:13), Linn

Co., Palisades-Kepler St. Pk. (ISUI:1), Madison Co., Pammel State Park

(ISUI:1), Page Co., Clarinda (ISUI:2), Shenandoah (ISUI:2), Scott Co.,

Pleasant Val. (CNCI:2), Story Co., Ames (ISUI:38, LACM:2), Webster Co.,

Dolliver Mem. St. Pk. (ISUI:1), Winnebago Co., Forest City (ISUI:2),

(DEFW:2, ISUI:16, MCZC:2]; KANSAS: Douglas Co., Baldwin (JSCC:1),

Lawrence (CNCI:31, EMUS:1), (LACM:1), Franklin Co., Ottawa (EMUS:2),

Riley Co. (USNM:2), Shawnee Co., Topeka (USNM:1), [CNCI:!]; KENTUCKY:

Christian Co., 5 mi. W. Hopkinsville (CNCI:1), Henderson Co., Henderson

(LACM:1), Rowan Co., Morehead (0SUC:2); LOUISIANA: Tangipahoa Par.,

Hammond (USNM:1); MARYLAND: Montgomery Co., Glen Echo (USNM:39),

Prince George1s Co., Bladensburg (UMDC:4, USNM:1), College Park (UMDC:3,

USNM:1), (UMDC:193; MASSACHUSETTS: Bristol Co., Fall River (MCZC:1),

Hampden Co., Springfield (MCZC:1), Middlesex Co., Arlington (MCZC:2),

Cambridge (MSUC:1), Framingham (MCZC:1), Lincoln (MCZC:2), Sherborn

(MCZC:1), Stoneham (CUIC:10), Suffolk Co., Dorchester (MSUC:2),

Roxbury (MCZC:3), CCUIC:13; MICHIGAN: Allegan Co ., Fennville (MSUC:4),

Bay C o .', (MSUC:2), Berrien Co., Galieu (MSUC: 18), Ingham Co., Okemos

(MSUC:13), Lapeer Co., Deerfield twp. (UCRC:1), Kalamazoo Co., Gull

Lake (MSUC:9), Midland Co. (MSUC:3), Monroe Co., Monroe (MSUC:1),

Oakland Co., Orion Lake (FMNH:1, MSUC:1), (MSUC:13]; MINNESOTA: Aitkin

Co., Tamarack (DEFW:4), Anoka Co. (DEFW:13), Benton Co. (DEFW:1), 150

Crow Wing Co. (DEFW:26), Pekota Co. (DEFW:1), Hennepin Co. (DEFW:2),

Houston Co., Mississippi Bluff (DEFW:21), Winnebago Creek (DEFW:14),

Kandiyohi Co. (DEFW:7), LeSueur Co., LeSueur, Minnesota River (DEFW:1),

Mille Lacs Co., Mille Lacs (DEFW:40), Mower Co., Austin (DEFW:2),

Polk Co., Crookston (DEFW:1), Ramsey Co., St. Paul, University Farm

(DEFW:2), St. Paul (DEFW:1), Rice Co., Fairbault (CNCI:1), Steel C o.,

Owatonna (CNCI:3, USNM:1), Wabasha Co., Lake City (CNCI:1, DEFW:6),

Washington Co. (DEFW:8), CDEFW:7, UCRC:1]; MISSOURI: Greene Co.,

Willard (ANSP:3), Montgomery City (CUIC:2), Phelps Co., Devils Elbow

(EHSC:39; Neotype:USNM), St. Louis Co., Ranken (JSCC:3), St. Louis

(USNM:1), [ANPS:8, DEFW.-13); MISSISSIPPI: Adams Co., Natchez (USNM:1);

