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A systematic study of the leafhopper tribe Hecalini (Homoptera: Cicadellidae: Deltocephalinae) of North America

Ma, Ning, Ph.D.

The Ohio State University, 1989

UMI 300 N. Zeeb Rd. Ann Arbor, MI 48106 A SYSTEMATIC STUDY OF THE LEAFHOPPER TRIBE

HECALINI (HOMOPTERA: CICADELLIDAE: DELTOCEPHALINAE)

OF NORTH AMERICA

DISSERTATION

Presented in the Partial Fulfillment of the

Requirements for the Degree Doctor of

Philosophy in the Graduate School of

the Ohio State University

by

Ning Ma, M. S.

* * * * *

•*

The Ohio State University

1989

Reading Committee: Approved by:

Charles A. Triplehorn

Norman F. Johnson

David J. Horn Advisor Department of Entomology To my wife, Xin

ii ACKNOWLEDGMENTS

I would like to express my appreciation to Dr..

Charles A. Triplehorn, my principle advisor, who was always encouraging, supporting and understanding during my stay at The Ohio State University. I also cherish the memory of the late Dr. D. M. DeLong, my original advisor, who played a key role in choosing this project for me.

OSU faculty, Drs. Norman F. Johnson and David

J. Horn kindly criticized this manuscript. Drs. Paul

H. Freytag, University of Kentucky, and William P.

Morrison, Texas Agricultural Extension Service, have been very helpful with some questions referring to this study. Many graduate students, faculty and research scientists have given tremendous assistance and help on the systematic theory in general and microscope work enabling me to complete this project. These include: Drs. John Wilcox, Paul Cwikla, Shawn Clark, and Richard Miller.

I also thank Paul W. Oman, Oregon State University,

iii K. G. A. Hamilton, Biosystematics Research Institute,

Canada, Robert W. Brooks, University of Kansas, Paul

H. Freytag, University of Kentucky, Raymond J. Gagne, and James Kramer, U.S. National Museum (Natural History),

W. J. Knight, British Museum, Frank W. Mead, Florida

Department of Agriculture, Norman D. Penny, California

Academy of Sciences, and H. Derrick Blocker, Kansas

State University, who kindly loaned the specimens in their care for this study.

A special thanks to my wife, Xin and my family, who are always supportive and understanding about my frequent absence. VITA

May 9, 1955 ------Born - Shenyang City, P.R.

China

1978 ------Graduated, Shenyang Agricultural

University, Shenyang City,

P.R. China

1981 ------M.S., N.W. University of Agriculture,

Shaanxi Province, P.R. China

1981-1983 ------Teaching Associate, Shenyang

Agricultural University,

Shenyang

City, P.R. China

1984-1989 ------Research/Teaching Associate,

Department of Entomology,

Ohio State University

PUBLICATIONS

Chou, I. and N. Ma, 1981. On some new species and new records of Typhlocybinae (Homoptera: Cicadellidae) from China. Entomotaxonomia, 1981, 111(3):191-210,

Ma, N. , 1983. A new species of Tautoneura and a new record from China (Homoptera: Cicadellidae: Typhlocybinae). Entomotaxonomia, 1983, V(2):149-150.

v Ma, N., 1984. A new record of Typhlocybinae from China (Homoptera: Cicadellidae). Entomotaxonomia, 1984, VI(1):157.

Cwikla, P. and N. Ma, 1986. A new species of Cariancha Oman Homoptera: Cicadellidae: Deltocephalinae) with a discussion on the tribal placement of the genus. J. New York Entomol. Soc. 94(1) 115-117.

Freytag, P. and N. Ma, 1987. An unusual new genus and species of Hecalini leafhoppers from Venezuela (Homoptera: Cicadellidae). Ent. News 99(3):153-156.

Ma, N., 1988. A review of the genus Spangbergiella (Homoptera: Cicadellidae: Deltocephalinae). J. Kansas Entomol. Soc. 61(4):000-000.

Ma, N., A revision of the genus Dicyphonia Homoptera: Cicadellidae: Deltocephalinae). In press (JKES).

Ma, N., A revision of the genus Memnonia (Homoptera: Cicadellidae: Deltocephalinae) with a phylogenetic analysis of species. In press (JKES).

Ma, N., Systematic and phylogenetic study of genus Hecalus (Homoptera: Cicadellidae: Deltocephalinae) of North America. In press (JKES).

Ma, N., The cladistic study of of Cicadellidae subfamilies (Homoptera). In manuscript.

FIELDS OF STUDY

Major Field: Systematic Entomology Studies in Leafhopper Systematics

vi TABLE OF CONTENTS

ACKNOWLEDGEMENTS ...... iii

VITA ...... v

LIST OF TABLES ...... x

LIST OF FIGURES ...... xi

INTRODUCTION ...... 1

Taxonomic History of Hecalini ...... 4 Natural History of Hecalini ...... 6 Morphology and Terminology ...... 7 Material and Methods ...... 10

CHAPTER

I. SYSTEMATIC STUDY OF HECALINI ...... 12

Check list of Hecalini ...... 12 Key to genera ...... 19 Phylogeny of Hecalini genera ..... 20

II. SYSTEMATIC STUDY OF GENUS MEMNONIA 26

key to species ...... 31 Phylogeny of Memnonia species .... 52

III. SYSTEMATIC STUDY OF GENUS DICYPHONIA ...... 56

Key to species ...... 59 Phylogeny of Dicyphonia species ... 77

IV. SYSTEMATIC STUDY OF GENUS HECULLUS 82

Key to species ...... 83

V. SYSTEMATIC STUDY OF GENUS NEOHECALUS ...... 87

Key to species ...... 89

VI. SYSTEMATIC STUDY OF GENUS

vii SPANGBERGIELLA ...... 92

Key to species ...... 94 Phylogeny of Soangbergiella species ...... 105

VII. SYSTEMATIC STUDY OF GENUS HECALUS ...... 108

Key to species ...... Ill Phylogeny of Hecalus species ..... 146

LIST OF REFERENCES ...... 151

APPENDIX A: ILLUSTRATIONS ...... 156

viii LIST OF TABLES

TABLES PAGE

1. Characters and their polarized conditions of tribe Hecalini ...... 23

2. Data matrix of characters or character states of tribe Hecalini, the numbers referring to Table 1 ...... 25

3. Characters and their polarization for Memnonia species ...... 51

4. Data Matrix of Memnonia species, numbers refer to characters used in table III. O=plesiomorphic, l=apomorphic ...... 52

5. Characters of Dicyphonia used in the cladistic analysis and their polarized conditions by outgroup comparison. Codes (6* etc.) represent the same characters but different states ...... 79

6. Data matrix of polarized characters or character states of Dicyphonia for cladistic analysis. 1 = apomorphic state, 0 = plesiomorphic state ..... 81

7. Characters used in the phylogenetic analysis of the species of Spangbergiella and their polarized condition ...... 106

8. Data matrix of the species of Spangbergiella. l=apomorphic states, plesiomorphic states ...... 107

9. Characters used in phylogenetic study of Hecalus and their polarized conditions ...... 148

10. Data matrix of the polarized characters of Hecalus. 1 = apomorphic condition, 0 = plesiomorphic conditions ...... 150

ix LIST OF FIGURES

FIGURES PAGE

1-7. Hecalus flavidus. 1) male, dorsal aspect; 2) face, ventral aspect; 3) right forewing, dorsal aspect; 4) pygofer, lateral aspect; 5) first abdominal segment and apodemes, ventral aspect; 6) connective, dorsal aspect; 7) aedeagus, lateral aspect ...... 157

8 . Hypothesized phylogenetic relationship among species of Memnonia Ball, the numbers equal to those of characters in Table 1, 2 and x = reverse condition ...... 159

9-15. Memnonia ornata. 9) male head, pronotum and scutellum, dorsal aspect; 10) male right forewing; 11) connective; 12) pygofer, ventral aspect; 13) apex of aedeagus; 14) aedeagus, lateral aspect; 15) male first abdominal segment ...... 161

16-22. Memnonia albolinea. 16) male right forewing; 17) male head, pronotum and scutellum, dorsal aspect; 18) pygofer, lateral aspect; 19) male first abdominal segment; 20) aedeagus, posterior aspect; 21) aedeagus, lateral aspect; 22) connective. Fig. 23 M. acuta. 23) male head, pronotum and scutellum ...... 163

24-31 . Memnonia consobrina. 24) male genital plate; 25) male right forewing; 26) male head, pronotum and scutellum, dorsal aspect; 27) male first abdominal segment; 28) apex of aedeagus; 29) aedeagus, lateral aspect; 30) connective; 31) pygofer, ventral aspect ..... 165

32-39. Memnonia minuta n. sp. 32) male head, pronotum and scutellum, dorsal

x aspect; 33) male right forewing; 34) apex of aedeagus; 35) aedeagus, lateral aspect; 36) male genital plate; 37) pygofer, ventral aspect; 38) connective; 39) male first visible abdominal segment ...... 167

40-46 Memnonia fraterna. 40) male right forewing; 41) male head, pronotum and scutellum, dorsal aspect; 42) connective; 43) male first visible abdominal segment; 44) aedeagus, posterior aspect; 45) aedeagus, lateral aspect; 46) pygofer, lateral aspect ...... 169

47-54 Memnonia brunnea. 47) male head, pronotum and scutellum; 48) male first two visible abdominal segments; 49) male right forewing; 50) connective; 51) aedeagus, lateral aspect; 52) aedeagus, posterior aspect; 53) pygofer, lateral aspect; 54) 6th abdominal tergite of male...... 171

55-61 Memnonia fenestrella. 55) male head, pronotum and scutellum; 56) male right forewing; 57) male first visible abdominal segment; 58) aedeagus, lateral aspect; 59) aedeagus, posterior aspect; 60) pygofer, lateral aspect; 61) connective ...... 173

62-68 Memnonia snadix. 62) male head, pronotum and scutellum; 63) male right forewing; 64) male first visible abdominal segment; 65) aedeagus, lateral aspect; 66) aedeagus, posterior aspect; 67) pygofer, lateral aspect; 68) connective ...... 175

69-74 Memnonia nigrafasciata sp. n. 69) male head, pronotum and scutellum; 70) male right forewing; 71) aedeagus, posterior aspect; 72) aedeagus, lateral aspect; 73) pygofer, lateral aspect; 74) first abdominal segment and apodemes, ventral aspect .... 177

xi 75-80 Memnonia trinlehorni sp. n. 75) male head, pronotum and scutellum; 76) male forewing, dorsal aspect; 77) pygofer, lateral aspect; 78) first abdominal segment and apodemes; 79) aedeagus, lateral aspect; 80) aedeagus, lateral aspect ...... 179

81-86 Memnonia hainiltoni sp. n. 81) male head, pronotum and scutellum; 82) male forewing, dorsal aspect; 83) aedeagus, posterior aspect; 84) aedeagus, lateral aspect; 85) male first abdominal segment and apodemes; 86) pygofer, lateral aspect .... 181

87 Hypothesized cladogram of Memnonia species. Numbers refer to characters listed in Tables 3, 4 ...... 183

88-100. 88-91 Dicyphonia plana. 88) head, pronotum and scutellum of male, dorsal aspect. 89) aedeagus, posterior aspect. 90) aedeagus, lateral aspect. 91) right forewing of male. 92-95 Dicyphonia plura. 92) head, pronotum and scutellum of male, dorsal aspect. 93) aedeagus, lateral aspect. 94) aedeagus, posterior aspect. 95) pygofer, lateral aspect. 96-100 Dicyphonia ornata. 96) head, pronotum and scutellum of male, dorsal aspect. 97) aedeagus, lateral aspect. 98) aedeagus, posterior aspect. 99) right forewing of male. 100) pygofer, lateral aspect ...... 185

101-112. 101-105 Dicyphonia zamorana. 101) head, pronotum and scutellum of male, dorsal aspect. 102) connective, dorsal aspect. 103) aedeagus, lateral aspect. 104) aedeagus, posterior aspect. 105) pygofer, lateral aspect. 106-112 Dicyphonia nigrita. 106) head, pronotum and scutellum, dorsal aspect. 107) right forewing of male. 108) connective, dorsal aspect. 109) aedeagus, lateral aspect. 110) aedeagus, posterior aspect. Ill) pygofer, ventral aspect. 112) sternal apodemes and first abdominal

xii segment 187

113-124. 113 Dicyphonia cona, female, dorsal aspect. 114-119 Dicyphonia delongi., new species. 114) aedeagus, posterior aspect. 115) aedeagus, lateral aspect. 116) head, pronotum and scutellum, dorsal aspect. 117) connective, dorsal aspect. 118) pygofer, ventral aspect. 119) sternal apodemes and first abdominal segment. 120-124 Dicyphonia mexicana. new species. 120) male face, ventral aspect. 121) aedeagus, posterior aspect. 122) aedeagus, lateral aspect. 123) connective, dorsal aspect. 124) head, pronotum and scutellum of male, dorsal aspect ...... 189

125. Hypothesized phylogenetic relationships among the species of Dicyphonia Ball (numbers refer to characters, listed in Tables 4, 5 and x = reverse states) ...... 191

126-133. Hecullus bracteatus. 126) female anterior half body, dorsal aspect; 127) male first abdominal segment and apodemes; 128) male 6th notum of abdomen; 129) aedeagus, posterior aspect; 130) aedeagus, lateral aspect; 131) male head, pronotum and scutellum; 132) left genital plate, ventral aspect; 133) pygofer, lateral aspect ...... 193

134-138. Hecullus balli. 134) female anterior half body, dorsal aspect; 135) aedeagus, posterior aspect; 136) aedeagus, lateral aspect; 137) male first abdominal segment and apodemes, ventral aspect; 138) male 6th notum of abdomen ...... 195

139-144. Neohecalus lineatus. 139) male head, pronotum and scutellum; 140) male forewing, dorsal aspect; 141) pygofer, lateral aspect. 142-144. Neohecalus apicalis. 142) male head, pronotum and scutellum; 143) male forewing, dorsal aspect; 144) pygofer, lateral aspect ...... 197

xiii 145-151. Neohecalus apicalis. 145) connective; 146) male first abdominal segment and . apodemes. 147-151. Neohecalus lineatus. 147) aedeagus, posterior aspect; 148) aedeagus, lateral aspect; 149) connective; 150) style, ventral aspect; 151) male first abdominal segment and apodemes ...... 199

152-153. Neohecalus apicalis. 152) aedeagus, lateral aspect; 153) aedeagus, posterior aspect ...... 201

154-155 Spangbergiella reticulata. 154) left forewing of female; 155) head, pronotum and scutellum of female, dorsal aspect; 156-162. Spangbergiella mexicana. 156) head, pronotum and scutellum of male, dorsal aspect; 157) left forewing of male; 158) left style of male genitalia, dorsal aspect; 159) left lateral aspect of pygofer; 160) aedeagus, left lateral aspect; 161) aedeagus, posterior aspect; 162) right genital plate, ventral aspect ...... 203

163-169 Spangbergiella quadripunctata. 163) head, pronotum and scutellum of male, dorsal aspect; 164) right genital plate, ventral aspect; 165) right forewing of male; 166) left style of genitalia, dorsal aspect; 167) left lateral aspect of pygofer; 168) left aspect of aedeagus; 169) posterior aspect of aedeagus ...... 205

170-176 Spangbergiella erratica♦ 170) head, pronotum and scutellum of male, dorsal aspect; 171) left aspect of pygofer. 172) left style of genitalia, dorsal aspect; 173) left aspect of aedeagus; 174) right forewing of male; 175) ventral aspect of left genital plate; 176) posterior aspect of aedeagus ...... 207

177-184 Spangbergiella vulnerata. 177, 179, 185) head, pronotum and scutellum of male, dorsal aspect; 178) head, pronotum and scutellum of female, dorsal aspect; 180) left lateral

xiv aspect of pygofer. 181) right style of genitalia, dorsal aspect; 182) left lateral aspect of aedeagus; 183) posterior aspect of aedeagus; 184) ventral aspect of left genital plate ...... 209

186 Cladogram for hypothesized relationship among the species of Spangbergiella Signoret, the numbers equal to those of Table 7 and 8 ...... 211

187-193 Hecalus planus. 187) head, pronotum and scutellum of female; 188) head, pronotum and scutellum of male; 189) connective; 190) aedeagus, lateral aspect; 191) aedeagus, posterior aspect; 192) pygofer, lateral aspect; 193) last several abdominal segments of female ...... 213

194-196 Hecalus major. 194) pygofer, lateral aspect; 195) sternum of female 7th abdominal segment; 196) notum of male 7th abdominal segment. 197 Hecalus planus. first male abdominal segment and apodemes ...... 215

198-204 Hecalus kansiensis. 198) aedeagus, posterior aspect; 199) head, pronotum and scutellum of male; 200) aedeagus, lateral aspect; 201) pygofer, lateral aspect; 202) connective; 203) sternum of female 7th abdominal segment; 204) first male abdominal segment and apodemes ...... 217

205-210 Hecalus rotundus. 205) head, pronotum and scutellum of male; 206) pygofer, lateral aspect; 207) aedeagus; lateral aspect; 208) connective; 209) aedeagus, posterior aspect; 210) right forewing of male ...... 219

211-217 Hecalus inflatus. 211) head, pronotum and scutellum, dorsal aspect; 212) connective; 213) pygofer, lateral aspect; 214) aedeagus, lateral aspect; 215) aedeagus, posterior aspect; 216) male right forewing; 217) first abdominal segment and sternal apodemes 221

xv 218-224 Hecalus continuus. 218) head, pronotum and scutellum, dorsal aspect; 219) connective; 220) aedeagus, lateral aspect; 221) aedeagus, posterior aspect; 222) style, dorsal aspect; 223) pygofer, lateral aspect; 224) first abdominal segment and sternal apodemes ...... 223

225-229 Hecalus atascaderus. 225) head, pronotum and scutellum of male; 226) head, pronotum and scutellum of female; 227) sternum of 7th abdominal segment of female; 228) aedeagus, lateral aspect; 229) aedeagus, posterior aspect ...... 225

230-233 Hecalus atascaderus. 230) pygofer, lateral aspect; 231) right forewing of male; 232) connective; 233) male first abdominal segment and apodemes ...... 227

234-237 Hecalus ma.ior. 234) head, pronotum and scutellum of male; 235) face of male, ventral aspect; 236) aedeagus, lateral aspect; 237) aedeagus, posterior aspect ...... 229

238-244 Hecalus viridis. 238) head, pronotum and scutellum of male; 239) face of male, ventral aspect; 240) aedeagus, posterior aspect; 241) aedeagus, lateral aspect; 242) pygofer, lateral aspect; 243) connective; 244) right genital plate ...... 231

245-251 Hecalus hepneri. 245) head, pronotum and scutellum of male; 246) face of male, ventral aspect; 247) pygofer, lateral aspect; 248) aedeagus, lateral aspect; 249) aedeagus, posterior aspect; 250) sternum of female 7th abdominal segment; 251) male first abdominal segment and apodemes ...... 233

252-258 Hecalus curtus. 252) head, pronotum and scutellum of male; 253) face of male, ventral aspect; 254) pygofer, lateral aspect; 255) aedeagus, lateral aspect; 256) aedeagus, posterior

xv i aspect; 257) male first abdominal segment and apodemes; 258) connective ...... 235

259-264 Hecalus flavidus. 259) head, pronotum and scutellum of male;- 260) aedeagus, posterior aspect; 261) aedeagus, lateral aspect; 262) pygofer, lateral aspect; 263) connective; 264) male first abdominal segment and apodemes ...... 237

265-271 Hecalus attenuatus. 265) head, pronotum and scutellum of male; 266) male first abdominal segment and apodemes; 267) pygofer, lateral aspect; 268) connective; 269) aedeagus, lateral aspect; 270) aedeagus, posterior aspect; 271) left style, dorsal aspect ...... 239

272-281 Hecalus grandis. 272) head, pronotum and scutellum of male; 273) head, pronotum and scutellum of female; 274) last several segments of female; 275) connective; 276) first male abdominal segment and apodemes; 277) sternum of 7th female abdominal segment; 278) aedeagus, posterior aspect; 279) aedeagus, lateral aspect; 280) left genital plate, ventral aspect; 281) pygofer, lateral aspect ...... 241

282-285 Hecalus nimbosus. 282, 283, 284) head, pronotum and scutellum of males; 285) face of male, ventral aspect. 286-288 Hecalus nigrafasciatus 286) head, pronotum and scutellum of male; 287) head, pronotum and scutellum of female; 288) sternum of 7th female abdominal segment ...... 243

289-293 Hecalus nigrafasciatus. 289) aedeagus, lateral aspect; 290) aedeagus, posterior aspect; 291) pygofer, lateral aspect; 292) connective; 293) first abdominal segment and apodemes ...... 245

294-297 Hecalus nimbosus. 294) aedeagus,

xvii lateral aspect; 295) aedeagus, posterior aspect; 296) right forewing of male; 297) first male abdominal segment and apodemes ...... 247

298-299 Hecalus elongatus. 298) female, dorsal aspect; 299) sternum of 7th female abdominal segment .... 249

300-301 Hecalus constrictus. 300) female, dorsal aspect; 301) sternum of 7th female abdominal segment .... 251

302-303 Hecalus montanus. 302) female, dorsal aspect; 303) sternum of 7th female abdominal segment .... 253

304 Cladogram of the hypothesized relationship among the species of Hecalus: the numbers refer to those in Tables 9, 10 and the x = reverse conditions ..... 255

Plate I. Screen Electron-Microscope pictures. A,B Hecalus flavidus: A) upper surface of male forewing; B) upper-lateral surface of male pygofer; C,D Hecalus major; C) upper surface of male forewing; D) upper-lateral surface of male pygofer; E,F Memnonia albolinea: E) upper surface of male forewing; F) upper-lateral surface of male pygofer ...... 257

xviii INTRODUCTION

Systematics is an old but fundamentally important branch of biological sciences. It consists basically of two aspects: (1) discovering the nature of the biological world around us, and (2) explaining how these entities arose and what kinds of relationships exist among them. These tasks are the very basis of all biological studies. To take leafhoppers (Insecta:

Homoptera: Cicadellidae) as an example, there are over four thousand recognized leafhopper species in the world, and some people have estimated that the number probably only represents the half of what we have in the nature. Some of these insects are serious pests of our crops, their feeding, oviposition and mechanical damage to crop causes million dollar losses in food production each year in the world. The leafhoppers usually feed on only one or a few species of plants, and the coevolution between these animals and their hosts is important. Leafhoppers also play a major role in the transmission of viral diseases which cause even more damage than feeding alone. Viruses and other pathogenic microorganisms are very specific in their vectors, and again we see the importance of coevolution between them. If the relationships

1 among leafhoppers are better known, we will be better

able to understand what has happened in the past and

to predict what may happen in the future. Therefore,

systematics will continue to play an active role in

today’s scientific endeavors.

Hecalini Distant is probably the most primitive

tribe in the subfamily Deltocephalinae (Homoptera:

Cicadellidae). It consists of 30 genera and more

than 500 species worldwide. They are most common

in tropical regions. Since its establishment in 1908,

there has been a great deal of confusion in the group

at various levels, even in its basic definition.

Only Morrison (1973) and Linnavuori (1975) have attempted

revisionary work in this group. As more species and

genera have been discovered and described, the problems

have become ever more obvious. Additionally, no phylogenetic

studies have been attempted in the Hecalini.

This study has focussed on two aspects of the

systematics of Hecalini. First, some of the problems

associated with the fauna of North America have been

addressed, including the tribal and generic definitions

and limits. Second, the phylogenetic relationships among genera and species have been investigated.

Many conclusions of this study remain preliminary 3 because of limitations of material, but it is my hope that they will help to resolve some of the fundamental problems of the tribe and provide useful information to facilitate further study. 4

Taxonomic History of Hecalini Distant

Hecalini as a group is characterized by the produced and flattened head, sexual dimorphism, long carina along the lateral margins of pronotum and deep sinuation on the genal margins under the eyes. It was established by Distant in 1908 as a division: Hecalusaria. Since its establishment, there has been a great deal of controversy over at which taxonomic level this group of leafhoppers should be placed. Haupt (1929) recognized the group as a subfamily with one tribe, Hecalini.

Lindberg (1948) first considered it a family. However, since Cicadellidae is considered a family by most leafhopper workers except Ossiannilsson (1946), Ishihara (1953) and Metcalf (1965), the argument about this group has been basically focused on whether to recognize it as a subfamily or a tribe. The people who consider

Hecaline a subfamily include Oman (1949), Young (1950),

China (1950), Evans (1951, 1954, 1955), Nast (1952),

Strickland (1953), Dlabola (1954), Lindberg (1954) and Morrison (1973). The opinion that Hecaline should be a tribe under Deltocephalinae was proposed by Linnavuori

(1959). In his study, the character of the anterior tentorial branches which defined Deltocephalinae was 5 examined for Hecaline and it was found that, as in all Deltocephalinae, the branches were bifurcate.

