GREAT BASIN NATURAUST MEMOIRS Brigham Young University Number 12 MC2 LIBRARY hm 1 ;^ iQQi Research inthe mrd Auchenorrhynchai'"'''' Homoptera: A Tribute to Paul W, Oman GREAT BASIN NATURALIST Editor. Stephen L. Wood, Department of Zoology, 290 Life Science Museum, Brigham Young University, Provo, Utah 84602. Editorial Board. Kimball T. Harper, Chairman, Botany and Range Science; Perron L. An- dersen, Zoology; James R. Barnes, Zoology; Hal L. Black, Zoology; Jerran T. Flinders, Botany and Range Science; Stanley L. Welsh, Botany and Range Science. All are at Brigham Young University. Ex Officio Editorial Board Members include Bruce N. Smith, Dean, College of Biological and Agricultural Sciences; Norman A. Darais, University Editor, University Publications; Stephen L. Wood, Editor, Great Basin Naturalist. The Great Basin Naturalist was founded in 1939. The journal is a publication of Brigham Young University. Previously unpublished manuscripts in English pertaining to the biological natural history of western North America are accepted. The Great Basin Naturalist Memoirs series was established in 1976 for scholarly works in biological natural history longer than can be accommodated in the parent publication. The Memoirs appears irregularly and bears no geographical restriction in subject matter. Manuscripts for both the Great Basin Naturalist and the Memoirs series will be accepted for publication only after exposure to peer review and approval of the editor. Subscriptions. Annual subscriptions to the Great Basin Naturalist are $25 for private individuals and $40 for institutions (outside the United States, $30 and $45, respectively), and $15 for student subscriptions. The price of single issues is $12. All back issues are in print and are available for sale. All matters pertaining to subscriptions, back issues, or other business should be directed to the Editor, Great Basin Naturalist, Brigham Young University, 290 Life Science Museum, Provo, Utah 84602. The Great Basin Naturalist Memoirs may be purchased from the same office at the rate indicated on the inside of the back cover of either journal. Scholarly Exchanges. Libraries or other organizations interested in obtaining either journal through a continuing exchange of scholarly publications should contact the Brigham Young University Exchange Librarian, Harold B. Lee Library, Provo, Utah 84602. Manuscripts. See Notice to Contributors on the inside back cover. 10-88 1100 36917 ISSN 017-3614 GREAT BASIN NATURALIST MEMOII Brigham Young University dumber 12 Research in the Auchenorrhyncha, Homoptera: A Tribute to Paul W. Oman CONTENTS Preface. Mervin W. Nielson 1 Paul W. Oman — an appreciation. John D. Lattin 3 Some new cicadellids from grasslands of Karachi, Pakistan (Homoptera: Cicadellidae). Manzoor Ahmed, A. Qadeer, and K. F. Malik 10 Classification of the subgenus Athijsanella , genus Athysanella Baker (Homoptera: Cicadellidae: Deltocephalinae). H. Derrick Blocker and James W. Johnson 18 A new genus, lleopeltus, related to Chlorotettix (Homoptera: Cicadellidae). Paul S. Cwikla 43 Some aspects of the biology, morphology, and evolution of leaflioppers (Homoptera: Cicadelloidea and Membracoidea). J. W. Evans 61 Revision of the genus Calliscarta (Homoptera: Cicadellidae: Neobalinae). Paul H. Freytag 67 A new species oiPazu Oman from eastern California (Homoptera: Cicadellidae: Delto- cephalinae). Raymond J. Gill 82 Evidence for an Indo-Pacific origin of Hawaiian endemics in Balcltitha and related genera (Cicadellidae: Macrostelini). W. J. Knight and M. D. Webb 86 Colladonus and related genera of Mexico and Central America with new taxa and synonymy (Homoptera: Cicadellidae). M. W. Nielson 103 New World Fulgoridae, Part I: genera with elongate head processes. Lois B. O'Brien . . 135 A numerical taxonomic analysis of interspecific morphological differences in two closely related species of Cicada (Homoptera, Cicadidae) in Portugal. J. A. Quartau 171 Revision of the Nirvaninae (Homoptera: Cicadellidae) of the Indian subcontinent. C. A. Viraktamath and C. S. Wesley 182 Genus Flexa7nia: new species, phylogeny, and ecology. Robert F. Whitcomb and Andrew L. Hicks .' 224 Afronisia, a new African genus of Meenoplidae (Homoptera: Fulgoroidea). Michael R. Wilson 324 Delphacidae of Alaska (Homoptera: Fulgoroidea). Stephen W. Wilson 335 Taxa in the Homoptera, Auchenorrhyncha described by Paul W. Oman 344 Index 349 Great Basin Naturalist Memoirs Research in the Auchenorrhyncha, Homoptera: A Tribute to Paul W. Oman No. 12 Brigham Young University, Provo, Utah 1988 PREFACE A committee was formed late in 1986 to organize and develop a volume of taxonomic papers on the Auchenorrhyncha, Homoptera to honor Dr. Paul W. Oman for his outstand- ing contributions in leafhopper systematics. This volume is the result of efforts by the committee and by contributors to the work. The 15 papers form not only a dedicatory trea- tise of Auchenorrhyncha systematics, but also a printed emblem of esteemed appreciation to Dr. Oman for his untiring efforts and many years of dedicated service in promoting leaf- hopper systematics in particular and the ento- mological sciences in general. The biography of Dr. Oman, prepared by his long-time associate John Lattin, and a spe- cial review of the biology, morphology, and evolution of leaftioppers by the Australian tax- onomist, John Evans, comprise part of the volume. The remainder consists of a selected group of 10 taxonomic papers on leaffioppers, three on planthoppers, and one on cicadas. The scope of these papers ranges from the description of a single taxon to a full taxonomic treatment of a higher category (subfamily). Each paper offers important data on the bio- geography of the respective group. Others cover host plant associations and relation- ships, phylogeny, and numerical taxonomy. Authors of the volume are long-time friends and associates of Dr. Oman, including a for- mer graduate student. Dr. C. A. Viraktamath. A symposium organized by H. Derrick Blocker and Paul H. Freytag was held at the 1988 Annual Meeting of the Entomological Society of Louisville, Kentucky. The culmina- tion of this symposium was the keynote ad- dress by Robert F. Whitcomb, the presenta- tion of papers, and the presentation of abound copy of this volume to the honoree by Mervin W. Nielson. The committee expresses appreciation to Dr. Stephen L. Wood, editor of The Great Basin Naturalist, Brigham Young University, for his efforts in editing and preparing the volume for publication. We also acknowledge Great Basin Naturalist Memoirs No. 12 Mrs. Terry Simmons and Mrs. Karen Jensen, Monte L. Bean Museum, Brigham Young University, for their invaluable assistance in typing those manuscripts that required con- version to the IBM WordPerfect system. Mervin W. Nielson, Coordinator H. Derrick Blocker Leon W. Hepner Robert F. Ruppel Paul H. Freytag James P. Kramer Organizing Committee PAUL W. OMAN— AN APPRECIATION John D. Lattin' Abstract — The contributions to professional entomology made by Paul W. Oman are reviewed. A bibliography of his published contributions to this field from 1930 to 1987 is included. I first met Paul Oman in December 1950 in Denver, Colorado, at the national meeting of the Entomological Society of America. He was in the uniform of the U.S. Army with the rank of major, having been called up again to serve in the Korean War (or "ruckus" as Paul preferred to call it). I was a graduate student at the University of Kansas, working with H. B. Hungerford. Dr. Hungerford encouraged me to attend the meeting, as did the other faculty members. He took special care to introduce the graduate students to other entomologists at the meeting, including Paul Oman, himself a graduate of the University of Kansas. My recollection of that meeting was that Paul took special interest in each student he met, even though his time was limited and he was quite busy with society affairs. He still takes time to meet and work with students. Now, thirty- eight years later, we share the same depart- ment, coffee room. Systematic Entomology Laboratory (OSU), and a good many hours of collaboration and discussion in and out of the field. Paul Wilson Oman was born in Garnett, Anderson County, Kansas, on 22 February 1908. He grew up on a farm about five miles outside of Garnett. His early education was in a rural school close to his home. He had to commute the five miles when he later entered Garnett High School. While growing up, he made the usual collections of natural history objects, including insects. A high school biol- ogy teacher was particularly influential in keeping alive Paul's interest in the natural sciences. He first attended the University of Kansas at Lawrence to strengthen his background in mathematics and English, since he was con- sidering the possibility of a career in the Navy, including attending Annapolis. He took a course in entomology to satisfy a biological science requirement and soon transferred to that department. Among the departmental faculty were H. B. Hungerford, chairman, K. C. Doering, P. B. Lawson, R. H. Beamer, and P. A. Readio. It is interesting to note that Hungerford (my own major professor in 1950) worked on aquatic Hemiptera, Readio on the Reduviidae, Kathleen Doering was a mor- phologist but worked on Homoptera, and both Lawson and Beamer worked not only on Homoptera but also on leafhoppers. Not sur- prisingly, Paul's interest in this group of in- sects was kindled at K.U., and he has contin- ued to work on the family during his entire scientific career. Paul Oman made a fine academic record at the University of Kansas, being elected to Sigma Xi, Phi Sigma, and Phi Beta Kappa in 1930. Prior to that date he had been elected to the Pen and Scroll in 1927 — an honorary soci- ety in the English Department. Those who have read Paul's papers or corresponded vdth him know that this award was well deserved and know too that he likes to communicate via the written memo. He graduated from the University of Kansas with an A.B. in entomol- ogy in the spring of 1930. Paul joined the recently organized Taxo- nomic Unit of the Bureau of Entomology, U.S. Department of Agriculture, in October 1930. This was to be a long association, for he retired from the USDA in 1967 when he joined the faculty of the Department of Ento- mology at Oregon State University. His re- sponsibilities included the auchenorrhync- hous Homoptera and the Psyllidae. Thus, his interests in the Homoptera, fostered by the faculty at the University of Kansas, provided Systematic Entomology Laboratory, Department of Entomology, Oregon State University. Corvallis, Oregon 97331. Great Basin Naturalist Memoirs No. 12 Fig. 1. Paul W. Oman. his entree into his profession. The need for a speciahst in the "hard" Homoptera was prompted by the extensive ecological work being done on the beet leafliopper and associ- ated insects, especially in the Twin Falls, Idaho, region. This fortuitous association with Circulifer tenellus (Baker), a vector of "curly- top of beets in the western United States, also provided a basis of his lifelong interest in applied systematics — the application of sys- tematic techniques toward the solution of problems of concern to society. His ultimate discovery that this insect was native to the Middle East (rather than South America as some thought) (Oman 1936, 1948) lead to ex- plorations for biological control agents from the Middle East. Further, this discovery strengthened his conviction that systematics should play an important role in biological control. Some years later, when he assumed charge of the entomological taxonomic unit of the US DA, he was instrumental in adding the biological control unit, resulting in the joint organization we see today. During his early years in Washington, D.C., he attended evening classes at George Washington University where he majored in entomology. He had extra courses from the University of Kansas, and these, together with his course work transferred to Lawrence from George Washington, plus a thesis, re- sulted in an M.A. degree in entomology from the University of Kansas in 1935. Not surpris- ingly, his thesis was on the leafhoppers and entitled "A Generic Revision of American Bythoscopinae and South American Jassinae," later published in the well-known University of Kansas Science Bulletin (Oman 1938). He continued his academic work at George Washington University as time permitted and received a Ph.D. degree from that institution in 1941. His thesis, somewhat revised, ap- peared under the title, "The Nearctic Leaf- hoppers (Homoptera: Cicadellidae), a Gener- ic Classification and Check List" (Oman 1949). As Z. P. Metcalf stated (1950, Ann. Entomol. Soc. Amer. 43: 458), "This is one of the most outstanding recent contributions to the study of one of the most difficult families of the Homoptera, and indeed it was. Paul pro- duced this landmark publication by working quietly, steadily, and efficiently, and accom- plishing a great deal — a Paul Oman hallmark. He was working as a taxonomic specialist at this time, and so the courses and thesis work were done largely after hours. He was the scientist responsible for research on the Auchenorrhyncha. The hiatus between the completion of his dissertation and the appear- ance of the published version was caused by World War II. Paul Oman entered the U.S. Army as a first lieutenant in the Medical Corps in October 1942 and was assigned to the South Pacific and the Far East commands. He served in that area, being promoted to captain, until he re- turned and left active service in February 1946. While in the South Pacific, he was re- sponsible for the organization and direction of entomological activities on Guadalcanal (No- vember 1943-February 1945) and Okinawa (April-June 1945). Although other medical entomology problems were involved, the con- trol of malaria vectors was the chief effort in both locales. The details of some of these ac- tivities can be found in Oman and Christen- son (1947) and Harper, Downs, Oman, and 1988 Lattin: PaulW. Oman Levine (1963). At the time of his discharge from the U.S. Army and his return to the Division of Insect Identification, US DA, he could hardly anticipate that the experience gained in medical entomology would be used again — in 1950, when he was recalled into the Army to serve three years in the Korean War. During the four years following his return from the South Pacific, Paul was deeply in- volved in the auchenorrhynchous Homop- tera. He became project leader in the Hem- iptera and ultimately assistant division leader of the Division of Insect Identification. He published a number of papers, including his 253-page generic classification of the Nearctic leaflioppers, as well as an account of some of the medical entomology work done in the South Pacific. His interest in applied system- atics is reflected in the papers of this period. In September 1950 he was reactivated to regular duty in the U.S. Army at the rank of major and assigned to the Far East during the Korean War. His first assignment was as ento- mologist. Headquarters, 3rd Army. Later he became chief of the Department of Entomol- ogy, 406th Medical General Laboratory in Tokyo, and then commanding officer. Far East Medical Research Unit, the position he held until his discharge in August 1953. He published several papers dealing with medi- cal entomology during this time, and, of spe- cial note to homopterists, a paper describing three new species of Errhornus with a key to the species. I doubt that he ever thought he would devote many of his later years to an intensive study of the genus Errhomus (Oman 1987), but we are glad he did. Once again he was discharged from the U.S. Army and again returned to the U.S. Department of Agriculture, this time as the head of what became the Insect Identification and Parasite Introduction Research Branch, Agricultural Research Service. His systematic work continued, as did his deep involvement in all aspects of entomology, entomological administration, and the Entomological Soci- ety of America. Gradually, his writing began to reflect his increased responsibilities and his ever-broadening interests, but always there was a deep interest in the leaflioppers. Part of his administrative responsibilities involved increased international activities. He was the leader of the United States ento- mological delegation to the USSR in 1959 un- der the U.S. -USSR Scientific and Cultural Exchange Program. In October 1960 he moved to New Delhi, India, to become the director. Far East Regional Research Office, Foreign Research and Technical Programs Division, ARS, USDA. There he was respon- sible for all technical and administrative as- pects of agricultural research studies in Asiatic countries where Public Law 480 funds were available for research of mutual interest to the United States and the foreign country. Paul returned home from India in Decem- ber 1962 to become assistant to the director. Entomology Research Division, ARS, USDA. A year later he was appointed assistant direc- tor of the Entomology Research Division, a position he retained until he retired from the U.S. Department of Agriculture in 1967. Paul's responsibilities included all extramural research programs, chairman of the Division Committee for the evaluations of research personnel, administration of divisional labora- tories, and an active role in the planning and development of programs and policies of the Entomology Research Division. Still he con- tinued to publish, chiefly on topics related to his responsibilities with the Division, but he found time to initiate and participate in the first conference under the new U.S. -Japan Cooperative Science Program. This confer- ence, held in Japan, dealt with arthropod- borne plant viruses (Maramorosch and Oman 1966). He was responsible for another U.S.