AN ABSTRACT OF THE THESIS OP Gerasimos Cassis for the degree of Doctor of Philosoph in Entomolozv presented on Narch 28, 1984 Title: A Systematic Study of the Subfamily DicyDhinae (Heteroptera: Miridae). / Abstract approved: John D. La t: A systematic study of the subfamily Dicyphinae Reuter (Heteroptera: Hiridae) resulted in a redefinition of the subfamily and included genera) and a revision of the genus Dicyphus Fieber in the Western Hemisphere.The study entailed examination of about 14,000 specimens, including numerous noudicyphine, cimicomorphan taxa to assess character transformations. New character sources, such as the thoracic pleura, were investigated to test current classifications, establish the present classification, and perform a preliminary phylogenetic analysis with included consideration of host associations and biogeography. In the first chapter, general information about the family Hiridae is given and the systematic problems of the Dicyphinae and the objectives of the study are e Inc ida ted. In the second chapter, a diagnosis of the subfamily is provided with some discussion of the intrafamilial relationships. Also the genera of the subfamily are redefined, redescribed, and included species for each genus are listed. Sixteen genera are recognized and one new genus, Glarisia n. gen., and one new subgenus, Uhlere].ia n. sgen., of the genus Dicyphus, are described. The six subgenera of Cyrtoeltis Fieber: Cyrtopeltis, Engytatus Reuter, Nesidocoris Kirkaldy, Singhaiesia China and Carvaiho, Tupiocoris China and Carvalbo, and UsingerellaChina and Carvalbo are elevated to generic rank, whereas the subgenera of Dicypbus are retained. Chapter three is a revision of the genus Dicyphus in the Western Hemisphere which contains nine species. There are eight endemic species belonging to the endemic subgenus D. (Ublerella), including two new species nigracorium n.sp. andoccidentalis n. sp.. A key is given to the species. A Systematic Study of the Subfemily Dicyphivae (Heteroptera: Miridae). by Gerasimos Cassis A THESIS submitted to Oregon State University in partial fulfillment of the requirments for the degree of Doctor of Philosophy Completed March 28, 1984 Commencement June 1984 APPROVED: Prof es4orof Entomology in charge of niajor Head of Department of Entomology Date thesis is presente4 March 28 1984 Typed by the author ACKNOWLEDGEMENTS I wish to express my sincere thanks to John D. Lattin, my major professor, who initiated my interests in the systematics of the family Miridae. Throughout the duration of this work he has been liberal with his time, encouragement, and advice. His influence over the last three and half years has been a stabilizing one and his friendship and guidance are greatly appreciated. I extend my warmest thanks to my committee: Paul W. Oman whohad a profound effect on mythinking,was never discouraged by my constant tapping on his door, his generosity seemed to be without limit; Kenton L. Chambers, my minor professor, was most supportive and offered much knowledge and literature pertaining to angiosperm pbylogeny; Gerald W. Krantz, who at times had half of his library and equipment in my office, and was a vital moral support and remains a good friend; Arthur 3. Boucot, who besides hijacking me into three fossil classes, has been a significant peer, extremely supportive, and was willing to be hostage to numerous cladistic discourses, Ray Siedler, my