Linnavuori (1959) did not use the anterior tentorial branches to define Deltocephalinae because he thought another subfamily, Noecoelidiinae, also had this character.

In my previous studies, this character has been examined for many species from all subfamilies, including Neocoelidiinae, and only Deltocephalinae and Hecalinae had the bifurcate tentorium branches; the character in Noecoelidiinae is not bifurcate but stick-like and bent at the apex.

Based on the evidence mentioned above and Linnavuori’s conclusion, the dorsal tentorial branches are the definitive apomorphic character for Deltocephalinae, and Hecaline is treated as a tribe under Deltocephalinae in this study. 6

Natural History of Hecalini Distant

Hecalini is world-wide in distribution. In North

America, these leafhoppers are more common in southern regions (Mexico and the southern states) than in the north. Only three genera, Hecalus, Neohecalus. and

Memnonia have been recorded from Canada.

The host plants for Hecalini are various grasses.

There are no records thus far that any leafhoppers have been collected from woody hosts. They seldom occur on cultivated plants and are of little or no economic importance. These leafhoppers are not known as vectors of any plant diseases.

The hecaline leafhoppers are often parasitized by Hymenoptera, Strepsiptera and Diptera, which form a dark-colored "bag" extending out from the abdomen.

The parasites sometimes deform the genitalia, which makes identification more difficult in some groups. 7

Morphology and Terminology

Head: Hecaline leafhoppers usually have the head produced (Fig. 1). The dorsal region of the head is referred to as the vertex or crown. The head length is the distance from mid-anterior margin to the pronotum, and the head width is the largest transverse distance of the head including the eyes. The ocellus is located from at the anterior margin of the head near the eye to the half way from the eye to mid-anterior point of the vertex. There is a coronal suture in the middle of the vertex. The face (Fig. 2) consists of clypeus, lorum, gena, eye, antenna, anteclypeus, and several sutures. The lateral clypeal sutures extend dorsally beyond the antennal pits to the ocelli.

The genal margins under the eyes are usually strongly sinuate. In many males and all females, the anterior margin of the vertex is strongly carinate.

Thorax: Pronotal length is the medial longitudinal distance from the anterior to posterior margin. The lateral margins of the pronotum are always carinate.

The scutellum is triangular, with a transverse suture in the middle. Forewings: In Hecalini, the venation is similar

to that of typical Deltocephalinae (Fig. 3), except

for the genera Dicyphonia Ball and Spangbergiella

Signoret, and many females, the latter usually have

rather reduced forewing. There are two claval veins

in all genera except Spangbergiella. which has one.

The other terms used in venation in this study follow

Oman (1949) (Fig. 3).

Hindwings: The hindwings are usually hyaline

or light brown. When at rest the hindwings are folded

under the forewing, and because of the difficulty

in viewing, they are rarely used in the classification

of Hecalini.

Abdomen: Very few external characters of visceral

segments have been used in Hecalini except color.

One internal character, sternal apodemes, however,

is used very often. These apodemes are usually developed

in both sexes and in many genera there are two pairs.

The form of the seventh sternum of the female is useful

and often mentioned.

Male genitalia: Although I am not willing to agree with, the male genitalia are probably the most often used and most important characters in not only 9

Hecalini, but many other leafhopper groups because a great number of taxa are based exclusively upon the male genitalia. The male genitalia consist of the following main parts: pygofer, aedeagus, connective, styles, genital plates and genital valve. The pygofer

(Fig. 4) is the dorsal-lateral cover of the male genitalia.

Its form varies from species to species in Hecalini, and is usually covered by setae on the posterior surface.

There is a pygofer rift on the ventral basal corner which is also common all Deltocephalinae tribes.

The aedeagal form is more primitive compared with the other tribes of Deltocephalinae and similar to that of Dorycephalinae or Gyponinae. The aedeagus usually consists of several parts: terminal process(es), shaft, preatrium, and phragma (Fig. 7), all of which vary in shape or form. Styles are paired structures, probably originated from the appendages of the 10th abdominal segment. In Hecalini, the styles are all basically the same. The connective (Fig. 6) is a

V-shaped structure, and its "apex" is connected with the aedeagus and its "foot" with the styles, with processes on the "foot" region, and sometimes another pair in between. The genital plates and valve show little variation among the taxa and are thus of less value in classification in Hecalini. 10

Material and Methods

Material: More than two thousand specimens of

Hecalini from North America were examined in this study and they were obtained from the following collections:

American Museum of Natural History, New York,

New York (ANHM)

Biosystematics Research Centre, Ottawa Canada

(BRI)

British Museum (Natural History), London, England

(BMNH)

California Academy of Sciences, San Francisco,

California (CAS)

Florida Department of Agriculture DPI, Gainesville,

Florida (FDAC)

Kansas State University Collections, Manhattan

Kansas (KSUC)

National Museum of Natural History, Washington

D .C. (NMNH)

Naturihistoriska Riksmeseet Sektionen for Entomologi,

Stockholm Sweden (NRSE)

North Carolina State University, Raleigh North

Carolina (NCSUC)

Ohio State. University Collection, Columbus Ohio 11

(OSUC)

Oregon State University Collection, Corvalis,

Oregon (ORSUC)

Snow Museum of University of Kansas, Lawrence

Kansas (SEMC)

University of Kentucky Collections, Lexington

KY (UKC)

University of Missouri-Columbia Collections (UMCC)

Methods: Because of the time and material limitations, only morphological characters and their comparison were used in this study. All figures were drawn by using a camera lucida except the cladograms. Male genitalia were dissected and soaked in 10 % KOH about

24 hours. The male genitalia were all kept in genitalia vials attached to the specimens after study. All cladistic analyses were based on the principles of phylogenetics and they were conducted by using PAUP (Phylogenetic

Analysis by Using Parsimony), version 2.40, developed and distributed by Swofford in 1985. CHAPTER I

SYSTEMATIC STUDY OF THE TRIBE

HECALINI OF NORTH AMERICA

Hecalini Distant

Hecalusaria Distant, 1908:273.

Hecalinae Haupt, 1929:1074.

Hecalini Linnavuori, 1975:54.

There are six recognized genera of Hecalini from

North America. The following is the check list of

North America Hecalini.

Check list of Hecalini from North America

Memnonia Ball

Memnonia Ball, 1900:66, type: consobrina under

Memnonia. by original designation.

Memnonia acuta Beamer

Memnonia acuta Beamer, 1938:80.

Memnonia albolinea Ball

Memnonia albolinea Ball, 1937:131.

Memnonia brunnea (Ball) New Combination

12 Parabolocratus brunneus Ball, 1900:71.

Memnonia consobrina Ball

Memnonia consobrina Ball, 1900:66.

Memnonia fossitia Ball, 1937:132. New Synonym.

Memnonia fenestrella (Ball) New Combination

Parabolocratus fenestrellus Ball, 1937:130.

Memnonia fraterna Ball

Memnonia fraterna Ball 1900:67.

Memnonia hamiltoni New species

Memnonia minuta New species

Memnonia nigrafasciata New species

Memnonia ornata Linnavuori & DeLong

Memnonia ornata Linnavuori & DeLong, 1978: 52-53.

Memnonia soadix (Ball) New Combination

Parabolocratus spadix Ball, 1937:130.

Memnonia triplehorni New species

Dicyphonia Ball

Dicyphonia Ball, 1900:69, type: ramentosa under

genus Dicyphonia which is a synonym of Platymetopius

ornatus Baker, 1900.

Jiutepeca Linnavuori and Delong, 1978:51-52,

type: zamorana, under Jiutepeca. by original

designation. New Synonym. Dicyphonia cona (Shaw)

Parabolocratus conus Shaw, 1932:45.

Dicyphonia conica [sic] Beamer, 1936:69.

Dicyphonia cona DeLong and Knull, 1946:27.

Dicyphonia delongi new species

Dicyphonia mexicana new species

Dicyphonia nigrita Ball

Dicyphonia nigrita Ball, 1937:132.

Dicyphonia ornata (Baker)

Platymetopius ornatus Baker, 1900:49. By

Dicyphonia pamentosa Baker[sic], by Ball,

1900:69.

Dicyphonia ornata Van Duzee, 1908:158.

Dicyphonia plana Beamer

Dicyphonia plana Beamer, 1936:69.

Dicyphonia minuta Beamer, 1936:70. New synonym.

Dicyphonia plura Beamer

Dicyphonia plura Beamer, 1936:68.

Dicyphonia zamorana (Linnavuori and DeLong)

Jiutepeca zamorana Linnavuori and DeLong,

1978:51-52. Reinstated.

Hecullus Oman

Hecullus Oman, 1949:31, type: bracteatus Ball,

under genus Hecalus by original designation. 15

Hecullus balli (Beamer)

Hecalus balli Beamer, 1937:12.

Hecullus bracteatus (Ball)

Hecalus bracteatus Ball, 1901:4-5.

Neohecalus Linnavuori

Neohecalus Linnavuori, 1975:55, type: lineatus

Uhler, under genus Glossocratus* . by original

designation.

Neohecalus apicalis (Van Duzee)

Hecalus apicalis Van Duzee 1909: 217.

Neohecalus lineatus (Uhler)

Glossocratus lineatus Uhler 1877: 463.

Hecalus lineatus (Uhler), Signoret 1879:

267.

Spangbergiella Signoret

Spangbergiella Signoret, 1879:273, type: vulnerata

Uhler, under genus Glossocratus by original

designation.

Bergiella Baker, 1897:157, type: vulnerata Uhler,

under genus Glossocratus by original designation.

By Evans, 1947:140. 16

Spangbergiella erratica Linnavuori & DeLong

Spangbergiella erratica Linnavuori & DeLong,

1978:196.

Spangbergiella mexicana Baker

Spangbergiella mexicana Baker, 1897:157.

Spangbergiella quadripunctata Lawson

Spangbergiella quadripunctata Lawson, 1932:

120.

Spangbergiella reticulata Linnavuori & DeLong

Spangbergiella reticulata Linnavuori & DeLong,

1977:184.

Spangbergiella vulnerata (Uhler)

Glossocratus vulnerata Uhler, 1877:464.

Spangbergiella vulnerata (Uhler), Signoret,

1879:274.

Spangbergiella lacerdae Signoret, 1879:274,

by Linnavuori, 1957:154.

Spangbergiella Felix (sic) Berg, 1884:34,

35. New Synonym

Spangbergiella punctato-guttata Berg, 1884:

35. New Synonym

Parabolocratus uruguayensis Berg, 1884:

36. New Synonym

Hecalus lynchii (Berg), 1897:268; Linnavuori

1957:153.

Spangbergiella vulnerata (Uhler), Lawson, 1932:117. New Synonym

Spangbergiella vulnerata convexa Lawson,

1932:118. New Synonym

Spangbergiella vulnerata lativittata Lawson,

1932:118. New Synonym

Hecalus Stal

Hecalus Stal, 1864:65, type: paykulli Stal under

genus Petalocephala. by original designation.

Parabolocratus Fieber, type: glaucescens Fieber

under genus Parabolocratus. by Stal 1866:55

Columbanus Distant, 1916:22, type: misranus Distant,

under genus Columbanus. by Morrison, 1973:410.

Linnavuoriella Evans, 1966:134, type: arcuatus

Motschulsky, under genus Parabolocratus.

by Morrison, 1973:410.

Hecalus atascaderus (Ball)

Parabolocratus atascaderus Ball, 1933:223.

Hecalus attenuatus (Shaw)

Parabolocratus attenuatus Shaw, 1932:40.

Hecalus constrictus (DeLong)

Parabolocratus constrictus DeLong, 1938:302.

Hecalus continuus (DeLong)

Parabolocratus continuus DeLong, 1938:301.

Hecalus curtus (Shaw) 18 Parabolocratus curtus Shaw, 1932:41.

Hecalus elongatus (DeLong)

Parabolocratus elongatus DeLong, 1938:302.

Hecalus flavidus (Signoret)

Parabolocratus flavidus Signoret, 1879:276. Hecalus grandis (Shaw)

Parabolocratus grandis Shaw, 1932:44.

Hecalus hepneri (Beamer)

Parabolocratus hepneri Beamer, 1948:63.

Hecalus inflatus (DeLong)

Parabolocratus inflatus DeLong, 1938:302.

Parabolocratus continuus. Oman, 1949:33.

Hecalus inflatus DeLong, Reinstated.

Hecalus kansiensis (Shaw)

Parabolocratus kansiensis Shaw, 1932:47.

Hecalus ma.ior (Osborn)

Parabolocratus ma.ior Osborn, 1915:110.

Hecalus montanus (Ball)

Parabolocratus viridis montanus Ball, 1900:71.

Parabolocratus montanus Van Duzee, 1916:69.

Parabolocratus viridis montanus Van Duzee,

1917:625.

Parabolocratus montanus Oman, 1949:33.

Hecalus nigrafasciatus (Beamer)

Parabolocratus nigrafasciatus Beamer, 1938:82.

Hecalus nimbosus (Ball) 19

Parabolocratus nimbosus Ball, 1937:129.

Hecalus planus (Shaw)

Parabolocratus planus Shaw, 1932:48.

Hecalus rotundus (DeLong)

Parabolocratus rotundus DeLong, 1938:302.

Hecalus viridis (Uhler)

Gypona reverta Uhler, 1872:472 [nom.nud.].

Glossocratus viridis Uhler, 1877:462.

Parabolocratus viridis Signoret, 1879:275.

♦Glossocratus is still a good genus according to Morrison (1973).

The males of Hecalini can be separated by the following key, but the females are difficult or impossible to distinguish in some genera.

Key to males of Genera of Hecalini

1. Forewing with one claval vein; head and pronotum

with red or orange stripes arranged in pattern

shown in Fig. 155 ...... Spangbergiella Signoret

Forewing with two claval veins; head and pronotum

without red or orange stripes or with them in

different pattern than above ...... 2

2. 3rd apical cell opened outwardly; a V-shaped stripe

at anterior margin of vertex (Fig. 88) ...... 20

Dicyphonia Ball

3nd apical cell opened posteriorly; vertex without

V-shaped stripe ...... 3

Body length < 5 mm; forewing color usually light

to dark brown (Fig. 10)...... Memnonia Ball

Body length > 5 mm; forewing color yellow or

pale green ...... 4

Connective long and fused (Fig. 130), female

forewing only covered two or three abdominal

segments ...... Hecullus Omen

Connective V-shaped (Fig. 6) female forewing

covered at least five abdominal segments

...... 5

Head, pronotum and scutellum with red or brown

longitudinal stripes (Fig. 139); setae on pygofer

in group (Fig. 141) ...... Neohecalus Linnavuori

Head, pronotum and scutellum without red or brown

longitudinal stripes; setae on pygofer not arranged

in group ...... Hecalus Stal

Phylogeny of Genera of Hecalini

The phylogenetic study of Hecaline genera of

North America is mainly based on the morphological

analyses. 21

The reason only morphological characters or information are used in the phylogenetic study of hecalini is that for Hecalini there are little historical, biological or ecological information available and there are not enough specimens or material for me to conduct a geographic analysis.

Character analysis: Several characters were used by previous workers (Oman, 1949 and Linnavuori,

1975) in taxonomic study of the groups, such as the very reduced forewing of females and setae in a group on the pygofer of males. However, these characters are autapomorphic and deal largely with such problems, as how to separate the groups rather than the relationships among them. Therefore, more characters or character states shared by two or more genera were examined in this study. The total of 14 characters or character states and their polarized conditions are listed in

Table 1, and the data matrix obtained is shown in

Table 2.

Outgroup: Since there have been no phylogenetic studies of the subfamily Deltocephalinae, and based on the author’s own opinion that Hecalini is the most primitive tribe in the subfamily, no sister group can be selected with confidence. Therefore, two possibl

candidate genera were chosen from two tribes of the

subfamily to represent the tribe respectively, namely,

Deltocephalus from Deltocephalini, and Osborne!lus

from Euscelini. Since they gave the same results,

only Deltocephalus is listed in Table 2.

Results: The options or commands, BANDB, and

UNORDER ALL were used in the analysis, and a single

most parsimonious tree which represents the hypothesized

relationship among Hecalini genera was generated with

a consistency index = 0.913 and tree length = 23.000.

The cladogram is shown in Figure 8.

From the cladogram several things can be seen

which are either interesting or different from the

previous hypotheses: 1) Memnonia is more closely related

to Dicyphonia neither than Hecalus : 2) Hecalini mainly

is divided into two groups, each is supported by only

one character. 3) Neohecalus has a strong relationship with Hecullus, even if it were separated from Hecalus . 23

Table 1: Characters and their polarized conditions of tribe Hecalini

Plesiomorphic Apomorphic Characters condition condition

1. Forewing color of yellow or pale black male green or smoky 2. Claval vein two one 3. Forewing of female covering most of not covering sternal segments most of sternal segments 4. Setae on pygofer scattered in groups 5. Second apical cell faces posterior faces of male outwardly 6. Body size of male usually larger usually smaller than 6 mm than 5 mm 7. Head of female produced, length well produced, in middle slightly length in longer than length middle much between eyes longer than length between eyes 8. Connective form branched fused 9. Head of male wider than pronotum not wider including eyes than pronotum 10. Sinus under eye angle larger than angle equal on gena 90* to 90 * 11. Appendix extended to 2nd extended apical cell to 1st apical=l; to 3rd apical = 2 12. Cross vein of near apex of claval distant outer anteapical region from apex cell 13. Length of male shorter than longer than head in middle pronotum pronotum but longer than width between eyes=l; longer than pronotum=2 14. Venation of male basic pattern like like Fig. forewing Deltocephalinae 157 = 1, (Fig. 3) like Fig. 91 = 2 Table 1(continue), 15. Pygofer process absent present on ventral margins 16. long carina absent present along sides of pronotum 17. sexual dimorphism not obvious obvious 25

Table 2: Data Matrix of characters

or character states of tribe Hecalini

Characters or character states

111111

Taxa 123456789012345

Deltocephalus 000000000000000

Memnonia 100000001010101

Spangbergiella 010001001001210

Hecalus 000001001020100

Hecullus 001001110101200

Neohecalus 000101100101200

Dicyphonia 000010001000221 CHAPTER II

SYSTEMATIC STUDY OF THE GENUS

Memnonia BALL

Memnonia was described by Ball in 1900 based

on the species M. fraterna and M. consobrina: the

latter was the designed type species. Van Duzee (1917)

made the first addition to the genus, M. simplex.

but this species was later considered a synonym of

Friscanus friscanus (Ball) by Oman (1938). In 1937,

Ball described two species, M. albolinea and M. fossitia.

without diagnostic illustrations. One year later,

Beamer described M. acuta which is apparently related

to M. albolinea. The most recent addition, M. ornata.

was by Linnavuori and DeLong (1978). Since the establishment,

no generic revision has occurred, even though its generic

limits and position in Cicadellidae have been questioned

(Oman, 1949; Linnavuori, 1959). In this study, four new species are described, M. fossitia Ball is considered

a synonym of M. consobrina Ball, and three species are transferred into Memnonia from Hecalus, the reason will be discussed in each species description.

Genus Memnonia Ball

Memnonia Ball 1900:66. Type species: consobrina Ball,

26 27

1900, under Memnonia. by original designation.

Usually small leafhoppers (male < 4 mm in length).

Sexual dimorphism obvious, as other genera of Hecalini.

General color dark brown in males, sordid yellow in females; forewing of male always dark. Head in both sexes produced, length sub-equal to width between eyes in males, longer than width in females. Crown not very flattened, as occurs in many species of Hecalus. usually no carina on anterior margin. Ocellus separated from eye by distance equal to 2X ocellar diameter.

Pronotal length equal to or shorter than length of crown, with long carinae along lateral margins, and posterior margin always angulate at middle. Forewing usually developed in males and reduced in females, as in many genera of Hecalini. Male forewing venation with normal pattern of Hecalini (Fig. 10) but sometimes with more accessory cross-veins. In most cases, spinulation of hind femur of both sexes 2:2:1. Male abdomen not very long, usually with two pairs of sternal apodemes.

Male genitalia relatively simple compared with other genera, such as Dicyphonia and Hecullus. Pygofer usually as in other members of Hecalini, but sometimes elongated, always with one or two pairs of processes on ventral margins. Aedeagus cylindrical, curved upwards, with one or two pairs of apical processes extending in various directions. Processes on arm

of connective variable among species.

There has been question concerning the identification

of Memnonia. especially as it is difficult to separate

from Hecalus. Actually, Memnonia is a genus which

is easily sorted from misellancous leafhoppers but

not easy to characterize literally. Oman (1949) gives

three characters to separate Memnonia from other Hecalini

genera: small size, dark colored males, and the lack

of carina on the anterior margin of the crown. In

this study, I found that none of those characters

is unique for Memnonia. and this genus is not closely

related to Hecalus, but rather to Dicyphonia (see Chapter

1). Therefore, some characters are added to the diagnosis such as the shape of the posterior margin of pronotum, which is shared by all members of Memnonia and none of Hecalus. to make the definition clearer.

Diagnosis: Species of Memnonia Ball can be separated from Hecalus and other genera of Hecalini by the following characters: 1) small size (male under 4 mm in length), 2) dark forewing and usually dark-colored body in males, 3) no carina on anterior margin of crown, 4) posterior margin of pronotum angulate, and

5) processes present on ventral margin of pygofer.

Hecalus brunneus, H. fenestrellus. and H. spadix are also small and have dark forewings. In addition, these three species have no carina on the anterior margin of crown in the male, and with posterior margin of pronotum angulate. Because these characters are more consistent with Memnonia than Hecalus. they are hereby transferred to Memnonia.

The fact that Memnonia is a valid Hecalini genus can also be supported by electronmicroscopy. Two species from Hecalus, H. major, H. flavidus. and one species from Memnonia. M. albolinea were chosen as the representative of each genus, and two or three individuals of each species have been examined. Two characters from each individual, namely the upper surface of the male forewing and dorsol-lateral surface of the male pygofer were used since these body parts of body are easy to treat and photograph. In addition, the forewing can be returned to the individual specimen after study. Besides the above species, H. viridis and M. consobrina were also examined as the replications, 30

and Dorycephalus (D.) platyrhychus Osborn from Dorycephalinae

was used as outgroup. The electron microscope results

are shown on Plate I. For Hecalus. the male forewing

has tiny bump-like projections on the entire surface.

The density of the projections varied among species,

while for Memnonia albolinea. the surface is smooth.

As for the second character, in Hecalus. species had

horn-like projections, while in Memnonia the projections

were small and comb-like. Only the pictures of the

former three species are shown in Plate I because

the replicates and outgroup showed basically the same

character pattern as Hecalus. Even though not all

species of both genera were examined, it is likely

that the species should belong to the different genera,

because the characters in M. albolinea definitely

are apomorphic. Since the data are incomplete, these

two characters were not included in the phylogenetic

study.

In addition, there has been some concern about

in which subfamily Memnonia should be put, and there

are mainly two schools of thought. Evans (1947) and

Metcalf (1965) placed it in the subfamily Aphrodinae;

Oman (1949), however, treated it as a member of the subfamily Hecalinae. Neither Evans nor Metcalf gave any details as to why they put Memnonia in Aphrodinae. 31

The main characters of Aphrodinae are its ocellar area with a distinct ledge or carina above the antennal pit, and with the episternum largely exposed. Memnonia does not match these characters. In addition, male genitalia pattern of Memnonia are quite different from Aphrodinae. On the contrary, Memnonia has not only the critical characters of subfamily Deltocephalinae such as branched dorsal tentotial arm, but also all four characters of Hecalini, namely a produced head, anterior margin of gena sinuate under eye, a long carina on lateral margins of pronotum and sexual dimorphism.

Therefore, I agree with Oman and consider Memnonia

Ball a member of Hecalini under subfamily Deltocephalinae.

Key to males of Memnonia Ball

1. General color of head and pronotum pale green

to light brown 2

General color of head and pronotum dark brown

8

2. Forewing reduced, having two or more abdominal

tergites exposed fraterna Ball

Forewing not reduced 3

3. Terminal processes of aedeagus extended caudally

fenestrella (Ball) Terminal processes of aedeagus not extended caudally

...... 4

Pronotum with longitudinal stripes ......

...... brunnea (Ball)

Pronotum without stripes ...... 5

Shaft of aedeagus swollen at middle in lateral view and upper rami longer than lower ones

...... spadix (Ball)

Shaft of aedeagus not swollen at middle in lateral view and upper rami not longer than lower ones

6

Forewing without any white spots around apical cross veins, and terminal processes of aedeagus as thick as shaft ...... nigrifasciata n. sp.

Forewin'g with some white spots around apical cross veins, and terminal processes of aedeagus not as thick as shaft ...... 7

Terminal processes of aedeagus shorter than 1/2 length of shaft ...... hamiltoni n. sp.

