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Biological and Morphological Studies of The AN ABSTRACT OF THE THESIS OF CAROL ANN MUSGRAVE for the DOCTOR OF PHILOSOPHY (Name) (Degree) in ENTOMOLOGY presented on May 6, 1974 (Major) (Date) Title: BIOLOGICAL AND MORPHOLOGICAL STUDIES OF THE SCAPHYTOPIUS ACUTUS COMPLEX (HOMOPTERA: CICADELLIDAE) Abstract approved: Redacted for Privacy Dr. Paul Oman Reinterpretation of the leafhopper tribe Scaphytopiini reveals it to be primarily a New World taxon with only limited representation in the Palearctic, Oriental and possibly Ethiopian regions.Approxi- mately 75 species assigned to three genera are known from the United States and Canada with at least another 75 species representing an undetermined number of genera and subgenera in Mexico, Central America and South America. The Scaphytopius acutus complex is the dominant element of the tribe in temperate North America in terms of number, distribution and economic significance.Proper identification of elements in this complex is important because of its implication as a vector of various plant pathogens.Using structural and distributional evidence, the morphologically diverse S. acutus complex can be subdivided into four subspecies.In the past, three of these, acutus, dubius and delongi, have been considered as either distinct species or synonyms of acutus; the fourth subspecies, cirrus,is described as new.Cross breeding experiments with laboratory cultures from three of these subspecies established that interpopulation fertility exists in varying degrees and apparently is not directly correlated with geographic proximity. Morphometric analyses of specimens from field collections and laboratory cultures indicated that both inter- and intrapopulation variation were consistent with the subspecies concept ofthe acutus complex. Several color and pattern morphs have been found amongthe nymphs of some field populations.Laboratory investigations of five of these morphs show that these are genetically determinedbut not sex limited; also, adult females produced by the five morphs differin fitness and development rates in the laboratory.Both field and laboratory studies suggest that the complex exists asseveral biotypes with respect to host utilization for feeding and ovipositing.Field populations have been collected from a wide variety of habitat types and altitudes ranging from sea level to nearly 7000 ft(2134 m). Suitable host plants include various members of the Ericaceae, Rosaceae, Leguminosae and Compositae and Rhamnaceae. Biological and Morphological Studies of the Scaphytopius acutus Complex (Homoptera: Cicadellidae) by Carol Ann Musgrave A THESIS submitted to Oregon State University in partial fulfillment of the requirements for the degree of Doctor of Philosophy June 1974 APPROVED: Redacted for Privacy Professor of Entomology in charge of major Redacted for Privacy Acting Head of Department of Entomology Redacted for Privacy Dean of Graduate School Date thesis is presented May 6, 1974 Typed by Mary Jo Stratton forCarol Ann Musgrave ACKNOWLEDGEMENTS I am grateful to my major professor, Dr. Paul Oman, for suggesting this problem, collecting some of the specimens and offering much advice and constructive criticism during the study. I also wish to thank Drs. Knud G. Swenson, James A. Kamm and Kenton L. Chambers for their helpful comments and other assistance. They and Dr. Paul 0. Ritcher also reviewed the manuscript, suggest- ing several improvements.Special thanks also go to Dr. Robert Whitcomb, Insect Pathology Laboratory, Beltsville, Maryland, and Mr. Steve Koinzan, Department of Entomology, University of Nebraska at Lincoln, for collecting and shipping two cultures of Scaphytopius acutus used in the laboratory phases of this study. The following institutions loaned specimens of the S. acutus complex for this investigation:U. S. National Museum, University of Kansas, California Academy of Science, South Dakota State Univer- sity, North Dakota State University, Texas A and M University, Illinois Natural History Survey, Ohio State University, University of Idaho, Washington State University, Montana State University, Los Angeles County Museum of Natural History, University of California at Riverside, University of California at Berkeley and University of California at Davis.Special thanks also go to Dr. Leon W. Hepner of Mississippi State University for loaning his personal collection of the acutus complex. This study was supported in part by an NDEA Fellowship awarded to the author.National Science Foundation Grants GB-7706 and GB-28292 "Phylogeny and Distribution of Nearctic and Palearctic Leafhoppers, " to Dr. Paul Oman contributed to the field and labora- tory phases of the investigation. TABLE OF CONTENTS Page L INTRODUCTION 1 II.LITERATURE REVIEW 3 III.MATERIALS AND METHODS 9 Morphological