NEBRASKA: Cuming Co., West Pt. (DEUN:1, USNM:1), Douglas Co., Omaha

(DEUN:3), Lancaster Co., Lincoln (DEUN:26), Otoe Co., Nebraska C'y

(DEUN:6); NORTH CAROLINA: Columbus Co., Chadbourn (USNM:3), Macon Co.,

Highlands (CNCI:3), Wake Co., Raleigh (JSCC:2, NCSU:19), Wilkes Co.,

Wilkesbora (NSCU:1), CMCZC.-2, USNM:3); NORTH DAKOTA: Grand Forks Co.,

University (USNM:2), Ransom Co., Lisbon (DEFW:1); NEW JERSEY: Bergen Co.,

Palisades (FMNH:1), Ridgewood (CUIC:7), Camden Co., Haddon Hts (CNCI:2),

Glouster Co., Wenonah (MCZC:3), [FMNH:31, SDSU:2]; NEW YORK: Cattaraugus

Co., Eden (CUIC:1), Monroe Co., Rochester (CUIC:1, LACM:2), Nassau Co.,

Flushing, L.I. (CNCI:5), Niagara Co., Olcott (CUIC:1), Onondaga Co.,

Elbridge (NMDC:2), Orange Co., West Point (USNM:18), Queens Co., Sea

Cliff, L.I. (FMNH:1), Tompkins Co ., Freeville (CUIC:1), Ithaca (CUIC:2S),

Van Natta*s Dam (CUIC:1), Westchester Co., Peekskill (PSUC:3) CAMNH:1,

CUIC:1, FMNH:1, MCZC:1, USNM:2); OHIO: Franklin Co., Columbus (0SUC:6), 151

Hancock C o . (0SUC:1), Logan Co. (DEFW:1); PENNSYLVANIA: Adams C o .,

Arendtsville (CUIC:5, PSUC:!), Allegheny Co., Pittsburgh (ICCM:7),

Armstrong Co., Lane (PSUC:2), Dauphin Co., Clarks Valley (EUBC:3),

Harrisburg (PADA:7)j 7 mi. E o£ Holg. (PSUC:1), Manada Hill (PADA:7),

Rockville (PSUC:1), Centre Co., Pine Grove Mills (PSUC:4), State

College (PSUC:21), Franklin Co., Chambersburg (PADA:1), Lehigh Co.,

Lehigh Water Gap (PADA:1), Montgomery C o ., Abington (MCZC:3),

Philadelphia C o ., Angora (MCZC:3), Philadelphia (FMNH:6), Snyder C o .,

Mt. Pleasant Mills (EUBC:1), Westmoreland C o ., Jeannette (ICCMrll,

PADA:2), CLACM:2, MCZC:41, OSUC:10, PADA:5}; SOUTH CAROLINA: [CNCI:5];

SOUTH DAKOTA: Beadle C o ., Huron (SDSU:2), Brookings Co., Brookings

(SDSU:4), Volga (MSUC:2, SDSU:12), Brown Co., Aberdeen (SDSU:3),

Browns Valley (SDSU:3), Hecla (SDSU:1), Brule Co., Chamberlain (SDSU:2),

Buffalo C o ., Ft. Thompson (SDSU:3), Fall River C o ., Hot Springs

(SDSU:5), Jackson C o ., Cottonwood (SDSU:23), Pennington Co.,Rapid City

(SDSU;1), Stanley C o., Pierre (SDSU:1), Union Co., Alcaster (SDSU:1),

Canton (SDSU:1), Elk Point (SDSU:3), CSDSU:43; TENNESSEE: Shelby C o .,

Memphis (MCZC:1), (MCZC:1); VIRGINIA: Fairfax Co., Arlington (USNM:1),

Loudoun Co. (USNM:5), [DEFW:1, USNM:I]; WISCONSIN: Brown Co., Green

Bay (USNM:1), Dane Co., Madison (CUIC:1, MCZC:3, 0SU0:1, UWEM:9),

Milwaukee Co., Milwaukee (UWEM:2), Racine C o ., Racine (CUIC:1, MCZC:1,

UWEM:3), Waupaca Co ., Waupaca (MCZC:1, UWEM:6); WYOMING: Yellowstone

Pk. (CNCI:3). CANADA: ALBERTA: Alabaster (FMNH:1), Edmonton (ANSP:2);

MANITOBA: Bodin (MCZC:14), Mile 214 Hudson Bay R.R. (MCZC:1), Rosebark 152

(CNCX:1); NORTHWEST TERRITORY: Aklavik (CNCI:2), Ft. McPherson (CNCI:6),

Yellow Knife (CNCI:4); ONTARIO: Aldershot (CNCI:8), Britannia (CNCI:2),

Bradleys (CNCI:1), Bothwell (CNCI:1), Chatham (CNCI:17), Eberts

(CNCI:1), Kingsville (CNCI:4), Leamington (CNCI:1), Manotick (CNCI:2),

Merivale (CNCI:1), Ottawa (CNCI:4), Pelee Island (CNCI:7), Pr. Edw.