Terminal processes of aedeagus longer than 1/2 length of shaft triplehorni n. sp.

With reticular stripes on forewing

...... ornata Linnavuori & DeLong Without such stripes on forewing ...... 9

Aedeagal rami twisted at apex (Fig. 21) 33

Aedeagal rami not twisted at apex (Fig. 29)

11

10. Pronotum with stripes ...... albolinea Ball

Pronotum without stripes ...... acuta Beamer

11. Sternal apodemes relatively smaller, not longer

than half length of first visible abdominal segment

(Fig. 39) ...... minuta n. sp.

Sternal apodemes normal size, longer than half

of first visible abdominal segment (Fig. 27)

...... consobrina Ball

Memnonia ornata Linnavuori & DeLong

(fig. 9-15)

Memnonia ornata Linnavuori & DeLong 1978:52-53.

Female unknown.

Male: Body length: 3.5 mm. General color brown

with some dark-brown stripes on crown, pronotum and

scutellum. Head produced and length slightly shorter

than width between eyes. Crown with two pairs of

stripes, lateral pair short, middle pair crescent

shaped. Pronotum longer than length of head, with

three pairs of longitudinal stripes. Scutellum with

lateral angles and posterior portion brown. Forewing

fully developed, with veins dark colored, forming reticular color-pattern. Two pairs of sternal apodemes present, both longer than half of first visible abdominal segment and middle pair long and slender. Male genitalia

Pygofer with one pair of processes on ventral margin.

Style and genital plate as described for the genus.

Aedeagus with two pairs of apical processes, both pairs directed anteriorly then turned ventrally.

Aedeagal preatrium much longer than half the length of shaft. Connective processes small.

Diagnosis: This species is easy distinguished from other species by its dark-colored body and brown stripes on pronotum and forewing.

Distribution: Mexico.

Type and Material Examined: 1 male, holotype,

1 male, paratype, Toluca, D.F., Mexico, IX-29-1941,

DeLong, Good, Caldwell & Plummer (OSUC).

Memnonia albolinea Ball

(figs. 16-22)

Memnonia albolinea Ball 1937:131.

Female: Body length: 5.3 mm. General color sordid yellow. Head not produced, length less than width between eyes, having one slender stripe along anterior 35

margin of crown. Ocellus far from eye, the distance

longer than twice its diameter. Forewing reduced,

leaving two or more abdominal segments exposed.

Male: Body length: 3.1-3.7 mm. General color

brown with several longitudinal light colored stripes

on pronotum. Head not much produced, length less

than width between eyes. Anterior margin of crown

light-colored, ocellus position as in generic description.

Forewing developed, color dark brown with several

white spots around apical cross-veins. Two pairs

of sternal apodemes present and almost as long as

first visible abdominal segment. Male genitalia:

Pygofer short and of usual form, with two pairs of

processes on lower margin. Aedeagus slightly curved at middle, two pairs of apical processes with twisted

apex. Connective process large, as long as length of arm.

Diagnosis: This species can be distinguished

from other species by the dark-colored body and light-colored stripes on pronotum.

Distribution: Arizona. Type and Material Examined: 1 female (holotype),

Santa Catalina Mts., Arizona, VII-2-1933, E.D.

Ball. 4 males (allotypes), 5 M. east Old Bnldy, Arizona,

VI-18-1936, E.D. Ball. 1 male, 1 female, Chiricahua

Mts., Arizona, VI-9-1933, P.W. Oman. 2 males, labeled

as allotypes. All specimens were obtained from NMNH.

Memnonia acuta Beamer

(fig. 23)

Memnonia acuta Beamer, 1938:80.

This species is very similar to M. albolinea

Ball and there is no difference between their male

genitalia and sternal apodemes. The only, differences

are that the head of M. acuta Beamer is more produced

and the length is equal to the width between the

eyes while in M. albolinea Ball the head is less than the distance between the eyes. In addition,

there are no light-color stripes on the pronotum

of M. acuta Beamer.

Remarks: Originally, I thought that M. acuta

Beamer was a synonym of M. albolinea Ball, but the distinct head shape and pronotal stripes convinced me that they are separate species. 37

Distribution: Colorado.

Type and Material Examined: 1 male, 1 female (paratypes),

Little Beaver Cr., Colorado, VII-11-1937, R.H. Beamer

(NMNH).

Memnonia consobrina Ball

(fig. 24-31)

Memnonia consobrina Ball 1900:66.

Memnonia fossitia Ball 1937:132, New Synonym.

Female: Body length: 3.7-4.5 mm. General color sordid yellow. Head produced and length slightly longer than width between eyes. Pronotum about as long as length of crown. Forewing reduced, two or more abdominal tergites exposed. Venation reduced and apical region very short. Seventh abdominal sternite with one median process on posterior margin. Ovipositor extended past pygofer and exposed.

Male: Body length: 2.9-3.2 mm. General color dark brown, color on crown and pronotum varying in individuals from dark to light brown with some light-colored spots. Head produced, length slightly longer than width between eyes. Pronotum as long and wide as 38

head. Forewing developed and with normal venation,

several white spots around apical cross-veins. Two

pairs of sternal apodemes present and both almost

as long as first visible abdominal segment. Male

genitalia: Pygofer not very long, with two pairs

of processes on ventral margin. Aedeagus curved dorsally,

with one large preatrium and two pairs of apical processes

whose dorsal rami are large and lobe-like. One pair

of spur-like tiny processes present near gonopore.

Connective process developed, as long as length of

arm.

Remark: In this study, the types of both consobrina

and fossitia were examined. Beamer did mentioned

the head shape difference between these two "species",

but there were some variation among all individuals

and the head shape difference was not justified enough

to put them into two species when compared with the variation among the other species. Besides, no differences

between their male genitalia, sternal apodemes, and

even their body color and size were observed. Therefore,

M. fossitia Ball is here considered a synonym of M. consobrina Ball.

Diagnosis: This species can be distinguished

from other species by the dark-colored body, without 39 a stripe on the pronotum and having their sternal apodemes longer than the half length of the first visible abdominal segment.

Distribution: Arizona, Colorado, Kansas, New

Mexico; Mexico.

Type and Material Examined: 1 males (lectotype chosen from cotypes), N. Colorado, IV-28-1898, and

V-17-1898, no collector. 1 male (paralectotype chosen from cotypes), labeled same as lectotype. 1 female

(paralectotype chosen from cotypes), Ft. Collins,

Colorado, V-27-1898, no collector. 1 female, Clay

Co., Kansas, VIII-1-1906, D.M. DeLong. 3 males, Tombstone,

Arizona, VIII-1-1931, E.D. Ball. 1 male, Alamagordo,

New Mexico, VI-30-1932, R.H. Beamer. 1 male, Prescott,

N.F. Arizona, VII-14-1940, D.J. & J.K. Knull. 1 male,

Delphos Kans., 1 male, Pueblo, Colorado, 2 males,

Clay Co., Kans., collected by Osborn, no date. 1 female (lectotype of M. fossitia chosen from 3 female cotypes on one pin), Douglas Arizona, VI-10-1936,

E.D. Ball. 2 females (paralectotypes chosen from 3 female cotypes on one pin), 3 males (allotypes on one pin), 3 males (paratypes), and 2 females (paratypes), all labeled as holotype (NMNH). 1 female, Clay Co.,

Kans. VIII-1-1906, D.M. DeLong, 1 female, Eaton, Colo. VIII-27-1914, H. Morrison, 1 male Saltillo Coah.,

Mexico, IX-23-1941, D.M. DeLong, Good, Caldwell and

Plummer (OSUC).

Memnonia minuta New Species

(fig. 32-39)

Description: Female unknown. Male: body length:

2.4-2.5 mm. Small, elongate leafhoppers. General

color dark brown, without any other marks except

some white spots around apical cross-veins of forewing

Head slightly wider than pronotum, crown produced,

length as long as width between eyes. Face narrow

and long, with apex and area below eyes light-colored.

Forewing developed, dark brown, not transparent. Vente

color as in rest of body; spinulation of hind femur

2:2:1. Abdomen short, without light-colored band on tergites. Two pairs of sternal apodemes which are not well developed and much shorter than half of first visible abdominal segment. Male genitalia:

Pygofer not very long, with apical half covered with

setae, having two pairs of processes on lower margin.

Genital plate long and narrow, 3-4 setae on lateral margin. Style as in generic description. Aedeagus like M. consobrina but smaller and more slender, and without spur-like processes near gonopore. In addition 41

lower rami not swollen near apex, and aedeagal preatrium

not as large as that of M. consobrina. Connective

processes small, not longer than half length of arm.

Diagnosis: This species is closely related to

M. consobrina but can be separated from it by the

smaller size, less developed connective processes,

and the sternal apodemes which are much shorter than

the first visible abdominal segment.

Holotype: Male, Garden of the Gods, Colorado,

Collected by Osborn, no date.

Paratypes: 6 males, labeled same as holotype.

All types are deposited in OSUC.

Memnonia fraterna Ball

(fig. 40-46)

Memnonia fraterna Ball 1900:67.

Female individuals not available in the study.

According to Ball (1900), female looks like male, which is the exception in this genus.

Male: Body length: 3.7-3.9 mm. General color sordid yellow, without any other marks except ones

on apical margin of forewing. Head produced, as

long as width between eyes. Forewing and venation

reduced, apical region short, like females of other

species of Memnonia. One or two brown spots present on apical-external margin of forewing. Two pairs

of sternal apodemes, these reduced, length not longer

than half length of first visible abdominal segment.

Male genitalia: Pygofer short, with two pairs of

processes on ventral margin, and posterior half covered

by setae. Style and plate like M. consobrina. Aedeagus

like M. consobrina. but subapical rami not swollen

near apex and preatrium not as large. Connective

process small, not longer than half length of arm.

Diagnosis: This species can be distinguished

from the other species by its reduced male forewing and the light-colored body.

Distribution: Colorado.

Type and Material Examined: 1 male, (lectotype chosen from two cotypes), N. Colorado, V-18-1898, no collector. 2 males (one is paralectotype chosen from two cotypes), Ft. Collins, Colorado, V-25-1898, 43 no collector. All specimens are from NMNH.

Memnonia brunnea (Ball) New combination

(fig. 47-54)

Parabolocratus brunneus Ball 1900:71.

Female: Body length: 5.8-6.4 mm. General color light yellow without other markings. Head produced, length slightly longer than width between eyes. Forewing and venation reduced, two or more abdominal segments exposed. Ovipositor long, with apex brown.

Male: Body length: 3.5-3.7 mm. Head color sordid yellow. Pronotum color sordid yellow with two faint brown stripes. Head produced, length as long as width between eyes. Forewing developed, brown, and not transparent. Venation normal. Abdomen brown, one light colored band in middle of dorsal surface.

Two pairs of sternal apodemes present, both longer than first visible abdominal segment with middle pair longer and more slender. Male genitalia: Pygofer long, length twice as long as height, posterior half covered by setae. One pair of processes present on ventral margin of pygofer. Style and plate as in generic description. Aedeagus stout and straight, one pair of branched apical processes which extend

downward. Connective straight and process small,

not reaching half length of arm.

Diagnosis: This species can be distinguished

from the other species by its light-colored body,

the stripes on the pronotum and aedeagal shaft not

swollen at the middle in lateral aspect.

Distribution: Colorado, Kansas, New Mexico.

Material Examined: 1 female, Eaton, Colo., VIII-

27-1914, H. Morrison (OSUC). 1 female, Olney, Colorado,

VIII-14-1927, Beamer-Lawson. 1 female, Belen, N.

Mexico, VIII-19-1927, R.H. Beamer. 1 female, Scott

Co., Kansas, VIII-23-1927, R.H. Beamer. 1 female,

Rosewell, N. Mexico, VII-16-1936, D.R. Lindsay. 7 males, 2 females, Springer, N. Mexico, VII-17-1936,

R.H. Beamer. 4 males, Belen, N. Mexico, Vii-20-1936,

R.H. Beamer (SEMC). All types are also deposited in SEMC.

Memnonia fenestrella (Ball) New Combination

(fig. 55-61)

Parabolocratus fenestrellus Ball 1937:130. 45

Female: Body length: 6.8-7.0 mm. General color

sordid white. Head not produced, length slightly

shorter than width between eyes. Forewing and venation

reduced, with apical region shortened, leaving one

or more abdominal tergites exposed. Ovipositor long,

extending past apex of pygofer.

Male: Body length: 3.6-3.8 mm. General color

sordid white to sordid brown, forewing brown with

apical portion of venation dark brown. Head not produced,

length slightly shorter than width between eyes.

Some individuals with light brown forewing and faint

brown marks on posterior half of pronotum. Forewing

developed, venation normal, sometimes with accessory

cross-veins on basal half. Two pairs of sternal apodemes

developed, middle pair twice as long as lateral pair

but more slender. Male genitalia: Pygofer long,

length twice the height in lateral aspect, posterior

half covered by setae. Aedeagus straight, narrow

at middle, upper half of shaft triangular in cross

section. One pair of branched apical processes directed ventrally, anterior branch slightly longer than posterior branch. Connective process not large, shorter than half length of arm.

Diagnosis: This species can be separated from 46

the other species by the light-colored body and the

reticular stripes on the male forewing.

Distribution: California.

Material Examined: 1 female, Palmdale, Calif.,

VII-6-1933, R.H. Beamer. 1 male, 3 females, Newberry

Spgs, Calif., VII-30-1936, D.R. Lindsay. 1 male,

Campo. Calif., VII-18-1940, D.E. Hardy. All specimens

examined are from SEMC and types are also deposited

there.

Memnonia spadix (Ball) New Combination

(fig. 62-68)

Parabolocratus spadix Ball 1937:130.

Female: Body length: 5.4-5.8 mm. General color

sordid green, without other markings. Head produced,

length slightly shorter than width between eyes.

Pronotum as long as crown. Forewing slightly reduced,

leaving only one abdominal tergite exposed. Venation

conspicuous, slightly reduced at apical region. Ovipositor extended past pygofer, apex brown.

Male: Body length: 3.7-3.9 mm. General color of head, pronotum, scutellum yellow green, and forewing light brown. Head produced, length slightly longer than width between eyes. Forewing developed normally, venation normal. Abdomen with two pairs of sternal apodemes, middle pair longer and more slender than lateral pair. Male genitalia: Pygofer not elongate, length less than one and half height, and posterior half covered by setae. Only one pair of processes on ventral margin of pygofer. Aedeagus stout, swollen at middle in lateral aspect, with one pair branched apical processes, posterior pair twice as thin as anterior pair in posterior aspect. Connective large, with process not much produced but almost as long as length of arm.

Diagnosis: This species can be distinguished by the light-colored body and the aedeagal shaft which is swollen at the middle in lateral aspect.

Distribution: New Mexico, Oklahoma.

Material Examined: 3 males, N. Forest of Oklahoma,

VI-28-1936, R.H. Beamer. 5 males, 3 females, Silver

City, New Mexico, VII-22-1936, R.H. Beamer. 1 male,

1 female, Roosevelt, Oklahoma, VI-27-1936, R.H. Beamer.

All specimens examined are from SEMC and types are also deposited there. 48

Memnonia nitfrifasciata New species

(Fig. 69-74)

Description: Female unknown. Male: body length:

3.2-3.3 mm. General color dark brown. Head produced,

with length in middle slightly shorter than width

between eyes. One black band present along anterior

margin of crown, two crescent spots beside middle

suture. Pronotum wider and longer than head, with

posterior half rugose. Forewing developed normally,

consistent dark brown, with apical margin much darker.

Two pairs of sternal apodemes present, first pair

twice longer and much more slender than second pair.

First pair of sternal apodemes also longer than first

visible abdominal segment. Male genitalia: Pygofer

short, covered with setae at posterior half, with

one pair processes on lower margins. Aedeagus short,

shaft swollen at basal 2/3 dorsal-ventrally. Terminal

processes of aedeagus very stout, lower rami longer

than upper ones.

Diagnosis: This species is similar to M. consobrina.

triplehorni and hamiltoni. but can be separated from them by the following characters: 1) a black band along the anterior margin of crown, 2) forewing without 49

white spots around apical cross-veins, and 3) terminal

processes of aedeagus stout, lower rami stouter than

shaft.

Distribution: Canada.

Holotype: Male, Manyberries, Alta, Canada, VI-3-1952,

collected by A.R. Brooks.

Paratypes: 1 male, labeled same as holotype.

2 males, Manyberries, Alta, Canada, V-27-1952 and

VI-3-1952, L.A. Konotopetz.

All types are deposited in BRI.

Memnonia triplehorni New species

(Fig. 75-80)

Description: Female unknown. Male: body length:

4.1-4.2 mm. General color yellow green on head,

pronotum and scutellum, forewing smoky brown. Head

produced, length in middle longer than width between

eyes. Ocellar position as in generic description.

Pronotum wider than head, as long as head, posterior half rugose. Forewing developed normally, normal venation. Veins dark brown at basal half and lighter color around apical cross veins. More cross vein present along outer margin of forewing. Abdomen dark brown to black. Two pairs of sternal apodemes developed, first pair longer and more slender than second one, but all shorter than first visible abdominal segment.

Male genitalia: Pygofer somewhat triangular in form in lateral view, covered with setae at posterior half, one pair of processes present on ventral margin.

Aedeagus long, swollen in apical half. One pair of ridges present at apical half of shaft on posterior surface, with several tiny teeth on dorsal lateral surface near gonopore. Terminal processes long, with a common stem, and upper rami sickle-like, shorter than lower one, lower one very sharp at apex.

Distribution: California.

Holotype: Male, 1000 Palms Oasis, Thousand Palms,

California, 111-18-1955, collected by W.R.M. Mason.

Paratype: 1 male, data the same as holotype.

All types are deposited in BRI.

This species is named after Dr. Charles A. Triplehorn 51 in recognition of his many contributions in entomology and education.

Memnonia hamiltoni New Species

(Fig. 81-86)

Description: Female: unknown. Male: body length:

3.3-3.4 mm. General color dark brown, with head lighter brown and several white spots around apical cross-veins of forewing. Head not much produced, pronotum wider than head, posterior half rugose. Forewing fully developed, venation normal. Abdcmen dark brown. Two pairs of sternal apodemes developed middle pair twice longer than lateral one. Male genitalia: Pygofer brown, covered with setae on posterior half, and with one pair of processes on ventral margin near basal corner.

Aedeagus slender, long and flattened laterally, having one pair of branched terminal processes, upper rami shorter than lower ones. Styles and genital plates as in generic description.

Diagnosis: This species is similar to M. consobrina and minuta. but the following characters can separate them: 1) lighter color of the head, 2) middle pair of sternal apodemes twice longer than lateral one, and 3) terminal processes of aedeagus with a common stem and upper rami shorter than lower ones.

Distribution: Canada.

Holotype: Male, Val Marie, Sask, Canada, VI-14-1955,

collected by A.R. Brooks.

Paratype: 1 male, labeled same as holotype.

All types are deposited in BRI.

This species is named for K. G. A. Hamilton for

his kindness in loaning specimens for this study.

Phylogeny of Memnonia Ball

A hypothesis of the relationship among the Memnonia

species has been constructed by using phylogenetic

systematics. In this study, 15 characters were examined

for 10 OTUs. A list of the characters which were polarized according to the parsimony principle is presented in Table 3, with the resulting data matrix 5 3

presented in Tables 3, and 4. Because M. consobrina

and M. minuta, M. albolinea and M. acuta are closely

related, which is strongly supported by the similarity

of their aedeagus and external appearance, 1 chose

to analyze those four taxa as only two OTUs, namely,

acuta+ and consobrina+. Actually, the combination

reduced the complexity of the results, without any

basic changes in the topology of the tree.

According to the study mentioned earlier, Memnon i a

is most closely related to Picyphonia, and this can

be supported by the fact that they have the processes on the ventral margin of the male pygofer and they are all fairly small. Therefore, Picyphonia ornata . the type species, has been chosen as the outgroup

in this analysis. Only a single most parsimonious tree was found in the analyses by using the BANDB option, and the cladogram is shown in Fig. 87. The consistency index is 0.833 and tree length 18.00.

From the cladogram we can see that Mejnnjp_ni_a is a monophyletic group which is supported by the characters

11, 12 and 13. There are two subgroups which are strongly supported by this analysis, that is, the subgroup including consobrina, minuta, a cut. a and albolinea, and that including ornata, trip1ehorni and fenestrella■ 54

TABLE 3: Characters and their polarization for Memnonia species, character numbers refer to characters used in Table 4.

Characters Plesiomorphic Apomorphic

1.Forewing normal size reduced 2.Apex of terminal not twisted twisted processes 3.Head without stripes with longitudinal stripes 4.Shaft of not swollen in swollen in Aedeagus lateral view lateral view 5.Forewing without white spot with the spot around cross-veins around apical cross-veins 6.Middle pair of longer than lateral shorter or sternal apodemes pair equal to lateral pair 7.General color pale yellow very dark brown of forewing 8.Upper half of without ridge with ridges aedeagus shaft 9.Length of at least 1.5 times as long as pygofer as height height 10.Body pale yellow partially brown 11.Posterior curved not angularly curved angularly margin of (Fig. 9) prontum 12.Front margin with carina without carina of crown 13.Ventral margin without processes with processes of pygofer 14.Forewing without stripes with reticular stripes 15.Forewing color pale yellow consistently brown TABLE 4: Data matrix of Memnonia

species, numbers refer to

characters used in table 3.

O=plesiomorphic,

l=apomorphic

Characters

111111

Taxa 123456789012345

P. flavidus 0 0 0 .0 0 0 0 0 0 0 0 0 0 0 0 consobrina+ 0 0 0 0 1 1 1 0 0 0 1 1 1 0 0 acutat 0 1 0 0 1 1 1 0 0 0 1 1 1 0 0 ornata 0 0 1 0 0 0 0 1 1 1 1 1 1 1 0 fenestrella 0 0 0 0 0 0 0 1 0 1 1 1 1 1 0 triplehorni 0 0 0 0 1 0 0 1 1 1 1 1 1 1 0 hamiltoni 0 0 0 0 1 0 0 0 0 1 1 1 1 0 1 nigrifasciat 0 0 0 1 0 0 0 0 0 1 1 1 1 0 1 spadix 0 0 0 1 0 0 0 0 1 1 1 1 10 1 brunnea 0 0 0 1 0 0 0 0 1 1 1 1 1 0 1 fraterna 1 0 0 0 0 1 0 0 0 0 1 1 1 0 0 CHAPTER III

SYSTEMATIC STUDY OF THE GENUS

Dicyphonia BALL

The genus Dicyphonia was described by Ball in

1900 for his species D. ramentosa. Van Duzee (1908)

recognized D. ramentosa Ball as a synonym of Platvmetopius

ornatus Baker (1900) which he transferred to Dicyphonia.

Therefore, Dicyphonia ornata (Baker) is the generic

type of Dicyphonia. Beamer (1936) published the first

revision for the group, and described three species,

D. minuta. D. plana. and D. plura. In addition, he

transferred Parabolocratus conus Shaw to Dicyphonia

and provided a key to the 5 species of Dicyphonia.

Ball (1937) described an additional species, D. nigrita.

bringing the total of species to six. Osborn (1924)

described D. picturata which was transferred to Scaphytopius

(Cloanthanus) by DeLong and Linnavuori (1978). Linnavuori

and DeLong (1978) established a new genus, Jiutepeca.

based on Dicyphonia nigrita Ball and described an

additional species, J. zamorana.

In this study eight species are recognized in

Dicyphonia. including two new species, D. delongi and D. mexicana. D. minuta Beamer is found to be a

56 synonym of D. plana Beamer, and the genus Jiutepeca

Linnavuori and DeLong is found to be a synonym of

Dicyphonia Ball.

Genus Dicyphonia Ball

Dicyphonia Ball, 1900:69. Type species: ramentosa

Ball, 1900:69 under genus Platymetopius. by original

designation, which is a synonym of Platymetopius

ornatus Baker now placed in Dicyphonia. Van Duzee

1908:158.

Jiutepeca Linnavuori and DeLong, 1978:49-50. New Synonym

Generic diagnosis: Dicyphonia can be distinguished

from other genera of Hecalini by the following characters

1) second outer-anteapical cell open outwardly but

not posteriorly (Fig. 91); 2) only one pair of sternal apodemes present; 3) with V-shaped stripe medially

on anterior margin of crown (Figs. 88, 92); 4) males usually with narrow stripe along dorsal margin of

face and various stripes on crown, pronotum and scutellum;

5) males usually less than 5 mm long, dark or with dark stripes.

Description: Small (males < 5 mm), and sexual 58

dimorphism obvious. Males pale yellow to dark brown

or black with various stripes, females usually light-colored.