graduate representative, showed unforseen encouragement and interest and his participation is warmly appreciated. Many thanks go to William P. Stephen, who was most encouraging, allowed me to use his I. B. H. word processor, and despite a lack of good Greek jokes, I will concede that he has had a major influence on my education. Words are insufficent to express my gratitude to Gary M. Stonedahi who was an ear for almost every idea I came up with. He edited a good deal of this thesis, was always encouraging and is a greatly valued friend. Past and present members of the Oregon State University Systematic Entomological Laboratory, including Vincent P. Razafimahatratra, Michael D. Schwartz, Rathy M. Phillips, and John D. Oswald have all assisted in many ways and their efforts are appreciated. The entomology graduate students are a group of people that contributed greatly to my mental stability, and provided a relaxed, but intense professional habitat. I thank all of them, and I particularly wish to thank Peter Follett, Joe Furnish, Jeff Witcosky, Russell Messing, and Paul Hanson. I would also like to thank the technical assistance of Scott Wood, Sandy Potter, Mark Morris, Stuart Sloane, and Gerry Baker. This thesis would not have been completed without the emotional, spiritual, and financial support of my mother, my father, my sister, my cousin, George E. Kosmides, and my friends, Theodore Montgomery, Fred Young, and William Trela. To these people I make a humble dedication in the realization that ink could not balance the debt I owe. TABLE OF CONTENTS Page CHAPTER 1. GENERAL INTRODUCTION 1 CHAPTER 2. A Generic Reclassification of the Subfamily DicyphinaeReuter (Heteroptera:Miridae) INTRODUCTION 6 MATERIALS 9 METHODS 11 TERMINOLOGY 16 NOMENCLATURE 18 HISTORICAL REVIEW .19 BIOLOGY 22 SUBFAMILY DICYPHINAE REUTER 25 SYNOPSIS OF SUBFAMILYDICYPHINAB REUTER. 34 KEY TO GENERA OF SUBFAMILY DICYPEINAE 36 TAXONOMY OF THE GENERA OF DI CYPHINAE. ..42 Genus Cainpyloneuropsis Poppius . 42 Genus Canyloneura F.eber 49 Genus Chjus Distant 55 GenusCyrtopeltis Fieber 58 Genus Dicyphus Fieber.....................64 KEY TO TILE SUBGENERA OF THE GENUSDICYPHUS FIEBER 71 Pare Subgenus Brachyceraea Fieber 72 Subgenus Dicyphus Fieber 79 Subgenus Idolocoris Douglas and Scott 88 Subgenus Mesodicyphus Wagner 94 Subgenus Uhierejia Cassis 95 Genus Engytatus Reuter 101 Genus Glarisia Cassis 112 Genus Macrolophus Fieber 117 Genus Nesidocoris Kirkaldy 128 Genus Setocoris China and Carvalho 137 Genus Singhalesia China and Carvalho 143 Genus Tupiocoris China and Carvalho 148 Genus Usjngerella China and Carvalho 160 GENERA RETAINED IN SUB FAMILY (NOT EXAMINED) 164 GENERA REMOVED FROM THE SUBFAMILY DICYPHINAE 166 PRYLOGENETIC ANALYSIS OF THE GENERA OF THE DICYPHI.NAE ..................1 73 CHAPTER 3. A Systematic Study of the Genus Dicyphus Fieber in the Western Hemisphere INTRODUCTION 289 MATERIALS 291 Page METHODS ...294 SYNOPSIS OF THE GENUS DICYPHUS IN ThE WESTERN HEMISPHERE 294 KEY TO SPECIES OF DICYPHUS FOR WESTERN HEMISPHERE 295 TAXONOMY 299 Dicyphus (Uhierella) discrepans Knight 299 Dicyphus (Ub].e;ella) famelicus (Uhier) 306 Dicyphus(Uhlere],la) gaci1entus Parshley 310 Dicyphus (Uhierella) hesperus Knight 314 Dicvpbus (Uhierella) nigracorium Cassis 320 Dicyphus (Uhlere]21a) occidentalis Cassis 325 Dicyphus (Uhierella) paddocki Knight 329 Dicyphus (Uhierella) vestitus Uhier 336 Dicyphus(Idolocoris) pallicornis (Neyer-Dur) 338 LITERATURE CITED 359 List ofFigures Page CHAPTER 2. 