Co. (JSCC: 2), Til bury e (CNCI: 3), Toch River (CNCI: 1,) , Toronto (CUIC:1)

QUEBEC: Ilede Montreal (CNCI:1), Montreal (DEFW:1, FMNH:5, MCZC:1);

YUKON TERRITORY: mi. 1192 Alaska Hwy, Nr. Snag. Juct. (CNCI:2).

NO DATA: CCNCI:3, DEFW:1, ISUI:1, PSUC:3, USNM:2, WSUC:2). SPECIES CONSIDERED NOMINA. DUBIA

PhyllQtreta alberta Chttn.

Phyllotreta alberta Chittenden, 1927. Entomologica Americana, 8(1):28. / Holotype: Female, USNM type #28791.

Type locality: Edmonton, Alberta, Canada.

Contrary to Chittenden (1927), this species is known only from a series of females. This species is most similar to P_: zimmermanni.

The following are illustrated: beetle (Fig. 172), antenna (Fig. 175), and spermatheca (Fig. 178).

Phyllotreta obtusa Chttn.

Phyllotreta obtusa Chittenden, 1927. Entomologica Americana, 8(1):30.

Holotype: Female, USNM type #28793

Type locality: Breckenridge, Colorado. —

Contrary to Chittenden (1927), the unique type is a female. This species is most similar to P. zimmermanni. The following are illustra­ ted: beetle (Fig. 173), antenna (Fig, 176), and spermatheca (Fig. 179).

Phyllotreta perspicua Chttn.

Phyllotreta perspicua Chittenden, 1927. Entomologica Americana, 8(1):29.

153 Holotype: Female^ USNM type #28812.

Type locality: Klamath Lake, Oregon.

Contrary to Chittenden (1927), the unique type is a female. This species is most similar to ;P^. zimmermanni. The following are illustrated: beetle (Fig. 174), antenna (Fig. 177), and spermatheca

(Fig. 180). 15S

Figure 1. Male 5th abdominal sternum and pygidium, ventral view.

Figure 2. Female 5th abdominal sternum, ventral view.

Figure 3. Label of lectotype of Phyllotreta armoraciae.

Figure 4. Male genitalia with structures labeled, dorsal view.

Figure 5. Male genitalia with structures labeled, lateral view.

Figure 6. Female spermatheca with structures labeled. 156

median lab*

median linear tm preii Sort

median longitudinal /* im prettion 25

lclerotized ring collar receptacle

pump / » \ 159817^241494^39

ledian lob* non-tdtro t i led • permathecol duct

•clerati xed spermat hacal duet

gland 157

Figure 7. Phyllotreta arcuata, holotype, dorsal.

Figure 8. Phyllotreta armoraciae, dorsal.

Figure 9. Phyllotreta attenuata, holotype, dorsal view.

Figure 10. Phyllotreta bipustulata, dorsal view.

•Figure 11. Phyllotreta bipustulata, right elytron showing color pattern variation, dorsal view. 8

L 159

Figure 12. Phyllotreta bisinuata, holotype, dorsal view.

Figure 13. Phyllotreta constricta, holotype, dorsal.

Figure 14. Phyllotreta decipiens, dorsal view.

Figure 15. Phyllotreta denticomis, dorsal view. ,

Figure 16. Phyllotreta denticomis, right elytron showing color pattern variation, dorsal view.

161

Figure 17. Phyilotreta dolichophalla, holotype, dorsal view.

Figure 18. Phyilotreta emarginata, holotype, dorsal view.

Figure 19. Phyilotreta lepidula, dorsal view.

Figure 20. Phyilotreta liebecki, holotype, dorsal ,view.

163

Figure 21. Phyllotreta oblonga, holotype, dorsal view.

Figure 22. Phyllotreta oregonensis, lectotype, dorsal view.

Figure 23. Phyllotreta oregonensis, right elytron showing color pattern variation, dorsal view.

Figure 24. Phyllotreta ramosa, dorsal view.

Figure 25. Phyllotreta ramosoides, holotype, dorsal view. n II 165

Figure 26. Phyllotreta robusta, dorsal view.

Figure 27. Phyllotreta robusta, right elytron showing color pattern variation, dorsal view.