Head generally produced in both sexes, length of head

longer than distance between eyes. Crown usually

with V-shaped stripe near anterior margin, which in

male is connected with usually irregularly shaped

stripes, and pair of fine stripes near lateral margin

anterior to eyes. Face slightly convex in lateral

aspect, with black band along dorsal margin and sometimes

some small, short stripes on clypeus in male. Pronotum

elongated, usually with some irregularly shaped stripes

in males and some females. Forewings of male fully

developed, with variously shaped stripes. Forewings

of female usually without stripes, slightly reduced,

and apical region short, sometimes leaving several

abdominal segments exposed. Most individuals with

nine rows of large setae on metatibiae. Only one

pair of abdominal-sternal apodemes developed. Male

genitalia: Relatively simple, pygofer short, covered with setae in apical half, with processes on ventral margins; aedeagus usually with developed shaft and

terminal processes which vary in number and shape, preatrium usually short. This genus is known only

from the Nearctic Region.

Notes: Jiutepeca. established by Linnavuori 59 and DeLong in 1978, was thought to be separable from other genera of Hecalini by the forewing venation, genital plate, connective and stout aedeagus. However, there is no difference in venation between D. nigrita and other species of Dicyphonia (Figs. 91, 107).

Furthermore, the differences in the genital plate, connective and aedeagus among D. nigrita and the other species of Dicyphonia are not great enough to justify

Jiutepeca as a separate genus, especially when compared with differences among other genera in Hecalini. Therefore,

Jiutepeca Linnavuori and DeLong is considered a synonym of Dicyphonia Ball.

Types of all species were examined.

Key to the males of Dicyphonia Ball

1. Forewing pale yellow basally, apical 2/3 black

4.

Forewing entirely covered by stripes ...... 2.

2. Head rounded, short, length beyond anterior angle

of eyes equal to the remainder; aedeagus cylindrical,

with one pair of apical processes ... plana Beamer

Head acute, length beyond anterior angle of eyes

twice longer than the remaining; aedeagus subtriangular 60 in cross-section, with two pairs of apical processes

...... 3.

Color dark brown to black, face with several oblique stripes ...... ornata (Baker)

Color pale yellow or pale brown, face without oblique stripes except for band along dorsal margin ...... plura Beamer

Median longitudinal stripe on head irregularly-shaped

(Fig. 101); aedeagus with one pair of apical processes ...... zamorana (Linnavuori and DeLong)

Median longitudinal stripe on head straight (Figs.

106, 116, 124); aedeagus with two pairs of apical processes ...... 5.

Face with several oblique stripes, thorax black ventrally mexicana. n. sp.

Face without oblique stripes except for band along dorsal margin, thorax never black ventrally

6 .

Stripe on pronotum widening posteriorly (Fig.

106), aedeagus large, apical processes equal to 1/2 length of aedeagal shaft

...... nigrita Ball

Stripe on pronotum not widening posteriorly (Fig.

116), apical processes almost reaching 1/3 length of aedeagal shaft ...... delongi. n. sp. 61

Key to the females of Dicyphonia Ball

1. Forewings short, several abdominal tergites exposed

2 .

Forewings long, usually only ovipositor exposed

...... 4.

2. Without markings on pronotum ..... cona (Shaw)

With brown pronotal markings ...... 3.

3. Vertex shorter than length between eyes

...... plana Beamer and plura Beamer*

Vertex longer than length between eyes

...... ornata (Baker)

4. Pronotum with markings ......

...... zamorana (Linnavuori and DeLong)

Pronotum without markings ......

nigrita Ball, delongi and mexicana n. spp.*

* Females of these species are difficult to separate.

Dicyphonia plana Beamer

(Figs. 88-91)

Dicyphonia plana Beamer, 1936:69.

Dicyphonia minuta Beamer, 1936:70. New Synonym

Female: 5.8-6.2 mm. General color light brown. Head with two or four dark spots on anterior margin

of vertex; forewing covered with reticular stripes.

Forewings reduced, leaving several abdominal segments

exposed.

Male: Body length: 2.9-3.1 mm. Male body dark

brown. Head, pronotum, scutellum, and forewing with

dark stripes or spots (Figs. 88, 91), stripes vary

in darkness. Ocellus on anterior angle of eye, with

a short black stripe inside it. Head round, length

beyond anterior angle of eye equal to remainder of

head. Face with one wide black band along dorsal

margin. Male genitalia: aedeagus short, stout, and

cylindrical, with one pair lobe-like terminal processes,

each with a small spine near base. Pygofer with distinct

processes on ventral margin. Sternal apodemes short,

widely separated.

Remarks: The holotypes and paratypes of Beamer*s

species, D. plana and D. minuta, were examined, and no difference between these specimens was found except

D. minuta has slightly longer head which can not be justified as species. Because these two species were described in the same paper, and D. plana was published on the page before D. minuta. D. plana has priority and D. minuta is considered a synonym. 63

Diagnosis: This species can be separated from the other species by the following male characters:

1) forewing with reticular stripes; 2) only one pair of terminal aedeagal processes present.

Distribution: Arizona, Kansas, New Mexico,

Wyoming.

Type and Material Examined: Male holotype, Cochise,

Arizona, VIII-24-1935, R.H. Beamer. 1 female, Willcox,

Arizona, X-16-1931, E.D. Ball. 1 male (holotype of

D. minuta). 2 males (paratypes of D. minuta), Laramie,

Wyo., VI-23-1935, R.H. Beamer. 1 male, 3 females,

Willcox, Arizona, VI-11-1936, E.D. Ball. 2 females

(paratypes of D. minuta). Laramie, Wyo., VII-13-1937,

C.L. Johnston. 1 male, Masgrav's Black R., Arizona,

VII-14-1936, E.D. Ball (NMNH). 1 female, Moriarty,

N. Mexico, VI-20-1940, D.E. Hardy. 1 male, 2 females,

Moriarty, N. Mexico, VI-23-1941, E.L. Todd. 1 male,

Redwing, Kansas, VIII-6-1945, R.H. Beamer. 1 male,

Moriarty, N. Mexico, VI-20-1940, D.E. Hardy (SEMC).

2 males, Chiricahua Mt. Ari., VII-26-1952, D.J. and

J.N. Knull (OSUC). Dicyphonia plura Beamer

(Figs. 92-95)

Dicyphonia plura Beamer, 1936:68.

Female: Body length: 5.6-7.0 mm. General color much lighter than male. Thorax pale yellow ventrally.

Forewing reduced, leaving several abdominal segments exposed.

Male: body length: 4.0-4.4 mm. General color pale yellow. Head, pronotum, scutellum and forewing with color pattern as in D. plana. but lighter and forewing with narrower stripes. Lateral margins of head with slender, black stripes connecting anteriorly.

Apex of forewing black or dark brown. Abdomen black with a light band on genital segment in males. Male genitalia: Aedeagus stout with two pairs of pincer- like terminal processes. Apices of upper rami sharp, that of lower ones stout. Aedeagal shaft with wing-like processes on dorsal margins, subtriangular in cross section. Style and connective as D. plana. Pygofer with processes on ventral margins. Sternal apodemes short and widely separated.

Diagnosis: This species can be separated from the other species as follows: 1) male forewing with reticular stripes; 2) two pairs of terminal aedeagal

processes; 3) face without markings except stripe

along dorsal margin.

Distribution: Arizona, New Mexico.

Type and Material Examined: 1 male (paratype),

Cochise, Arizona, VIII-24-1935, Jean Russell. 1

female (paratype), labeled as above, but collected

by R.H. Beamer (NMNH). 1 female, Willcox, Ari., VI-

11-1936, E.D. Ball (OSUC). 57 males, Belen, N. Mexico.

VII-26-1936, R.H. Beamer (SEMC). The female holotype

is deposited in SEMC.

Dicyphonia ornata (Baker)

(Figs. 96-100)

Platymetopius ornatus Baker, 1900:49.

Dicyphonia pamentosa [sic] Ball, 1900:69.

Dicyphonia ornata (Baker), Van Duzee, 1908:158.

Female: Body length: 5.4-7.4 mm. General color

light brown. Head very produced, with length in middle much longer than width between eyes. Forewing reduced and leaving several abdominal segments exposed. Male: Body length: 3.9-4.1 mm; General color dark brown to black. Vertex acute, pattern on vertex, pronotum and scutellum similar to D. plana but much darker. Thorax and abdomen brown to black ventrally and face with several small oblique stripes and one band along dorsal margin. Male genitalia: Aedeagus short and robust with two pairs of pincer-like terminal processes, upper rami longer than lower ones. Aedeagus flattened laterally, wing-like processes on dorsal margins developed dorsally. Pygofer with dark processes on ventral margins.

Diagnosis: This species can be distinguished from the other species as follows: 1) male forewing with reticular stripes; 2) two pairs of terminal processes of aedeagus present; 3) several oblique stripes on face.

Distribution: Arizona, Colorado, Kansas, Nebraska,

South Dakota, Utah.

Type and Material Examined: 1 male (Holotype),

Horace, Kansas, VII-28-1891, C.F. Baker (NMNH). 1 male, 1 female, Cheyenne, Kansas, VIII-14-1932, D.A.

Wilbur (KSUC). 1 female, Salt Lake City, Utah, V-6-1934, E.W. Davis. 1 male, Salt Lake City, Utah, VII-13-1935,

P. Oman (NMNH). 1 female, Liberal, Kansas, VI-23-1941,

E.L. Todd (SEMC). 1 male, Meade, St. L. Kansas,

VII-22-1944, R.H. Beamer. 1 female, Scott, S. Pk.

Kansas, VIII-9-1945, R.H. Beamer. 1 male, Meade,

Kansas, VII-14-1945, R.H. Beamer. 1 female, Cheyenne,

Kansas, VII-7-1949, R.H. Eeamer (NMNH). 3 females,

Kinsley, Kansas, IX-1-1950, H.C. Severin (OSUC). 1

male, 3 females, Johns Town, Colorado, VII-26-1967,

P. Oman. 4 females, Stuart, Nebr. VIII-26-1967, P.

Oman. 1 male, Wash., Colorado, VIII-8-1973, J. Sawbridge.

1 female, Sahuarita, Arizona, VIII-6-8-1967, L.A.

Kelton. 3 males, 1 female, Denver, Colorado, VIII-11-1968

P. Oman (ORSU). 1 male, Apache Co., Arizona, VIII-25-1971

Harris and Harris. 2 males, 2 females, Stuart, Nebr.

VIII-6-1979, H.D. Blocker and R.A. Sweet. 4 males,

Yankton, S. Dakota, VIII-7-1979, H.D. Blocker and

R.A. Sweet. 1 female, Cherry Co., Nebr. VIII-10-1979,

H.D. Blocker and R.A. Sweet (KSUC). 3 males, Pueblo,

Colo., by H. Osborn, no date (OSUC).

Dicyphonia cona (Shaw)

(Fig. 113)

Parabolocratus conus Shaw, 1932:45.

Dicyphonia conica [sic] (Shaw), Beamer, 1936:69. Dicyphonia cona (Shaw), DeLong & Knull, 1946:27.

Female: Body length: 7.4-7.6 mm. General color

green. Head acute, length of head beyond eyes at

least twice as long as width between eyes. Vertex

with V-shaped stripe which is indistinct in posterior

1/2, with slender stripes on lateral margin. Face

and venter of body without stripes. Forewing short,

reaching 5th abdominal segment. Ovipositor extending

beyond pygofer, apex brown.

Male unknown.

Diagnosis: Females of this species are quite

distinct when compared to those of other species;

it can be separated by the short forewing and absence

of body stripes except for those on the head.

Remarks: This species was originally described

from a single female by Shaw (1932). Beamer (1936) moved it to Dicyphonia with the misspelled name,

conica. DeLong and Knull (1946) emended the name,

cona. In the present study, more female specimens were found in the Snow Museum collection.

Distribution: New Mexico. 69

Type and Material Examined: 1 female (Holotype),

Torrance, N. Mexico VI-12-1931, J.G. Shaw; 24 females,

Moriarty, N. Mexico, V-23-1941, R.H. Beamer. 2 females,

Estancia, N. Mexico, VII-30-1941, E.L. Todd (SEMC).

Dicyphonia zamorana (Linnavuori and DeLong)

(Figs. 101-105)

Jiutepeca zamorana Linnavuori and DeLong, 1978:51-52.

Dicyphonia zamorana (Linnavuori and DeLong), Reinstated

Length of male: 4.9-5.0 mm; Female: 5.0-5.5

mm. Linnavuori and DeLong (1978) gave a detailed

description, which was obviously based on the holotype.

Diagnosis: This species can be distinguished

from the following three species as follows: 1) distinctly

shaped black stripe on head and pronotum (Fig. 101);

2) female with stripe pattern similar to male of D.

plana; 3) aedeagus with one pair of terminal processes;

4) connective form distinctive (Fig. 102).

Remark: Six of the paratypes designated by Linnavuori

and DeLong are moved to new species, D. delongi and

D. mexicana respectively.

Distribution: Mexico. 70

Type and Material Examined: 1 male (holotype),

Zamora, Mich., Mexico, X-2-1941, DeLong, Good, Caldwell

and Plummer. 1 female (paratype), Guadalajara, Mexico,

X-3-1941, DeLong, Good, Caldwell and Plummer. 13

males (paratypes), 10 females, same data as holotype.

1 female, Saltillo, Coah., Mexico, IX-23-1941, DeLong,

Good, Caldwell and Plummer. 1 female, Lk. Chapala,

Jal., Mexico, X-3-1941, DeLong, Good, Caldwell and

Plummer. 1 male, Cuernavaca, Mor., Mexico, IX-25-1945,

Plummer, DeLong, Hershberger and Elliott. 1 female,

Mexico City, Mexico, IX-25-45, Plummer, DeLong, Hershberger

and Elliott.

Dicyphonia nigrita Ball

(Figs. 106-112)

Dicyphonia nigrita Ball, 1937:132.

Jiutepeca nigrita (Ball), Linnavuori and DeLong, 1978:50-51.

Dicyphonia nigrita Ball, Reinstated

Female: Body length: 4.8-5.5 mm. General color pale yellow, without stripes except on head. Forewing

fully developed and covering all abdominal segments.

Sternal apodemes with flattened terminals. Male: Body length: 3.8-4.0 mm. General color

pale yellow. Head, pronotum and scutellum with one

medial black stripe, widening posteriorly on pronotum.

Lateral margins of stripe on scutellum parallel or

narrowing posteriorly. Forewing black in apical 2/3

and inner margin of claval area. Face without stripes

apices of tibiae and tarsomeres brown. Male genitalia

Aedeagus long, with two pairs of sickle-like terminal

processes which are more than 1/2 length of shaft.

Upper pair of terminal processes longer than lower

ones and extending posteriorly. Aedeagal preatrium

equal to 1/2 length of shaft. Pygofer with processes

on ventral margins and with fewer than 20 large setae.

Connective form distinct (Fig. 108).

Remark: Ball (1937) did not describe the male

genitalia. Linnavuori and DeLong (1978) made an error

when they described the male genitalia, which was

based on a specimen that did not belong to D. nigrita.

Diagnosis: This species can be separated from

the other species as follows: 1) forewing black in apical 2/3; 2) medial stripe on pronotum widening

posteriorly; 3) terminal aedeagal processes equal

to 1/2 length of shaft. Distribution: Mexico.

Type and Material Examined: 4 males, 1 female

(Paratypes), Cuernv-Acap. Road, Mexico, VIII-29-1936,

Ball and Stone (NMNH). 1 male, Jiutepec, Morelos,

Mexico, IX-6-1939, DeLong. 3 males, Iguala, G ’ro.,

IX-11-1939, DeLong. 2 males, Chilpancingo, Guerrero,

Mexico, IX-10-1939, DeLong. 1 male, Buena Vista,

G'ro., Mexico, X-23-1941, DeLong, Good, Caldwell and Plummer. 1 male, Mazacian, G ’ro., Mexico, X-3-1945,

Balock, DeLong and Hershberger. 1 male, Yucatan Valladol

Mexico, IX-16-1986, J.A. Shuey. 5 males without labels

(OSUC). The holotype is deposited in NMNH.

Dicyphonia delongi. New Species

(Figs. 114-119)

Jiutepeca zamorana Linnavuori and DeLong, 1978:51-52

(in part).

DESCRIPTION: Length of male: 4.1-4.3 mm; Female:

5.0-5.5 mm. Body pale yellow. Head, pronotum and scutellum of male with one black medial stripe. Face pale yellow, without stripes except for band along dorsal margin. Thorax pale yellow ventrally. Forewing as in D. zamorana. Abdomen black, with light colored 73

band on genital segments. Fore and middle legs pale

yellow, hind legs with several brown stripes at apices

of tibiae and protarsomeres. Female pale yellow,

body without stripes except on head; forewings fully

developed; ovipositor hardly exposed, with apex brown.

Aedeagus cylindrical, with two pairs of terminal processes

extending posterolaterally, upper pair subtriangular

and longer than lower pair. Aedeagal preatrium short,

about 1/3 length of shaft. Connective narrow and

long. Pygofer brown, with more than 20 setae on each

side. Sternal apodemes narrow, close together and

rounded apically.

Diagnosis: This species is similar to D. nigrita.

The distinguishing characters are as follows: 1) stripe

on pronotum and scutellum (Figs. 116); 2) relatively

smaller size of aedeagus (Figs. 115); 3) subtriangular

shape of upper aedeagal terminal processes (Figs.

114, 115); 4) relatively narrower sternal apodemes

(Figs. 119); 5) connective form (Figs 117); 6) with

more setae on pygofer.

Distribution: Mexico.

Holotype: Male, Jiutepec, Morelos, Mexico, IX-6-1939,

DeLong and Plummer. 74

Paratypes: 40 males, 53 females, same data as

holotype. 1 male, Taxco, Guerrero, Mexico, IX-9-1939,

DeLong and Plummer. 2 males, Mexico City, Mexico,

IX-1-1939, DeLong. 4 males, Acapulco, Guerrero, Mexico,

IX-10-1939, DeLong and Plummer. 1 male, Iguala, G ’ro.,

Mexico, IX-11-1939, DeLong. 1 male, Chilpancingo,

Guerrero, Mexico, DeLong. 1 male (paratype of Jiutepeca

zamorana). Zamora, Mich. Mexico, X-2-1941, DeLong,

Good, Caldwell and Plummer. 1 male, Cuernavaca, Mexico,

XI-25-1945, DeLong, Hershberger and Elliott. 2 males,

Mazacian, G ’ro., Mexico, X-3-1945, Balock, DeLong

and Hershberger. 2 males, Tierra Colorado, G ’ro.,

Mexico, X-5-1945, without collector name.

All types are deposited in OSUC.

This species is named in honor of the late D.

M. DeLong in recognition of his many contributions to the study of leafhoppers.

Dicyphonia mexicana, New Species

(Figs. 120-124)

Jiutepeca zamorana Linnavuori and DeLong, 1978:51-52

(in part). 75

DESCRIPTION: Length of male: 4.4-4.6 mm; Female:

4.1-4.3 mm. Male body pale yellow. Head, pronotum

and scutellum with stripe pattern similar to D. delongi.

except for one brown band along posterior margin of

pronotum and one pair of brown spots above the band.

Also, scutellum brown under suture instead of black

and with one pair of spots at corners. Face with

several oblique stripes and wide, cross band ventral

to eyes (Fig. 120). Thorax black ventrally. Legs

and abdomen with color pattern similar to D. nigrita

and D. delongi. Forewing similar to D. nigrita except

for stripe in claval area which extends more anteriorly.

Female with fully developed forewing, and ovipositor

hardly exposed. Unlike the above two species, female

head, pronotum and scutellum with stripe pattern similar

to that of male but color slightly lighter. Male

genitalia: Aedeagus short, straight, with two pairs

of terminal processes extending posterolaterally, upper pair less than or equal to length of lower pair.

Connective short, branches widely separated. Sternal apodemes similar to D. delongi.

Diagnosis: This species can be easily distinguished

from the other species as follows: 1) male forewing black in apical 2/3; 2) pronotum with median longitudinal black stripe; 3) stripes on face and black band ventral

to eye; 4) venter of the thorax black.

Distribution: Mexico.

Holotype: male, Zimipan, Hdg., Mexico, X-3-1945,

DeLong, Hershberger and Elliott.

Paratypes: 1 male, Jiutepec, Morelos, Mexico,

IX-6-1939, DeLong and Plummer. 9 males (including

one of the paratypes of Jiutepeca zamorana). Saltillo,

Coah., Mexico, IX-23-1941, DeLong, Good, Caldwell

and Plummer. 3 males (paratypes of J. zamorana).

Zamora, Mich., Mexico, X-2-1941, DeLong, Good, Caldwell

and Plummer. 1 male, Lk. Chapala, Mexico, X-3-1941,

DeLong, Good, Caldwell and Plummer (paratype of J.

zamorana). 2 males, Guadalajara, Jal., Mexico, X-3-1941,

DeLong, Good, Caldwell and Plummer. 2 males, Bosencheve,

Mexico, IX-28-1945, Plummer, DeLong, Hershberger

and Elliott. 2 males, Morelia, Mich., Mexico, IX-30-1945

Plummer, DeLong, Hershberger and Elliott. 1 male,

Mazacian, G ’ro., Mexico, X-3-1945, Balock, DeLong

and Hershberger. 1 male, Mexico City, Mexico, X-22-

1945, Plummer, DeLong, Hershberger and Elliott. 1 male, Tulancingo, Hdg., Mexico, X-25-1945, Stone,

DeLong, Hershberger and Elliott. 2 females, Chilpancingo 77

Guerrero, Mexico, X-10-1939, DeLong and Plummer.

1 female, Iguala, Guerrero, Mexico, IX-11-1939, DeLong.

1 female, Buena Vista, Guerrero, Mexico, IX-11-1939,

DeLong and Plummer.

All types are deposited in the OSUC.

Phylogeny of Dicyphonia Species

The phylogenetic relationships among species of Dicyphonia were studied using the PAUP (Phylogenetic

Analysis Using Parsimony), version 2.4, developed by Swofford (1985). A total of 14 characters have been examined, and their descriptions and polarized conditions by comparison with the outgroup are listed in Table 1 and the data matrix derived is shown in

Table 2. Eight taxa including the outgroup were used.

There are some multiple-state characters, but in this study they are polarized into either 1 (apomorphic) or 0 (plesiomorphic) instead of the multiple-states.

The reason for that is that it is difficult to prove the way the evolution happened, namely 1 to 2, or 1 to 3 or otherwise, and the UNORDER option of PAUP can not be used in this case since obviously the other states are from the first apomorphic state (1) but not the plesiomorphic one (0) as to the characters

6, 7 and 14 (Table 1). According to the study mentioned 78

in Chapter I, Dicyphonia Ball is most closely related

to the genus Memnonia Ball, because they share two

synapomorphic characters: small (males < 5 mm in length)

and all have processes on the ventral margins of the

male pygofer. Therefore, M. consobri na Bal], generic

type, has been chosen as the outgroup in this analysis.

Since most of the characters for D.icyphonia

classifications are from male individuals and the

male of the species, D. cona (Shaw), is unknown,

this species is not included in the cladistic analysis.

In the analysis, PAUP option, BAN'DB is used,

and only one most parsimonious tree (Fig. 125) was

found, which represents the hypothesized phylogenetic

relationships among the species of Dicyphonia. The total consistent index = 0.792, and tree length =

24.000. The outgroups are not shown on figure 125 because it is not necessary.

From the cladogram, Dicyphonia is divided into two groups which is strongly supported by the analysis, and also by the current distribution data in this sudy.

Besides, the relationship among' species is also clear which is also strongly supported by the characters. 79

TABLE 5. Characters of Dicyphonia used in the cladistic analysis and their polarized conditions by outgroup comparison. Codes (6* etc.) represent the same characters but different states

Characters Plesiomorphic Apomorphic

1. V-shaped stripe on absent present anterior margin of crown 2. Second apical cell open posteriorly open outwardly 3. Pronotum no colored spots with many spots 4. Forewing color unicolorous black in apical 2/3 5. Aedeagal shaft not swollen swollen in middle 6. Terminal aedeagal one pair two pairs, pincer processes like (6’), or upper rami triangular (6 *’) 7. Middle stripe on none covers pronotum whole pronotum ( 7 ’ ) i or narrow (Fig. 106) (7 * *) 8. Black band ventral absent present to eyes on gena 9. Female forewing short, several long, abdominal segments ovipositor exposed exposed

10.Ventral color of as other part of different thorax body from other part of body 11.Abdomen color brown black except genital segments 12.Male forewing unicolorous with reticular stripes Table 5 (continue), 13.Aedeagus curved ventrally curved in S-shape 14.First pair of absent present, connective processes (Fig. 98) (14 ’ ) TABLE 6. Data Matrix of polarized characters

or character states of Dicyphonia for

cladistic analysis. 1 = apomorphic state,

0 = plesiomorphic state

Characters or character states

111111

Taxa 1 2 3 4 5 6 6 ’6"7 7 * 7 " 8 9 0 1 2 3 4 4*

Outgup* 0 0 0 0 0 0 0 000000000000 plana 1110 0 0 0 000000101000 ornata 1110 111 000000101000 plura 1110 111 000000101000 zamorana 1 1 0 10 0 0 011011110100 mexicana 1 1 0 10 10 110111110010 nigrita 1 1 0 10 10 010101010111 delongi 1 1 0 1 0 1 0 110101010111

* outgroup = Memnonia consobrina CHAPTER IV

SYSTEMATIC STUDY OF THE GENUS

Hecullus OMAN

Genus Hecullus Oman

Hecullus Oman, 1949:31. Type: bracteatus Ball, 1901:4-

5, under Hecalus. by original designation.