- Japan Conference held in Washington, D.C., in 1967. This time the subject matter was "Systematics in Relation to the Geographical Distribution of Insects in the Pacific. " Paul Oman and Karl V. Krombein were the orga- nizers for the United States. It was obvious that Paul was held in high regard by his Japanese colleagues. I was pleased to have been a participant in that gathering; it was a special occasion. By this time Paul had retired after 37 years with the U.S. Department of Agriculture and had joined the faculty of the Department of Entomology at Oregon State University, and yet another chapter had be- gun in his productive career. He still main- tains active connections with the USDA, and, even as this is being written, he is assisting them to resolve a problem dealing with the Homoptera. Paul W. Oman, now Professor Paul W. Oman, joined the departmental faculty in the Great Basin Naturalist Memoirs No. 12 fall of 1967. He assumed many of my responsi- bilities in systematic entomology when I moved into administration. Characteristi- cally, he launched into his new career with great energy. He developed his own course in systematic entomology, developed a course in advances in pest management, and began to direct graduate student studies. His enor- mous experience in all aspects of entomology made him a highly valued member of the department, and his advice was sought (and still is) on many topics. As program director at Oregon State University, he participated in the NSF-funded Inter-University Program in Pest Population Ecology that ran from July 1969 until August 1975, a forerunner of the IPM programs at OSU we know today. Most of the leading ecologists of the world were brought to our campus (and other campuses as well) under this program. It was obvious that pest problems were universal problems, and solid science was required to solve them. During this time educational turmoil was ev- erywhere, but these well-organized programs provided some academic stability during un- stable times. The Pest Population Ecology program provided a solid core upon which was built a revised curriculum. It took him less than a year to obtain fund- ing from the National Science Foundation for his work on the systematics of the lealhop- pers. This funding extended from 1968 until he retired in 1975. Again, his publications reflected his efforts and interests as he pub- lished a series of leaffiopper papers. At the same time, he was publishing papers dealing with other aspects of his activities. He also renewed his interest in the genus Errhomus and spent many seasons in the field carefully documenting the complex systematic and bio- geographical problems associated with that taxon. The result, of course, is the superb monograph on Errhomus that appeared in 1987. While many people would be content to stop there, even before the publication was in press he took up the world catalog project with Bill Knight and Merv Nielson. Compla- cency is not a characteristic of Paul Oman. Paul was curator of the Entomology Mu- seum from 1967 to 1971. It must have seemed like child's play after having been responsible for the entire systematic operation of the USDA, and yet he took it very seriously and made major strides in establishing sound pro- tocol for the management of the collection. He added many needed books and sought out reprints from major systematists. An active effort was made to acquire, mount, label, and accession thousands of specimens. He made a special effort to develop the Homoptera col- lection that had languished. During the years since his release as official curator, he has added literally tens of thousands of specimens to the collection, and still continues to add more. These specimens are all mounted, la- beled, and, if leaflioppers, identified, and cu- rated. Talk about a curator's dream! We all know that collections become major scientific resources because of the work of many people with a common goal. Paul is the epitome of such a person. During the ensuing years he served ento- mology in many ways. He was on the editorial board of the Annual Review of Entomology from 1972 to 1976. He was chairman of the ad hoc Committee on Entomological Collections in the United States. He was a member of the Entomological Society of America Advisory Committee on Systematic Resources in Ento- mology from 1973 to 1975. He was a member of the National Policy Advisory Committee (NSF) for the National Drosophila Species Re- source Center in 1975. He was secretary of a study team that prepared a 138-page report in 1978 entitled "Biological Agents for Pest Con- trol: Status and Prospects ' for the USDA in cooperation with land-grant universities, the State Department of Agriculture, and the Agricultural Research Institute. When his long-time friend and colleague, Knud Swenson, then chairman of our depart- ment, was forced to step down because of a tragic illness, Paul took over as acting chair- man. He served in that capacity from 1973 to 1974 while the department sought a new de- partment chair. His long experience in sci- ence administration was most evident during that time, and we were able to experience yet another facet of his skills. He was a low-key, but a most efficient and effective, administra- tor. He paid attention to detail and provided leadership and stability that created an atmo- sphere of professionalism. He rarely com- plained and favored working out a solution to a problem. Because his dedication to the job at hand was always evident, people responded in a similar fashion. He was especially effec- tive in getting the most out of the resources available to him. 1988 Lattin: Paul W. Oman Paul Oman retired in 1975 — again. There was only an imperceptible change before and after this date — occasionally he would head for the golf course if the afternoon was espe- cially nice. Although I have never played golf with Paul, I know some who have — be pre- pared for a fierce competitor and don t wager very much on each hole. During the past 13 years of his "retirement" he has averaged six to eight hours a day in his office, has spent hun- dreds of hours in the field throughout the West, has given many reading and conference courses to many students, has served on nu- merous graduate student committees, has hosted foreign scientists and freely shared with them his vast experience with the Ci- cadellidae, and has identified thousands of leafhoppers for many individuals and institu- tions. One of my most recent interactions with Paul occurred on 16 March 1988, when he participated in the program planning session for Adam Asquith, one of my new doctoral students. Paul had given Adam a reading and conference course in zoological nomenclature during the preceding three months. In fact, one of the other graduate committee mem- bers brought a particularly knotty problem on nomenclature to the meeting. Ultimately, the problem was resolved by Paul and Adam. Paul brought several current articles from science that were appropriate to the graduate pro- gram and the proposed thesis topic. He has always given help freely to the students and faculty, and he did so again on 16 March 1988. His eightieth birthday had been celebrated three weeks earlier, and he had supposedly retired in 1975, but for Paul Oman it was just a regular day. As one who has known him since 1950, 1 can attest to his many contributions and deep devotion to entomology. He joined the ESA in 1929, served as president in 1959, was elected an honorary member in 1975, and received the Woodworth Award from the Pacific Branch in 1982. He joined our faculty in 1967, and while he holds the title Emeritus Professor of Entomology, his efforts are indis- tinguishable from those of regular faculty members. He is one of those rare scientists whose influence has extended far beyond his office and laboratory. I cherish our friendship. Paul Wilson Oman Bibliography L Oman, P. W. 1930. A new Paracoelidia (Homoptera: Cicadellidae). J. Kansas Entoniol. Soc. 3; 78-79. 2. Oman, P. W. 193L Some new Neocoelidia with notes on other species (Homoptera: CicadelHdae). J. Kansas Entomol. Soc. 4: 62-68. 3. Oman, P. W. 1931. Newjassinae, with notes on other species. J. Washington Acad. Sci. 21: 430-436. 4. Oman, P. W. 19.32. Two new Laevicephalus from CaUfornia. Proc. Entomol. Soc. Washington 34: 90-92. 5. Oman, P. W. 19.33. A classification of North American agallian leaflioppers. U.S. Dept. Agric. Tech. Bull. 372. 93 pp. 6. Oman, P. W. 1933. Phlepsius ishidae Matsumura in North America. Proc. Entomol. Soc. Washington 35: 205-206. 7. Oman, P. W. 19.34. New species and a new genus of deltocephaline leaflioppers (Hemiptera: Ho- moptera). Proc. Entomol. Soc. Washington 36: 75-81. 8. Oman, P. W. 1934. South American leafhoppers of the genus Agalliana. Rev. de Entomol. 4: 333-340. 9. Oman, P. W. 1934. The agallian leafhoppers of the Biologia material. Ann. Entomol. Soc. Amer. 27: 445-461. 10. Oman, P. W. 1935. New North American agallian leafhoppers with notes on other species. J. Kansas Entomol. Soc. 8: 9-16. 11. Oman, P. W. 1936. New Neotropical empoascan leaflioppers. J. Washington Acad. Sci. 26: 34-40. 12. Oman, P. W. 1936. A new Poblicia from Texas (Ho- moptera: Fulgoridae). J. Kansas Entomol. Soc. 9: 105-107. 13. Oman, P. W. 1936. Two new leaflioppers from tropi- cal America. Pan-Pac. Entomol. 12: 116-119. 14. Oman, P. W. 1936. Distributional and synonymical notes on the beet leafhopper, Eutettix tenellus (Baker). Proc. Entomol. Soc. Washington 38: 164-165. 15. Oman, P. W. 1936. South American leafhoppers of the genus Xerophloea Gerniar (Homoptera). Rev. de Entomol. 6: 394-401. 16. Oman, P. W. 1937. Fossil Hemiptera from the Fox Hills Sandstone (Cretaceous) of Colorado. J. Pale- ontol. 11:38. 17. Oman, P. W. 1937. The leaflioppers of the genus Scaphoidula (Homoptera: Cicadellidae). Rev. de Entomol. 7: 92-96. 18. Oman, P. W. 1937. The cinerosus group of the genus Laevicephalus (Homoptera: Cicadellidae). J. Washington Acad. Sci. 27: 474-479. 19. Oman, P. W. 1937. New eupterygine leafhoppers from Puerto Rico (Homoptera: Cicadellidae). Univ. Puerto Rico J. Agric. 21: 567-.572. 20. Oman, P. W. 1938. A generic revision of the Ameri- can Bythoscopinae and South American Jassinae. Univ. Kansas Sci. Bufl. 24: 343-420. 21. Oman, P. W. 1938. A contribution to the classification of South American agallian leafhoppers. Ann. Carnegie Museum 25: 351-460. 22. Oman, P. W. 1938. Revision of the Nearctic leafliop- pers of the tribe Errhomenellini (Homoptera: Great Basin Naturalist Memoirs No. 12 Cicadellidae). Proc. U.S. National Museum 85: 16.3-180. 23. Oman, P. W., and Nancy H. Wheeler. 19.38. The North American empoascan leafhoppers of the as- persa group (Homoptera; Cicadellidae). Proc. En- tomol. Soc. Washington 40: 1,33-147. 24. Oman, P. W. 1939. Revision of the genus Cprailis group with description of a new species (Homoptera: C^icadellidae). Proc. Entomol. Soc. 1988 Lattin: Paul W. Oman Washington 72; 30-32. 62. Oman, P. 1970. The Oregon State University insect collection. Bull. Entomol. Soc. of Amer. 16: 165. 63. Oman, P. 1971. A new Agallia from the western United States. J. Kansas Entomol. Soc. 44: 325- 372. 64. Oman, P. 1971. The iema\e oiThatuna gilletti Oman, with biological notes. Proc. Entomol. Soc. Washington 73: 368-372. 65. Oman, P. 1972. Documentation of biological control operations. International Organization for Biologi- cal Control, Western Hemisphere Region Sec. Newsletters: 1-2. 66. Oman, P. 1972. A new megophthalmine leafhopper from Oregon, with notes on its biology and behav- ior (Homoptera: Cicadellidae). Royal Entomol. Soc. London, J. Entomol. (B)41: 69-76. 67. Oman, P. 1972. The lealliopper subfamily Koebeli- inae (Homoptera: Cicadellidae). Published as part of a Festschrift honoring Professor K. Yasumatsu of Japan, pp. 129-139 (dated 1971, issued 1972). 68. Oman, P. 1974. Curtis P. Clausen ('32). (A vignette.) Cosmos Club Bull. 27: 3-5. 69. Oman, P. 1974. Identification and classification in pest management and control. Pages 77-86 in F. G. Maxwell and F. A. Harris, eds., Proc. Summer Institute on Biological Control of Plant Insects and Diseases. University of Mississippi Press, Jack- son. 647 pp. 70. Oman, P., and C. A. Musgrave. 1975. The Nearctic genera of Errhomenini (Homoptera: Cicadelli- dae). Melanderia21: 1-14. 71. Oman, P. 1976. Review of World catalogues of the Homoptera: Auchenorrhyncha. Bull. Entomol. Soc. Amer. 22: 161-164. 72. Gill, R. J., and Paul Oman. 1982. A new species and new distributional records for megophthalmine leafhoppers, genus Tiaja (Homoptera: Cicadelli- dae). Entomography 1; 281-288. 73. Klein, M., B. Raccah, and P. W. Oman. 1982. The occurrence of a member of the Circulifer tenellus species complex (Homoptera: Cicadellidae: Eu- scelini) in Israel. Phytoparasitica 10(4): 237-240. 74. Converse, R. H., R. G. Clarke, P. W. Oman, Sr., and G. M. Milbrath. 1982. Witches' broom disease of black raspberrv in Oregon. Plant Disease 66(10): 949-951. 75. Lattin, J. D., and P. Oman. 1983. Where are the exotic insect threats? Pages 93-137 in Charles L. Wilson and Charles L. Graham, eds.. Exotic plant pests and North American agriculture. Aca- demic Press. 498 pp. 76. Oman, P. 1985. A synopsis of the Nearctic Dorycephalinae (Homoptera: Cicadellidae). J. Kansas Entomol. Soc. 58: 314-336. 77. Oman, P., and R. 1. Sailer. 