Figure 28. Phyllotreta spatulata, holotype, dorsal view.

Figure 29. Phyllotreta striolata, dorsal view.

Figure 30. Phyllotreta striolata, right elytron showing color pattern variation, dorsal view.

Figure 31. Phyllotreta striolata, right elytron showing color pattern variation, dorsal view.

Figure 32. Phyllotreta striolata, right elytron showing color pattern variation, dorsal view.

167

Figure 33. Phyllotreta striolata, right elytron showing color pattern variation, dorsal view.

Figure 34. Phyllotreta striolata, right elytron showing color pattern variation, dorsal view.

Figure 35. Phyllotreta striolata, right elytron showing color pattern variation, dorsal view.

Figure 36. Phyllotreta utana, dorsal view.

Figure 37. Phyllotreta utanula, holotype, dorsal view.

Figure 38. Phyllotreta ziiranermanpi, dorsal view.

Figure 39. Phyllotreta zimmermanni, right elytron showing color pattern variation, dorsal view. I

35 169

Figure 40. Male antenna of Phyllotreta arcuata, holotype, . anterodorsal view of right.

Figure 41. Female antenna of Phyllotreta arcuata, allotype, anterodorsal view of right.

Figure 42. Male antenna of Phyllotreta armoraciae, anterodorsal view of right.

Figure 43. Female antenna of Phyllotreta armoraciae, anterodorsal view of right.

Figure 44. Male antenna of Phyllotreta attenuata, holotype, anterodorsal view of right.

Figure 45. Female antenna of Phyllotreta attenuata, allotype, anterodorsal view of right.

Figure 46. Male antenna of Phyllotreta bipustulata, anterodorsal view of right.

Figure 47. Female antenna of Phyllotreta biputulata, anterodorsal view of right.

Figure 48. Male antenna of Phyllotreta bisinuata, holotype, anterodorsal view of right.

Figure 49. Female antenna of Phyllotreta bisinuata, allotype, anterodorsal view of right.

Figure 50. Male antenna of Phyllotreta constricta, holotype, anterodorsal view of right. 170

40 41 4 2 4 3 44 4 5

46 47 48 49 50 51 171

Figure SI. Female antenna of Phyllotreta constricta, allotype, anterodorsal view of right.

Figure 52. Male antenna of Phyllotreta decipiens, anterodorsal view of right.

Figure 53. Female antenna of Phyllotreta decipiens, anterodorsal view of right.

Figure 54. Male antenna of Phyllotreta denticomis, anterodorsal view of right.

Figure 55. Male antennal segments 5-8 of Phyllotreta denticomis, anterior view of right.

Figure 56. Female antenna of Phyllotreta denticornis, anterodorsal view of right.

Figure 57. Male antenna of Phyllotreta do1ichophalla, holotype, anterodorsal view of right.

Figure 58. Female antenna of Phyllotreta dolichophalla, allotype, anterodorsal view of left-

Figure 59. Male antenna of Phyllotreta emarginata, holotype, anterodorsal view of right.

Figure 60. Female antenna of Phyllotreta emarginata, allotype, anterodorsal view of right.

Figure 61. Male antenna of Phyllotreta lepidula, anterodorsal view of right.

Figure 62. Female antenna of Phyllotreta lepidula, anterodorsal view of right. 172 173

Figure 63. Male antenna of Phyllotreta liebecki, holotype, anterodorsal view of right.

Figure 64. Male antennal segments 4-6 of Phyllotreta liebecki, holotype, anterior view of right.

Figure 65. Female antenna of Phyllotreta liebecki, anterodorsal view of right.

Figure 66. Male antenna of Phyllotreta oblonga, anterodorsal view of right.

Figure 67. Female antenna of Phyllotreta oblonga, holotype, anterodorsal view of right.

Figure 68. Male antenna of Phyllotreta oregonensis, anterodorsal view of right.

Figure 69. Female antenna of Phyllotreta oregonensis, anterodorsal view of right.

Figure 70. Male antenna of Phyllotreta ramosa, anterodorsal view of right.

Figure 71. Female antenna of Phyllotreta ramosa, anterodorsal view of right.

Figure 72. Male antenna of Phyllotreta ramosoides, holotype, anterodorsal view of right.

Figure 73. Female antenna of Phyllotreta ramosoides, allotype, anterodorsal view of right. 174 175

Figure 74. Male antenna of Phyllotreta robusta, anterodorsal view of right.