Hecullus was established by Oman in 1949 based

on Hecalus bracteatus Ball. He gave a detailed description

of the genus in his paper. Hecullus is a very distinctive

genus and can be separated from other genera of Hecalini

by the female head form and short forewing (Fig.

126), according to Oman (1949). In this study, the

following male characters were found useful in identifying

the genus: 1) head without carina on anterior margin;

2) small aedeagus with a long fused connective (Fig.

130); and 3) genital plate with about 10 setae on

lateral margin. Hecullus is found only in the United

States.

Hecullus Oman is represented by only two species,

H. balli (Beamer) and H. bracteatus (Ball), and they

82 83 can be distinguished by following key:

Key to species of Hecullus Oman

1. Female head long, at least 3 times as long as

width between eyes; aedeagus stout, with basal

processes swollen at base; sternal apodemes not

well developed ...... balli (Beamer)

Female head about 1.5 times as long as width between

eyes; aedeagus acute apically, with basal processes

not swollen at base; sternal apodemes large

...... bracteatus (Ball)

Hecullus bracteatus (Ball)

(fig. 126-133)

Hecalus bracteatus Ball, 1901:4-5.

Female: Body length: 6.50-6.75 mm. General color pale yellow, with some brown longitudinal stripes on dorsal surface of head, pronotum and abdomen, and forewing. Head broadly produced, thin, foliaceous, with length 1.5 times the width between eyes. Pronotum short, less than 1/2 length of crown. Forewing and venation much reduced, barely reaching second visible abdominal segment. Ovipositor not extending past pygofer. Male: Body length: 4.55-4.90 mm. General color

pale, with some brown longitudinal bands on dorsal

surface of head and pronotum and forewing. Head not

much produced, length shorter than width between eyes.

No carina at anterior margin of crown. Forewing well

developed, with most margins of distal cells light

brown and veins pale. Venation as Hecalini pattern

except some additional cross-veins. Male genitalia:

Pygofer long, surface smooth, with a few setae on

dorsal-posterior corner. Genital plates with about

10 setae on lateral margin. Aedeagus small, with

two large basal processes and shaft curved dorsally.

Connective distinctive, fused and linear, like many

genera of Deltocephalinae but not genera of Hecalini.

Distribution: Arizona, California, New Mexico,

Texas.

Type- and Material Examined: 1 female, Rockey

Ford, VI-16-1900; 1 male, L. Animas, [probably N.M.],

VII-17-1901; 1 male, 2 females, Ashland, VIII-16-1929, without collector name. 2 males, Hurley, N. Mexico,

VIII-25-1935, R.H. Beamer (NMNH). 2 females, Hualpai

Mts., Ari., VII-4-1937, D.J. and J.N. Knull. 1 male,

Huachuca Mts. VII-20-1937, D.J. and J.N. Knull. 1 85

male, Wickenburg, Ari., VIII-20-1938, D.J. and J.N.

Knull. 2 females, Lincoln Co., N.M. VII-9-1940, D.J.

and J.N. Knull. 3 females, Cottonwood Sp. Ari., VIII-

22-1940, D.J. and J.N. Knull. 2 females, Davis Mts.

Tex., V-9-1941, D.J. and J.N. knull. 9 males, 50

females, Santa Rosa Mts. California, V-27-1946, D.J.

and J.N. Knull. 4 females, Dragoon Mts. Ari., IX-

10-1947, D.J. and J.N. Knull. 8 males, 1 female,

Chiricahua M., Arizona, IX-4-1962, D.J. & J.N. Knull

(OSUC). 1 female, People’s Valley, VIII-18-1967,

L.A. Kelton. 1 male, 1 female, Blue Water Village,

N. Mexico, VIII-25-1970, Haris & Haris, G.L. 1 male,

Santa Cruz, Arizona, VII-28-1972, D. Baine (BRI).

Types are deposited in SEMC.

Hecullus balli (Beamer)

(fig. 134-138)

Hecalus balli Beamer, 1937:12.

Female: Body length: 8.30-9.05 mm. General color

pale yellow, with very light brown stripes on head and pronotum. Head well produced, length longer

than 5 times width between eyes, with middle longitudinal ridge. Pronotum short and narrower than head including eyes. Forewing and venation very reduced, leaving most abdominal tergites exposed. Ovipositor not extending beyond pygofer.

Male: Body length: 4.85-5.30 mm. General color pale, with forewing sordid white. Head produced more broadly and roundly than H. bracteatus. with length slightly longer than width between eyes.

Forewing well developed, with normal Hecalini venation pattern plus additional cross-veins. Two pairs of small sternal apodemes present, not extending more than half length of first visible abdominal segment.

Male genitalia: Much like H. bracteatus but phragma of aedeagus elongated, as long as shaft, and basal processes enlarged at base.

Distribution: Arizona.

Type and Material Examined: 4 males, Benson,

Arizona, IX-10-1935, E.D. Ball. 1 male, Tucson, Arizona,

IV-24-1936, E.D. Ball. 1 male, 3 females, Arizona,

IX-6-1936, E.D. Ball (NMNH). Types are also deposited in NMNH. CHAPTER V

SYSTEMATIC STUDY OF GENUS

Neohecalus LINNAVUORI

The oldest argument in Hecalini involves the genera Hecalus Stal and Parabolocratus Fieber. Stal

(1897) claimed that Parabolocratus was a synonym of his genus Hecalus. After Linnavuori (1961) confirmed that these two genera actually shared the same generic types, Parabolocratus has been treated as a synonym of Hecalus. However, there is still some confusion about this group of leafhoppers from the Nearctic region because they are easily recognized as two groups or genera, one of which was named Hecalus and the other

Parabolocratus. In order to solve the problem, Evans

(1966) described a genus Linnavuoriella. to contain the species without produced heads in the males.

Morrison (1973) synonymized Linnavuoriella with Hecalus because there were too many intermediate states among the male heads. However, the species originally under

Hecalus are quite distinctive from the real Hecalus species. For them, Oman (1949) gave a very clear definition. Therefore, Linnavuori (1975) established a new genus for them, Neohecalus.

87 Neohecalus Linnavuori

Neohecalus Linnavuori, 1975:55. Type: lineatus Uhler

1877:463, under genus Glossocratus. by original

designation.

DESCRIPTION: Elongate, broad and flat-headed

leafhoppers. Sexual dimorphism obvious.

FEMALE: General color from sordid yellow to

sordid green, and usually without markings. Head

broadly produced, thin and foliaceous, with carina

at anterior margin. Ocellus usually near eye and

distance between them shorter than or equal to ocellus

diameter. Eye usually small and cross diameter much

less than distance from it to middle suture. Pronotum

slightly narrower than head including eyes, and long

carinae present at lateral margins. Episternum exposed

slightly. (It is completely covered by gena in other genera of Hecalini.) Forewing subbrachypterous, apical

region usually reduced. Abdomen long, equal to or

longer than rest of body. Ovipositor extending well beyond pygofer which is well developed and longer than or equal to combined length of last two pre-genitalic segments. MALE: General color like females, but usually with some dark-colored spots or stripes. Head broadly produced, foliaceous, with carina at anterior margin.

Ocellus location usually like female. Pronotum slightly narrower than head including eyes. Forewing usually well developed, venation different from Hecalus by

R vein which connects with outer anteapical cell at inner corner instead of outer one. This character is shared only by Dicyphonia Ball in Hecalini. Abdomen usually dark-colored and with two pairs of sternal apodemes. MALE GENITALIA: Pygofer long, at least

1.5 times longer than length of aedeagus, with setae usually arranged in definite groups. Aedeagus relatively simple with one pair of terminal processes, Style, genital plate and connective like Hecalus.

There are only two species in Neohecalus. and they are recorded from North America. These two species can be distinguished by the following key.

Key to species of Neohecalus

1. Red longitudinal stripes on head, pronotum and

scutellum; male forewing with two black bands

near disc cells and apical region ......

...... lineatus (Uhler) 90

Brown longitudinal stripes on head, pronotum and

scutellum; male forewing without black band .....

...... apicalis (Van Duzee)

Neohecalus lineatus (Uhler)

(Fig. 139-141, 145-148)

Glossocratus lineatus Uhler 1877:463.

Hecalus lineatus (Uhler), Signoret 1879:267.

Female: Body length: 11.0-12.1 mm. General

color pale yellow, with several red, longitudinal

stripes on head, pronotum and scutellum. Head thin

and produced, with length 1.5 times as long as width

between eyes. Forewing fully developed and only ovipositor

exposed.

Male: body length: 5.6-6.1 mm. General color pale yellow, with several red, longitudinal stripes on head, pronotum and scutellum. Two black bands on disc cells area and apical region of forewing.

Head produced, length slightly longer than width between eyes. Male genitalia: Pygofer long, with two groups of setae, one near apex and one near base. Aedeagus flattened laterally, with one pair of terminal processes near apex. Two pairs of sternal apodemes, with lateral 91 pair very large (Fig. 146).

Distribution: Illinois, Iowa, Kansas, Nebraska,

Oklahoma.

MATERIAL EXAMINED: 3 males, 4 females, Ames,

Iowa, VIII-14-1916, H. Osborn, 1 male,, 1 female,

Lincoln, Nebr., VI-30-1931, D.M. Johnson, 1 male,

Evergreen 111. IX-9-1934, D.M. DeLong. 1 female,

Tulsa Okla. VII-22-1937, Standish-Kaiser, 7 males,

25, females, Valley, Nebr. VI-30-1938, D.J. and J.N.

Knull, 1 male, Douglas Co., Kans., by F.H. Snow, no date (OSUC).

Neohecalus apicalis (Van Duzee)

(Fig. 142-144, 149-153)

Hecalus apicalis Van Duzee 1909: 217.

Female: Body length: 14.6-15.5 mm. General color pale green. Head very produced, with length

2.5 times as long as width between eyes. Forewing reduced, only reaching 5th abdominal segment. Ovipositor long, with exposed part longer than pygofer. PLEASE NOTE:

Duplicate page number(s); text follows. Filmed as received.

UMI Male: Body length: 6.4-6.8 mm. General color light brown, with several brown, longitudinal stripes on head, pronotum and scutellum. Head produced, with length equal to width between eyes. Forewing developed, with base of claval area and apical region dark brown.

Male genitalia: Pygofer long, with three groups of setae (Fig. 144). Connective processes obvious.

Aedeagus swollen near apex of shaft, with one pair of terminal processes, which are longer than aedeagal shaft. Two pairs of sternal apodemes, lateral pair larger than middle one.

Distribution: Florida.

Material Examined: 1 male, Daytona Florida,

IX-5-1926, E.D. Ball. 1 female, Sanford Florida,

IV-22-1926, E.D. Ball. CHAPTER VI

SYSTEMATIC STUDY OF THE GENUS

Spangbergiella SIGNORET

Spangbergiella Signoret

Spangbergiella Signoret, 1879:273. Type: vulnerata

Uhler, 1877:464, under genus Glossocratus. by

original designation.

Bergiella Baker, 1897:157.

Spangbergiella was erected by Signoret (1879)

for the species Glossocratus vulnerata which was

described by Uhler (1877). Berg (1884) added three

species, S. felix. S. punctato-guttata. and S. uruguayensis.

all from South America. Baker (1897) described another

species, S. mexicana from Mexico, and transferred one of Berg’s species, Hecalus lynchii. to Spangbergiella.

Lawson (1932) published the first revision of

Spangbergiella for North America. In that paper, he described S. quadripunctata and two subspecies,

S. vulnerata convexa and S. vulnerata lativittata.

Linnavuori (1957) treated S. lynchii (Berg) as a synonym 93 of the subspecies S. vulnerata lacerdae (Signoret).

Since S. lacerdae Signoret was synonymized with S. vulnerata by Signoret himself (1879), S. lynchii (Berg) is here treated as a synonym of S. vulnerata (Uhler).

Oman’s (1938) and Linnavuori’s (1957) treatment of

S. lacerdae Signoret is in error as S. lynchii is not distinct from S. vulnerata. More recently, Linnavuori and DeLong (1977, 1978) added two additional species,

S. reticulata and S. erratica. respectively, bringing the total to eight species in Spangbergiella.

Spangbergiella has been redescribed by Oman (1938,

1949) and Linnavuori (1957). There are three distinct characters that separate Spangbergiella from other genera of Hecalini. These are: 1) distinctive venation of the forewing which differs from other Hecalini in the reduced pattern and in having only one claval vein (Figs. 154, 157, 165); 2) red or orange stripes which are usually arranged in a distinctive pattern on the head and pronotum, and occasionally on the forewing of both males and females (Fig. 157); and

3) the female head which is always foliaceous with the length as long as the width between the eyes.

The members of this genus occur only in the New World.

The five species presently recognized in Spangbergiella 94 for the Nearctic and Neotropical regions may be distinguished by the following key.

Key to the species of Spangbergiella

Forewing with reticular veins (Fig. 154) ......

reticulata Linnavuori and DeLong

Forewing without reticular veins (Fig. 157)

2

Forewing usually without red stripes; if stripes

exist, they are orange; abdomen pale green

vulnerata (Uhler)

Forewing with red stripes at least in males;

abdomen usually dark brown ...... 3

Aedeagus with two terminal processes (Fig. 173)

...... erratica Linnavuori & DeLong

Aedeagus with three terminal processes (fig.

160) ...... 4

Aedeagus with all terminal processes of equal

length (Fig. 160) mexicana Baker

Aedeagus with median process much shorter than

lateral processes (Fig. 168) ......

...... quadripunctata Lawson 95

Spangbergiella reticulata Linnavuori & DeLong

(Figs. 154-155)

Spangbergiella reticulata Linnavuori & DeLong, 1977:184.

Female: Length of body: 6.75 mm. Body pale green, without ventral markings. Head, pronotum, and forewing with red stripes arranged in distinct pattern (Fig.

154). Forewing slightly shorter than in females of other species; venation reticulate. Abdomen long, ovipositor exposed. Seventh sternum with posterior margin straight.

Male unknown.

Diagnosis: This species can be separated by the reticular venation on forewing from the following four species.

Distribution: Chile.

Type and Material Examined: 1 female, holotype,

Pailaco, Chile, 1-3-1968, D.M. DeLong (OSUC). 96

Spangbergiella mexicana Baker

(Figs. 156-162)

Spangbergiella mexicana Baker, 1897:157.

Female: Length of female: 6.0-6.1 mm. General

color of female slightly lighter than male. Head

always produced with anterior margin very thin. Forewing

with or without distinct red or orange stripes.

Male: Length of male: 5.1-5.7 mm; General color

pale green or pale yellow, Head with pair of red,

posteriorly diverging stripes. Pronotum with pair

of red, posteriorly diverging stripes and short median

red stripe. Forewing always with distinct red or

orange stripes; Male genitalia: Aedeagus short,

stout, curved posteriorly, bearing three terminal

processes of approximately equal length.

Diagnosis: This species is easy to separate

from the others by three terminal aedeagal processes;

they are all of equal length, and the aedeagal shaft

is not triangular in cross-section.

Species identification and the above description are based on the original description and that of Linnavuori (1957) in which he provided a detailed illustration and description of the types, and on specimens determined by Linnavuori and DeLong from OSUC.

Distribution: Kansas, Tennessee, Texas; Mexico.

Type and Material Examined: 2 males, Nashville,

Tenn., IX-11-1915, D.M. DeLong, 1 male, 1 female,

Knoxville, Tenn., IX-13-1915, D.M. DeLong, 1 male,

Donarin, Tenn., XI-19-1915, D.M. DeLong, 16 males and 5 females, Clarksville, Tenn., VII-4-1917, D.M.

DeLong, 1 male, Alhajueln, C. Z. 1-11-1921, J.G. Sanders,

1 male, 1 female, Verge, Chia., Mexico, V-22-1935, no collector, 1 male, Guadalajara, Mexico, VIII-22-

1937, W.E. Stone,^1 male, Valles, S.L.P.K. 475, XI-

5-1939, DeLong, Hershberger and Elliott, 1 male, 5 females, Mex., D.F. 8200ft. S. 43 kms, IX-3-1941,

D.M. DeLong, 1 male, Saltillo, Coah., Mex., IX-23-

1941, DeLong, Caldwell and Plummer, 1 female, Mexico

City, Mex., X-22-1945, no collector, 1 male, 2 females,

Tamazunchale, S.L.P. XI-22-1945, 3 males, Mex., Mor.,

7 mi S Tres Cumbres, VII-7-1975, C. Martinson (OSUC).

The types are deposited in NMNH.

Spangbergiella quadripunctata Lawson 98

(Figs. 163-169)

Spangbergiella quadripunctata Lawson, 1932:120.

Female: Length of body: 5.1-5.3 mm. General color pale green. Head and forewing with distinct red or orange stripes. Forewing fully developed and only ovipositor exposed.

Male: Length of body: 4.6-4.8 mm. General color pale brown. Head and pronotal stripes identical or similar to S. mexicana. Vertex slightly more acute than that of S. mexicana. Male genitalia: Aedeagus long, cylindrical, with three apical processes, median process only one-fifth as long as lateral ones.

Diagnosis: This species can be distinguished as follows: 1) aedeagal shaft not triangular in cross- section; 2) three terminal aedeagal processes; 3) middle terminal aedeagal process much shorter than lateral ones.

Distribution: Arizona, Kansas, Mississippi,

North Carolina, Texas.

Type and Material Examined: Six male and two female paratypes from SEMC were examined, 3 males, 4 females, Tupelo, Miss, XI-1920, D.J. and J.N. Knull,

5 males, Greenville, Miss., May, 1930, D.W. Grimes,

3 males, 6 females, LeLand, May, 1930, D.W. Grimes,

2 males, 1 female, Wiggins, Miss., VI-10-1933, D.W.

Grimes, 2 males, 1 female, Swing, N.C. VIII-20-1934,

C. Mayer, 1 female, Dayton, Tex., V-21-1939, D.J.

and J.N. Knull, 1 female, Val Verde Co., Tex. VI-14-

1949, D.J. and J.N. Knull (OSUC). According to Lawson

(1932), the holotype and some paratypes are deposited

in SEMC.

Spangbergiella erratica Linnavuori & DeLong

(Fig. 170-176)

Spangbergiella erratica Linnavuori & DeLong, 1978:196.

Female: unknown.

Male: Length of body: 4.9-5.0 mm. Body form

and color pattern similar to S. mexicana Baker except

slightly smaller. Male genitalia: Aedeagus (Figs.

173, 176) with only two terminal processes which are more slender than those of S. mexicana.

Diagnosis: This species can be separated from

the other species as follows: 1) aedeagal shaft not 100

triangular in cross-section; 2) only two terminal

aedeagal processes.

Distribution: North Carolina, Texas; Mexico.

Type and Material Examined: 1 male, holotype,

Tamazunchale, S.L.P. IX-14-1939, D.M. DeLong, 1 male,

paratype, date same as holotype, 1 male, paratype,

labelled as M.B. 286, Mexico, by A. Dampf (OSUC).

Spangbergiella vulnerata (Uhler)

(Figs. 177-184)

Glossocratus vulnerata Uhler, 1877:464.

Spangbergiella vulnerata (Uhler), Signoret, 1879:274.

Spangbergiella lacerdae Signoret, 1879:274.

Spangbergiella Felix (sic) Berg, 1884:34, 35. NEW

SYNONYM

Spangbergiella punctato-guttata Berg, 1884:35. NEW

SYNONYM

Parabolocratus uruguayensis Berg, 1884:36. NEW SYNONYM

Hecalus lynchii (Berg), 1897: 268; Linnavuori 1957:153.

Spangbergiella vulnerata vulnerata (Uhler), Lawson,

1932:117. NEW SYNONYM

Spangbergiella vulnerata convexa Lawson, 1932:118.

NEW SYNONYM 101

Spangbergiella vulnerata lativittata Lawson, 1932:118.

NEW SYNONYM

Female: Body length: 7.25-10.4 mm. General

color pale green. Female head produced, foliaceous,

anterior margin very thin. Forewing without any red

or orange stripes.

Male: Body length: 5.0-6.5 mm. General color

pale green. Anterior margin of male head round, not

thin, and without carina. Head of not much produced,

length less than width between eyes. Vertex and pronotum

of both sexes with pair of red stripes diverging posteriorly,

occasionally with short median stripe. Pattern of

stripes on head highly variable. Red or orange stripes

usually absent from forewing of both sexes. Male genitalia: Aedeagus short, stout, and in cross-section

somewhat triangular-shaped, bearing a pair of lobe-like terminal processes.

Remarks: In 1957, Linnavuori suggested that

S. uruguayensis Berg might be a subspecies of S. vulnerata

(Uhler). In this study, the male lectotype of S. uruguayensis. the male lectotype (from Uruguay) and male paralectotype (from Argentina) of S. punctato-guttata. the male lectotype of S. felix. which were all obtained 102

from LPUA, and female lectotype and female paralectotype

of S. vulnerata (Uhler) from USNM were studied.

Besides the type specimens, over two hundred specimens

from the United States, South America and Caribbean

region were examined. Although the male lectotype

of S. punctato-guttata Berg has a slightly longer

head, the three species have the same general appearance

in both males and females and the male genitalia

identical to S. vulnerata (Uhler). This is more

obvious when comparing the variation of characters

of other species of Spangbergiella. In addition,

Berg (1884) defined his three species based on the

stripe pattern and the length of the head, however,

the specimens in the study show a great variation

in body length, color and shape of the head stripe

(Figs. 177, 178, 179, 185). Therefore, S. felix.

S. punctato-guttata and S. uruguayensis are synonymized

under S. vulnerata (Uhler).

Lawson (1932) divided S. vulnerata into three

subspecies, and provided a key to them based on the variation of the head length and the stripe pattern.

After examining the holotypes of Lawson’s subspecies and hundreds of other specimens from North and South

America, I found that all males of S. vulnerata lack a carina on the anterior margin of the crown. Furthermore, the variation of the stripe pattern on the head and

pronotum is so great with so many intermediate states

that the establishment of taxa based on these characters

is not warranted. Therefore, the three subspecies

of S. vulnerata are treated as synonyms of S. vulnerata

(Uhler).

Diagnosis: This species can be separated from

the other species by the triangular aedeagal shaft

in cross-section and absence of red stripes on the

forewing.

Distribution: Arizona, Arkansas, Florida, Kansas,

Maryland, Mississippi, Oklahoma, South Carolina, North

Carolina, Tennessee, Texas, Virginia; Argentina; Brazil;

British Guiana; Cuba; Peru, Uruguay.

Type and Material Examined: Types are deposited

in Argentina, 1 female, Bartica, Br. Guiana, III-

20-30-1901, H. Osborn, 2 males, Raleigh, N. C. late

July, 1909, Z.P. Metcalf, 1 female, Orangeburg, S.C.

VIII-20-1914, F.H. Lathrop, 1 male, Knoxville, Tenn.,

IX-15-1915, D.M. DeLong, 1 male, New Smyrna, Fla.

V-1920, M. Wright, 2 males, Cape Charles, Va., VIII-

1-1920, D.M. DeLong, 1 male, LaBelle Fla. IV-19-1921,

D.M. DeLong, 8 males, Paradise Key. Fla. IV-501921, 104

D.M. DeLong, 3 males, 2 females, Orlando, Fla. IV-

29-1921, D.M. DeLong, 10 males, 12 females, Ospr.

Miss., 11-14-1922, H. Osborn, 1 female, Paeston, Cuba,

1-12-1925, H. Osborn, 2 males, Aguirra, P.R. 1-18-

1929, H. Osborn, 3 males, 2 females, Salinas, P.R.

111-12-1929, H. Osborn, 2 males, 1 female, Brownsville,

Tex., VI-1-1934, J.N. Knull, 1 female, Eufaula, E.A.

Sooter, VII-6-1935, no collector, 2 females, Lebanon,

Okla. VII-2-1937, Standish-Kaiser, 1 female, Oswalt,

Okla. VII-3-1937, Stadish-Kaiser, 1 female, Rolf,

Okla. VII-15-9137, Stadish-Kaiser, 1 female, Ada,

Okla., VII-16-1937, Stadish-Kaiser, 1 male, Brownsville,

Tex., VIII-8-1937, D.J. and J.N. Knull, 2 males, 1 female, Fort Davis, Texas, IX-20-1938, D.J. and J.N.