1986. The role of catalog- ing in the advancement of systematics and bio- geography. Tymbal (Auchenorrhyncha Newslet- ter) 7: 16-21. 78. Oman, P. 1987. Alary polymorphism in the Cicadelli- dae and its ecological implications. Pages 55-63 in M. R. Wilson and L. R. Nault, eds.. Proceedings of the 2nd International Workshop on Leafhop- pers and Planthoppers of Economic Importance. C.A.B. International Institute of Entomology. 368 pp. 79. Oman, P. 1987. The leafhopper genus Errhomus (Homoptera: Cicadellidae: Cicadellinae): system- atics and biogeography. Oregon State University, Department of Entomology, Systematic Entomol- ogy Laboratory Misc. Pub. No. 1: 1-72. SOME NEW CICADELLIDS FROM GRASSLANDS OF KARACHI, PAKISTAN (HOMOPTERA: CICADELLIDAE) Manzoor Ahmed', A. Qadeer', and K. F. Malik' Abstract. — A brief survey of grassland lealhoppers of Karachi during 1985 revealed the existence of five new species of the family Cicadellidae. These include Exitianus major, E. minor. Chiasmus karachiensis , C. lobosa, and Goniagnathiis bifurcatus . Recent studies of grassland leafhoppers of Pakistan by Ahmed (1986), Ahmed and Rao (1986), and Ahmed and Yunus (1986) showed that leafhoppers form a very important com- ponent of the phytophagous fauna of our grasslands. The leafhopper fauna described by these and other workers, e.g.. Blocker et al. (1972) from USA and Theron (1982) from South Africa, appears to be widely diversified. Andrezejewska (1984) believed that species composition of Auchenorrhyncha in grass- lands depends to a high degree on immigrant species, which indicates that most of the spe- cies encountered in grasslands move in from the neighboring crops when the latter are har- vested. Thus, grasslands serve not only as alternate feeding grounds for leafhopper pests of a number of crops but also as breeding places for many species of leafhoppers during their off-crop seasons. Studies of grassland leafhoppers could therefore be useful from many aspects. The leafhoppers described herein have been studied following the methodology out- lined by Ahmed (1985). Exitianus Ball The genus Exitianus Ball is known from all major regions of the world. Oman (1949) re- described the genus and listed the Nearctic species. Metcalf (1967) listed 35 world spe- cies, and Ahmed and Rao (1986) recorded £ . capicola (Kb.) and described E . peshawaren- sis from Pakistan. During the present study two more new species of the genus were col- lected from Karachi, Pakistan, and are de- scribed below. Exitianus m^jor Ahmed & Qadeer, n. sp. (Figs. 1 A-K) Length. — Male 4.2 mm, female 4.8 mm; head broad, rounded at anterior apex; median length of crown nearly equal to or slightly less than the interocular width; transocular width of head slightly more than midlength of crown; ocelli close to mesal margin of eyes on anterolateral margin; crown with dim brown markings, transverse band of dark brown color in middle, rest of crown pale browTi; frontoclypeus narrowing anteriorly; ante- clypeus longer than broad, narrowing anteri- orly; lora broad; pronotum possessing tiny spots of brown color on disc; scutellum dim brown, with two brownish spots anterolater- ally. Forewings with apex smoothly rounded; maximum width slightly prior to midlength; appendix broad, extending around wing apex up to 4th apical cell; apical cells 5; subapical cells 3; veins in the basal region possessing brownish markings all along; hind wings with 4 closed apical cells. Male. — Plate in ventral view broad in basal half, narrowed in apical half, possessing 6-7 macrosetae on ventrolateral surface; pygofer with posterior margin extended into narrow posterodorsal lobe, with one long and one small blackish macrosetae on lobe, style in dorsal view massive, sclerotized in marginal areas, membranous in middle, with broad, bluntly rounded lobe, apical extension spinose, curved laterad; connective Y-shaped with arms quite close to each other; aedeagus with preatrium reduced, dorsal apodeme well Department of Zoology, University of Karachi, Karachi-32, Pakistan. 10 1988 Ahmed et al. : New Grassland Cicadellidae from Pakistan 11 FiPs 1 A K Exitianus maior n sp.: A. head and thorax, dorsal view; B, head, ventral view; C, forewing; D, hind winfE gentf clpLTe S view' F, male plate, ventral view; G, style, dorsal v.ew; H, connective, dorsal v,ew; I, aedeagus, lateral view; J, aedeagus, dorsal view; K, female seventh sternum. developed, shaft in lateral view broad in basal 2/3, narrowed apically, smoothly curved dor- sally at apex. Female. — Seventh sternum narrowed pos- teriorly, posteromedian margin sinuate. Exitianus major appears quite close to E . capicola (Stal) in venation and shape of aedea- gus, but differs in its smaller size, shape ot connective, chaetotaxy of male plate, and to some extent in the shape of female seventh sternum. Holotype (male).— Pakistan: Karachi, grass, 12 Great Basin Naturalist Memoirs No. 12 Fig. 2 A-K. Exitianus mmor, n. sp.: A, head and thorax, dorsal view; B, head, ventral view; C, forewing; D, hind wing; E, genital capsule, lateral view; F", male plates, ventral view; G, style, dorsal view; H, connective, dorsal view; I, aedeagus, lateral view; J, aedeagus, dorsal view; K, female seventh sternum. 26. VI. 85, Qadeer. Seven paratypes, .same head projected in front, broadly rounded at data as holotype, all in Zoological Museum, University of Karachi, Karachi, Pakistan. Exitianus minor Ahmed & Qadeer, n. sp. (Figs. 2 A-K) Length. — Male 4.00 mm, female 4.5 mm; anterior apex; median length of crown less than interocular width; transocular width of head nearly ecjual to transverse width of pronotum; coronal suture present up to more than midlength; ocelli on anterolateral margin 1988 Ahmed ETAL.: New Grassland Cicadellidae from Pakistan 13 of crown, close to eyes; crown with brownish markings on disc and transverse brownish band at the level of anterior margin of eyes; ground color of crown more or less pale brown; frontoclypeus nearly parallel sided, narrowed at apex; anteclypeus narrowed ante- riorly, much longer than broad; lora broad; pronotum with some tiny spots in middle in anterior region, with posterior margin nearly transverse; scutellum pale brown, with oval- shaped, brownish spots anterolaterally. Forewing with apex broadly rounded; api- cal part slightly broader than basal part; ap- pendix broad, extending up to 5th apical cell; apical cells 5; subapical cells 3; hind wing with 4 closed apical cells. Male. — Plate in ventral view, broad basally, narrowed apically, possessing 4-5 macrosetae near apex on lateral margin, long row of 9-10 macrosetae from near base to apex on ventral surface; pygofer with poste- rior margin directed into posterodorsal broad and rounded lobe, lobe possessing 4-5 brownish macrosetae; style in dorsal view massive, membranous in middle, sclerotized along margins, preapical lobe flattened, al- most truncate mesally, possessing 3-4 macro- setae, apical extension prominent, spinose, extreme apex directed caudolaterad; connec- tive Y-shaped, with arms smaller than the stem, converging at their apex; aedeagus in lateral view, with preatrium reduced, dorsal apodeme present, appearing broad in dorsal view, shaft tubular, curved smoothly dorsad, apical part narrowed. Female. — Seventh sternum with posterior margin possessing three lobes, lateral lobes more projected than the median. Exitianus fninor appears close to E. ka- rachiensis Ahmed and E. major described hitherto in its venation and general pattern of male genitalia, but differs in its shape of con- nective and female seventh sternum. Holotype (male). — Pakistan: Karachi, grass, 12. V. 1985, Qadeer. Thirty paratypes, all in Zoological Museum, University of Karachi, Karachi, Pakistan. Chiasmus Mulsant and Rey Distant (1908) recorded Chiasmus uzelli Melichar from Ceylon. Pruthi (1930) de- scribed three new species from various parts of India and reassigned Kartwa mustelina Distant to the genus Chiasmus . The present description of two new species in Chiasmus is the first record of the genus from Pakistan. Chiasmus karachiensis Ahmed & Qadeer, n. sp. (Figs.3A-J) Length. — Male 2.3 mm; head strongly projected in front, apex of crown narrowed and rounded; median length of crown much less than interocular width; transocular width of head slightly more than transverse width of pronotum; coronal suture present up to more than midlength of crown; ocelli on anterolat- eral margin of crown, close to eyes; color of eyes blackish; crown possessing blackish brown dots throughout; ground color of crown pale yellow; pronotum yellowish, its posterior margin nearly straight; scutellum dim yellow; frontoclypeus narrowing anteriorly, blackish in color throughout, except median, longitu- dinal stripe of yellow color; anteclypeus black in middle; lora broad, expanded. Forewing with apex smoothly rounded; maximum width of wing slightly prior to mid- length; appendix broad, extending to 3rd api- cal cell; apical cells 4; subapical cells 2; hind wing with 4 closed apical cells. Male. — Plate in ventral view broad at base, middle and apical parts much narrowed, macrosetae 2-3 on ventral surface in apical half, apex of plate rounded and directed cau- dad; pygofer in lateral view with posterior margin narrowed into angular, posterodorsal, much-projected lobe, processes and setae usually absent; style in dorsal view membra- nous in middle, sclerotized along margins, preapical lobe short, narrow, and angular, apical extension dark brown in color, spinose and curved laterad; connective Y-shaped, with arms longer than the stem, arms apposed most of their anterior part, forming nearly rounded space in middle; aedeagus in lateral view with preatrium reduced, dorsal apo- deme absent, shaft tubular, somewhat curved dorsad towards apex, processes absent. C . karachiensis appears close to C . alata Pruthi in general appearance as well as pat- tern of venation and genitalia, but differs in the shape of pygofer. Holotype (male). — Pakistan: Karachi, grass, 1.XI.85, Qadeer. One paratype, same data as holotype, both in the Zoological Museum, University of Karachi, Karachi, Pakistan. 14 Great Basin Naturalist Memoirs No. 12 Figs. 3 A-J. Chiasmus karachiensis , n. sp.: A, head and thorax, dorsal view; B, head, ventral view; C, forewing; D, hind wing; E, genital capsule, lateral view; F, male plates, ventral view; G, style, dorsal view; H, connective, dorsal view; I, aedeagus, lateral view; J, aedeagus, dorsal view. Chiasmus lohata Ahmed & Qadeer, n. sp. (Figs. 4 A-J) Length, — Male 2.5 mm, female 3.0 mm; head projected and narrowed in front, apex rounded; median length of crown nearly equal to or slightly less than interocular width; transocular width of head equal to transverse width of pronotum; coronal suture present up to more than midlength; ocelli present on anterolateral margin of crown, close to eyes; crown possessing few, dim, brownish spots in middle along posterior margin; frontoclypeus broad, narrowed in apical 1/3; anteclypeus longer than broad; lora broad; pronotum pos- sessing rows of brownish markings. Forewing with apical margin smoothly rounded; apical cells 4; appendix broad, reaching 3rd apical cell; subapical cells 2; 1988 Ahmed et al.: New Grassland Cicadellidae from Pakistan 15 Figs. 4 A-J. Chiasmus lobata, n. sp.: A, head and thorax, dorsal view; B, head, ventral view; C, forewing; D, hind wing; E, genital capsule, lateral view; F, male plate, ventral view; G, style, dorsal view; H, connective, dorsal view; I, aedeagus, lateral view; J, aedeagus, dorsal view. veins with brownish shades along their length; hind wing with 3 closed apical cells. Male. — Plate in lateral view broad at base, narrowed in apical half, apex rounded, pos- sessing row of 2-3 macrosetae on ventral sur- face in apical half, another row of 7-8 micro- setae on lateral margin from near base to mid- length, length of plate more than the pygofer, apposed mesally at base; pygofer in lateral view with posterior margin broad, dented, more projected posterodorsally, disc with dorsoventral row of 4 macrosetae; anal tube moderately developed; style in dorsal view with preapical lobe narrowed, angular, apical extension thin, spinose, directed laterad; aedeagus in lateral view with preatrium re- duced, dorsal apodeme present, shaft tubu- lar, somewhat curved dorsad in middle, apex rounded and thin, directed caudad; connec- tive Y-shaped, with arms much longer than stem, arms confluent again forming space in middle. Chiasmus lobata appears quite close to C . karachiensis as well as to C . niger Pruthi and C . jagdishi Pruthi in the pattern of male geni- talia, but differs from them in the peculiar 16 Great Basin Naturalist Memoirs No. 12 shape of pygofer, particularly the posterodor- sal margin. HoLOTYPE (male). — Pakistan: Karachi, 15. XII. 85, grass, Qadeer. One paratype, same data as holotype, both in Zoological Museum, University of Karachi, Karachi, Pakistan. Goniagnathus Fieber The genus Goniagnathus Fieber was re- described by Dlabola (1954). Distant (1918) described three species of the genus from In- dia. Ahmed and Yunus (1986) reported G. guttulinervis Kirsch. from Pakistan. Another species of the genus is described here from grass. Goniagnathus bifurcattis Ahmed & Qadeer, n. sp. (Figs. .5 A-I) Length. — Male 5.0 mm; head with ante- rior margin broadly convex, smoothly rounded in front; median length of crown much less than the interocular width; transoc- ular width of head slightly more than the transverse width of pronotum; coronal suture up to more than midlength; ocelli on antero- lateral margin of head, close to apex; ante- clypeus longer than broad; lora broad; prono- tum with few brownish black dots along anterior margin and on posterior margin; scutellum grayish black throughout. Forewing with apex broad and rounded; maximum width of wing in basal half; ap- pendix broad, extending to 4th apical cell; apical cells 5; subapical cells 3; black markings present in appendix, 2nd, 3rd, and 4th apical cells, in some discal subapical cells and along all the veins; hind wing with 4 closed apical cells. Male. — Genital segment strongly sclero- tized, dark brown in color; plates in ventral view fused completely to form a single ovid structure, width of plate decreasing smoothly to rounded apex, with few blackish macrose- tae and marginal microsetae in apical part; pygofer in lateral view with posterior margin broad, nearly truncate, slightly sinuate pos- terodorsally, with at least three prominent groups of stout macrosetae, one closely placed group posterodorsally, one mixed group of short and long in middle of posterior margin in middle, and one group in posteroventral re- gion of disc, microsetae dispersed throughout pygofer; anal tube moderately long, possess- ing fine microsetae; posterior margin of pygofer finely dentate; style in dorsal view with preapical lobe minutely developed, api- cal part expanded, with extreme apex obliquely truncate; connective in dorsal view rod-shaped, arms indistinct, connective curved in middle; aedeagus in lateral view with preatrium reduced, dorsal apodeme well developed, closely associated with connec- tive; shaft of aedeagus short, tapered apically, long processes arising from atrium, subparal- lel to shaft, exceeding shaft in length; gono- pore subterminal on dorsal surface. Goniagnathus bifurcatiis is close to G. gut- tulinervis Kirschbaum in the general pattern of male genitalia, particularly the male plate and aedeagus, but differs in the shape of style, pygofer, and connective. HoLOTi'PE (male). — Pakistan: Karachi, grass, 10. XI. 85, Qadeer. One paratype, same data as holotype, both in Zoological Museum, Uni- versity of Karachi, Karachi, Pakistan. Literature Cited Ahmed. M 1985. Typhlocybinae of Pakistan. Pakistan Agric. Res. Council, Islamabad. 