11.1-11.16. Dorsal and lateral view of head andpronotum 237 11.17-11.32. Dorsal and lateral view of head and pronotum 239 11.33-11.47. Thoracic structures 241 11.48-11.77. Femoral trichobothria 243 11.78-11.92. Wings 245 11.93-11.110. Wings 247 11.111-11.127. Pygophore 249 11.128-11.167. Left and right claspers ........... 251 11.168-11.186. Aedeagus. ....... 253 11.187-11.201. Aedeagus 255 11.202-11.221. Female bursa copulatrix ........... 257 11.222-11.225. Thoracic pleura 259 11.226-11.230. Thoracic pleura 261 11.231-11.236. Thoracic pleura 263 II 237-Il 242 Thoracic pleura 265 II 243-lI 248 Pretarsus 267 11.249-11.256. Pretarsus 269 11.257-11.262. Pygophore 271 II 263-Il 268Pygophore 273 11.269-11.271. Pygophore ...275 Page 11.272. Babitus of Dicyphus (Idolocpris) regulus . 277 11.273. Habitus of Dicyphus (Uhierella) addocki ..., . ..279 11.274. Habitus ofTuDiocorip californica 281 11.275. Cladogram showing the distribution of synapomorphies inDicyphinae genera 283 II 276 C].adogram showing theregional distribution of Dicyphinae genera 285 11.277. Cladogramshowingthe host plant family associations forDicypbivaegenera 287 CHAPTER 3.Diagnostic characters of Dicyphus (Uhierella)species and .. (Idolocoris) pallicornis 111.1-111.25. 357 List of Tables Page CHAPTER 2. Characters and character states for the genera of Dicyphinae 174 Raw data f or the cladistic analysis of the genera of the Dicypbinae 180 Regional distribution of the Dicyphinae genera, and the outgroup Feliscus 193 List of Maps Page CHAPTER 2. 11.1 Canpy1oneuropsis 203 11.2 Campyloneura 205 11.3 Chius .. ..... 11.4 Cyrt.ppeltis ............... .209 11.5 Dicyphus (Bracbycerae) ......... .211 11.6 Dicyphus (Dicyphxs) 213 11.7 Dicvphus (Id9locoris) 215 11.8 DicyTbuL (Mesodicyphus) 217 11.9 Dicyphus (Uhierella) 219 11.10 Engytatus 221 11.11 Glarisja 223 11.12 Macrolophus ...... 225 11.13 Nesidocoris 227 11.14 Setoc9rj .229 11.15 Singlialesia 231 11.16 Tupiocoris ...233 11.17 tlsingerella 235 CHAPTER 3. 111.1 p. (flQ.ere11a) discrepans 339 III 2 D (Uh1ere11a famelLcus 341 III 3 B (Ublerella) gracilentus 343 III 4 D (Uhierella) hesperus 345 111.5 D. (Uhierella) nigracoriurn 347 Page 111.6 D. (Uhierella) occidentalis .......349 111.7 D. (Uhierella) paddocki . 351 111.8 D. (U11ere1la) vestitus .353 111.9 U. (Idolocoris) pallicornis 355 A Systematic Study of the Subfamily Dicyphinae (Reteroptera: Miridae). CHAPTER 1 GENERAL INTRODUCTION The Miridae is the most speciose family in the Heteroptera and exhibits a wide diversity in morphology, biological habits, and host plant associations. Many of the species are pbytophagous, although mycetophagy (e.g Cylapinae, Schuh, 1976) and predation (Deraeocorinae, Razafiinahatratra, 1980) are known, and it is quite probable that oligophagy, which is common in the Dicyphinae (Cobben, 1968), is prevalent in other mind taxa. At present, the systematic knowledge of the family is based chiefly on the fauna of the Palearctic and eastern Nearctic regions. Knowledge of the minds of the Ethiopian (Odhiambo, 1962; Linnavouri, 1975), Australasian (Carvaiho and Gross, 1979), and Oceanic (Carvalbo, 1956) regions is limited Carvalbo and his students (1945-present) have contributed greatly to the description of the Neotropical mind fauna, however, much remains to be accomplishedThe family is cosmopolitan in distribution.