Knull, 1 male, Dade Co., Fla. V-12-1939, D.J. and

J.N. Knull, 2 males, Bentonville, Ark., IX-4-1940,

H. Osborn, 6 males, 6 females, Chiricahua Mts. Ari.,

VIII-28-1940, D.J. and J.N. Knull, 1 male, Amazonas,

Brazil, 11-19-1943, L. Berg Jr., 2 males, Huachuca

Mts. Ari., VIII-10-1953, D.J. and J.N. Knull, 2 males,

Piracicaba, Sao Paulo, Brazil, II-8-1965, C.A. Triplehorn,

9 males, Machu Picchu, Peru, XI-25-1967, D.M. DeLong

(OSUC). 105

Phylogeny of Spangbergiella Species

PAUP was used to conduct the phylogenetic study

of the species of Spangbergiella. According to the

analysis mentioned in Chapter I, one outgroup is chosen

from the genus Hecalus: H. flavidus. There are 14

characters examined in the study; the polarized resuits

are shown in Tab]e 7, and the data matrix obtained

is listed in Table 8.

There is a single most parsimonious tree found

by using command, BANDB, and the consistent index

is 0.929, with free length is 14. The hypothesized

relationship between the species of Spangbergiella,

represented by the tree or cladogram is shown on figure

1 86 .

From the tree we can see that ret.i culata, rnj^xi.cana, erratica. and quadri punctata are related, which has

strong' support from characters 5, 6, 7 and 10. Above

that point, each branch is supported only by a single character. 106

TABLE 7. Characters used in the phylogenetic analysis of the species of Spangbergiella and their polarized condition

Characters Apomorphic Plesiomorphic

1. red stripes on body present absent 2 . claval vein one two 3. reticulate venation present absent 4. cross-section of aedeagus triangular round 5. black spots on male forewing present absent 6. red stripes on male forewing four none 7. male head produced pointedly roundly 8. black stripes along anterior present absent margin of crown 9. aedeagus shaft curved curved dorsally ventrally 10 . red or orange stripes on present absent female forewing 11 . light brown stripes on present absent male forewing 12 . posterior margin of male without with pronotum angle angle 13 . female head produced pointedly roundly TABLE 8. Data matrix of the species

of Spangbergiella. l=apomorphic

states, O=plesiomorphic states

Characters

Taxa 1 2 3 4 5 6 7 8 9 10 11 12 1

H. flavidus 0 0 0 0 0 0 0 0 0 0 0 0 0 reticulata 1 1 1 0 9 9 9 9 9 1 9 9 0 mexicana 1 1 0 0 1 1 1 0 1 1 0 1 1 erratica 1 1 0 0 1 1 1 1 0 1 1 0 9 vulnerata 1 1 0 1 0 0 0 0 0 0 0 1 0 quadri- 1 1 0 0 1 1 1 1 0 1 0 1 1 CHAPTER VII

SYSTEMATIC STUDY OF THE GENUS

Hecalus STAL

Genus Hecalus Stal

Hecalus Stal, 1864:65, Type: paykulli Stal, 1854:252,

under genus Petalocephala. by original designation.

Parabolocratus Fieber 1866:502.

Columbanus Distant, 1916:22.

Linnavuoriella Evans, 1966:34.

Hecalus was established by Stal in 1864 based

on the female of Petalocephala paykulli Stal, which

is deposited in the Naturihustoriska Riksmuseet, Stockholm.

Fieber (1866) described the genus Parabolocratus from

a female of P. glaucescens Fieber which is deposited

in the Museum d'Histoire Naturella in Geneva, Switzerland.

Stal (1870) declared that Parabolocratus was a synonym

of his Hecalus. However, his statement was ignored until Linnavuori (1961) examined the both types of the two genera and found "the generotype of the genus

Hecalus Stal is congeneric with species of Parabolocratus 109

Fieber." Morrison (1973) also examined the types

of both genera and reached the same conclusion.

After the establishment of the synonymy of Parabolocratus

with Hecalus. Evans (1966) described the genus Linnavuoriella

to contain the species whose heads are not spatulate.

However, Morrison (1973) considered Linnavuoriella

a synonym of Hecalus because he was convinced that

there is too much variation and intermediate states

among the heads of Hecalus. and the other characters

of both genera are essentially the same.

It is a judgement that Morrison (1973) mistreated

the genus Thomsoniella Signoret as synonym of Hecalus.

After examining several species of Thomsoniella. it

is obvious that Thomsoniella is a good genus and has

very close relationship, not with Hecalus but, with

Spangbergiella Signoret, because these two genera

have one claval vein on the forewing and a distinct

pattern of stripes on the head, pronotum and forewings.

As to the only two species which originally were thought

to be members of Hecalus from the Nearctic region,

it was observed that their characters, such as head

form, male pygofer and aedeagus, are not only rather different from those of Hecalus. but also distinctive

in Hecalini. Therefore, a new genus, Neohecalus 110

was established by Linnavuori which was preciously

mentioned for these two species, namely N. lineatus

and N. apicalis.

Generic Description: Body form usually elongated

and flattened. General color from sordid brown to

pale yellow, a few species with distinct stripes

or other markings on body and forewing. Male usually

slightly darker than female. Head usually produced

either roundly or sharply, with carina at anterior

margin. Ocellus usually near eye but distance between

them larger than ocellus diameter. Face convex or

rather flattened, with or without tiny hairs. Pronotum

usually as long as crown, with long carina along lateral

margin. One omega-shaped line present across pronotum

and part after which rufous in male. Forewing usually

well developed in male and reduced variously in female.

Venation generalized, with four apical cells and three

anteapical cells, as in many genera of Deltocephalinae.

In the case of reduced forewing, apical region usually

very short, and discal veins also reduced. Hind femur

spinulation usually 2-2-1. Abdomen usually long in

female, and two pairs of sternal apodemes present

in both male and female. Male genitalia: Male genitalia

form generalized. Pygofer usually elongated, with dark-colored band or stripe on dorsal surface in some Ill species, having posterior half covered by setae.

Style and genital plates as in description of tribe and with little value in separation of species. Aedeagus simple, cylindrical shaft curved upward, with one or two pairs of terminal processes. In some species preatrium enlarged and elongated. Several species share one common aedeagal pattern, so-called "major" pattern, shown in Figs. 236, 237. In this group, the variation of the aedeagus is not as great as in other genera of Hecalini.

Hecalus is a global genus. In the Nearctic region

18 species are recognized presently, and their males can be distinguished by the following key:

Key to males of Hecalus

1. Aedeagus "major" pattern (Fig. 236) ...... 2

Aedeagus not "major" pattern ...... 10

2. Body with obvious markings ...... 3

Body without obvious markings ...... 4

3. Crown beyond anterior angle of eye and pronotum

black nigrafasciatus (Beamer)

Crown yellow grey with anterior margin brown and

pronotum with two faint bands at middle ..

...... nimbosus (Ball) Apical region of forewing very short

...... inflatus (DeLong)

Apical region of forewing normal ...... 5

Aedeagal shaft swollen near apex, pygofer with posterior margin round ...... rotundus (DeLong)

Aedeagal shaft not swollen near apex, pygofer with posterior margin shape distinct (Fig. 190)

6

Aedeagus with extended preatrium and phragma shorter than 1/2 length of shaft...... 7

Aedeagal preatrium not extended and phragma longer than 1/2 length of shaft...... 9

Sternal apodemes longer than 1/2 length of first visible abdominal segment and aedeagal preatrium plus phragma longer than shaft planus (Shaw)

Sternal apodemes shorter than 1/2 length of first visible abdominal segment and aedeagal preatrium plus phragma shorter than shaft ...... 8

Crown length equal to width between eyes and aedeagal preatrium longer than phragma ... atascaderus (Ball)

Crown much shorter than width between eyes and preatrium shorter than phragma ... continuus (DeLong)

Crown length equal to width between eyes and pygofer without sinus on posterior-ventral angle

...... kansiensis (Shaw)

Crown much shorter than width between eyes and 113

pygofer with sinus on posterior-ventral angle ....

...... ma.ior (Osborn)

10. Face light brown or with black band along dorsal

margin and aedeagus with one pair of apical processes

11

Face color like other part of body and aedeagus

with two pairs of apical processes ...... 13

11. Crown with light brown band along anterior margin

and aedeagal processes extended upward ....

...... hepneri (Beamer)

Crown without band and aedeagal processes extended

downward ...... 12

12. Aedeagal processes having common stem with two

small lateral processes ...... curtus (Shaw)

Aedeagal processes without common stem and without

small processes ...... viridis (Uhler)

13. Sternal apodemes longer than first visible abdominal

segment and aedeagus not longer than 5 times width

of widest part of shaft ...... grandis (Shaw)

Sternal apodemes shorter than first visible abdominal

segment and aedeagus longer than 5 times width

of widest part of shaft ...... 14

14. Pygofer length twice its height, with less than

50 setae ...... attenuatus (Shaw)

Pygofer length slightly longer than height, with

more than 50 setae ...... flavidus (Signoret) 114

The following seven species of Hecalus are close

related since they share the same aedeagal pattern,

termed "major" pattern, and they are called "major"

group here.

Hecalus planus (Shaw)

(Fig. 187-193, 197)

Parabolocratus planus Shaw, 1932:48, 49.

Female: Body length: 7.40-8.90 mm. General color

sordid yellow to yellowish green, without other marks

except tip of ovipositor which is reddish or brown.

Head broadly produced, flattened, with length equal

to width between eyes. Anterior margin of crown strongly

reflexed and foliaceous. Pronotum short, posterior

half rugose. Forewing reduced, apical region short,

leaving pygofer and ovipositor exposed. Seventh sternite

produced caudally, with two middle sinuses.

Male: Body length: 5.40-6.40 mm. General color

sordid yellow to yellowish green, with no marks except

one on abdomen. Head not very produced and length

less than width between eyes. Crown flattened and

reflexed, with anterior margin very thin. Pronotum

length almost equal to head, with posterior half rugose.

Forewing fully developed but sometimes leaving male genitalia slightly exposed, with normal venation except some cross-veins on costal area. Abdomen black or

dark brown on middle part of tergites, one black band

usually present on middle part of IV, V, and VI sternites

Two pairs of sternal apodemes present, with middle

one longer than half of first abdominal segment and

lateral one very short. Male genitalia: Pygofer

short, quite similar to H. atascaderus except no black

pigments. Aedeagus similar to 'major' pattern, but

terminal processes thinner, longer and extended more

laterally. Connective large, wide with black band

in middle of its arms.

Diagnosis: This species is close related to

H. kansiensis since they share wide open terminal

aedeagal processes (Fig. 191) and the posterior margin

of the pygofer is curved angularly. It can be separated

from H. kansiensis by the form of preatrium (Fig. 190).

Distribution: California, Oregon, Wyoming.

Type and Material Examined: 1 female, Lone Tree,

Wyo., VII-30-1904, H. Osborn. 2 females, Cane Tree,

Wyo., VIII-15-1904, H. Osborn. 1 female (paratype),

Orange Co., Cal., VII-14-1929, R.H. Beamer (OSUC).

1 female (paratype), Orange Co., Calif., VII-14-1929,

R.H. Beamer; 1 female (paratype), Bakersfield, Calif., 116

VII-24-1929, R.H. Beamer; 1 male (paratype), Lemmon

Cave, Calif., VII-26-1929, R.H. Beamer; 1 female,

Mint Canyon, Calif., VII-6-1933, R.H. Beamer; 5 males,

4 females, Lafayette, Calif., VII-14-1933, R.H. Beamer;

1 female, Lapine, Oregon, VII-2-1935, R.H. Beamer;

4 males, 1 female, Cuyamaca, Ranch, Calif., VII-25-1935,

R.H. Beamer (SEMC). Holotype and more paratypes are also deposited in SEMC.

Hecalus kansiensis (Shaw)

(Fig. 198-204)

Parabolocratus kansiensis Shaw, 1932:47.

Female: Body length: 8.50-9.30 mm. General color sordid yellow to pale green, without any marks.

Head very produced and foliaceous, with length much longer than width between eyes. Ocellus far from eye, distance much larger than diameter of ocellus.

Forewing reduced and two or one terminal abdominal segments exposed. Hind femur spinulation 2-2-2.

Seventh sternite wave-like on posterior margin. Only one row of setae present along margin of pygofer and ovipositor tip color same as other parts of body. Male: Only one metatype was examined in this

study. Body length: 6.45 mm. General color pale

green. Head produced and foliaceous, with length

longer than width between eyes. Front end of crown

black. Distance between eye and ocellus greater than

diameter of ocellus. Face unmarked. Forewing fully

developed, with common pattern of venation. Two short

pairs of sternal apodemes present. Male genitalia:

Pygofer with posterior margin directed downward, having

some setae on posterior half. Aedeagus 'major’ pattern

with two long and curved terminal processes, extending

cephalad, phragma very long, longer than half length

of shaft.

Diagnosis: This species is closely related

the H. planus and can be separated by the non-extended

preatrium of aedeagus.

Distribution: California, Kansas, Michigan.

Type and Material Examined: 1 male (metetype),

3 females, Pottawatomie Co., Kansas, V-18-?, D.A.

Wilbur; 6 females, Cheboygan Co., Mich., VII-27-1933,

H.B. Hungerford; 1 female, Cuyamaca Ranch, Calif.,

VII-25-1935, R.H. Beamer; 6 females, Medora, Kansas,

VI-24-1936, R.H. Beamer (SEMC). Holotype is deposited 118

in SEMC.

Hecalus rotundus (DeLong)

(Fig. 205-210)

Parabolocratus rotundus DeLong, 1938:302, 303.

Female unknown.

Male: Body length: 6.50-6.70 mm. General color

sordid yellow to yellowish green, without other marks

except one brown band along upper margin of face.

Head flattened, slightly produced, length much less

than width between eyes. Front crown round, with

margin thin. Pronotum flattened, longer than head,

with rear half rugose. Forewing reduced slightly,

apical region short with more cross veins in costal

area. Abdomen same color as other parts of body apex

usually exposed. No sternal abdominal apodemes developed.

Male genitalia: Pygofer large and wide, having less

than 20 setae on posterior half and posterior margin

round, not produced. Aedeagus 'major' pattern but

shaft short, stout, slightly curved at middle, apex swollen laterally. Terminal processes long, almost as long as shaft. Connective small, straight with arm processes black. 119

Diagnosis: This species can be distinguished

from the closest relative, H. inflatus, by the normally

developed apical region of the forewing.

Distribution: Ohio, Wisconsin.

Type and Material Examined: 1 male (holotype),

1 male (paratype), Madison, Wis., IX-2-1916, D.M.

DeLong; 1 male (paratype), Castalia, Ohio, VII-25-1917,

D.M. DeLong (OSUC).

Hecalus inflatus (DeLong) Reinstated

(Fig. 211-217)

Parabolocratus inflatus DeLong, 1938:301, 302.

Parabolocratus continuus. Oman, 1949:33.

Female unknown (there are no females of H. inflatus

or H. continuus in OSUC even though DeLong, 1938, mentioned

several female paratypes from Yellowstone National

Park and Idaho).

Male: Body length: 4.90 mm. General color

pale yellow, without other marks. Head produced,

length equal to width between eyes. Crown convex,

anterior margin thin. Pronotum shorter than head, without rugose region on posterior half. Forewing 120 reduced slightly, with apical region short. Abdomen color slightly darker than head and thorax, with very undeveloped sternal apodemes. male genitalia: Pygofer short, with posterior margin produced broadly and only several setae on apical half. Aedeagus 'major* pattern except smaller preatrium. Connective form distinct (Fig. 212).

Remarks: Oman (1949) considered this species as a synonym of Parabolocratus continuus Delong.

After examining the holotypes of both species, I am convinced that H. inflatus (DeLong) is actually a good species, and it can be separated from H. continuus by the following characters: 1) reduced apical region of forewing, which is quite special in Hecalus: 2) head length equal to width between eyes but that of

H. continuus is much shorter; 3) undeveloped sternal apodemes; and 4) posterior margin of pygofer not produced sharply and with fewer setae. This species is closely related to H. rotundus but can be separated by the reduced apical region of forewing. DeLong

(1938) mentioned several paratypes of this species but in this study only the holotype was located from the tray of H. continuus in DeLong*s collection.

Distribution: Wyoming. 121

Type and Material Examined: 1 male (holotype),

Yellowstone Nat. Pk., Wyo., V-4-1930, D.M. DeLong

(OSUC).

Hecalus continuus (DeLong)

(Fig. 218-224)

Parabolocratus continuus DeLong, 1938: 301.

Female unknown.

Male: Body length: 5.60 mm. General color sordid yellow or tawny. Face and venter of thorax light brown, abdomen dark brown. Head not very produced, length much less than width between eyes, margin near apex thin and curved upward. Ocellus not near eye, distance between them greater than diameter of ocellus. Pronotum wider and longer than head, posterior half rugose. Forewing fully developed and with normal venation. Only one pair of sternal apodemes present which are short and wide. Male genitalia: Pygofer shape distinct, upper-posterior angle of posterior margin produced sharply, and covered with less than

20 setae on apical half. Aedeagus not very long but straight in apical half, flattened laterally with one pair of terminal processes and one large preatrium.

Style wide, large and connective slender and large. 122

Diagnosis: This species is closely related

to H. atascaderus but can be distinguished by the

less produced male head (Fig. 218) and pygofer form

(Fig. 223).

Distribution: Wyoming; Canada.

Type and Material Examined: 1 male (holotype),

Farewell Ck., S. Sask, Canada, no collecting date;

3 males (paratype), Yellowstone Nat. Pk., Wyo., VII-4-1930,

DeLong.

Hecalus atascaderus (Ball)

(Fig. 225-233)

Parabolocratus atascaderus Ball, 1933:223.

Female: Body length: 7.1 mm. General color sordid

yellow without other marks. Head broadly produced,

length much longer than width between eyes, anterior margin thin. Ocellus not very near eye, distance between them greater than diameter of ocellus. Forewing reduced, apical region short, leaving pygofer and ovipositor exposed. Seventh sternite broadly produced posteriorly, apex truncate. Male: Body length: 5.20-5.60 mm. General color

sordid green with venter of thorax, legs and tergites

of abdomen black-colored. Head produced, length equal

to width between eyes, anterior margin of crown thin

and slightly curved upward. Pronotum wider than head,

posterior half rugose. Scutellum large, having two

sordid white spots on corners. Forewing developed

fully, with normal venation but more cross-veins.

Hindwing light brown with venation dark brown. Abdomen

large, black dorsally, sordid yellow ventrally, with

one longitudinal black band at middle of sterna.

Two sternal apodemes present, lateral one wider and

shorter. Male genitalia: Pygofer not very long,

anterior half black-colored and posterior-dorsal angle

produced. Some short setae on posterior half of pygofe

Aedeagus "major" pattern but preatrium black-colored.

Diagnosis: This species is closely related

to H. continuus and can be separated by the produced male head and pygofer form (Fig. 230).

Distribution: California.

Type and Material Examined: 1 male, 1 female,

Goed Run, Calif., VI-18-1958, G.F. Knowlton; 1 male,

Lk. Tahoe, Calif., VI-19-1948. D.J. & J.N. Knull (OSUC) 124

The identification and description are based

on specimens determined by Ball and the type specimens

are deposited in NMNH.

Hecalus ma.ior (Osborn)

(Fig. 194-196, 234-237)

Parabolocratus ma.ior Osborn, 1915: 110.

Female: Body length: 7.55-7.70 mm. General color

sordid yellow without other marks. Head produced

and foliaceous, length slightly less than width between

eyes. Ocellus not near eye, distance between them

greater than diameter of ocellus. Forewing fully

developed and covering entire abdomen. Abdomen color

slightly darker than other parts of body, seventh

abdominal segment and ovipositor exposed. Seventh

sternite with one shallow sinus in middle of rear

margin. Hind femur spinulation 2-2-1.

Male: Body length: 6.70 mm. General color like

female but face with one cross band along upper margin.

Head produced foliasciously and length equal to width

between eyes. Ocellus not near eye, distance greater

than diameter of ocellus. One Omega-shaped line present

on pronotum, posterior region rufous. Forewing fully developed and covering entire body. Abdomen dark

brown or black, black portion of notum T-shaped.

Male genitalia: Pygofer not very long, black colored

on dorsal part, and truncate on posterior margin.

Aedeagus long, with one pair of terminal processes,

preatrium very large with one long phragma.

Diagnosis: This species can be separated from

the above six species by the posterior margin of pygofer

which is curved angularly and not sinuate on the ventral

margin of pygofer.

Distribution: Kansas, Michigan, Ohio.

Type and Material Examined: Female holotype,

Steubenville, Oh. VIII-1940, H. Osborn. 5 females

(paratype), same date as holotype. 1 female (paratype),

Castalia, 0. VII-29-1940, H. Osborn. 1 female (paratype)

Castalia, H. Osborn, no date. 5 females (paratype),

Orono, Me., VI-18-1913, H. Osborn (OSUC). 1 male,

labelled as cuw 2027, C.F. Baker collection; 1 female,

Douglas L. Mich., VII-20-1925, H.B. Hungerford; 1

female, labeled as Ft. Collins, Col. 93098, University

of Kansas, Lot 925; 1 female, Cheboygan, Co., Mich.,

VII-14-1931, J. Brennan (SEMC). 126

Hecalus viridis (Uhler)

(Fig. 238-244)

Glossocratus viridis Uhler, 1887:462.

Female: Body length: 6.2-7.3 mm. General color sordid yellow to green, without other marks except one band along upper margin of face. Head produced and foliaceous, length longer than width between eyes.

Ocellus not very near eye, distance between them greater than ocellus diameter. Pronotum equal to head in length. Forewing not much reduced, not covering ovipositor.

Only one row of setae along pygofer ventral margin, tip of ovipositor red. Hind tibiae spinulation formula

2- 2- 0 .

Male: Body length: 5.4-5.8 mm. General color like female, some individuals slightly darker. Head also produced and foliaceous, length equal to width between eyes. Ocellus position like female. Pronotum length equal to head, with omega-shaped mark in middle.

Two faint markings on basal corners of scutellum.

Forewing fully developed, with one small spot on end of clava. Male genitalia: Pygofer form distinct

(Fig. 242), with rear margin truncate, ventral-posterior angle extended, and one deep sinus on ventral margin. Many setae present along ventral 127 and posterior margins. Plate with two rows of setae.

Aedeagus straight, long, with two terminal processes.

Connective quite like many other Hecalus species.

Diagnosis: This species can be distinguished from the other species of Hecalus by its aedeagus form (Fig. 240, 241).

Distribution: Arizona, California, Colorado,

Iowa, Kansas, Montana, Nebraska, New Mexico, Ohio,

Tennessee, Texas, Washington, Wyoming.

Material Examined: 1 male, Hodgman Co., Kansas,

VII-17-25- 1917, J.H. Shaw; 1 male, Reno Co., Kansas,

VIII-13-20-1917, J. H. Shaw; 3 females, Cheyenne

Co., Kansas, VII-5-1923, H.D. Deay; 1 female, Montgomery

Co., Kansas, VIII-3-1923, Beamer-Lawson; 1 male, 6 females, Morton Co., Kansas, VII-27-1924, C.O. Bare;

1 female, Decater Co., Kansas, VII-6-1925, H.O. Deay;

1 female, Cheyenne Co., Kansas, VII-1-1926, R.H. Beamer;

1 female, Slos, Colo., VII-17-1929, P. W. Oman; 1 female, Oroville, Wash., VIII-6-1931, R. H. Beamer;

2 females, Baboquivari Mt., Arizona, VII-19-1932,

R.H. Beamer; 1 male, Laramie, Wyo., VI-23-1935, J.

Beamer; 1 female, Seymour, Tex., VI-30-1936, R.H.

Beamer; 1 male, N. Mexico, VII-16-1936, R.H. Beamer; 2 males, Spronger, VII-17-1936, R.H. Beamer; 1 male,

Rocky Mt ., Colo., VIII-5-1947, R.H. Beamer; 1 male,

Hugoton, Kansas, F.X. Williams, without date; 2 males,

1 female, Ottawa Co., Kansas, P.B. Lawson, without date (SEMC). 5 females, Omaha, Nebr., VII-14-1934,

E.J. Johnson; 3 males, Pine Valley, Cal., VII-27-

1940, D.J. and J.N. Knull; 13 females, Santa Fe.

Co., N.M. VII-10-1940, E.J. Johnson; 6 females, Bishop

Cal., VII-30-1940, E.J. Johnson; 1 female, Grt. Smoky

Mts. Tenn., VI-7-1942, D.J. and J.N. Knull; 1 female,

Scioto Co., 0., VI-1-1945, D.J. and J.N. Knull; 10 males, 6 females, Huachuca Mts. Ari., VIII-19-1950,

D.J. and J.N. Knull; 4 males and 1 female, Jeff Davis

Co., Tex., VII-4-1953, D.J. and J.N. Knull; 1 female,

Ames, Iowa, No. 51296, H. Osborn; 1 female, Ft. Collins,

Colo., No. 7150, H. Osborn; 1 female, Bismarck, N.D.