270 pp. 1986. Some investigations of leafhoppers of grass- lands and allied crops in Pakistan (in press). 1986. Some investigations of lealhoppers of grass- lands in Lower Sind, Pakistan (in press). Ahmed. M , andS Rao. 1986. Some commonly found leaf and planthoppers on vegetable plants in the sub- urbs of Peshawar. NWFP, Pakistan (in press). Andrezejewska. L. 1984. Ecological structure of Auchenorrhyncha meadow communities under increasing cultivation. Mitt. Sch. Entomol. Ges. 57(4): 40,5-406. Blocker, H D . T L Harvey, and J L Launchbaugh. 1972. Grassland leafhoppers. 1. Leafhopper popu- lations of upland seeded pastures in Kansas. Ann. Entomol. Soc. Amer. 6.5(1); 166-172. Distant. \V L 1908. The fauna of British India including Ceylon and Burma. Rh\nchota4: 1.56-419. 1918. The fauna of British India including Ceylon and Burma. Rhynchota 7; 1-109. Dlabola, J 1954. Fauna CSR Svazek-Homoptera. Prace Ceskoslocenske Akad. 1954: 7-339. Metcalf, Z. P 1967. General catalogue of the Ho- moptera. Fasc. VI(IO). 2695pp. C)nl\n, P W 1949. The Nearctic le;iflioppers (Homop- tera: Cicadellidae). A generic classification and check list. Mem. Entomol. Soc. Washington 3: 1-253. Pruthl H. S 1930. Studies on Indian Jassidae (Ho- moptera). Mem. Indian Mns. 11(1): 1-68. Theron. J G 1982. Grassland le;iflioppers (Hemiptera: Cicadellidae) from Natal, South Africa, with de- scriptions of new genera and species. Phytophy- lactica 14; 17-.30. 1988 Ahmed et al.; New Grassland Cicadellidae from Pakistan 17 Figs. 5 A-I. Goniagnathus bifurcatus , n. sp. : A, head and thorax, dorsal view; B, head, ventral view; C, forewing; D, hind wing; E, genital capsule, lateral view; H, aedeagus and connective, lateral view; I, aedeagus, dorsal view. CLASSIFICATION OF THE SUBGENUS ATHYSANELLA, GENUS ATHYSANELLA BAKER (HOMOPTERA, CICADELLIDAE, DELTOCEPHALINAE)^ H. Derrick Blocker^ and James W. Johnson^ Abstract. — The subgenus Athysanella Baker is reviewed and a possible phylogeny of the group is discussed. A total of 35 species is recognized. Descriptions, illustrations, and a key to males are presented. Twelve new species are described: aphoda, cursa, deserta,furnaca, hemijona, krameri, pastora, strobila, stylosa, vativala, and whitcombi are from the western United States, and marthae is from Mexico. The subgenera of Athysanella Baker have been reviewed by Blocker and Johnson (1988). The nominate subgenus revised herein contains 35 species, of which 12 are new. Distribution of this subgenus is primar- ily in the western half of the United States, with 4 species recorded from Mexico and 4 from western Canada. The hosts are primarily shortgrasses. Blocker (1984) has described the morpho- logical variation that occurs in both sexes be- cause of incompletely developed genital structures. Undeveloped and partially devel- oped specimens are often difficult to identify. We have attempted to illustrate sufficient characters to allow identification of males, but the user must attempt to make identifications from a series of specimens that appear to be fully developed. As an example, the hind tib- ial spur of partially developed males may be absent or reduced; the genitalia also may be reduced in size. Females are particularly difficult to deter- mine. It is best to try to associate them with a male and then check the shape of the posterior margin of sternum VII. There is much in- fraspecific variation in specimens of both sexes. Ball and Beamer (1940) should be con- sulted for additional and original descriptions; Blocker and Wesley (1985) and Wesley and Blocker (1985) should be consulted for distri- bution in Canada- Alaska and Mexico, respec- tively. Depositories Persons and institutions (acronyms are used in the text) furnishing material for this study include (the authors regret any omissions): UAz - University of Arizona, F. Werner. CAS - California Academy of Science, P. Arnaud and N. Penny. CDA - California Department of Agriculture, R. Gill and J. Sorensen. CNC - Canadian National Collection, K. G. A. Hamil- ton. IPL - Insect Pathology Laboratory, US DA, Belts- ville, A. L. Hicks and R. F. Whitcomb. KSU - Kansas State University. KU - Snow Entomological Museum, University of Kansas, R. W. Brooks. MWN - Collection of M. W. Nielson. OhSU - Ohio State University, P. S. Cwikla. OkSU - Oklahoma State University, R. A. Drew. OrSU - Oregon State University, P. W. Oman. UBC - University of British Columbia, G. Scudder and S. Cannings. USNM - U.S. National Museum, Natural History, J. P. Kramer. Athysanella (Athysanella) Baker Athysanella Baker, 1898; 185. Type species: Athysanella magdalena Baker, 1898, by original designation. This subgenus is characterized by the pres- ence of a hind tibial spur and the absence of a pygofer process (Blocker and Johnson 1988). It is a sister of the subgenus Gladionura. Species of the subgenus Athysanella: aphoda Blocker, n. sp. Nevada. aspera Ball and Beamer, 1940. California. bifida Ball and Beamer, 1940. Colorado, Montana, New Mexico, North Dakota, South Dakota, Texas, Wyoming, Canada. cursa Blocker, n. sp. Arizona, Colorado, New Mexico, Utah. deserta Blocker, n. sp. California. foeda Ball and Beamer, 1940. Wyoming. fredonia Ball and Beamer, 1940. Arizona, Colorado, New Mexico, Nevada, Utah. furnaca Blocker, n. sp. California. gardenia Osborn, 1930. Colorado, Wyoming. 'Contribution 88-.312-] from the Kansas Agriculture Experiment Station. ^Department of EntomoloKy, Kansa.s State University. Manhattan, Kansas 66506 Department of Entomology, University of Missouri, Columbia, Missouri 65221. 18 1988 Blocker, Johnson: Subgenus Athysanella 19 frigida Oshorn, 1930. glohosa Ball and Beanier, 1940. Arizona. hemijona Blocker, n. sp. California. incerta Ball and Beamer, 1940. Colorado, Kansa.s, Mon- tana, Nebraska, New Mexico, Wyoming. incongrua Baker, 1898. Colorado, Iowa, Kan.sas, Ne- braska, New Hampshire, New Mexico, Okla- homa, South Dakota, Texas, Wyoming. kadokana Knull, 1951. Colorado, Montana, Nebraska, South Dakota, Wyoming, Canada. kanahana Ball and Beamer, 1940. Arizona, Kansas, New Mexico, Utah. krameri Blocker, n. sp. California. laeta Ball and Beamer, 1940. Arizona, New Mexico. magdalena Baker, 1898. Arizona, California, Colorado, Montana, New Mexico, Nevada, Oklahoma, Texas, Wyoming, Mexico. inarthae Blocker, n. sp. Mexico. parca Ball and Beamer, 1940. Colorado, New Mexico. pastora Blocker, n. sp. New Mexico. plana Ball and Beamer, 1940. California. planata Ball and Beamer, 1940. California. robusta Baker, 1898. Colorado, Montana, Nebraska, North Dakota, Wyoming, Canada. montana Osborn, 1930. rostrata Ball and Beamer, 1940. California. salsa Ball and Beamer, 1940. Kansas, Texas, Mexico. strobila Blocker, n. sp. California, Idaho, Nevada, Ore- gon. stylosa Blocker, n. sp. New Mexico. supina Ball and Beamer, 1940. Texas. tenera Ball and Beamer, 1940. Nevada. (ereforans (Gillette and Baker), 1895. Colorado, Montana, North Dakota, Nebraska, Utah, Wyoming, Can- ada. utahna Osborn, 1930. Arizona, California, Colorado, Idaho, Nevada, Utah, Oregon, Washington, Canada, Mexico. vativala Blocker, n. sp. Nebraska, North Dakota. whitcombi Blocker, n. sp. Nevada. yumana Osborn, 1930. Arizona, California. Key to Athysanella Males, sensu stricto 1. Style rounded apically; pygofer variable; plates embrowned apically 2 — Style widened, usually bifid apically; pygofer usually broadly rounded apically; plates vari- able . ' ' 5 2(1). Pygofer extended apically; connective, plates, and vertex variable 3 — Pygofer truncate apically; connective shorter than style; plates truncate apically; vertex usually with fuscous spots .... gardenia Osborn 3(2). Aedeagus with basal process, unique; plates truncate apically; connective as long as style; vertex without fuscous spots .... robusta Baker — Aedeagus simple or with ventral serrations; connective shorter than style; vertex usually with fuscous spots 4 4(3). Aedeagus with shaft simple, elongate; plates rounded apically, embrowned . . pastora, n. sp. — Aedeagus with shaft shortened, expanded and serrate ventrally; plates truncate apically . . . globosa Ball & Beamer 5(1). Aedeagus with conspicuous apical hook; style exceeding apex of plate; vertex with or with- out fuscous spots 6 — Aedeagus with poorly developed or no apical hook (apical serrated flanges may be present); style variable; vertex with or without fuscous spots 22 6(5). Aedeagus with single apical hook 7 — Aedeagus with semicircular apical hook .... 20 7(6). Aedeagus not widest basally (may have crests at midlength), not tapered apically; plates rounded apically; vertex with or without fus- cous spots 14 — Aedeagus widest in basal 1/2, shaft tapered in apical 1/2, apical hook conspicuous; plates usually truncate apically; vertex with or with- out fuscous spots 8 8(7). Aedeagus with distinct lateral tubercles at base of shaft; style with ventral apical arm widened, dorsal arm slender; vertex without fuscous spots planata Ball & Beamer — Aedeagus without distinct tubercles; style with ventral and dorsal arms variable; vertex with or without spots 9 9(8). Style without apical lateral flange, dorsal arm short, wide; plates rounded apically krameri, n. sp. — Style with distinct apical lateral flange or ven- tral arm expanded laterally, dorsal arm vari- able, elongate; plates truncate apically 10 10(9). Aedeagus with apex of dorsal apodeme ex- panded; style with dorsal arm elongate 11 — Aedeagus with dorsal apodeme rounded; style variable 12 11(10). Style with dorsal arm slender; vertex with no fuscous spots; smaller in overall size whitcombi, n. sp. — Style with dorsal arm more robust; vertex usually with fuscous spots; larger in size .... plana Ball & Beamer 12(10). Aedeagus with shaft robust, short; style with dorsal apical arm longer, variable apically . . 13 — Aedeagus with shaft less robust; style with dorsal arm rounded apically aspera Ball & Beamer 13(12). Style with dorsal arm bulbous apically; aedea- gus with apical hook conspicuous, with slight processes at base aphoda, n. sp. — Style with dorsal arm acute apically; aedeagus with apical hook less developed, no process at base strobila, n. sp. 14(7). Style bifid apically; aedeagus with apical hook short; vertex with or without fuscous spots . . 15 — Styles not bifid apically or dorsal arm elon- gate; aedeagus with apical hook variable; ver- tex with or without fuscous spots 17 15(14). Aedeagus short, shaft serrate ventrally, widest at midlength; style without distinct lateral apical flange; vertex without fuscous spots supina Ball & Beamer 20 Great Basin Naturalist Memoirs No. 12 — Aedeagus longer, shaft not serrate or widened; style variable; vertex with or with- out fuscous spots 16 16(15). Style with conspicuous apical lateral flange; vertex without fuscous spots foeda Ball & Beamer — Style without apical lateral flange; vertex with fuscous spots parca Ball & Beamer 17(14). Style with dorsal arm elongate; vertex without fuscous spots rostrata Ball & Beamer — Style without elongate dorsal arm; vertex usu- ally with fuscous spots 18 18(17). Aedeagus with shaft shortened, widest in api- cal 1/2, apical hook short; style with conspicu- ous lateral flange 19 — Aedeagus with shaft longer, not widest in api- cal 1/2, apical hook longer; style without lat- eral flange fredonia Ball & Beamer 19(18). Aedeagus with pronounced keels on ventral margin of shaft, shorter; hind tibial spur 1/2 1st tarsomere; from SW cursa, n. sp. — Aedeagus without pronounced keels on shaft, longer; hind tibial spur longer; from Califor- nia (east face of Sierras) hemijona, n. sp. 20(6). Style with apical dorsal arm truncate; vertex commonly with fuscous spots; plates rounded apically incerta Ball & Beamer — Style with apical dorsal arm rounded or acute; vertex usually without fuscous spots; plates variable 21 21(20). Plates rounded apically; style with ventral apical arm thickened, well developed kadokana KnuU — Plates truncate apically; style with apical lat- eral flange; ventral apical arm not as well de- veloped; vertex occasionally with fuscous spots salsa Ball & Beamer 22(5). Vertex usually without fuscous spots; aedea- gus with serrated ventral margin; plates not elongate 23 — Vertex usually with fuscous spots, or if absent, then aedeagus without ventral serrations; plates may be elongate 29 23(22). Style distinctly bifid apically; connective shorter than style; aedeagus without apical crests on shaft 25 — Style with only dorsal arm developed; con- nective variable in length; aedeagus with crests on shaft 24 24(23). Aedeagus with apical crests on shaft; connec- tive equal to style in length; vertex occasion- ally with fuscous spots kanabana Ball & Beamer — Aedeagus with medial crests on shaft; connec- tive shorter than style; vertex without spots furnaca, n. sp. 25(23). Style with apical lateral flange, apical arms equal in length; aedeagus with shaft elongate '. 26 — Style without flange, dorsal arm longer; aedeagus with shaft shortened, acute apically ! 27 26(25). Style with conspicuous lateral flange; plates short, truncate apically; vertex length less than interocular width nmrthae, n. sp. — Style with lateral flange smaller; plates rounded apically, vertex length may equal interocular width bifida Ball & Beamer 27(25). Style conspicuously bifid, dorsal arm slender, tapered iitahna Osborn — Style less bifid, dorsal arm robust 28 28(27). Style with dorsal arm acute apically; vertex occasionally with spots yumana Osborn — Style with dorsal arm bulbous apically; vertex without spots deserta, n. sp. 29(22). Tibial spur 2/3 length of 1st tarsomere; plates elongate, acute apically, exceeding apex of styles; outer arm of style acute apically inagdalena Baker — Tibial spur variable; plates shorter; style equal to or exceeding apex of plate, outer arm variable 30 30(29). Style with dorsal arm acute apically 31 — Style with dorsal arm rounded or truncate . . 32 31(30). Aedeagus with shaft approximately 1 1/2X length of dorsal apodeme, with minute serra- tions on ventral margin . . . laeta Ball & Beamer — Aedeagus with shaft 2X length of dorsal apodeme, with conspicuous serrations on ventral margin sttjlosa, n. sp. 32(30). Style with outer arm rounded; aedeagus with shaft only slightly widened or tapered in api- cal 1/2, ventral margin with or without serra- tions 33 — Style with outer arm truncate; aedeagus with shaft widest in apical 1/2, ventral margin not serrated incongrua Baker 33(32). Aedeagus with shaft short, widest at mid- length; style with large lateral apical flange tenera Ball & Beamer — Aedeagus with shaft longer, even-margined or slightly enlarged apically; style without conspicuous flange 34 34(33). Style deeply bifid apically, ventral arm widened, dorsal arm even-margined; aedea- gus with shaft slightly widened apically .... vativala, n. sp. — Style less bifid, with inconspicuous apical lat- eral flange, dorsal arm tapered; aedeagus with shaft even-margined terebrans (Gillette & Baker) Character.s Used in Character Code Character]. 0 no fuscous spots on vertex. 1 fuscous spots may be present. Character 2. 0 hind tibial spur 1/2 1st tarsomere or less. 1 = spur approximate]} length of 1st tar- somere. 1988 Blocker, Johnson: Subgenus Athysanella 21 character 3. Character 4. Characters. Character 6. Character 7. Character 8. 0 = ocellus approximately its diameter from eye. 1 = ocellus more remote. 0 --^ male plates unicolorous. 1 male plates embrowned apically. 0 ^ male plates elongate. 