Figure 75. Female antenna of Phyllotreta robusta, anterodorsal view of right.

Figure 76. Male antenna of Phyllotreta spatulata, holotype, anterodorsal view of right.

Figure 77. Female antenna of Phyllotreta spatulata, allotype, anterodorsal view of right.

Figure 78. Male antenna of Phyllotreta striolata, anterodorsal view of right.

Figure 79. Female antenna of Phyllotreta striolata, anterodorsal view of right.

Figure 80. Male antenna of Phyllotreta utana, paratype, anterodorsal view of right.

Figure 81. Female antenna of Phyllotreta utana, paratype, anterodorsal view of right.

Figure 82. Male antenna of Phyllotreta utanula, holotype, anterodorsal view of right.

Figure 83. Male antenna of Phyllotreta zimmermanni, anterodorsal view of right.

Figure 84. Female antenna of Phyllotreta zimmermanni, anterodorsal view of right. 176

74 75 7 6 7 7 7 8 79

80 81 82 83 84 Figure 85. Male genitalia o£ Phyllotreta arcuata, paratype, dorsal view.

Figure 86. Male genitalia of Phyllotreta arcuata, paratype, lateral view.

Figure 87. Male genitalia of Phyllotreta armoraciae, dorsal view.

Figure 88. Male genitalia of Phyllotreta armoraciae, lateral view.

Figure 89. Male genitalia of Phyllotreta attenuata, holotype, dorsal view.

Figure 90. Male genitalia of Phyllotreta attenuata, holotype, lateral view.

Figure 91. Male genitalia of Phyllotreta bipustulata, dorsal view.

Figure 92. Male genitalia of Phyllotreta bipustulata, lateral view.

Figure 93. Male genitalia of Phyllotreta bisinuata, paratype, dorsal view.

Figure 94. Male genitalia of Phyllotreta bisinuata, paratype, lateral view.

Figure 95. Male genitalia of Phyllotreta constricta, paratype dorsal view.

Figure 96. Male genitalia of Phyllotreta constricta, paratype lateral view. 178

\ ! % \

8 86

87

8 9

233248

#-* \

91 92 93 94 95 96 Figure 97. Male genitalia of Phyllotreta decipiens, dorsal view.

Figure 98. Male genitalia of Phyllotreta decipiens, lateral view.

Figure 99. Male genitalia of Phyllotreta denticomis, dorsal view.

Figure 100. Male genitalia of Phyllotreta denticomis, lateral view.

Figure 101. Male genitalia of Phyllotreta dolichophalla, holotype, dorsal view.

Figure 102. Male genitalia of Phyllotreta dolichophalla, holotype, lateral view.

Figure 103. Male genitalia of Phyllotreta emarginata, paratype, dorsal view.

Figure 104. Male genitalia of Phyllotreta emarginata, paratype, lateral view.

Figure 105. Male genitalia of Phyllotreta lepidula, dorsal view.

Figure 106. Male genitalia of Phyllotreta lepidula, lateral view.

Figure 107. Male genitalia of Phyllotreta liebecki, dorsal view.

Figure 108. Male genitalia of Phyllotreta liebecki, lateral view. 180

> V- -✓ 'V / «'■

97 98 99 100

1 0 1 102

il

103 104 105 106 107 108 Figure 109. Male genitalia of Phyllotreta oblonga, dorsal view.

Figure 110. Male genitalia of Phyllotreta oblonga, lateral view.

Figure 111. Male genitalia of Phyllotreta oregonensis dorsal view.

Figure 112. Male genitalia of Phyllotreta oregonensis lateral view.

Figure 113. Male genitalia of Phyllotreta ramosa, dorsal view.

Figure 114. Male genitalia of Phyllotreta ramosa, lateral view.

Figure 115. Male genitalia of Phyllotreta ramosoides, holotype, dorsal view.

Figure 116. Male genitalia of Phyllotreta ramosoides, holotype, dorsal view.

Figure 117. Male genitalia of Phyllotreta robusta, dorsal view.

Figure 118. Male genitalia of Phyllotreta robusta, lateral view.

Figure 119. Male genitalia of Phyllotreta spatulata, holotype, dorsal view.