No. 7209, H. Osborn; 4 females, Bozeman, Mont. No.

7037, H. Osborn(OSUC).

Hecalus hepneri (Beamer)

(Fig. 245-251)

Parabolocratus hepneri Beamer, 1948:63.

Female: Body length: 6.60-7.05 mm. General color sordid yellow. Head not very produced, length much less than width between eyes. Crown round, foliaceous 129

at front margin. Ocellus distant from eye, distance

between them more than twice ocellus diameter. Face

light brown on clypeus and slightly lighter along

upper margin. Forewing developed, with normal venation

but leaving terminal segment exposed. Abdomen dark

colored but rear 1/4 and dorsal part of each segment

lighter. Formula of hind femur 2-2-0. Seventh sternite

with a small sinus on posterior margin. Ovipositor

exposed, with brown tip.

Male: Body length: 4.90-5.35. General color

sordid yellow to brown, slightly darker than females.

Head not very produced, length slightly longer than

1/2 width between eyes. Crown round, foliaceous,

dark brown at anterior margin. Distance of ocellus

from eye greater than diameter of ocellus. Face dark

brown or black on clypeus and area above antenna ridge.

Forewing reduced slightly, apical region short, leaving

2 or 3 terminal segments exposed. Abdomen black, rear

1/5 notum of each segment pale yellow. Two pairs of sternal apodemes developed, both longer than 1/2 of first abdominal segment. Formula of hind femur same as female. Male genitalia: Pygofer short, with several rows of setae along posterior margin and deep sinus on ventral margin. Aedeagus straight, with one branched terminal process, bending posterior-laterally. 130

Diagnosis: This species can be distinguished

from the other species of Hecalus by its aedeagus

form (Fig. 248, 249).

Distribution: Kansas.

Type and Material Examined: 12 males, 9 females,

paratypes, Garden City, Kansas, VI-12-1946, R.H. Beamer

(SEMC). The holotype is also deposited in SEMC.

Hecalus curtus (Shaw)

(Fig. 252-258)

Parabolocratus curtus Shaw, 1932:41.

Female: Body length: 6.90-7.55 mm. General color

pale green without other marks. Head not produced,

length less than width between eyes, but front end

quite foliaceous. Ocellus not very near eyes, distance

between them equal to diameter of ocellus. Face with

one cross black band along upper margin. Forewing developed, only ovipositor exposed but apical region short. Seventh sternum square and without sinus on rear margin. Ovipositor tip red, only one row of 131 setae present along margin of pygofer.

Male: Body length: 4.55-4.90 mm. General color

pale green without other marks. Head, unlike other

members of Hecalus. not produced, anterior margin

round. Ocellus large and very near eyes. Face with

one black cross band along upper margin. Pronotum

with one dim Omega-shaped line in middle, the remainder

rugose, posterior margin deeply convex. Only one

pair of sternal apodemes developed. Male Genitalia:

Pygofer not long, covered by setae on rear half, posterior

end tapering, ventral margin deeply situate. Aedeagus

shape distinctive, with one terminal process branched

laterally. Two tooth-like processes present beside

genital opening.

Diagnosis: This species is easily recognized

by the black band across the face, tooth-like processes

beside genital opening and distinctive terminal processes

of aedeagus.

Distribution: Kansas.

Type and Material Examined: 11 males, 5 females,

Hutchinson, Reno Co., Kansas, VII-14-1940; 3 females,

Hutchinson, Reno Co., Kansas, VI-18-1949, C. D. Michener 132

(SEMC). The types are also deposited in SEMC.

Hecalus flavidus (Signoret)

(Fig. 259-264)

Parabolocratus flavidus Signoret, 1879:276.

Females not available.

Male: Body length: 5.05-5.50 mm. General color

pale green, without other markings. Crown flattened,

angularly produced, length equal to width between

eyes. No carina on anterior margin of crown. Ocellus

near eye, distance between them shorter than ocellus

diameter. Forewing fully developed, with normal venation.

Two pairs of sternal apodemes developed, with middle

paii' very long, almost equal to length of first abdominal

segment, lateral pair diverging laterol-posteriorly.

Male genitalia: Pygofer tapering posteriorly, with many setae on posterior half. Aedeagus long, straight, with one pair of bifurcate terminal process, upper rami lobe-like. Top of shaft swollen slightly, with one ridge on dorsal side.

Diagnosis: This species can be distinguished from the other species of Hecalus by its aedeagus form (Fig* 260, 261). 133

Distribution: Alabama, California, Florida,

Georgia, Kansas, Mississippi, Nebraska, North Carolina,

Ohio, South Dakota, Texas, Virginia, Washington D.C.,

Wyoming.

Type and Material Examined: female holotype, labeled as: ins. Philipp., Semper, 487-71, 70-88 (NRSE); 1 male, Beauregard Par., La., VIII-16-1928, A.M. James;

1 male, plesiotype, Douglas Co., Kansas, VIII-26-1928,

P.R. Lawson; 5 males, Fulton, Miss., VII-14-1930,

P.W. Oman; 1 male, Shuqualak, Miss., VII-16-1930,

P.W. Oman; 3 males, Gallion, Alabama, VII-16-1930,

P.W. Oman; 1 male, Pfattsburg, Georgia, VII-25-1930,

R.H. Beamer; 4 males, Wildwood, Florida, VIII-2-1930,

L.D. Tuthill; 2 males, Hilliard, Florida, VIII-19-1930,

J. Nottingham; 3 males, Hilliard, Florida, VIII-31-1930,

R.H. Beamer; 1 male, Natchitoches Par., La., VIII-19-1932,

R.H. Beamer; 2 males, Natchitoches Par., La., VIII-16-1938,

A.M. James (SEMC). 1 female, Vinton, Ohio, VI-19-

22-1901, H. Osborn; 1 female, Ch. Bridge, Va. VIII-

13-1905, J.G. Sanders; 1 male, Wrightville Beach,

N.C. VII-27-1919, Osborn and Metcalf; 3 females,

New Smyrna, Fla., V-1920, M. Wright; 1 female, Omaha

Nebr., VIII-29-1930, D.E. Johnson; 2 males and 6 females, Biloxi Miss., VI-11-1934, D.W. Grimes; 2 males, Dade Co., Fla. V-12-1939, D.J. and J.N. Knull; 134

3 males, 1 female, Rusk Co., Tex., VI-18-1940, D.J. and J.N. Knull; 1 male, Gillespie Co., Tex., VI-23-

1940, D.J. and J.N. Knull; 3 males, Brookings, S.D.

H. Osborn, no date; 1 female, Fairfield Co., 0. VIII-

15-1945, D.J. and J.N. Knull; 1 male, 2 females,

Washington D.C. 8-10, No other data; 2 males and 3 females, Bay Ridge Md. VII-1, no year, D.J. and J.N.

Knull; 1 male, Yellowstone Pk. Wyo., H. Osborn, no date (OSUC).

Hecalus attenuatus (Shaw)

(Fig. 265-271)

Parabolocratus attenuatus Shaw, 1932:40.

Female: Body length: 6.15-6.40 mm. General color pale green, without other markings. Head quite produced, length much longer than width between eyes. Crown flattened, apex rounded, without carina on anterior margin. Ocellus distant from eyes, with slight depression in front of it. Pronotum flattened, with relatively longer lateral margin. Posterior margin of pronotum convex, with angle at middle. Forewing fully developed, with common venation pattern. Ovipositor and pygofer exposed slightly, tip of ovipositor red. Face and body venter with same color as other parts of body. Male: Body length: 4.70-5.20 mm. General color

like female, without special markings. Head angularly

produced, length equal to width between eyes. Crown

flattened, without carina on front margin. Ocellus

distant from eye. Pronotum with relatively long

lateral margin, posterior margin angularly convex.

Forewing fully developed, with common venation pattern.

Face and venter of body without markings. Two pairs

of sternal apodemes present, with middle one longer

and narrower. Male genitalia: Pygofer very long,

covered by large setae over entire surface. Aedeagus

very long, with one pair of branched terminal processes,

extended downward, upper rami smaller and more slender

than lower one. Connective narrow at upper part and

style usual.

Diagnosis: This species can be easily determined

by the long pygofer and the form of the aedeagus (Fig.

269, 270).

Distribution: Arizona.

Type and Material Examined: 1 male, 1 female,

Sabino, Cah., Arizona, VII-14-1932, R.H. Beamer; 2 females, Santa Rita Mts., Arizona, VII-17-1932, R.H.

Beamer; 4 males, 3 females, Baboquivari Mts., Arizona, R.H. Beamer; 1 female, Santa Rita Mts., Arizona,

VIII-18-1935, R.H. Beamer; 1 female, Hereford, Arizona,

VIII-22-1935, R.H. Beamer; 1 female, Mustang Mts.,

Arizona, VIII-22-1935, R.H. Beamer. All material are from SEMC, and the types also are deposited in

SEMC.

Hecalus grandis (Shaw)

(Fig. 272-281)

Parabolocratus grandis Shaw, 1932:44-45.

Female: Body length: 7.4-9.1 mm. Flattened and elongated. General color pale yellow without other markings. Head flattened, foliaceous and broadly produced. One middle longitudinal ridge on crown, front margin of crown very thin. Ocellus not very near eye, distance between them larger than ocellus diameter. Forewing reduced, reaching to middle of

6th tergite or slightly beyond it. Venation reduced, apical region very short. Posterior margin of 7th sternite broadly produced, with one small middle process.

Ovipositor not very long, extended slightly beyond pygofer.

Male: Body length: 5.0-5.7 mm. General color 137 pale yellow, without other markings. Head sharply produced but not foliaceous, with length longer than one and half of distance between eyes. Ocellus near eye, with distance between them shorter than ocellus diameter. Pronotum long, posterior half rugose.

Forewing fully developed, with normal venation. Abdomen color pale green, genital segments slightly lighter.

Two pairs of sternal apodemes present, middle pair longer and narrower. Male genitalia: Pygofer long, length twice its height, and covered with setae at apical half. Aedeagus short, stout, shaft sides nearly parallel and swollen preapically then narrowed at apex. Two pairs or one pair of branched terminal process present, with upper rami shorter than lower ones.

Diagnosis: This species can be distinguished from the other species of Hecalus by its aedeagus form (Fig. 278).

Distribution: Illinois, Kansas, South Dakota,

Texas.

Type and Material Examined: 7 females, Douglas

Co., Kansas, VTII-11-1919, R.H. Beamer; 1 female(paratype),

Douglas Co., Kansas, VII-17-1921, R.H. Osborn; 1 female, 138

Douglas Co., Kansas, IX-28-1932, M.W. Sanderson; 1 female, Evergreen Park, Illinois, VII-23-1934, DeLong

& Ross; 2 males, Blanco Co., Tex., VI-20-1940, D.J.

& J.N. Knull (SEMC). 1 male, 1 female, Delphos, Kansas,

VIII-31-1930, H. Osborn; 5 males, 2 females, Blanco

Co., Tex., VI-20-1940, D.J. and J.N. Knull; 2 females,

Brookings S. D. H. Osborn, no date (OSUC). Types are deposited in SEMC.

The following two species also share the ‘major’ pattern aedeagus, but they both have black stripes or bands on the body.

Hecalus nigrafasciatus (Beamer)

(Fig. 286-293)

Parabolocratus nigrafasciatus Beamer, 1938:82.

Female: 7.35-8.05 mm. General color brown, with some large black spots on head, pronotum and forewing.

Head roundly produced and foliaceous, length longer than width between eyes. Crown black beyond eyes.

Ocellus distant from eye, distance between them greater than ocellus diameter. Face unmarked except faint black band along upper margin. Pronotum with one large, square black spot on disk. Scutellum with 139 no markings above suture and black under suture. Forewing not well developed, leaving three abdominal tergites exposed, with claval region and subcostal region black.

Abdomen black except front margin and middle part of each segment. Ovipositor extending beyond pygofer, same color as pygofer. Hind femur spinulation 2-2-1.

Male: Body length: 5.40-5.70 mm. General color like female, also with some large black spots on head, pronotum and forewing, pattern similar to female.

Head foliaceously produced, length longer than width between eyes. Face faint . black but area around eyes light-colored. Forewing well developed, with some additional cross-veins on apical area. Abdomen dark brown, with two pairs of sternal apodemes. Male genitalia:

Pygofer tapering caudally, with less than 20 setae on each apical half. Style and plate as in generic description. Aedeagus •major* pattern but apical processes larger and dorsal apodeme longer.

Diagnosis: This species is very easy to identify by the distinctive color pattern of both male and female (Fig. 289, 290).

Distribution: Colorado. 140

Type and Material Examined: 6 females (paratypes),

2 females, Creede Colorado, VII-6-1937, L.D. Tuthill.

10 males, 5 females, Florrisant Colorado, VII-6-1949,

R.H. Beamer (SEMC).

Holotype is also deposited in SEMC.

Hecalus nimbosus (Ball)

(Fig. 282-285, 294-297)

Parabolocratus nimbosus Ball, 1937:129.

Female: Body length: 7.20-8.00 mm. General color

sordid brown to dark grey, with middle black band

on head, pronotum and scutellum. Head roundly produced,

length longer than width between eyes. Forewing reduced,

leaving two or three abdominal tergites exposed.

Some individuals with black band along claval suture.

No markings on ventral side of body.

Male: Body length: 5.20-5.60 mm. General color

dark brown to dark grey, with one median longitudinal

black band on head, pronotum and scutellum. Ventral

color of body darker than dorsal. Forewing well developed, also with dark brown or black band along claval suture.

Male genitalia ‘major* pattern but shaft and apical processes longer than usual form.

Diagnosis: This speices can be separated from

the above species by the color pattern on the male

body (Fig. 294, 295).

Distribution: New Mexico.

Type and Material Examined: 9 males, 12 females,

Chama, New Mexico, VII-5-1937, L.D. Tuthill. All

material was studies from SEMC, and the types are

also deposited there.

Remark: Ball (1937) gave a good illustration

of this species by means of which identification of

the species is easy.

The following three species were described only

from female specimens. Since not all of the species mentioned before are known for their females, such as H. continuus DeLong, H. rotundus DeLong and H.

inflatus DeLong, it seems to me that there is little basis to judge if those species are valid or actually belong to the species described earlier based only on male individuals. However, from my knowledge of the known females, it is possible that these three 142

species are valid. Therefore, in this study, they

are treated as valid species. Hopefully the specimens

of opposite sex of the species described based on

both male or female will- be found eventually.

Hecalus elongatus DeLong

(Fig. 298-299)

Parabolocratus elongatus DeLong, 1938:302.

Female: Body length: 10.00-10.45 mm, the longest

females in Hecalus. General color sordid yellow without

other markings. Head very produced, length longer

than width between eyes. Crown convex, anterior margin

very thin. Pronotum narrower and shorter than head.

Forewing moderately reduced, barely reaching fourth

abdominal segment. Abdomen long, third abdominal

segment widest part of body. Seventh sternite with

posterior margin expended slightly in middle.

Male unknown.

Diagnosis: This species can be separated from known females by the elongated body and very short

forewing (Fig. 298).

Distribution: South Dakota. 143

Types and Material Examined: 1 female (holotype) and 1 female (paratype), Interior, S. D., VIII-28-1922,

H. C. Severin. The second paratype DeLong'mentioned, also from S. Dakota was not found in DeLong’s collection in OSUC.

Hecalus constrictus DeLong

(Fig. 300-301)

Parabolocratus constrictus DeLong, 1938:302.

Female: Body length: 7.40-8.00 mm. General color sordid yellow without other markings. Head very produced, concavely narrow, length much longer than width between eyes. Pronotum wider than head including eyes. Forewing developed, with quite normal venation and apical region not short, leaving only ovipositor exposed. Seventh sternite not very long, posterior margin with three small processes, middle one largest. Pygofer like other species, ovipositor extended well beyond pygofer.

Male unknown.

Distribution: Florida.

Type and Material Examined: 1 female (holotype),

Miami, Florida, IV-14-2921, DeLong; 4 females (paratypes),

Data same as holotype (OSUC). 144

Hecalus montanus (Ball)

(Fig. 302-303)

Parabolocratus viridis montanus Ball, 1900:71.

Parabolocratus montanus. Van Duzee, 1916:69.

Female: Body length: 6.80-7.45 mm. General color pale yellow with some fusco-piceous stripes or bands on forewing and abdomen. Head not produced, round and foliaceous on anterior margin. Head length much shorter than width between eyes. Pronotum as wide as head, head and pronotum without markings.

Forewing reduced, leaving 2-3 abdominal segments exposed.

One short fusco-piceous stripe on claval region and one long stripe on basal region just along costal margin. One pair of fusco-piceous spots on each notum of abdominal segments and on dorsal-lateral surface of pygofer. Seventh sternite with posterior margin slightly curved at middle. Ovipositor not extended much beyond pygofer.

Male: unknown.

Distribution: Colorado, New Mexico.

Type and Material Examined: 1 female, Victoria, 145

B.C. VII-16-1901, R.C. Osborn; 1 female, Sloss, Colo.,

VIII-17-1929, P.W. Oman; 6 female, Yellowstone N.P.,

Wyo., VII-12-1939, D.J. & J.N. Knull; 4 females,

Santa Fe. Co., N.M. VII-10-1940, D.J. & J.N. Knull

(SEMC). 4 females, Yellowstone N.P. VII-12-1939,

D.J. and J.N. Knull; 4 females, Santa Fe. Co., N.M.

VI-10-1940, -H. Osborn (OSUC). Type is deposited in

SEMC.

Remarks: Ball (1900) described this species as a subspecies of P. viridis. Van Duzee (1916) listed it as species in Parabolocratus without any explanation, and Oman (1949) also listed it as a species in Parabolocratus. even though some workers, such as Shaw (1932) and DeLong and Knull (1946), listed it as a subspecies. After examining Ball’s description and specimens, I found that this taxon has nothing in common with H. viridis in color pattern and other characters such as the seventh sternite, ovipositor and the head form are different from H. viridis.

Therefore, this taxon is treated as species. However, as there are still some species in which females are unknown, it is possible that the species may be the female of one of the species based only on males. 146

Phylogeny of Hecalus Species

There is a dilemma in the phylogenetic study of Hecalus species, that is, for some species only male individuals are known and for the others the opposite situation is true. Since many characters used in the classification of the group are based on males, there are either not enough female characters and many species have been described without female specimens, this study deals exclusively with the species in which male individuals are available. Therefore, the species H. montanus (Ball), H . constri ctus (DeLong), and H. elongatus (DeLong) are not included in the phylogenetic analysis.

The phylogeny of Hecalus species was studied by using cladistic analysis which is based upon outgroup comparison and parsimony principles to polarize the characters or character states and to generate the most parsimonious tree(s). In this study, a total of 23 characters or character states have been examined for the species of Hecalus, and their descript ions and polarized conditions are listed in Table 9. The data matrix obtained is shown in Table 10. The sister group of Hecalus in Hecalini, according to the analysis mentioned in Chapter I, is a group containing three genera, Neohecalus Linnavuori, Spanghergi e1la Signoret 14 7

and Hecullus Oman. Therefore, two most closely related

genera, Hecullus Oman and Spangbergiella Signoret

are chosen as the outgroups and two species H. breatectus

(Ball) and S. vulnerata (Uhler) as the representatives

of these two genera. Since the results obtained by

using the two outgroups are the same, only one outgroup,

S. vulnerata (Uhler) was shown on the data matrix in

Table 10.

The following PAUP options have been used in

the analysis: swap=global, maxtree=100, and mulpars,

and only one most parsimonious tree was found by using

the data in Table 10, which is shown in Fig. 304, with the tree length = 31.000, and the total consistent

index = 0.742.

From tha cladogram, Hecalus was separated into three major groups, which is strongly supported by the characters 3, 7, 8, 10, and 15. The cases of the further divisions in these three groups varied. For the first one, including grandis . flavidus and attermatus, and second one, including v i r i d i s. hepneri and curtus, the relationship seems clear and strong to me even though sometimes only one character appears on the branch. For the third group, the situation was more complex. 148

Table 9: Characters used in phylogenetic study of Hecalus and their polarized conditions

Characters Plesiomorphic Apomorphic

1. Terminal processes not sharing a sharing of aedeagus common stem common stem 2. Number of terminal one two processes of aedeagus 3. Aedeagus form various like Fig. 236 4. Horn-like structure at absent present top of aedeagal shaft 5. Connective processes one pair two pairs (Fig. 5) 6. Rami of two pairs of upper pair lower pair terminal processes of thicker thicker aedeagus than lower than upper pair pair 7. Pygofer ventral margin without sinus with sinus 8. Preatrium of aedeagus not wider than wider than in posterior aspect aedeagal shaft aedeagal shaft 9. Body color without black with black pigment stripes or spots 10.Posterior margin of round curved angulately pygofer (Fig. 213) (Fig. 194) 11.Apex of aedeagus not swollen swollen 12.condition of terminal curved upward curved upward processes of aedeagus only and medially 13.Apical region of male normal length reduced (Fig.216) forewing 14.Black band on face none with one along dorsal margin 15.Face color yellow or pale smoky or black green 16.Male forewing covering several abdominal entire abdomen segments exposed 17.Preatrium of aedeagus not extended extended ventrally ventrally (Fig. 237) 18.Bottom of aedeagal not convex convex preatrium 19.Terminal processes curved upward curved downward of aedeagus Table 9 (continue), 20.Aedeagal shaft not flattened flattened laterally laterally 21.Setae on pygofer more than 20 less than 15 22.Carina on front absent present margin of male head 23.Length of male head much shorter slightly than width shorter, between eyes equal to or slightly longer than width between eyes Table 10: Data matrix of the Polarized

characters of Hecalus. 1 = apomorphic

condition, 0 = plesiomorphic condition

Characters

1 1 1 1 1 1 1 1 1 2 2 2 2 2

Taxa 1 2 3 4 5 6 7 8 9 0 1 2 3 4 5 6 7 8 9 0 1 2 3

* 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

nim- 0 0 1 0 0 0 0 1 1 1 0 1 0 1 1 0 0 0 0 0 0 1 1

nitf- 0 0 1 0 0 0 0 1 1 1 0 1 0 1 1 0 0 0 0 0 0 1 1

pla- 0 0 1 0 0 0 0 1 0 1 1 0 0 0 0 0 1 0 0 1 0 1 1 ma.i- 0 0 1 0 0 0 1 1 0 1 0 0 0 1 0 0 0 0 0 0 0 1 1

inf- 0 0 1 0 0 0 0 1 0 0 1 1 0 1 0 0 0 0 0 0 1 1 1 rot- 0 0 1 0 0 0 0 1 0 0 1 1 1 0 0 0 0 0 0 0 1 1 1 con- 0 0 1 0 0 0 0 1 0 1 0 1 0 1 0 0 1 0 0 1 0 1 1 ata- 0 0 1 0 0 0 0 1 0 1 0 1 0 1 0 0 1 0 0 1 0 1 1 kan- 0 0 1 0 0 0 0 1 0 1 1 0 0 0 0 0 0 0 0 1 0 1 1 fla- 0 1 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 1 1 f?ra- 0 1 0 0 0 0 0 0 0 00000000000 0 1 1 att- 0 1 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 vir- 0 0 0 1 0 0 1 1 0 0 0 0 0 1 0 0 0 0 1 0 0 1 1 hep- 1 0 0 0 0 0 1 1 0 0 0 001110110011 cur- 1 0 0 0 1 0 1 1 0 0 0 0 0 1 0 0 0 1 1 0 0 1 1 LIST OF REFERENCES

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Baker, C F. 1900. Four new species of Platymetopius. Can. Entomol. 32:49-50.

Ball, E. D. 1900. Notes on the Acocephalina (Homoptera- Jassidae). Iowa Acad. Sci. Proc. 7: 64-72.

Ball, E. D. 1901. New Jassidae from the Rocky Mountain and Pacific region. Canada Ent. 33:4-11.

Ball, E. D., 1933. Some new genera and species of western leafhoppers. Brooklyn Entomol. Soc. Bui. 28:223-228.

Ball, E. D., 1937. New North American leafhoppers belonging to Parabolocratus and related genera. Proc. Biol. Soc. Washington 50:129-132.

Beamer, R. H. 1936. The genus Dicyphonia Ball. J. Kansas Entomol. Soc., 9:66-71.

Beamer, R. H. 1938. Some new species of leafhoppers (Homoptera-Cicadellidae). Kansas Entomol. Soc. Jour. 11: 77-84.

Beamer, R. H., 1948. A new Lonatura and Parabolocratus (Homoptera-Cicadellidae). Kans. Entomol. Soc. Jour. 21:62-64.

Berg, C. 1884. Addenda et emendan da Hemiptera Argentina. Ann. Soc. Cient. Argentina 17: 20-41.