1 ^ male plates rounded apically. 2 = male plates truncate apically. 0 = male pygofer with caudal margin rounded. 1 = male pygofer with caudal margin ex- tended. 0 ^ anal tube not reaching pygofer apex. 1 = anal tube equal to or exceeding pygofer apex. 0 = male pygofer not embrowned. 1 = male pygofer embrowned caudoven- trally. Character 9. 0 = male plates exceeding apex of pygofer. 1 = male plates equal to apex of pygofer. 2 = male plates not reaching apex of pygofer. Character 10. 0 = styles not reaching apex of pygofer. 1 = styles equal to apex of pygofer. 2 = styles exceed apex of pygofer. Character 11. 0 = styles not reaching apex of plates. 1 = styles equal to apex of plates. 2 = styles exceed apex of plates. Character 12. 0 = styles without lateral flange. 1 = styles with lateral flange. 2 = styles with ventral arm thickened. Character 13. 0 = connective shorter than style. 1 = connective as long as or longer than style. Character 14. 0 = aedeagus with caudal surface smooth. 1 = aedeagus with caudal surface troughlike. Character 15. 0 = shaft of aedeagus flared in caudal aspect. 1 = shaft even-margined in caudal as- pect. Character 16. 0 = aedeagus with no basal processes on shaft. 1 = shaft with basal processes. Character 17. 0 = aedeagus with caudal margin of shaft smooth. 1 = caudal margin serrate. Character 18. 0 = aedeagus with dorsal apodeme sim- ple. 1 = dorsal apodeme enclosing shaft basally. Athysanella gardenia Osborn Figs. 1-5 Athysanella gardenia Osborn 1930:701. Gladionura frigida Osborn 1930:709. Length of male 2.5 to 2.9 mm, female 3.2 to 3.4 mm; head width of male 1.0 to 1.05 mm, female 1.05 to 1.15 mm; pronotal width of male 0.95 to 1.05 mm, female 1.0 to 1.1 mm; interocular width of male 0.45 to 0.55 mm. female 0.5 to 0.55 mm; vertex length of male 0.35 to 0.4 mm, female 0.35 to 0.45 mm; pronotal length of male 0.3 to 0.4 mm, female 0.3 to 0.35 mm. Vertex length 0.76 to 0.8 interocular width; pronotal length 0. 85 to 0. 88 vertex length. Character code: 1-1-1-1-2-0-0-0-2-1-2-0-0- 1-0-0-0-0. Lectotype, male (No. 43183), and lectoallo- type, female (No. 43183), from Garden of the Gods, Colorado, (Webster) No. 7104, in USNM. This species is related to pastora but can be distinguished by the apex of the style, which is narrowed (Fig. 2), and the posterior margin of the plate, which is embrowned and has a me- dian projection. The style occasionally has macrosetae; long-winged specimens have been observed. This species has been col- lected in Colorado and Wyoming. Athysanella robusta Baker Figs. 6-9 Athysanella robusta Baker 1898:187. Athysanella montana Osborn 1930:700. Length of male 2.3 to 3. 1 mm, female 3.4 to 3.7 mm; head width of male 0.95 to 1. 15 mm, female 1.05 to 1.15 mm; pronotal width of male 0.9 to 1. 15 mm, female 1.05 to 1. 15 mm; interocular width of male 0.4 to 0.55 mm, female 0.5 to 0.55 mm; vertex length of male 0.35 to 0.5 mm, female 0.4 to 0.5 mm; prono- tal length of male 0.3 to 0.4 mm, female 0.35 to 0.4 mm. Vertex length 0.8 to 0.9 interocu- lar width; pronotal length 0.76 to 0.88 vertex width. Character code: 0-0-1-1-2-1-1-0-2-0-2-0-1- 0-0-1-0-0. The type of this species has not been stud- ied; identification is based on determined ma- terial from various collections and the litera- ture. Athysanella robusta is related to pastora but can be separated by the unique shape of the aedeagus (Fig. 6), the pronotum nearly as wide as the head, and the connective, which is as long as the style. There are no fuscous spots on the vertex of this species. It has been col- lected in Colorado, Montana, Nebraska, North Dakota, Wyoming, and Canada. Athysanella pastora Blocker, n. sp. Figs. 10-13 Length of male 2.5 to 2.7 mm, female 3.6 to 22 Great Basin Naturalist Memoirs No. 12 V^^ Figs. 1-1.3. Figs. 1-5. Athysanella gardenia: 1, aedeagus, lateral view; 2, style, broad aspect; 3, pygofer, lateral view; 4, plate, ventral view; 5, female sternum VII, ventral view. Figs. 6-9. Athysanella rohusta. 6, aedeagus, lateral view; 7, style, broad aspect; 8, pygofer, lateral view; 9, female sternum VII, ventral view. Figs. 10-13. Athysanella pastora: 10, aedeagus, lateral view; 11, style, broad aspect; 12, pygofer, lateral view; 13, female sternum VII, ventral view. 3.9 mm; head width of male 1.05 to 1.15 mm, female 1.2 to 1.25 mm; pronotal width of male 0.9 to 1.0 mm, female 1.05 to 1. 15 mm; inter- ocular width of male 0.4 to 0.5 mm, female 0.5 to 0.6 mm; vertex length of male 0.35 to 0.45 mm, female 0. 45 to 0. 5 mm ; pronotal length of male 0.3 to 0.4 mm, female 0.35 to 0.4 mm. Vertex length 0.84 to 0.95 interocular width; pronotal length 0.76 to 0.94 vertex length. Character code: 1-1-0-1-1-1-0-1-1-2-2-0-0- 0-0-0-0-0. Color stramineous; vertex with fuscous spots; vertex, pronotum, and abdomen with additional, conspicuous brown pattern; fore- wings commonly with darkened stripes; face with lateral fuscous marks; legs with various amounts of fuscous coloring. Forewings brachypterous, exposing 3 to 4 pregenital abdominal terga; ocellus approxi- mately 1.5 its diameter from eye; hind tibial spur 3/4 length of 1st tarsomere; female ab- dominal sternum VII (Fig. 13) variable but hind margin usually with conspicuous lateral lobes, medial third truncate or rounded, vari- ously embrowned. 1988 Blocker, Johnson: Subgenus Athysanella 23 Pygofer with caudal margin extended, slightly embrowned on caudoventral border, densely set with macrosetae (25+); anal tube attains apex of pygofer; valve with caudal mar- gin rounded; plates rounded and conspicu- ously embrowned apically, slightly separated at base; connective 2/3 or less length of style; styles rounded apically, with conspicuous preapical lobe, apex slightly exceeding apex of plates; aedeagus with shaft simple, slightly enlarged apically, elongate, with dorsal apodeme slightly overlapping base of shaft. Holotype, male, from Chaves Co., New Mexico, Bitter Lake NWR, 14 Aug 1984 (R. F. Whitcomb); 4 male and 8 female paratypes, same data. Holotype and paratypes in USNM; paratypes in IPL and KSU. Athysanella pastora is related to robusta and glohosa but can be separated from the former by the absence of an aedeagal process. It can be separated from glohosa by the shape of the apex of the style, which is not widened (Fig. 11), and by the shaft of the aedeagus (Fig. 10), which is elongate and not serrated. This species has been collected only on gyp grama, Bouteloua hreviseta, in gypsum flats of southeastern New Mexico. Athysanella glohosa Ball & Beamer Figs. 14-15 Athysanella glohosa Ball & Beamer 1940:18. Length of male 2.9 mm; head width 1.2 mm; pronotal width 1.1 mm; interocular width 0.45 mm; vertex length 0.35 mm; pronotal length 0.4 mm. Vertex length 0.78 interocular width; pronotal length 1.07 vertex length. Female unknown. Character code: 1-0-1-1-2-1-1-0-1-1-1-0-0- 1-0-0-1-0. Holotype, male, from Grand Canyon, Ari- zona, 4 Aug 1930 (E. D. Ball) in USNM. Male and female paratypes, same data, at KU. Athysanella glohosa is related to gardenia but can be separated by the shape of the shaft of the aedeagus, which is serrate at midlength (Fig. 14). Female paratypes examined are doubtfully conspecific with the male. This species is known only from its type locality in Arizona; a specimen from Utah is assigned to this species but may represent a new species. Athysanella planata Ball & Beamer Figs. 16-18 Athysanella planata Ball & Beamer 1940:24. Length of male 2.6 to 2.7 mm, female 3.4 to 4.0 mm; head width of male 1.0 to 1.05 mm, female 1. 1 to 1. 15 mm; pronotal width of male 0.9 to 1.0 mm, female 1.0 to 1.1 mm; interocu- lar width of male 0.35 to 0.4 mm, female 0.4 to 0.5 mm; vertex length of male 0.35 mm to 0.4 mm, female 0.4 to 0.45 mm; pronotal length of male 0.3 to 0.35 mm, female 0.3 to 0.35 mm. Vertex length 0.87 to 1.0 interocular width; pronotal length 0.8 to 0.86 vertex length. Character code: 0-1-0-0-1-0-0-0-1-1-1-2-0- 0-0-0-0-0. Holotype, male, and allotype, female, from Newberry Springs, California, 30 July 1936 (D. R. Lindsay and R. H. Beamer) in KU. Athysanella planata is related to krameri but can be separated by the presence of pro- cesses on the base of the shaft of the aedeagus (Fig. 16), a more slender apical arm of the style (Fig. 17), and the absence of fuscous spots on the vertex. This species has been collected only from the type locality in Cali- fornia. Athysanella krameri Blocker, n. sp. Figs. 19-21 Length of male 3.0 to 3. 1 mm, female 4.4 to 4.7 mm; head width of male 1.05 to 1.15 mm, female 1.2 to 1.25 mm; pronotal width of male 1.0 to 1.1 mm, female 1.1 to 1.2 mm; interocu- lar width of male 0.45 to 0.5 mm, female 0.55 to 0.6 mm; vertex length of male 0.35 to 0.45 mm, female 0.45 to 0.5 mm; pronotal length of male 0.3 to 0.35 mm, female 0.35 to 0.4 mm. Vertex length 0.83 to 0.95 interocular width; pronotal length 0.77 to 0.87 vertex length. Character code: 1-1-0-0-1-1-0-0-2-1-2-0-0- 0-0-0-0-0 . Color stramineous; vertex commonly with fuscous spots, often with orange dashes; male pygofer commonly with conspicuous fuscous spot; remainder of body mostly unicolorous; female without fuscous spots on vertex. Forewings brachypterous, exposing 3.5 to 4.5 pregenital abdominal terga; ocellus 1.5 its diameter (or less) from eye; hind tibial spur approximately 3/4 length of 1st tarsomere; fe- male abdominal sternum VII (Fig. 21) with elongate lateral lobes and rounded medial lobe on hind margin. Pygofer rounded apically, with about 20 macrosetae; valve rounded on apical margin; plates rounded apically, slightly separated or touching at base; connective less than 3/4 24 Great Basin Naturalist Memoirs No. 12 Figs. 14-27. Fig.s. 14-15. Athijsanella globosa: 14, aedeagus, lateral view; 15, style, broad aspect. Figs. 16-18. Athysanella planata. 16, aedeagus, lateral view; 17, style, broad aspect; 18, female sternum Vll, ventral view. Figs. 19-21. Athysanella krameri: 19, aedeagus, lateral aspect; 20, style, broad aspect; 21, female sternum Vll, ventral aspect. Figs. 22-23. Athysanella whitcombi: 22, style, broad aspect; 23, female sternum Vll, ventral view. Figs. 24-27. Athysanella plana: 24, aedeagus, lateral view; 25, aedeagus, anterodorsal view; 26, style, broad aspect; 27, female sternum VII, ventral view. length of style; styles (Fig. 20) bifid apically, dorsal arm short, robust, no preapical lobe, exceeding apex of plates, equal to apex of pygofer; aedeagus (Fig. 19) with shaft short, widest basally, and tapered to acute apical hook. Holotype, male, from Little Lake, Inyo Co., California, 7 June 1929 (E. P. Van Duzee); 5 male and 6 female paratypes, same data; 3 males from Olancha, Inyo Co., 8 June 1929; 1 male from Lone Pine, Inyo Co., 10 June 1929. Holotype and paratypes in CAS, paratypes in KSU. Athysanella krameri is related to planata but can be separated by the absence of projec- tions at the base of the aedeagus and the dorsal arm of the style, which is shorter and more robust. An undeveloped male specimen has 1988 Blocker, Johnson: Subgenus Athysanella 25 been observed. This species is known only from the above locahties in CaHfornia. Athysanella whitcombi Blocker, n. sp. Figs. 22-23 Length of male 2.6 mm, female 4.0 mm; head width of male 1.0 mm, female 1.15 to 1.2 mm; pronotal width of male 0.9 mm, female 1.0 to 1.05 mm; interocular width of male 0.4 mm, female 0.55 to 0.6 mm; vertex length of male 0.35 to 0.4 mm, female 0.5 mm; pronotal length of male 0.25 to 0.3 mm, female 0.45 mm. Vertex length 0.93 to 0.94 interocular width; pronotal length 0.73 to 0.74 vertex length. Character code: 0-1-1-0-1-0-0-0-1-1-1-1-0- 0-0-0-0-1. Color stramineous; vertex without fuscous spots; body virtually unicolorous. Forewings of male brachypterous, exposing 4.5 pregenital abdominal terga; ocellus ap- proximately 2X its diameter from eye; hind tibial spur about 1/2 length of 1st tarsomere; female abdominal sternum VII with lateral lobes, without medial lobe on caudal margin. Pygofer rounded, with about 20 macrose- tae; valve rounded on caudal margin; plates rounded apically, touching at base then diver- gent apically; connective 3/4 length of style; styles bifid apically, dorsal arm slender, no preapical lobe, exceeding apex of plates, equal to apex of pygofer; aedeagus with shaft and dorsal apodeme as plana. Holotype, male, from Austin, Nevada, 12 Aug 1940 (D. E. Hardy) and a long- winged female paratype, same data, in KU. Athysanella whitcombi is related to plana but can be separated by its smaller size, the absence of fuscous spots on the vertex, and the more slender dorsal apical arm of the style. This species is named for R. F. Whitcomb, who has made numerous contributions to this research; it has been collected in Nevada. Athysanella plana Ball & Beamer Figs. 24-27 Athysanella plana Ball & Beamer 1940:13. Length of male 3.0 to 3.3 mm, female 4.5 to 4.7 mm; head width of male 1.05 to 1.15 mm, female 1.2 to 1.25 mm; pronotal width of male 1.0 to 1.05 mm, female 1.15 to 1.2 mm; intero- cular width of male 0.45 to 0.5 mm, female 0.55 to 0.6 mm; vertex length of male 0.4 to 0.45 mm, female 0.45 to 0.5 mm; pronotal length of male 0.3 to 0.4 mm, female 0.35 to 0.4 mm. Vertex length 0.89 to 0.95 interocu- lar width; pronotal length 0.77 to 0.82 vertex length. Character code: 1-1-1-0-2-0-0-0-1-1-1-1-0- 0-0-0-0-0. Holotype, male, and allotype, female, from Califa, California, 12 June 1935 (P. W. Oman) in USNM; paratypes in USNM. Athysanella plana is related to aspera but can be separated by the expanded base of the dorsal apodeme of the aedeagus (Fig. 25) and the longer dorsal apical arm of the style (Fig. 26). This species has been collected only from the Central Valley (Fresno Co.) of California. Athysanella aspera Ball & Beamer Figs. 28-30 Athysanella aspera Ball & Beamer 1940:17. Length of male 2.5 to 3. 1 mm, female 4.3 to 5.0 mm; head width of male 0.95 to 1. 15 mm, female 1. 15 to 1.3 mm; pronotal width of male 0.9 to 1.1 mm, female 1.1 to 1.25 mm; interoc- ular width of male 0.4 to 0.5 mm, female 0.45 to 0.5 mm; vertex length of male 0.35 to 0.45 mm, female 0.4 to 0.5 mm; pronotal length of male 0.3 to 0.4 mm, female 0.35 to 0.45 mm. Vertex length 0.8 to 0.95 interocular width; pronotal length 0.76 to 0.93 vertex length. Character code: 1-1-1-0-2-1-0-0-2-1-2-1-0- 0-0-0-0-0. Holotype, male, and allotype, female, from Mojave, California, 1 July 1931 (E. D. Ball) in USNM. Athysanella aspera is related to plana but can be separated by the apex of the style, which has a shorter dorsal arm (Fig. 29), and the dorsal apodeme of the aedeagus, which is not widened apically. In one specimen, the vertex length equaled the interocular width. The shaft of the aedeagus has an indistinct pair of basal tubercles similar to planata. A male specimen parasitized by Pipinculidae and specimens with undeveloped genitalia have been observed. This species has been col- lected from the Mojave Desert in California. Athysanella aphoda Blocker, n. sp. Figs. 31-32 Length of male 3.5 mm; head width 1.2 mm; pronotal width 1.1 mm; interocular width 0.5 mm; vertex length 0.45 mm; prono- tal length 0.4 mm; female unknown. Vertex 0.9 interocular width; pronotal length 0.83 to 26 Great Basin Naturalist Memoirs No. 12 Figs. 28-39. Figs. 28-30. Athysanella aspera: 28, aedeagus, lateral view; 29, style, broad aspect; 30, female sternum VII, ventral view. Figs. 31-32. Athysanella aphoda: 31, aedeagus, lateral view; 32, style, broad aspect. Figs. 33-36. Athysanella strobila: 33, aedeagus, lateral view; .34, style, broad aspect; 35, pygofer, lateral aspect; 36, female sternum VII, ventral view. Figs. 37-.39. Athysanella supina: 37, aedeagus, lateral view; 38, style, broad aspect; 39, female sternum VII, ventral view. 0.84 vertex length. Female unknown. Character code: 1-0-1-0-2-0-1-0-2-1-2-2-0- 0-0-1-0-0. Color stramineous; vertex and pronotum with distinct, darker pattern; forewings with dark stripes; face with distinct lateral arcs; legs with slight fuscous coloring. Forewings brachypterous, exposing 3.5 pregenital abdominal terga; ocellus approxi- mately its diameter from eye; hind tibial spur slightly shorter than 1st tarsomere. Pygofer as strobila but larger; valve rounded posteriorly; plates truncate apically; styles (Fig. 32) with dorsal arm bulbous api- cally; aedeagus (Fig. 31) with apical hook more distinct than strobila , shaft with indis- tinct, small process at base. Holotype, male, from Lander Co. , Nevada, 1988 Blocker, Johnson: Subgenus Athysanella 27 30 mi N Austin, 24 Julv 1986 (H. D. Blocker) in KSU. Athysanella aphoda is closely related to and possibly conspecific with strohila but can be readily separated by its larger size, the bul- bous apex of the dorsal arm of the style, and the more robust aedeagus, which has a more pronounced apical hook. This species is repre- sented by the holotype from Nevada. Athysanella strohila Blocker, n. sp. Figs. 33-36 Length of male 2. 