Survey 9 Dissection of Male Genitalia 10 Biological Review of the Scaphytopius acutus Complex 14 Field Studies and Distributional Records 14 Source of Biotypes for Laboratory Cultures 16 Laboratory Culture Methods 18 Egg Handling 20 Dissections of Females 20 Development of Nymphs 21 Color Pattern Morphs 22 Oviposition Tests 23 Male Choice Experiments 24 Hybridization Experiments 26 IV,CHARACTERIZATION AND TAXONOMY OF THE TRIBE SCAPHYTOPIINI 29 Key to Genera in the Scaphytopiini 30 Key to Nearctic Subgenera of Scaphytopius Ball 32 Key to Species Groups of Scaphytopius (Cloanthanus) Ball 35 Key to Species in the trilineatus Species Group 38 V. MORPHOLOGICAL SURVEY OF THE SCAPHYTOPIUS ACUTUS COMPLEX 37 Color 43 Structural Features 44 Genitalia 45 Scaphytopius (Cloanthanus) acutus acutus (Say) 46 Scaphytopius (Cloanthanus) acutus dubius (Van Duzee) 48 Scaphytopius (Cloanthanus) acutus delongi Young 50 Page Scaphytopius (Cloanthanus) acutus cirrus new subspecies 52 Distribution of the Subspecies of acutus 54 Inter- and Intrapopulation Variation in the acutus Complex 56 VI.BIOLOGICAL STUDIES OF THE SCAPHYTOPIUS ACUTUS COMPLEX 65 Reproductive Compatability of the four S. acutus Laboratory Cultures 77 Definitive Hybridization Experiments 80 Color and Pattern Morphs in Nymphs 94 The Waldport Culture 104 VII.DISCUSSION AND CONCLUSIONS 112 BIBLIOGRAPHY 135 APPENDIX 140 LIST OF PLATES Figure Page PLATE I 1 Scaphytopius acutus acutus (Say).(a) Dorsal view of crown, pronotum and scutellum; (b) Face; (c) Profile view of head and prothorax. 118 2 Japananus hyalinus (Osborn).(a) Forewing; (b) Ventral view of connective and one style; (c) Ventral view of valve and plates. 118 3 Scaphytopius acutus acutus (Say).Forewing showing venation only. 118 4 Ascius triangularis De Long.(a) Dorsal view of aedeagus; (b) Female seventh sternum. 118 5 Scaphytopius (Convelinus) Ball, aedeagi rep- resentative of the species groups. (a) torridus group, S. torridus (Ball); (b) irroratus group, S. irroratus (Ball); (c) californiensis group, S. californiensis Hepner. 118 6 Scaphytopius fuscifrons (Van Duzee).Crown pattern. 118 7 Scaphytopius magdalensis group, connective and paraphyses.(a) S. magdalensis (Provancher); (b) S.fulvus (Osborn). 118 PLATE II 8 Scaphytopius angustatus (Osborn).Lateral view of aedeagus. 120 9 Scaphytopius cinereus (Osborn and Ball). Lateral view of paraphyses and aedeagus. 120 10 Scaphytopius acutus acutus (Say).Pygofer margin. 120 Figure Page 11 Scaphvtopius oregonensis (Baker).(a) Pygofer margin; (b) Dorsal view of connective and paraphyses; (c) Lateral view of aedeagus. 120 12 Scaphytopius argutus De Long.(a) Aedeagus, lateral view; (b) Ventral view of aedeagus. 120 13 Scaphytopius utahensis Hepner.(a) Dorsal view of connective and paraphyses; (b) Lateral view of aedeagus. 120 14 Scaphytopius graneticus (Ball).(a) Dorsal view of connective and paraphyses: (b) Lateral view of aedeagus; (c) Ventral view of aedeagus. 120 15 Scaphytopius canus Hepner.(a) Dorsal view of connective and paraphyses; (b) Lateral view of aedeagus. 120 16 Scaphytopius trilineatus (Ball).(a) Lateral view of connective and paraphyses; (b) Valve and plates; (c) Lateral view of aedeagus. 120 17 Scaphytopius trilineatus spicatus Hepner. (a) Dorsal view of connective and paraphyses; (b) Valve and plates; (c) Lateral view of aedeagus. 120 PLATE III 18 Scaphytopius acutus acutus (Say).Live insect photographed from the Maryland culture. 121 PLATE IV 19 Scaphytopius (Cloanthanus) acutus acutus (Say). (a) Dorsal view of connective, style and paraphyses; (b) Lateral view of aedeagus; (c) Ventral view of aedeagus; (d) Lateral view of style, connective and paraphyses. 123 Figure Page PLATE IV (cont'd) 20 Scaphy_topius (Cloanthanus) acutus dubius (Van Duzee).(a) Dorsal view of connective, style and paraphyses; (b) Lateral view of aedeagus; (c) Ventral view of aedeagus; (d) Lateral view of style, connective and paraphyses. 123 21 Scaphytopius (Cloanthanus) acutus delongi Young.(a) Dorsal view of connective, style and paraphyses; (b) Lateral view of aedeagus; (c) Ventral view of aedeagus; (d) Lateral view of style, connective and paraphyses. 123 22 Scaphytopius (Cloanthanus) acutus cirrus new subspecies.(a) Dorsal view of connective, style and paraphyses; (b) Lateral view of aedeagus; (c) Ventral view of aedeagus; (d) Lateral view of style, connective and paraphyses. 123 PLATE V 23 Scaphytopius acutus cirrus new subspecies. Laboratory reared from The Dalles culture. Dorsal view of the connective, style and paraphyses with lateral view of the aedeagus. 125 24 Scaphytopius acutus cirrus new subspecies. Laboratory reared from The Dalles culture. Dorsal view of the connective, style and paraphyses with lateral view of the aedeagus. 125 25 Scaphytopius acutus cirrus new subspecies. Laboratory
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