China, W . E. 1950. A check list of the British Hemiptera- Homoptera Auchenorhyncha. Ent. Monthly Mag. 87:243-251.

DeLong, D. M., 1938. Some new species of Parabolocratus (Homoptera: Cicadellidae). Ohio Jour. Sci. 38:301-303.

DeLong, D. M., and D. J. Knull. 1946. Check list of the Cicadellidae of America, north of Mexico. Ohio State Univ. Grad. School Stud., Biol. Sci. Ser. 1:1-102.

151 152

DeLong, I i. M., and R. E. Linnavuori. 1978. Studies on Neotropical leafhoppers. Entomol. Scand., 9(2):111-123.

Distant, W. L. 1908. Rhynchota -- Homotptera. The fauna of British India including Ceylon and Burma. V 4. 501 p.

Dlabola, J. 1954. Fauna CSR Svazek 1. Krisi -- Homoptera. Prace Ceskoclovanske Akad. Ved (Sekce Biol.) 1954:7-339.

Evans, J. W. 1947. A natural classification of leafhoppers (Jassidae, Homoptera). Entomol. Soc. London. Trans. 98: 105-271. Washington Entomol. Soc. Mem. 3: 1-253.

Evans, J . W. 1951. Some notes on the classification of leafhoppers with special reference to the Nearctic fauna. Societas Scientiarum Rennica Comn. Biol. 12(3):1—11.

Evans, J. W. 1954. Les Cicadellidae de Madagascar. Inst. Sci. Madagascar Mem. (E) 4:87-137.

Evans, J. W. 1955. Cicadellidae. Exploration du pare Nat. Albert 84:3-23.

Evans, J. W., 1966. The leafhoppers and froghoppers of Australia and New Zealand (Homoptera: Cicadellidae and Cercopidae). Australia Mus. Mem. 12:1-347.

Fieber, F . A., 1866. Nue gattungen und arten in Homoptera. Zool.-Bot. Gesell. Wien. Verh. 16:497-516.

Haupt, H. 1929. Neueinteilung der Homoptera Cicadin nach phylogenetisch zu wertenden Merkmalen. Internatl, Zool. Congr. Proc. 10:1071-1075.

Ishihara, H. 1953. A tentative check list of the superfamily Cicadelloidea of Japan. Matsuyama Agr. col. Sci. Rpt. 11:1-72.

Lawson, P . B. 1932. The genus Spangbergiella in America north of Mexico. J. Kansas Entomol. Soc. 4: 116-122.

Lindberg, H. 1948. On the insect faune of Cyprus. Results of the expedition of 1939 by Harld Haken and P.H. Lindberg. Comm. Biol. 10(7):23— 175. 153

Lindberg, H. 1954. Hemiptera Insularum Canariensium. Systematik, Okologie und Verbreitung der Kanarischen Heteropteran und Cicadinen. comm. Biol. 14( 1) :1 — 304.

Linnavuori, R. E. 1957. Neotropical Hecalinae. Ann. Entomol. Fenn. 23: 133-143.

Linnavuori, R., 1961. Hemiptera (Homoptera) Cicadellidae. South African Animal Life 8:452-486.

Linnavuori, R., 1975. Revision of Cicadellidae (Homoptera) of Ethiopian region 3. Deltocephalinae: Hecalini. Acta Zool. Fenn. 143:1-37.

Linnavuori, R. E., and D. M. DeLong. 1978. Genera and species of Hecalini known to occur in Mexico. Ann. Entomol. Fenn., 44(2):48-53.

Linnavuori, R. and D. M. DeLong. 1977. The leafhoppers known from Chile. Brenesia 12/13: 163-267.

Linnavuori. R., 1978. Seventeen new species and three new genera of Central and South America Deltocephalini. Brenesia 14/15: 195-226.

Metcalf, Z. P. 1963. General catalogue of the Homoptera. Fascicle VI, Cicadelloidea, part 9 and 10. Agr. Res. Serv., U.S. Dep. Agr.

Morrison, W. P., 1973. A revision of the Hecalinae (Homoptera: Cicadellidae) of the Oriental region. Pacific Insect 15(3-4):379-438.

Nast, J. 1952. Uwagi niektorych Ledrinae. Notes on some Ledrinae. Mus. Zool. Plonici Ann. 15: 33-42.

Oman, P. W. 1938. Revision of the Nearctic leafhoppers of the tribe Errhomenellini. U. S. Natl. Mus. Proc. 85: 163-180.

Oman, P. W. 1938. A generic revision of American Bythoscopinae and South American Jassinae. Kansas Univ. Sci. Bull. 24: 343-420. Oman, P. W., 1949. The Nearctic leafhoppers (Homoptera; Cicadellidae). A generic classification and check list.

Osborn, H. 1924. Neotropical Homoptera of the Carnegie 154

Museum. Part 4. Ann. Carnegie Mus., 15:297-462.

Osborn, H., 1915. Leafhoppers of Maine. Maine Agr. Expt. Sta. Bui. 238:81-159.

Ossiannilsson, F. 1946. 7 Halvvingar. Hemiptera. Stritar, Homoptera, Auchenorrhyncha. Svensk insektfauna. Entomologiska Foreningen I Stockholm 36:1-150.

Shaw, J. G., 1932. The genus Parabolocratus in North America (Homoptera, Cicadellidae). Kans. Entomol. Soc. Jour. 5:38-52.

Signoret, V. , 1879. Essai sur les Jassides Stal, Fieb. et plus particulierement sur les Acocephalides Puton. Soc. Entomol. de France Ann. 9(5):259- 280.

Signoret, V. 1879. Essai sur les Jassides Stal, Fieb. et plus particulierement sur les Acocephalides Puton. Ann. Soc. Entomol. France 9(5): 41-70.

Stal, C. I., 1864. Hemiptera nonnula nova vel minus cognita. Soc. Entomol. France Ann. 4:47- 68.

Stal, C. I., 1870. Hemiptera insularum Philippinarum. Bidrag till Philippinska Oarnes Hemiptera- fauna. Svanska Vetensk. -Akad. Ofv. Forh. 27:607-776.

Strickland, E. H. 1953. An annotated list of the Homoptera of Alberta. Canad. Ent. 85: 193-214.

Swofford, D. L. 1985. PAUP, Phylogenetic Analysis by Using Parsimony (2.4) User’s Manual. Natural and History Survey, Champaign, Illinois.

Uhler, P. R., 1872. Order VI.--Hemiptera. A list of Hemiptera collected in eastern Colorado and northeastern New Mexico by C. Thomas during the expedition of 1869. U.S. Geol. Surv., Prelim. Rpt. 1872:471-472.

Uhler, P. R., 1877. Report upon the insects collected by P. R. Uhler during the explorations of 1875, including monographs of the families Cydnidae and Saldae [sic] and the Hemiptera collected by A. S. Packard, Jr. U.S Geol. and Geog. Survey of the Terr. Bui. 3(2):355- 475. Van Duzee, E. P. 1908. List of Hemiptera taken by W. J. Palmer about Quinze Lake, P. Que. in 1907. Can. Entomol. 40:157-160.

Van Duzee , E. P., 1916. Suborder Homoptera Latr. 1810. Section Auchenorhyncha A. & S. 1843. Check list of Hemiptera of America north of Mexico. 1916:1-111.

Van Duzee , E. P., 1917. Catalogue of the Hemiptera of America north of Mexico excepting the Aphididae', Coccidae, and Aleurodidae. Calif. Agr. Expt. Sta., Entomol. Tech. Bui. 2:1- 902.

Van Duzee , E. P. 1917. Report upon a collection of Hemiptera made by Walter M. Giffard in 1916 and 1917, Chiefly in California. Calif. Acad. Sci. Proc. 7(4): 249-318.

Young, D. A. 1950. A prelininary list of Kentucky Cicadellidae. Kentucky. Acad. Sci. Trans. 13:54-67. ILLUSTRATIONS

Fig. 1-7 Hecalus flavidus. 1) male, dorsal aspect; 2) face ventral aspect; 3) right forewing, dorsal aspect; 4) pygofer, lateral aspect; 5) first abdominal segment and apodemes, ventral aspect; 6) connective, dorsal aspect; 7) aedeagus, lateral aspect.

156 157

ANTENNA

CLYPEUS GENA

PRONOTUM LORUM SCUTELLUM ANTECLYPEUS

FOREWING

COSTAL AREA OUTER ANTEAPICAL

CENTRAL ANTEAPICAL IIINDWING DISCAL CELLS 4TII APICAL

3RD APICAL

2ND APICAL CLAVAL SUTURE 1ST APICAL

APPENDIX HINDLEG CLAVAL VEINS \ INNER ANTEAPICAL DRACIIIAL CELL

SETAE

TERMINAL PROCESS

PYGOFER SHAFT PROCESSES PYGOFER RIFT

PIIRAGME

PREATRIUM LATERAL APODEMES

MIDDLE APODEMES Fig. 8 Hypothesized phylogenetic relationship among species of Memnonia Ball, the numbers equal to those of characters in Table 2 and x = reverse condition. MEMNONIA w

DIC YPHONIA

| - * C 0 vJOi W ■ > < ■ HECALUS w

05 SPANGBERGIELLA

tO • t- HECULLUS 00 w

GO :cd NEOHECALUS 159 160

Fig. 9-15 Memnonia ornata. 9) male head, pronotum and scutellum, dorsal aspect; 10) male right forewing; 11) connective; 12) pygofer, ventral aspect; 13) apex of aedeagus; 14) aedeagus, lateral aspect; 15) male first abdominal segment. 161

15 Fig. 16-22 Memnonia albolinea. 16) male right forewing 17) male head, pronotum and scutellum, dorsal aspect; 18) pygofer, lateral aspect; 19) male first abdominal segment; 20) aedeagus, posterior aspect; 21) aedeagus, lateral aspect; 22) connective. Fig. 23 M. acuta. 23) male head, pronotum and scutellum. 163 164

Fig. 24-31 Memnonia consobrina. 24) male genital plate; 25) male right forewing; 26) male head, pronotum and scutellum, dorsal aspect; 27) male first abdominal segment; 28) apex of aedeagus; 29) aedeagus, lateral aspect; 30) connective; 31) pygofer, ventral aspect. 991 166

Fig. 32-39 Memnonia minuta n. sp. 32) male head, pronotum and scutellum, dorsal aspect; 33) male right forewing; 34) apex of aedeagus; 35) aedeagus, lateral aspect; 36) male genital plate; 37) pygofer, ventral aspect; 38) connective; 39) male first visible abdominal segment. 167 Fig. 40-46 Memnonia fraterna. 40) male right forewing 41) male he.ad, pronotum and scutellum, dorsal aspect; 42) connective; 43) male first visible abdominal segment; 44) aedeagus, posterior aspect; 45) aedeagus lateral aspect; 46) pygofer, lateral aspect.

170

Fig. 47-54 Memnonia brunnea. 47) male head, pronotum and scutellum; 48) male first two visible abdominal segments; 49) male right forewing; 50) connective; 51) aedeagus, lateral aspect; 52) aedeagus, posterior aspect; 53) pygofer, lateral aspect; 54) 6th abdominal tergite of male.

172

Fig. 55-61 Memnonia fenestrella. 55) male head, pronotum and scutellum; 56) male right forewing; 57) male first visible abdominal segment; 58) aedeagus, lateral aspect; 59) aedeagus, posterior aspect; 60) pygofer, lateral aspect; 61) connective. 173 174

Fig. 62-68 Memnonia spadix. 62) male head, pronotum and scutellum; 63) male right forewing; 64) male first visible abdominal segment; 65) aedeagus, lateral aspect; 66) aedeagus, posterior aspect; 67) pygofer, lateral aspect; 68) connective. 175

64 176

Fig. 69-74 Memnonia nigrafassiata sp. n. 69) male head, pronotum and scutellum; 70) male right forewing; 71) aedeagus, posterior aspect; 72) aedeagus, lateral aspect; 73) pygofer, lateral aspect; 74) first abdominal segment and apodemes, ventral aspect.

Fig. 75-80 Memnonia triplehorni sp. n. 75) male head, pronotum and scutellum; 76) male forewing, dorsal aspect; 77) pygofer, lateral aspect; 78) first abdominal segment and apodemes; 79) aedeagus, lateral aspect; 80) aedeagus, lateral aspect. 179 Fig. 81-86 Memnonia hamiltoni sp. n. 81) male head, pronotum and scutellum; 82) male forewing, dorsal aspect; 83) aedeagus, posterior aspect; 84) aedeagus lateral aspect; 85) male first abdominal segment and apodemes; 86) pygofer, lateral aspect.

Fig. 87 Hypothesized cladogram of Memnonia species. Numbers refer to characters listed in Table 3. consobrina

minuta

albolinea

acuta

fraterna

• • • ornata J J J u r u j triplehorni

fenestrella

hamiltoni

nigrafassiata

spadix

brunnea 184

Fig. 88-100. 88-91 Dicyphonia plana. 88) head, pronotum and scutellum of male, dorsal aspect. 89) aedeagus, posterior aspect. 90) aedeagus, lateral aspect. 91) right forewing of male. 92-95 Dicyphonia plura. 92) head, pronotum and scutellum of male, dorsal aspect.- 93) aedeagus, lateral aspect. 94) aedeagus, posterior aspect. 95) pygofer, lateral aspect. 96-100 Dicyphonia ornata. 96) head, pronotum and scutellum of male, dorsal aspect. 97) aedeagus, lateral aspect. 98) aedeagus, posterior aspect. 99) right forewing of male. 100) pygofer, lateral aspect. 185 186

Fig. 101-112. 101-105 Dicyphonia zamorana. 101) head, pronotum and scutellum of male, dorsal aspect. 102) connective, dorsal aspect. 103) aedeagus, lateral aspect. 104) aedeagus, posterior aspect. 105) pygofer, lateral aspect. 106-112 Dicyphonia nigrita. 106) head, pronotum and scutellum, dorsal aspect. 107) right forewing of male. 108) connective, dorsal aspect. 109) aedeagus, lateral aspect. 110) aedeagus, posterior aspect. Ill) pygofer, ventral aspect. 112) sternal apodemes and first abdominal segment. 187 188

Fig. 113-124. 113 Dicyphonia cona. female, dorsal aspect. 114-119 Dicyphonia delongj. new species. 114) aedeagus, posterior aspect. 115) aedeagus, lateral aspect. 116) head, pronotum and scutellum, dorsal aspect. 117) connective, dorsal aspect. 118) pygofer, ventral aspect. 119) sternal apodemes and first abdominal segment. 120-124 Dicyphonia mexicana. new species. 120) male face, ventral aspect. 121) aedeagus, posterior aspect. 122) aedeagus, lateral aspect. 123) connective, dorsal aspect. 124) head, pronotum and scutellum of male, dorsal aspect. 189 Fig. 125. Hypothesized phylogenetic relationships among the species of Dicyphonia Ball (numbers refer to characters, see text, and x = reverse states). PLANA

ORNATA ro w

a 0 ) 01 PLURA • ••

ZAMORANA

III -*■ M E X 1 C A C A ►» C E* 00 ^ * w

o

Fig. 126-133. Hecullus breateatus. 126) female anterior half body, dorsal aspect; 127) male first abdominal segment and apodemes; 128) male 6th notum of abdomen; 129) aedeagus, posterior aspect; 130) aedeagus, lateral aspect; 131) male head, pronotum and scutellum; 132) left genital plate, ventral aspect; 133) pygofer, lateral aspect.

Fig. 134-138. Hecullus balli. 134) female anterior half body, dorsal aspect; 135) aedeagus, posterior aspect; 136) aedeagus, lateral aspect; 137) male first abdominal segment and apodemes, ventral aspect; 138) male 6th notum of abdomen. 195

137

138 Fig. 139-141. Neohecalus lineatus. 139) male head, pronotum and scutellum; 140) male forewing, dorsal aspect; 141) pygofer, lateral aspect. 142-144. Neohecalus apical is. 142) male head, pronotum and scutellum; 143) male forewing, dorsal aspect; 144) pygofer, lateral aspect. 197

W s Fig. 145-146. Neohecalus apicalis. 145) connective; 146) male first abdominal segment and apodemes. 147- 151. Neohecalus lineatus. 147) aedeagus, posterior aspect; 148) aedeagus, lateral aspect; 149) connective 150) style, ventral aspect; 151) male first abdominal segment and apodemes.

Fig. 152-153. Neohecalus apicalis. 152) aedeagus lateral aspect; 153) aedeagus, posterior aspect. 153 Figs. 154-155 Spangbergiella reticulata. 154) left forewing of female; 155) head, pronotum and scutellum of female, dorsal aspect; 156-162. Spangbergiella mexicana. 156) head, pronotum and scutellum of male, dorsal aspect; 157) left forewing of male; 158) left style of male genitalia, dorsal aspect; 159) left lateral aspect of pygofer; 160) aedeagus, left lateral aspect; 161) aedeagus, posterior aspect; 162) right genital plate, ventral aspect. i\ w Figs. 163-169 Spangbergiella quadripunctata. 163) head, pronotum and scutellum of male, dorsal aspect; 164) right genital plate, ventral aspect; 165) right forewing of male; 166) left style of genitalia, dorsal aspect; 167) left lateral aspect of pygofer; 168) left aspect of aedeagus; 169) posterior aspect of aedeagus. 205

f / Figs. 170-176 Spangbergiella erratica. 170) head, pronotum and scutellum of male, dorsal aspect; 171) left aspect of pygofer. 172) left style of genitalia dorsal aspect; 173) left aspect of aedeagus; 174) right forewing of male; 175) ventral aspectof left genital plate; 176) posterior aspect of aedeagus. 207

170

171 172

176

174 208

Figs. 177-184 Spangbergiella vulnerata. 177, 179, 185) head, pronotum and scutellum of male, dorsal aspect; 178) head, pronotum and scutellum of female, dorsal aspect; 180) left lateral aspect of pygofer. 181) right style of genitalia, dorsal aspect; 182) left lateral aspect of aedeagus; 183) posterior aspect of aedeagus; 184) ventral aspect of left genital plate. 209

178 177 185

179

180,

181

183 184 182 Fig. 186 Cladogram for hypothesized relationship among the species of Spangbergiella Signoret, the numbers equal to those of Table 8. vulnerata

reticulata

tnexicana in o>^4

erratica IT IT V

quadripunctata 212

Fig. 187-193 Hecalus planus. 187) head, pronotum and scutellum of female; 188) head, pronotum and scutellum of male; 189) connective; 190) aedeagus, lateral aspect; 191) aedeagus, posterior aspect; 192) pygofer, lateral aspect; 193) last several abdominal segments of female. 213 214

Fig. 194-196 Hecalus major. 194) pygofer, lateral aspect; 195) sternum of female 7th abdominal segment; 196) notum of male 7th abdominal segment. 197 Hecalus planus. first male abdominal segment and apodemes.

216

Fig. 198-204 Hecalus kansiensis. 198) aedeagus, posterior aspect; 199) head, pronotum and scutellum of male; 200) aedeagus, lateral aspect; 201) pygofer, lateral aspect; 202) connective; 203) sternum of female 7th abdominal segment; 204) first male abdominal segment and apodemes.

t 217

199

198

200

201 202 203

i Fig. 205-210 Hecalus rotundus. 205) head, pronotum and scutellum of male; 206) pygofer, lateral aspect; 207) aedeagus; lateral aspect; 208) connective; 209) aedeagus, posterior aspect; 210) right forewing of male. 219

205 206

208

207

209 210 220

Fig. 211-217 Hecalus inflatus. 211) head, pronotum and scutellum, dorsal aspect; 212) connective; 213) pygofer, lateral aspect; 214) aedeagus, lateral aspect; 215) aedeagus, posterior aspect; 216) male right forewing; 217) first abdominal segment and sternal apodemes. 221

213

211

212

215

214

216

217 Fig. 218-224 Hecalus continuus. 218) head, pronotum and scutellum, dorsal aspect; 219) connective; 220) aedeagus, lateral aspect; 221) aedeagus, posterior aspect; 222) style, dorsal aspect; 223) pygofer, lateral aspect; 224) first abdominal segment and sternal apodemes. 223

218 220

221

219

222 223

224 224

Fig. 225-229 Hecalus atascaderus. 225) head, pronotum and scutellum of male; 226) head, pronotum and scutellum of female; 227) sternum of 7th abdominal segment of female; 228) aedeagus, lateral aspect; 229) aedeagus, posterior aspect. 225

225 226

227 229 228 226

Fig. 230-233 Hecalus atascaderus. 230) pygofer, lateral aspect; 231) right forewing of male; 232) connective; 233) male first abdominal segment and apodemes. 227

230

231 232

A 228

Fig. 234-237 Hecalus major.. 234) head, pronotum and scutellum of male; 235) face of male, ventral aspect; 236) aedeagus, lateral aspect; 237) aedeagus, posterior aspect. 229

235

234

237

236 230

Fig. 238-244 Hecalus viridis. 238) head, pronotum and scutellum of male; 239) face of male, ventral aspect; 240) aedeagus, posterior aspect; 241) aedeagus, lateral aspect; 242) pygofer, lateral aspect; 243) connective; 244) right genital plate. 231

240

241

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239

242 243 Fig. 245-251 Hecalus hepneri. 245) head, pronotum and scutellum of male; 246) face of male, ventral aspect; 247) pygofer, lateral aspect; 248) aedeagus lateral aspect; 249) aedeagus, posterior aspect; 250) sternum of female 7th abdominal segment; 251) male first abdominal segment and apodemes. 233

247

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246 249

250 248

251 Fig. 252-258 Hecalus curtus. 252) head, pronotum and scutellum of male; 253) face of male, ventral aspect; 254) pygofer, lateral aspect; 255) aedeagus, lateral aspect; 256) aedeagus, posterior aspect; 257) male first abdominal segment and apodemes; 258) connective. 235

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258 257 Fig. 259-264 Hecalus flavidus. 259) head, pronotum and scutellum of male; 260) aedeagus, posterior aspect 261) aedeagus, lateral aspect; 262) pygofer, lateral aspect; 263) connective; 264) male first abdominal segment and apodemes. 237

259 260 261

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263

264 Fig. 265-271 Hecalus attanuatus. 265) head, pronotum and scutellum of male; 266) male first abdominal segment and apodemes; 267) pygofer, lateral aspect; 268) connective; 269) aedeagus, lateral aspect; 270) aedeagus, posterior aspect; 271) left style, dorsal aspect. 239

265 266

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269 270

271 240

Fig. 272-281 Hecalus grandis. 272) head, pronotum and scutellum of male; 273) head, pronotum and scutellum of female; 274) last several segments of female; 275) connective; 276) first male abdominal segment and apodemes; 277) sternum of 7th female abdominal segment; 278) aedeagus, posterior aspect; 279) aedeagus, lateral aspect; 280) left genital plate, ventral aspect; 281) pygofer, lateral aspect. 241 Fig. 282-285 Hecalus nimbosus. 282, 283, 284) head, pronotum and scutellum of males; 285) face of male, ventral aspect. 286-288 Hecalus nigrafasciatus. 286) head, pronotum and scutellum of male; 287) head, pronotum and scutellum of female; 288) sternum of 7th female abdominal segment. 243 Fig. 289-293 Hecalus nigrafasciatus. 289) aedeagus, lateral aspect; 290) aedeagus, posterior aspect; 291) pygofer, lateral aspect; 292) connective; 293) first male abdominal segment and apodemes. 245

290 289 291

292

293 Fig. 294-297 Hecalus nimbosus. 294) aedeagus, lateral aspect; 295) aedeagus, posterior aspect; 296) right forewing of male; 297) first male abdominal segment and apodemes.

Fig. 298-299 Hecalus elongatus. 298) female, dorsal aspect; 299) sternum of 7th female abdominal segment. 249 Fig. 300-301 Hecalus constrictus. 300) female, dorsal aspect; 301) sternum of 7th female abdominal segment.

Fig. 302-303 Hecalus montanus. 302) female, dorsal aspect; 303) sternum of 7th female abdominal segment. 302 Fig. 304 Cladogram of the hypothesized relationship among the species of Hecalus: the numbers refer to those in Tables 9, 10 and the x reverse conditions. grandis Havidus attenuatus viridis VI £ a hepneri ^ (c> curtus major Vi00"* atascaderus continuus ♦ 10 planus 00 xr ♦ xr -B-CN kansiensis M H inflatus VI H u> VI o Hs| -JttK -p- N N O H rotundus 04 nigrafasciatus H O* nimbosus 255 256

Plate I Scanning Electron-Micrographs. A,B Hecalus flavidus. A) upper surface of male forewing; B) upper-lateral surface of male pygofer. C,D Hecalus major* C) upper surface of male forewing; D) upper-lateral surface of male pygofer. E,F Memnonia albolinea. E) upper surface of male forewing; F) upper-lateral surface of male pygofer. PLEASE NOTE:

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