6 to 3. 9 mm, female 4. 2 to 5.3 mm; head width of male 1.05 to 1.3 mm, female 1.15 to 1.35 mm; pronotal width of male 0.95 to 1.25 mm, female 1.05 to 1.3 mm; interocular width of male 0.4 to 0.55 mm, female 0.45 to 0.65 mm; vertex length of male 0.4 to 0.55 mm, female 0.45 to 0.65 mm; pronotal length of male 0.3 to 0.4 mm, female 0.3 to 0.45 mm. Vertex length 0.89 to 1.0 interocular width; pronotal length 0.7 to 0.82 vertex length. Character code: 0-1-0-0-2-0-0-0-2-1-2-2-0- 1-0-0-0-0. Color stramineous, occasionally patterned; vertex without fuscous spots; abdomen occa- sionally with some darker pattern; other body areas occasionally with irregular, brown pat- tern. Forewings brachypterous, exposing 3.0 to 4.5 pregenital abdominal terga; ocellus ap- proximately its diameter from eye; hind tibial spur 3/4 length of 1st tarsomere; female ab- dominal sternum VII (Fig. 36) with conspicu- ous lateral lobes on posterior margin, with a broadly rounded medial lobe. Pygofer rounded on caudoventral margin, with a distinct lobe (Fig. 35), with 15 or fewer macrosetae; valve rounded posteriorly; plates truncate apically, nearly touching basally, then diverging; connective 3/4 length of style; styles bifid (Fig. 34) apically, with ventral arm thickened, exceeding apex of plates, equal to apex of pygofer; aedeagus with shaft short, widest in proximal 1/2, slightly tapered in api- cal 1/2 to a single apical hook. Holotype, male, from Lone Pine, Califor- nia, 28 July 1940 (R. H. Beamer); 4 male and 3 female paratypes, same data; male and fe- male, Fallon, Nevada, 12 Aug 1940 (L. C. Kuitert); 2 males and 8 females. New Hum- boldt Co., Nevada, 7 mi W Denio, 23 June 1971 (Oman); 6 males and 3 females. Eureka Co., Nevada, W. Eureka, 25 July, 1986 (H. D. Blocker); male and female, Lander Co., Nevada, 5 mi W Austin, 24 July 1986 (H. D. Blocker); male, Owyhee Co., Idaho, 10 mi W Bruneau, 9 July 1981 (H. D. Blocker, J. W. Johnson); numerous specimens from Albert Lake, Oregon, 5 mi N Valley Falls, 18 May 1969 (Oman), and 6 mi N Silver Lake, Ore- gon, 17 May 1969 (Oman). Holotype and paratvpes in KU; paratypes in KSU and OrSU. Athysanella strohila is closely related to as- pera but can be separated by the shape of the shaft of the aedeagus, which is more robust and shorter, and by the dorsal apical arm of the style, which is longer and acute apically. It can be recognized by the unique lobe on the caudal margin of the pygofer. An undevel- oped female specimen has been examined. This species has been collected in California, Idaho, Nevada, and Oregon. Athysanella supina Ball & Beamer Figs. 37-39 Athysanella supina Ball & Beamer 1940:22. Length of male 2.8 to 3.2 mm, female 4.3 to 4.6 mm; head width of male 1.0 to 1.1 mm, female 1.15 to 1.25 mm; pronotal width of male 1.0 to 1.05 mm, female 1.1 to 1.2 mm; interocular width of male 0.45 to 0.5 mm, female 0.55 to 0.6 mm; vertex length of male 0.45 to 0.55 mm, female 0.5 to 0.6 mm; prono- tal length of male 0.35 to 0.4 mm, female 0.35 to 0.45 mm. Vertex length 1.0 to 1. 15 interoc- ular width; pronotal length 0.68 to 0.75 vertex length. Character code: 1-1-1-0-1-0-0-0-2-1-2-1-0- 1-0-0-1-0. Holotype, male, and allotype, female, from Boca Chica, Texas, 30 May 1933 (P. W. Oman) in USNM; paratypes in USNM and KU. Athysanella supina is related to fredonia but can be separated by the shaft of the aedea- gus, which has a shorter crest on the ventral margin (Fig. 37), and the shape of the apex of the style, which is more bifid (Fig. 38). This species, described from Texas, has recently been collected there (A. L. Hicks) from shore- grass, Monathochloe littoralis. Athysanella foeda Ball & Beamer Figs. 40-43 Athysanella foeda Ball & Beamer 1940:19. 28 Great Basin Naturalist Memoirs No. 12 Figs. 40-.52. Figs. 40-43. Athysanella foeda: 40, head and prothorax, dorsal \'ievv; 41, aedeagus, lateral view; 42, style, broad aspect; 4.3, female sternum VII, ventral view. Figs. 44-46. Athysanella parca: 44, aedeagus, lateral view; 45, style, broad aspect; 46, female sternum VII, ventral view. Figs. 47-49. Athysanella rostrata: 47, aedeagus, lateral view; 48, style, broad aspect; 49, female sternum VII, ventral aspect. Figs. 50-52. Athysanella fredonia: 50, aedeagus, lateral view; 51, style, broad aspect; 52, female sternum VII, ventral view. Length of male 3.4 to 3.6 mm, female 4.9 to 5.1 mm; head width of male 1.2 to 1.3 mm, female 1.4 to 1.5 mm; pronotal width of male 1.2 to 1.3 mm, female 1.35 to 1.4 mm; interoc- ular width of male 0.55 to 0.6 mm, female 0.65 to 0.75 mm; vertex length of male 0.5 to 0.55 mm, female 0.55 to 0.6 mm; pronotal length of male 0.4 to 0.45 mm, female 0.4 to 0.45 mm. Vertex length 0.83 to 0.88 interocular width; pronotal length 0.76 to 0.85 vertex length. Character code: 0-1-1-0-1-0-0-0-1-2-2-1-0- 1-0-0-0-1. Holotype, male, and allotype, female, from Laramie, Wyoming, 23 June 1935 (R. H. and 1988 Blocker, Johnson: Subgenus Athysanella 29 J. Beamer) in KU; paratypes in KU. Athysanella foeda is related to parca but can be separated by its larger size and the presence of a conspicuous lateral lobe at the apex of the style (Fig. 42). It can be separated from fredonia by the shape of the apex of the style, which is distinctly bifid. This species has been collected in Wyoming. Athysanella parca Ball & Beamer Figs. 44-46 Athysanella parca Ball & Beamer 1940:15. Length of male 2.7 to 3.4 mm, female 3.9 to 4.4 mm; head width of male 1.0 to 1.15 mm, female 1. 15 to 1.4 mm; pronotal width of male 0.9 to 1.15 mm, female 1.05 to 1.25 mm; interocular width of male 0.4 to 0.5 mm, fe- male 0.5 to 0.65 mm; vertex length of male 0.4 to 0.45 mm, female 0.45 to 0.55 mm; pronotal length of male 0.3 to 0.4 mm, female 0.4 to 0.45 mm. Vertex length 0.89 to 0.95 interocu- lar width; pronotal length 0.82 to 0.88 vertex length. Character code: 1-1-0-0-1-0-0-0-2-1-2-2-0- 0-0-0-0-0. Holotype, male, and allotype, female, from Belen, New Mexico, 20 July 1936 (R. H. Beamer and D. R. Lindsay) in KU; paratypes inKU. Athysanella parca is related to foeda but can be separated by the absence of a lateral flange on the style and the usual presence of fuscous spots on the vertex. It can be sepa- rated from incerta by the shape of the apex of the shaft of the aedeagus, which has a single acute apical hook (Fig. 44). A long-winged female has been examined. This species has been collected in Colorado and New Mexico; in New Mexico, the host is saltgrass, Distichlis stricta. Athysanella rostrata Ball & Beamer Figs. 47-49 Athysanella rostrata Ball & Beamer 1940:21. Length of male 3.5 mm, female 5.0 to 5.1 mm; head width of male 1.15 to 1.25 mm, female 1.3 to 1.35 mm; pronotal width of male 1.1 to 1.15 mm, female 1.2 to 1.25 mm; intero- cular width of male 0.5 to 0.55 mm, female 0.6 to 0.65 mm; vertex length of male 0.45 to 0.5 mm, female 0.5 to 0.55 mm; pronotal length of male 0.35 to 0.4 mm, female 0.4 to 0.45 mm. Vertex length 0.86 to 0.91 interocular width; pronotal length 0.75 to 0.78 vertex length. Character code: 0-1-0-0-2-0-0-0-2-2-2-2-0- 1-0-0-0-0. Holotype, male, and allotype, female, from Perris, California, 5 June 1935 (P. W. Oman) in USNM; paratypes in USNM. Athysanella rostrata is related to foeda but can be separated by the distinctly elongate dorsal arm of the style (Fig. 48). A paratype with undeveloped genitalia has been exam- ined. This species has been collected only from its type locality in southern California. Athysanella fredonia Ball & Beamer Figs. 50-52 Athtjsanella fredonia Ball and Beamer, 1940:12 (in part). Length of male 2.6 to 2.9 mm, female 3.9 to 4.3 mm; head width of male 1.0 to 1.15 mm, female 1. 15 to 1.4 mm; pronotal width of male 0.95 to 1.1 mm, female 1.0 to 1.25 mm; intero- cular width of male 0.4 to 0.45 mm, female 0.45 to 0.6 mm; vertex length of male 0.35 to 0.4 mm, female 0.4 to 0.5 mm; pronotal length of male 0.3 to 0.35 mm, female 0.35 to 0.4 mm. Vertex length 0.83 to 0.94 interocu- lar width; pronotal length 0.75 to 0.87 vertex length. Character code: 1-1-0-0-1-0-0-0-1-1-1-0-0- 1-0-0-1-0. Color stramineous; vertex with fuscous spots anteriorly; pronotum commonly with small brown spots; other dorsal body pattern variable; face with lateral fuscous marks; legs with various amounts of fuscous coloring. Forewings brachypterous, exposing 3.5 to 4.0 pregenital abdominal terga; ocellus ap- proximately its diameter from eye; hind tibial spur over 3/4 length of 1st tarsomere; female abdominal sternum VII (Fig. 52) with lateral lobes extending posteriorly slightly more than median lobe. Pygofer rounded apically, densely set with short setae; valve rounded on apical margin; plates rounded apically, slightly separated at base; connective shorter than style; styles (Fig. 51) widened apically, nearly truncate, dorsal arm acute apically, curved, usually with conspicuous lateral flange, equal to apex of plates and pygofer; aedeagus (Fig. 50) with shaft not widened apically, finely serrate on ventral margin. Holotype, male, from Fredonia, Arizona, 6 Aug 1930 (E. D. Ball) in USNM. 30 Great Basin Naturalist Memoirs No. 12 Athysanella fredonia is related to cursa but can be separated by the longer shaft of the aedeagus, which is not widened apically. Ex- amination of the holotype has revealed that the male genitalia differ from the genitalia described by Ball and Beamer (1940). For this reason, fredonia is redescribed and a new species, cursa, which agrees with the pub- lished illustration, is described. It has been collected in Arizona, Colorado, New Mexico, Nevada, and Utah, where it occurs on galleta, Hilariajamesii. At the type locality, it occurs with cursa on the same host (R. F. Whit- comb). Athysanella cursa Blocker, n. sp. Figs. 53-55 Athysanella fredonia Ball & Beamer (in part). Length of male 2.5 to 3.0 mm, female 4.0 to 4.4 mm; head width of male 1.05 to 1. 15 mm, female 1.2 to 1.3 mm; pronotal width of male 0.95 to 1.05 mm, female 1.1 to 1.2 mm; intero- cular width of male 0.4 to 0.45 mm, female 0.5 to 0.6 mm; vertex length of male 0.35 to 0.4 mm, female 0.45 to 0.5 mm; pronotal length of male 0.3 to 0.35 mm, female 0.35 to 0.4 mm. Vertex length 0.88 to 0.94 interocular width; pronotal length 0.8 to 0.88 vertex length. Character code: 0-1-0-0-1-0-0-0-1-1-1-1-0- 1-0-0-1-0. Color stramineous with fuscous spots on vertex; vertex, pronotum, and abdomen vari- ously patterned with brown markings; forewings commonly with brown stripes; face with lateral fuscous marks, legs with various amounts of fuscous coloring. Forewings brachypterous, exposing 3.5 to 4.5 pregenital abdominal sterna; ocellus ap- proximately 1.5X its diameter from eye; hind tibial spur usually about 1/2 length of 1st tar- somere; female abdominal sternum VII (Fig. 55) with lateral lobes slightly longer than me- dian lobe. Pygofer rounded but slightly extended api- cally; densely set with short setae; valve with caudal margin rounded; plates rounded api- cally; connective shorter than styles; styles (Fig. 54) widened apically and shallowly bifid, dorsal arm narrowed and acute apically, usu- ally with a conspicuous lateral flange, equal to apex of plates and pygofer; aedeagus (Fig. 53) with shaft short, lightly serrate on ventral margin, with pronounced flanges on ventral margin, slightly widest in apical 1/2. Holotype, male, from Whites City, New Mexico, 13 July 1936 (R. H. Beamer). One male and two female paratypes from Flagstaff, Arizona, 27 July 1936 (R. H. Beamer); two males from Flagstaff, Arizona, 21 July 1936 (D. R. Lindsey); female from St. Johns, Ari- zona, 26 July 1936 (R. H. Beamer); two males from Estancia, New Mexico, 24 June 1940 (R. H. Beamer); five males and one female from Cuervo, New Mexico, 23 June 1940 (R. H. Beamer). Holotype in KU; paratypes in KU and KSU. Athysanella cursa is closely related to hemi- jona but can be separated by the shorter aedeagus with pronounced flanges on the ven- tral margin, the shorter tibial spurs, and the pygofer, which is slightly extended apically. The plates occasionally have a macroseta. Long-winged specimens and specimens para- sitized by Dryinidae have been observed. This species is illustrated as fredonia in Ball and Beamer (1940), but a study of the holo- type of fredonia reveals that this is a different species. It has been reported from the Desert Plains region of Arizona, Colorado, New Mex- ico, and Utah, where it occurs on galleta, Hilariajamesii. Athysanella he?nijona Blocker, n. Figs. ,56-58 sp. Length of male 3.2 to 3.3 mm, female 4.8 to 4.9 mm; head width of male 1.2 to 1.25 mm, female 1.3 to 1.4 mm; pronotal width of male 1. 1 to 1. 15 mm, female 1.25 to 1.3 mm; intero- cular width of male 0.45 to 0.55 mm, female 0.5 to 0.55 mm; vertex length of male 0.4 to 0.45 mm, female 0.5 to 0.55 mm; pronotal length of male 0.35 to 0.4 mm, female 0.4 to 0.45 mm. Vertex length 0.85 to 0.95 interocu- lar width; pronotal length 0.73 to 0.83 vertex length. Character code: 1-1-0-0-1-0-0-0-1-1-1-1-0- 1-0-0-1-1. Color stramineous; vertex with fuscous spots; vertex, pronotum, and abdomen with conspicuous, dark pattern; forewings with darkened stripes; face with lateral fuscous marks; legs with various amounts of fuscous coloring. Forewings brachypterous, exposing 3.5 to 4.5 pregenital abdominal terga; ocellus ap- proximately 1.5X its diameter from eye; hind tibial spur over 3/4 length of 1st tarsomere; female abdominal sternum VII (Fig. 58) with 1988 Blocker, Johnson: Subgenus Athysanella 31 Figs. 53-70. Figs. 53-55. Athysanella cursa: 53, aedeagus, lateral view; 54, style, broad aspect; 55, female sternum VII, ventral view. Figs. 56-58. Athysanella hemijona. 