Figure 120, Male genitalia of Phyllotreta spatulata, holotype, lateral view. 182 1 ir

18759166678^939186

109 110 1 112 113 114

\ I

7 115 116 117 118 119 120 Figure 121. Male genitalia of Phyllotreta striolata, dorsal view.

Figure 122. Male genitalia of Phyllotreta striolata, lateral view.

Figure 123. Male genitalia of Phyllotreta utana, dorsal view.

Figure 124. Male genitalia of Phyllotreta utana, lateral view.

Figure 125. Male genitalia of Phyllotreta utanula, holotype, dorsal view.

Figure 126. Male genitalia of Phyllotreta utanula, holotype, lateral view.

Figure 127. Male genitalia of Phyllotreta zimmermanni dorsal view.

Figure 128. Male genitalia of Phyllotreta zimmermanni lateral view. N

_---

K> 124 123 N 185

Figure 129. Female spermatheca of Phyllotreta arcuata, paratype, lateral view.

Figure 130. Female spermatheca of Phyllotreta armoraciae, lateral view.

Figure 131. Female spermatheca of Phyllotreta attenuata, allotype, lateral view.

Figure 132. Female spermatheca of Phyllotreta bipustulata, lateral view.

Figure 133. Female spermatheca of Phyllotreta bisinuata, allotype, lateral view.

Figure 134. Female spermatheca of Phyllotreta constricta, paratype, lateral view.

Figure 135. Female spermatheca of Phyllotreta decipiens, lateral view.

Figure 136. Female spermatheca of Phyllotreta denticomis, lateral view.

Figure 137. Female spermatheca of Phyllotreta dolichophalla, allotype, lateral view.

Figure 138. Female spermatheca of Phyllotreta emarginata, paratype, lateral view. 186

129 130

131 132

133 134

135 136

137 138 Figure 139. Female spermatheca of Phyllotreta lepidula, lateral view.

Figure 140. Female spermatheca of Phyllotreta liebecki, lateral view.

Figure 141. Female spermatheca of Phyllotreta oblonga, holotype, lateral view.

Figure 142. Female spermatheca of Phyllotreta oregonensis, lateral view.

Figure 143. Female spermatheca of Phyllotreta rainosa, lateral view.

Figure 144. Female spermatheca of Phyllotreta ramosoides, paratype, lateral view.

Figure 145. Female spermatheca of Phyllotreta robusta, lateral view.

Figure 146. Female spermatheca of Phyllotreta spatulata, paratype, lateral view.

Figure 147. Female spermatheca of Phyllotreta striolata, lateral view.

Figure 148. Female spermatheca of Phyllotreta utana, lateral view.

Figure 149. Female spermatheca of Phyllotreta zimmermanni, lateral view. 188

140 139

142

143 144

145 146

147

149 189

(30 170

o\

y -

120 I VO to

Figure 150. Distribution of Phyllotreta arcuata 6coleofMtI«

N 00 70 70

* -

I0U m

do n 120 MO

Figure 152. Distribution of Phyllotreta attenuata 192 70

£ o too ec<

A A ari V- /V

00 120 ICfl K T0

Figure 153. Distribution of Phyllotreta bipustulata 193 70 00 70

BO 170

120

Figure 154. Distribution of Phyllotreta bisinuata. 194

ficaled Mites

Figure 155. Distribution of Phyllotreta constricta. 195 70 7D

[170 CCi

no 110 100 to 7 0

Figure 156. Distribution of Phyllotreta decipiens. 196

Figure 157. Distribution o£ Phyllotreta denticomis. 70 d o 70

I/O

*> -

no m fla 10

Figure 158. Distribution of Phyllotreta dolichophalla 198 70 70

00 [170 tas ec,

&

30

9 -

120 70

Figure 159. Distribution of Phyllotreta emarginata. 199 70 eo 7 0

(70 too

30

• in n

u« 10 0 3

Figure 160. Distribution of Phyllotreta lepidula 200

70 BO 70

1 * ** -*1 B3I rro

X-.

too »0 *0

Figure 161. Distribution of Phyllotreta liebecki. 70 10

«0 & rco

fit

I1Q ICO BO

Figure 162. Distribution of Phyllotreta oblonga. 202 70 00 to 70

170 ‘SLJ®

m 90 n

Figure 163. Distribution of Phyllotreta oregonensis. 7 0

120 too

Figure 164. Distribution of Phyllotreta ramosa. 70 eo 64 a * eo 70

Figure 165. Distribution of Phyllotreta ramosoides. 2 OS 80 70

rtO

*> -

IEO ©o

Figure 166. Distribution of Phyllotreta robusta. 206 70 8 4 60

•c,

120 ttD 79

Figure 167. Distribution of Phyllotreta spatulata. 207 W

170 (00

..c

►— \ J

us I IB m M

Figure 168. Distribution of Phyllotreta striolata. 208 70

170 ■§*!»