56, aedeagus, lateral view; 57, style, broad aspect; 58, female sternum VII, ventral view. Figs. 59-61. Athysanella incerta: 59, aedeagus, lateral view; 60, style, broad aspect; 61, female sternum VII, ventral view. Figs. 62-65. Athysanella kadokana: 62, aedeagus, lateral view; 63, style, broad aspect; 64, 65, female sternum VII, ventral view. Figs. 66-70. Athysanella salsa: 66, head and pronotum, dorsal view; 67, aedeagus, lateral view; 68, style, broad aspect; 69, male plate, ventral view; 70, female sternum VII, ventral view. median lobe approximately same length as lateral lobes. Pygofer rounded apically, densely set with short, stubby setae; valve with caudal margin broadly rounded; plates broadly rounded, slightly separated at base; connective shorter than style; styles (Fig. 57) widened apically, shallowly bifid, dorsal arm constricted and curved; preapical lobe inconspicuous, with conspicuous apical lateral lobe, extending to apex of plates and pygofer; aedeagus (Fig. 56) with shaft short, slightly wider apically, ser- rated on ventral margin. Holotype, male, from Inyo Co., California, Westgard Pass, 14 June 1983, sweeping grasses (R. J. Gill and D. Blocker). Two male 32 Great Basin Naturalist Memoirs No. 12 and three female paratypes, same data. Holo- type and paratype in KSU; paratypes in CDA and KU. Athysanella hemijona is related to ctirsa but can be separated by the shaft of the aedeagus, which is longer, and with the ventral margin not as flared. This species is known only from its type locality in California. Athysanella mcerta Ball & Beamer Figs. 59-61 Athysanella incerta Ball & Beamer 1940:15. Length of male 2.7 to 3.3 mm, female 3.9 to 4.1 mm; head width of male 1.05 to 1.1 mm, female 1. 1 to 1.25 mm; pronotal width of male 1.0 to 1. 1 mm; female 1.0 to 1. 15 mm; interoc- ular width of male 0.45 to 0.5 mm, female 0.5 to 0.55 mm; vertex length of male 0.35 to 0.45 mm, female 0.45 to 0.5 mm; pronotal length of male 0.3 to 0.35 mm, female 0.35 to 0.4 mm. Vertex length 0.84 to 0.95 interocular width; pronotal length 0.75 to 0.82 vertex length. Character code: 1-1-0-0-1-0-0-0-2-1-2-2-0- 0-0-0-0-0. Holotype, male, and allotype, female, from Lamar, Colorado, 20 Aug 1936 (R. H. Beam- er) in KU; paratypes in KU, USNM, and CSU. Athysanella incerta is very closely related to kadokana but can be separated by the shape of the apex of the style (Fig. 60), which is not excavated as deeply, and with the dorsal arm neither expanded apically nor with an acute projection. Fuscous spots are some- times present on the vertex of incerta . Long- winged specimens have been examined. This species has been collected in Colorado, Kan- sas, Montana, Nebraska, New Mexico, and Wyoming, and appears to be a specialist on Distichlis spicata. Athysanella kadokana Knull Figs. 62-65 Athysanella kadokana Knull 1951:180. Length of male 2.7 to 3.4 mm, female 4.0 to 4.4 mm; head width of male 0.95 to 1.1 mm, female 1. 1 to 1.25 mm; pronotal width of male 0.9 to 1.05 mm, female 1.05 to 1.15 mm; interocular width of male 0.4 to 0.5 mm, fe- male 0.5 to 0.6 mm; vertex length of male 0.35 to 0.45 mm, female 0.45 to 0.55 mm; pronotal length of male 0.3 to 0.35 nmi, female 0.35 to 0.4 mm. Vertex length 0.84 to 1.0 interocular width: pronotal length 0.72 to 0.82 vertex length. Character code: 0-1-1-0-1-0-0-0-1-2-2-2-0- 1-0-0-0-1. Holotvpe, male, from Kadoka, South Da- kota, Badlands, 19 July 1950 (H. C. Severin) atOhSU. This species is closely related to salsa but can be separated by the shape of the male plate, which is rounded, and by the apex of the style, which has a more conspicuous ven- tral arm (Fig. 63). The shape of the posterior margin of the female sternum VII is variable in this species (Figs. 64, 65). Specimens para- sitized with Pipinculidae and Dryinidae and specimens with undeveloped genitalia have been examined; occasionally a specimen with faint spots on the vertex is seen. This species, collected in Colorado, South Dakota, Mon- tana, Nebraska, Wyoming, and Canada, ap- pears to be a specialist on Distichlis spicata. Athysanella salsa Ball & Beamer Figs. 66-70 Athysanella salsa Ball & Beamer 1940:20. Length of male 2.9 to 3.2 mm, female 3.6 to 4.4 mm; head width of male 1.0 to 1.15 mm, female 1.0 to 1.2 mm; pronotal width of male 0.9 to 1.05 mm, female 0.95 to 1.15 mm; interocular width of male 0.45 to 0.5 mm, female 0.45 to 0.55 mm; vertex length of male 0.4 to 0.5 mm, female 0.45 to 0.55 mm; prono- tal length of male 0.3 to 0.4 mm, female 0.3 to 0.4 mm. Vertex length 0.94 to 1. 1 interocular width; pronotal length 0.72 to 0.88 vertex length. Character code: 0-1-1-0-2-0-0-0-2-1-2-2-0- 1-0-0-0-0. Holotvpe, male, and allotype, female, from St. John,' Kansas, 11 Sept 1936 (R. H. Beamer) in KU. Athysanella salsa is closely related to kadokana but can be separated by the trun- cate apices of the male plates and the less conspicuous ventral apical arm of the style (Fig. 68). Specimens studied from Mexico have fuscous spots on the vertex and dark stripes on the forewings. Long-winged fe- males have been examined. Males and fe- males parasitized by Dryinidae and undevel- oped specimens have been examined. This species has been collected in Kansas, Texas, and Mexico, and appears to be a specialist on Distichlis spicata. 1988 Blocker, Johnson: Subgenus Atmysanella 33 ^-m^ \r^r^ Figs. 71-86. Figs. 71-73. Athysanella kanabana: 71, aedeagus, lateral view; 72, style, broad aspect; 73, female sternum VII, ventral view. Figs. 74-76. Athysanella furnaca: 74, aedeagus, lateral aspect; 75, style, broad aspect; 76, female sternum VII, ventral view. Figs. 77-79. Athysanella marthae: 11, aedeagus, lateral view; 78, style, broad aspect; 79, female sternum VII, ventral view. Figs. 80-83. Athysanella bifida: 80, aedeagus, lateral view; 81, 82, style, broad aspect; 83, female sternum VII, ventral view. Figs. 84-86. Athysanella utahna: 84, aedeagus, lateral view; 85, style, broad aspect; 86, female sternum VII, ventral aspect. Athysanella kanabana Ball & Beamer Figs. 71-73 Athysanella kanabana Ball & Beamer 1940:20. Length of male 3. 1 to 3.6 mm, female 4.4 to 4.7 mm; head width of male 1.1 to 1.2 mm, female 1.25 to 1.4 mm; pronotal width of male 1.05 to 1.15 mm, female 1.2 to 1.3 mm; inter- ocular width of male 0.5 to 0.55 mm, female 0.6 to 0.7 mm; vertex length of male 0.4 to 0.5 mm, female 0.5 to 0.55 mm; pronotal length of male 0.35 to 0.4 mm, female 0.35 to 0.45 mm. Vertex length 0.8 to 0.95 interocular width; pronotal length 0.76 to 0.88 vertex length. Character code: 0-1-1-1-1-0-0-0-2-1-2-2-0- 1-0-1-1-0. 34 Great Basin Naturalist Memoirs No. 12 Holotype, male, and allotype, female, from Kanab, Utah, 9 Aug 1936 (R. H. Beamer) in KU; paratypes in KU and USNM. Athysanella kanabana is related to tenera but can be separated by the embrowned area on the plates, the aedeagus with apical crests on the shaft (Fig. 71), and the unique shape of the apex of the style (Fig. 72). Occasionally, there are fuscous spots on the vertex. Unde- veloped specimens are represented in the paratype series. This species has been col- lected on Sporobolus airoides in Arizona, Kansas, New Mexico, and Utah. Athysanella furnaca Blocker, n. sp. Figs. 74-76 Length of male 3.1 mm, female 4.4 to 4.7 mm; head width of male 1. 15 mm, female 1.25 to 1.3 mm; pronotal width of male 1.1 mm, female 1.2 to 1.25 mm; interocular width of male 0.5 mm, female 0.5 to 0.55 mm; vertex length of male 0. 5 mm, female 0.5 to 0. 55 mm; pronotal length of male 0.4 mm, female 0.4 to 0.45 mm. Vertex length equal to interocular wddth; pronotal length 0.78 to 0.79 vertex length. Character code: 0-0-1-0-2-0-0-0-2-1-2-2-0- 1-0-0-1-1. Color stramineous, except for dark dots on abdomen and slight fuscous on legs. Forewings brachypterous, exposing 3.5 pregenital abdominal terga; ocellus approxi- mately 2X its diameter from eye; hind tibial spur 1/2 length (or less) of 1st tarsomere; fe- male abdominal sternum VII (Fig. 76) with short lateral lobes and slight median projec- tion. Pygofer with caudal margin extended, ex- ceeding anal tube, with approximately 25 se- tae; valve with caudal margin rounded; plates truncate apically, barely exceeding apex of valve; style with ventral arm short (Fig. 75), conspicuously flattened, dorsal arm elongate; connective 3/4 length of pygofer; aedeagus (Fig. 74) with conspicuous keels on shaft. Holotype, male, from Death Valley, Cali- fornia, 5.7 mi N Furnace Cr., 19 March 1971 (Oman); two female paratypes, same data. Holotype and paratype in OrSU; paratype in KSU. Athysanella furnaca is related to kanabana but can be distinguished by the medial crests on the shaft of the aedeagus and the connec- tive, which is shorter than the style. This spe- cies is known only from the type locality in the Mojave Desert of California. Athysanella marthae Blocker, n. sp. Figs. 77-79 Length of male 2.9 to 3.3 mm, female 4. 1 to 4.6 mm; head width of male 1.15 to 1.25 mm, female 1.25 to 1.35 mm; pronotal width of male 1.1 to 1.2 mm, female 1.1 to 1.2 mm; interocular width of male 0.45 to 0.5 mm, female 0.5 to 0.55 mm; vertex length of male 0.4 to 0.45 mm, female 0. 45 to 0. 5 mm; prono- tal length of male 0.35 to 0.4 mm, female 0.4 to 0.45 mm. Vertex length 0.89 to 0.95 intero- cular width; pronotal length 0.82 to 0.94 ver- tex length. Character code: 0-1-1-0-2-0-0-0-2-2-2-1-0- 1-0-0-1-1. Color uniformly stramineous, except for ir- regular, fuscous markings on venter and legs. Forewings brachypterous, exposing 3 to 4.5 pregenital abdominal terga; ocellus approxi- mately 2X its diameter from eye; hind tibial spur 2/3 length of 1st tarsomere; female ab- dominal sternum VII (Fig. 79) with conspicu- ous lateral lobes on hind margin. Pygofer broadly rounded apically, with fewer than 25 macrosetae, exceeding apex of plates; anal tube exceeding the apex of pygofer; valve with caudal margin rounded; plates short, truncate, with conspicuous mi- crosetae; style (Fig. 78) with conspicuous lat- eral flange and preapical lobe, deeply bifid apically, dorsal arm truncate, ventral arm rounded, exceeding apex of plates and pygofer; connective 3/4 length of style; aedea- gus (Fig. 77) with caudal margin of shaft ser- rate, troughlike in caudal view, with incon- spicuous process apically, dorsal apodeme simple. Holotype, male, from Cuatro Cienegas, Coahuila, Mexico, 9 June 1985, (A. L. Hicks) 1266B; five male and five females paratypes, same data. Holotype and paratypes in KSU; paratypes in IPL. Athysanella nuirthae is related to bifida but can be distinguished by the conspicuous lat- eral flange of the style and the short, truncate plates. This species has been collected only at the type locality in Mexico on Monanthochloe littoralis. I name this species for my wife. Athysanella bifida Ball & Beamer Figs. 80-83 Athysanella bifida Ball fit Beamer 1940:23. 1988 Blocker, Johnson; Subgenus Athysanella 35 Length of male 2.4 to 3.0 mm, female 3. 1 to 3.7 mm; head width of male 0.9 to 1.05 mm, female 0.95 to 1. 1 mm; pronotal width of male 0.8 to 1.0mm, female 0.9 to 1.0mm;interocu- lar width of male 0.4 to 0.45 mm, female 0.4 to 0.5 mm; vertex length of male 0.35 to 0.45 mm, female 0.4 to 0.5 mm; pronotal length of male 0.25 to 0.35 mm, female 0.3 to 0.35 mm. Vertex length 0.87 to 1.0 interocular width; pronotal length 0.7 to 0.82 vertex length. Character code: 0-1-1-0-1-0-0-0-1-2-2-1-0- 1-0-0-1-0. Holotype, male, and allotype, female, from Monument, Colorado, 19 Aug 1936 (R. H. Beamer) in KU; paratypes in KU and USNM. Athysanella bifida is related to utahna and ijwnana but can be separated by the elongate shaft of the aedeagus (Fig. 80), which is not widened at midlength. Specimens parasitized by Dryinidae and Strepsiptera, undeveloped specimens, and long-winged specimens have been examined. This species has been col- lected in Arizona, Colorado, Montana, New Mexico, North Dakota, South Dakota, Texas, Wyoming, and Canada. In the southern part of its range, at least, the host is blue grama, Bouteloua gracilis (Hicks et al. 1988). Athysanella utahna Osborn Figs. 84-86 Athysanella utahna Osborn 1930:705. Length of male 2.6 to 3.3 mm, female 3.9 to 4.9 mm; head width of male 0.9 to 1.05 mm, female 1.05 to 1.25 mm; pronotal width of male 0.85 to 1.0 mm, female 1.0 to 1.1 mm; interocular width of male 0.35 to 0.45 mm, female 0.45 to 0.55 mm; vertex length of male 0.4 to 0.45 mm, female 0.45 to 0.5 mm; prono- tal length of male 0.3 to 0.35 mm, female 0.3 to 0.4 mm. Vertex length 0.94 to 1. 13 interoc- ular width; pronotal length 0.7 to 0.82 vertex length. Character code: 0-1-0-0-1-0-0-0-2-1-2-0-0- 1-0-0-1-1. Lectotype, male, and lectoallotype, fe- male, from Ephraim, Utah, 20 July 1914 (E. D. Ball) in USNM. Athysanella utahna is related to yumana but can be separated by the more slender apical dorsal arm and less robust ventral arm of the style (Fig. 85). It can be separated from bifida by the shorter shaft of the aedeagus (Fig. 84). Rarely a specimen is seen with fus- cous spots on the vertex; the styles occasion- ally have a macroseta. Specimens with long wings and undeveloped genitalia and speci- mens parasitized by Dryinidae and Pipinculi- dae have been examined. This species has been collected in Arizona, California, Colo- rado, Idaho, Nevada, Utah, Oregon, Wash- ington, Canada, and Mexico. Reported from saltgrass, Distichlis spicata, in Mexico and Utah. Athysanella yumana Osborn Figs. 87-89 Athysanella yumana Osborn 1930:704. Length ofmale 2.8 to 3.1 mm, female 3.8 to 4.7 mm; head width ofmale 1.05 to 1. 15 mm, female 1. 1 to 1.25 mm; pronotal width ofmale 0.95 to 1.1 mm, female 1.0 to 1.2 mm; interoc- ular width ofmale 0.4 to 0.45 mm, female 0.45 to 0.55 mm; vertex length ofmale 0.4 to 0.45 mm, female 0.45 to 0.50 mm; pronotal length ofmale 0.3 to 0.35 mm, female 0.3 to 0.4 mm. Vertex length 0.94 to 1.0 interocular width; pronotal length 0.76 to 0.82 vertex length. Character code: 1-1-1-0-2-0-0-0-2-1-2-2-0- 1-0-0-1-0. Lectotype, male, and lectoallotype, fe- male, from Yuma, Arizona (H. Osborn) in USNM; paratypes in USNM and KU. Athysanella yumana is related to utahna but can be separated by the shape of the apex of the style (Fig. 88), which has more robust ventral and dorsal arms. Occasionally, there are fuscous spots on the vertex. The style may have an occasional macroseta. This species has been reported from saltgrass, Distichlis spi- cata; it has been collected in Arizona and Cali- fornia. Athysanella deserta Blocker, n. sp. Figs. 90-92 Length ofmale 3.0 to 3.3 mm, female 4.3 to 4.8 mm; head width ofmale 1.1 to 1.2 mm, female 1.2 to 1.3 mm; pronotal width ofmale 1.05 to 1.15 mm, female 1.15 to 1.25 mm; interocular width of male 0.45 to 0.5 mm, female 0.5 to 0.6 mm; head width ofmale 0.45 to 0.5 mm, female 0.5 to 0.6 mm; pronotal length ofmale 0.35 to 0.4 mm, female 0.35 to 0.45 mm. Vertex length 0.9 to 1.0 interocular width; pronotal length 0.75 to 0.83 vertex length. Character code: 0-0-1-0-2-0-1-0-2-2-2-2-0- 1-0-0-1-1. 36 Great Basin Naturalist Memoirs No. 12 Figs. 87-103. Figs. 87-89. Athijsanella ijmnana: 87, aedeagiis, lateral view; 88, style, broad aspect; 89, female sternum VII, ventral view. Figs. 90-92. Athijsanella deserta: 90, aedeagus, lateral view; 91, style, broad aspect; 92, female sternum VII, ventral view. Figs. 93-97. Athijsanella magdalena: 93, head and pronotum, dorsal view; 94, aedeagus, lateral view; 95, style, broad aspect; 96, male plate, ventral view; 97, female sternum VII, ventral view. Figs. 98-100. Athijsanella laeta. 