30

KQ

Figure 169. Distribution of Phyllotreta utana. 70 2 m

.X

170 ____

no M BO n

Figure 170. Distribution of Phyllotreta utanula. M r t i a (Scale <^Mjiea

Figure 171. Distribution of Phyllotreta ziironermanni 211

Figure 172. Phyllotreta alberta, holotype, dorsal view.

Figure 173. Phyllotreta obtusa, holotype, dorsal view.

Figure 174. Phyllotreta perspicua, holotype, dorsal view

Figure 175. Female antenna of Phyllotreta alberta, holotype, anterodorsal view of left.

Figure 176. Female antenna of Phyllotreta obtusa, holotype, anterodorsal view of left.

Figure 177. Female antenna of Phyllotreta perspicua, holotype, anterodorsal view of right.

Figure 178. Female spermatheca of Phyllotreta alberta, holotype, lateral view.

Figure 179. Female spermatheca of Phyllotreta obtusa, holotype, lateral view.

Female 180. Female spermatheca of Phyllotreta perspicua, holotype, lateral view. 212

174 172 173

178

175 176 177 179

180 213

Figure 181. Phenograms representing color pattern variation of 14 samples of Phyllotreta striolata; information given: state or province, n = sample size, X = mean, V = coefficient of vai'iation. J3 o Ii 3 0 0 u

O 0) o O m <0 V)O V) VV 214 2 ntio n *• II K II

i*. **. « Pi o m „ Pi PiO n O S N C I X > X I C c IX c > e lx > u n , n n u b 10 x

1T 0 X ix c > i nonH o £ 0; PI PI -MO 0; £ ■© o w ■n "» .« o 2 e ix > — pi— o m °* - i w. w m w o ^ Pi e IX > IX e II II II e IX > IX e c lx > lx c N II II ww m 0> (0

T T E x 5 c u U o •o 0 c o i i i 83* O <► x 7 7° ni X o i-■* o « « « C > |x e m o’« m # II fl c lx > I U HII IX> c Ii II J > X I C N II t] IX > IX

o> o pi nn*n n o

i" n i - i i i | « - | i i " “nr i 4 \ CO LITERATURE CITED

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______, and G.A. Samuelson (eds.). Directory of Coleoptera

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215 216

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______' 1845. Xii d*0rbigny, Dictionaire Universel

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______. 1920. A new species of Phyllotreta. J. Washington

Acad. Sci., 10:389-390; 1 fig.

______. 1923. Notes on the distribution and habits of North

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______. 1927. The species of Phyllotreta north of Mexico.

Entomologica Americana, 8(l):l-63; 8 figs. 217

Clavareau, H. 1913. Coleopterorum Catalogus, pars 53, Chrysomelidae:

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Crotch, G.R. 1873. Materials for the study of the Phytophaga of

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______' ' 1874. Descriptions of new species of Coleoptera from

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Duckett, A.B. 1920. Annotated list of Halticini. Univ. Maryland Agr.

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Duporte, M. 1914. The wavy striped flea-beetle. Canadian Ent., 46:

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Essig, E.O. 1926. Insects of western North America. MacMillan Co.,

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Fabricius, J.C. 1775. Systema Entomologiae. Lipsiae. 30+832 p.

______. 1787. Mantissa Insectorum ... 1. Hagniae. 348 p.

______. 1801a. Systema Eleutheratorura. Kiliae. 1, 506 p.

______. 1801b. Systema Eleutheratorum. Kiliae. 2, 687 p.

______. 1803. Index alphabeticus in J.C. Fabricii Systema

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Foudras, A.C.M.E. 1860. Altisides. Ann. Soc. Linn. Lyon, 6(ser.2):

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Frost, S.W. 1924. The leaf mining habit in the Coleoptera, Part I. 218

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