98, aedeagus, lateral view; 99, style, broad aspect; 100, female sternum VII, ventral view. Figs. 101-103. Athijsanella stijlosa: 101, aedeagus, lateral view; 102, style, broad aspect; 103, female sternum VII, ventral view. Color stramineous; faint orange pattern of- ten seen on vertex; faint fuscous dots may be present on abdominal terga; face usually with faint lateral lines; venter with various amounts of fuscous coloring. Forewings brachypterous, exposing 3.5 to 4.0 pregenital abdominal terga; ocellus ap- proximately 1.5 its diameter from eye; hind tibial spur 1/3 to 1/2 length of 1st tarsomere; female abdominal sternum VII (Fig. 92) with conspicuous lateral lobes, inconspicuous me- dial lobe rounded, with various amounts of fuscous coloring. Pygofer with caudal margin rounded, with 25 -h microsetae; plates separated at base, truncate apically, not reaching apex of pygo- fer; anal tube exceeding apex of pygofer; valve with caudal margin rounded; connective 2/3 length of style; styles (Fig. 91) slightly exceeding apex of pygofer with apical arm 1988 Blocker, Johnson: Subgenus Athysanella 37 enlarged at apex, ventral arm with small, acute process; aedeagus (Fig. 90) with shaft narrowed apically, finely serrate on ventral margin, approximately 2X length of dorsal apodeme. Holotype, male, from Death Valley, Cali- fornia, 36 mi S Furnace Cr., 19 March 1971 (Oman); 18 male and 15 female paratypes, same data. Holotypes and paratypes in OrSU; paratypes in KSU. Athysanella deserta is related to ijiwiana but can be separated by the shape of the apical arms of the style with the dorsal arm enlarged at the apex and the ventral arm with a small, acute process. This species has been collected only at the type locality in the Mojave Desert of California. Athysanella magdalena Baker Figs. 93-97 Athysanella magdalena Baker 1898:185. Length ofmale 2.2 to 2.7 mm, female 3.0 to 3.6 mm; head width ofmale 0.85 to 0.95 mm, female 0.95 to 1. 1 mm; pronotal width ofmale 0.75 to 0.9 mm, female 0.9 to 1.1 mm; interoc- ular width ofmale 0.3 to 0. 4 mm, female 0. 4 to 0.5 mm; vertex length ofmale 0.3 to 0.4 mm, female 0.35 to 0.45 mm; pronotal length of male 0.25 to 0.35 mm, female 0.3 to 0.35 mm. Vertex length 0.93 to 1.1 interocular width; pronotal length 0.8 to 0.91 vertex length. Character code: 1-1-0-0-0-0-0-0-0-2-0-1-0- 1-0-0-1-0. Lectotype, female, from Magdalena Mts., New Mexico, Aug 1894 (F. H. Snow), and lectoallotype, male, from Forresters Ranch, Laramie Co. , Colorado, 3 Aug 1896 (No. 2013 ofBaker)inUSNM. Athysanella magdalena is related to tenera but can be separated by the elongate plates and the shape of the apex of the style (Fig. 95), which is more clearly bifid. Plates occasionally have one or more macrosetae, and occasion- ally the male hind tibial spur is undeveloped. Numerous undeveloped specimens, speci- mens parasitized by Dryinidae and Strep- siptera, and long-winged males and females have been examined. This species, collected in Arizona, California, Colorado, Montana, New Mexico, Nevada, Oklahoma, Texas, Utah, Wyoming, and Mexico, is a common specialist of blue grama, Bouteloua gracilis (Whitcomb et al. 1987, Hicks et al. 1988). Athysanella laeta Ball & Beamer Figs. 98-100 Athysanella laeta Ball and Beamer, 1940:16. Length ofmale 2.7 to 2.9 mm, female 3. 8 to 4.1 mm; head width ofmale 1.0 to 1.1 mm, female 1.1 to 1.2 mm; pronotal width ofmale 0.95 to 1.05 mm, female 1.05 to 1.15 mm; interocular width of male 0.45 to 0.5 mm, female 0.5 to 0.55 mm; vertex length ofmale 0.35 to 0.4 mm, female 0.4 to 0.45 mm; prono- tal length ofmale 0.35 to 0.4 mm, female 0.35 to 0.45 mm. Vertex length 0.75 to 0.89 intero- cular width; pronotal length 0.87 to 1.0 vertex length. Character code: 1-1-1-0-2-0-0-0-2-1-2-0-0- 1-0-0-1-1. Holotype, male, from Huachuca Mts., Ari- zona, 15 July 1934 (E. D. Ball); allotype, fe- male, same data except 14 July, in USNM; paratypes in USNM and KU. Athysanella laeta is related to incongrua but can be separated by its usually smaller size and the shape of the shaft of the aedeagus (Fig. 98), which is shorter and not as avicephaliform apically. Some of the male paratypes have undeveloped genitalia. This species has been collected in New Mexico and Arizona. Athysanella stylosa Blocker, n. sp. Figs. 101-103 Length ofmale 2.4 to 2.8 mm, female 3.8 to 4.1 mm; head width ofmale 0.95 to 1.1 mm, female 1. 15 to 1.2 mm; pronotal width ofmale 0.9 to 1.0 mm, female 1.05 to 1.1 mm; interoc- ular width ofmale 0.35 to 0.45 mm, female 0.45 to 0.5 mm; vertex length ofmale 0.35 to 0.45 mm, female 0.4 to 0.5 mm; pronotal length ofmale 0.3 to 0.35 mm, female 0.3 to 0.4 mm. Vertex length 0.93 to 1.06 interocu- lar width; pronotal length 0.75 to 0.82 vertex length. Color stramineous with brown pattern on vertex and pronotum, vertex with fuscous spots, wings with brown stripes, face with brown arcs. Forewings brachypterous, exposing 3 to 4.5 pregenital abdominal terga; ocellus approxi- mately its diameter from eye; hind tibial spur 3/4 1st tarsomere; female abdominal sternum VII (Fig. 103) with well-developed lateral lobes, slightly concave medially. Pygofer broadly rounded apically, with fewer than 25 setae; anal tube exceeds apex of 38 Great Basin Naturalist Memoirs No. 12 pygofer; valve with caudal margin rounded; plates rounded apically; styles (Fig. 102) bifid, dorsal arm acute, ventral arm truncate, with- out lateral flange or preapical lobe, exceeding apex of plates and pygofer; connective 3/4 length of style; aedeagus (Fig. 101) with shaft serrate on caudal margin, troughlike in caudal view, with inconspicuous process apically. Holotype, male, from Roswell, Chaves Co., New Mexico, Bitter Lake Natl. WLR, 14 Aug 1984 (R. F. Whitcomb) 0868A; 17 male and 7 female paratypes, same data; 1 male paratype, same data except 21 Aug 1985, 01931. Holotype and paratypes in KSU; paratypes in USNM and IPL. Athysanella stylosa is related to supina but can be distinguished by the fuscous spots on the vertex and by the shaft of the aedeagus, which is not as widened medially. This species is different in that the vertex length is equal to the interocular width and fuscous spots are present. It has been collected on gyp drop- seed, Sporobolus nealleiji, in the gypsum flats of southeastern New Mexico. Athysanella incongrua Baker Figs. 104-106 Athysanella incongrua Baker 1898:188. Length of male 3.0 to 3.4 mm, female 4.2 to 4.8 mm; head width of male 1.1 to 1.2 mm, female 1.15 to 1.35 mm; pronotal width of male 1.05 to 1. 15 mm, female 1. 1 to 1.25 mm, interocular width of male 0.45 to 0.5 mm, female 0.55 to 0.65 mm; vertex length of male 0.35 to 0.45 mm, female 0.45 to 0.5 mm; pronotal length of male 0.35 to 0.4 mm, fe- male 0.35 to 0.45 mm. Vertex length 0.76 to 0.85 interocular width; pronotal length 0.88 to 1.0 vertex length. Character code: 1-1-0-0-2-1-0-0-2-1-2-2-0- 1-0-0-0-0. Lectotype, male, from Fort Collins, Colo- rado, 21 July (C. F. Baker) in USNM. Athysanella incongrua is related to laeta and terebrans but can be separated by the length and shape of the apex of the shaft of the aedeagus (Fig. 104). The fuscous spots on the vertex are commonly missing. Long-winged males and females have been examined. Spec- imens, collected in Colorado, Iowa, Kansas, Nebraska, New Hampshire, New Mexico, Oklahoma, South Dakota, Texas, and Wyo- ming, appear to be associated with little bluestem, Schizachyrium scroparium. The specimen from New Hampshire was collected by P. W. Oman at Wiley House, 20 Aug 1934. Athysanella tenera Ball & Beamer Figs. 107-109 Athysanella tenera Ball & Beamer 1940:17. Length of male 2.5 to 3.0 mm, female 3.9 to 4.0 mm; head width of male 1.0 to 1.1 mm, female 1.1 to 1.2 mm; pronotal width of male 0.9 to 1.0 mm, female 1.0 to 1.15 mm; interoc- ular width of male 0.4 to 0.45 mm, female 0.45 to 0.5 mm; vertex length of male 0.35 to 0.45 mm, female 0.4 to 0.45 mm; pronotal length of male 0.3 to 0.35 mm, female 0.35 to 0.4 mm. Vertex length 0.94 to 1.0 interocular width; pronotal length 0.76 to 0.8 vertex length. Character code: 1-1-0-0-1-0-0-0-1-0-2-2-0- 1-0-0-1-0. Holotype, male, and allotype, female, from Las Vegas, Nevada, 8 Aug 1936 (R. H. Beamer) in KU; paratypes in KU. Athysanella tenera is related to magdalena but can be distinguished by the shorter plates and the irregular shape of the outer apical arm of the style (Fig. 108), which is widened and appears to be trifid. This species is known only from the type locality in Nevada. Athysanella vativala Blocker, n. sp. Figs. 110-112 Length of male 2.9 to 3.2 mm, female 4.2 to 4.6 mm; head width of male 1.05 to 1.15 mm, female 1.25 to 1.35 mm; pronotal width of male 1.0 to 1.15 mm, female 1.15 to 1.3 mm; interocular width of male 0.4 to 0.55 mm, female 0.55 to 0.65 mm; vertex length of male 0.35 to 0.5 mm, female 0.45 to 0.55 mm; pronotal length of male 0.35 to 0.4 mm, fe- male 0.35 to 0.45 mm. Vertex length 0.78 to 0.9 interocular width; pronotal length 0.74 to 0.94 vertex length. Character code: 1-0-0-0-1-0-0-0-1-1-1-0-0- 1-0-0-1-0. Color stramineous; vertex with fuscous spots; vertex, pronotum, and abdomen with additional brown pattern; fore wings with darkened stripes; face with lateral fuscous marks; legs with fuscous areas. Forewingsbrachypterous, exposing 4 to 4. 5 pregenital abdominal terga; ocellus 2X its di- ameter from eye or less; hind tibial spur 1/2 length of 1st tarsomere; female abdominal 1988 Blocker, Johnson: Subgenus Athysanella 39 V^ 113 114 Figs. 104-117. Figs. 104-106. Athysanella incongrua: 104, aedeagus, lateral view; 105, style, broad aspect; 106, female sternum VII, ventral view. Figs. 107-109. Athysanella tenera: 107, aedeagus, lateral view; 108, style, broad aspect; 109, female sternum VII, ventral view. Figs. 110-112. Athysanella vativala: 110, aedeagus, lateral view; 111, style, broad aspect; 112, female sternum VII, ventral view. Figs. 113-117. Athysanella terebrans: 113, head and pronotum, dorsal view; 114, aedeagus, lateral view; 115, 116, style, broad aspect; 117, female sternum VII, ventral sternum VII (Fig. 1 12) with lateral lobes much longer than medial lobe. Pygofer rounded apically, fewer than 25 macrosetae; valve with caudal margin angu- late; plates separated at base, rounded api- cally; connective 3/4 length of style; styles (Fig. Ill) deeply bifid, dorsal arm elongate and slender, ventral arm thickened, no preapical lobe present; aedeagus, in lateral view, with shaft slightly widened in apical half (Fig. 110), troughlike on ventral margin, and slightly serrate. Holotype, male, from Sheridan Co., Ne- braska, 7 mi N Rushville, Hwy 87, 9 Aug 1979 (H. D. Blocker and R. A. Sweet); female paratype, same data; 4 male and 7 female paratypes, Cherry Co., Nebraska, S Valen- tine, Hwy 83, mi 202, 10 Aug 1979 (H. D. Blocker and R. A. Sweet); 2 males. Cherry Co., Nebraska, near Manker, Hwy 20, 9 Aug 1979 (H. D. Blocker and R. A. Sweet); 1 male. Bowman, North Dakota, 5 July 1968, GL 819 (Harris and Cooley); 2 males and 10 females, Tryon, McPherson Co., Nebraska, 7 Aug 40 Great Basin Naturalist Memoirs No. 12 1977, (R. F. Whitcomb) 0480. Holotype and paratypes in KSU; paratypes in CNC, IPL, USNM. Athysanella vativala is very closely related and possibly conspecific with terebrans but can be separated by the shape of the apex of the style, which has a more slender dorsal arm, a more patterned and longer vertex, and the ocelli usually more distant from the eye. Undeveloped specimens and specimens para- sitized with Strepsiptera have been exam- ined. This species has been collected in Ne- braska and North Dakota on Calanovilfa longifolia. Athysanella terebrans (Gillette & Baker) Figs. 113-117 Euttetix terebrans Gillette & Baker 1895:102. Length of male 2.7 to 3. 1 mm, female 4.2 to 4.6 mm; head width of male 1.0 to 1.15 mm, female 1.2 to 1.35 mm; pronotal width of male 0.95 to 1.05 mm, female 1.15 to 1.3 mm; interocular width of male 0.45 to 0.5 mm, female 0.55 to 0.6 mm; vertex length of male 0.35 to 0.45 mm, female 0.45 to 0.5 mm; pronotal length of male 0.3 to 0.4 mm, female 0.4 to 0.45 mm. Vertex length 0.75 to 0.89 interocular width; pronotal length 0.82 to 1.0 vertex length. Character code: 1-0-0-0-1-0-0-0-1-1-1-0-0- 1-0-0-1-0. Holotype, female, from North Park, Colo- rado, 30 July (Gillette) in USNM; allotype, male, from Wray, Colorado, 13 July 1899, at Colorado State University. Athysanella terebrans is related to incon- grua but can be distinguished by the dorsal apical arm of the style (Figs. 115, 116), which is rounded, and by the shaft of the aedeagus (Fig. 114), which is not conspicuously widened apically; the female abdominal ster- num VII (Fig. 117) may be variable in shape. Long-winged and undeveloped specimens were commonly examined. This species has been collected in Colorado, Montana, Ne- braska, North Dakota, South Dakota, Utah, Wyoming, and Canada. It is reported from Dystichlis striata in Manitoba. Phylogeny Phylogeny of the subgenera oi Athysanella is discussed in Blocker and Johnson (1988). The subgenus Athysanella is designated a sis- ter o{ Gladionura and can be separated by the absence of a pygofer process in the former. The presence of a pygofer process is a conver- gent character that occurs in two other less closely related subgenera. We selected 40 characters that show differences within the subgenus for analysis. An intuitive phylogeny based on the proposed sequence of occur- rence of major derived characters was gener- ated, as well as an analysis by PAUP (Swofford 1986). Results of these analyses are shown in Figures 118 and 119. PAUP generated five equally most parsimonious trees; the tree with no unresolved trifurcations is presented here. A hypothetical ancestor consisting of a uniform plesiomorphic character set was uti- lized. Character sequences are contained in the preceding descriptions; the entire PAUP data set is available on request. Overall, the PAUP cladogram and the intuitive phylogeny are in good agreement. Species groups 1, 3, 4, and 7 are consistently grouped together (Fig. 119), whereas the rostrata and incongrua- planata groups are widely fragmented. It is possible that this fragmentation of the groups is indicative of the high degree of homoplasy in the genus. The robusta group is characterized by re- tention of the rounded shape of the style apex; the male plates are embrowned in all species (2'); the styles are widened and often bifid apically in all other species (1'). This group is recognized in the PAUP output. The rostrata group has an extended apex of the male style (1"); PAUP has the species in this group occurring close together but none as sisters. The incerta group is characterized by the presence of a semicircular apex of the shaft of the aedeagus (3'). There is fairly good congru- ence in the cladogram, which includes planata as a sister o( incerta . The terebrans-bifida group is characterized by an aedeagal shaft that is serrate on the caudal margin (4'). The terebrans group has a shortened vertex (5'); five of the seven species are recognized by PAUP; the other two (laeta and marthae) split out sooner but are close. The bifida group retains an elongate vertex; they are represented in the last half of the cladogram, close together, but only bifida and magdalena arc sisters. The incongrua-planata group retains an aedeagus with a simple shaft. PAUP separates 1988 Blocker. Johnson: Subgenus Athysanella 41 ro 4- CO rf 4^ Q ^ O) pastora robust a glob OS a gardenia kadokana salsa incerta vativala terebrans lacta marthae fredonla cursa hemijona bifida utahna ■ yumana ■ deserta •stylos a ■ tenera ■ magdalena • supina foeda parca incongrua aphoda strobila aspera plana whitcombi krameri planata rostrata kanabana furnaca \ I q o~- en 4^ OJ t^ Si CQ CQ OQ OQ CO Ua -! '-J "1 "1 ^ ■-! o o o o o o c c; c c c c "X3 "1:3 13 'T3 "O "O •^ - ■^ - ^ ■ CO - ^ - CO CO I to — tf ro O) ro ro en c;i O) - Lcs: 4^ cn ^ - rC cn Oi 4^- Oulgroup robusla pastora globosa gardenia krameri aspera wbitcoinbi plana parca aphoda incerta planata salsa kadokana strobila rostrata incongrua kanabana deserta furnaca yumana tenera supina foeda marthae bifida magdalena laeta utahna •stylata • vativala terebrans fredonia cursa hemijona Fig. 118. Intuitive cladogram. the incongrua group to a greater degree than any other group in the intuitive cladogram. The planata group species are characterized by an ocellus that is more remote from the eye than other groups (6') and all split out in the first half of the PAUP cladogram. Four species {krameri, aspera, whitcombi, and p/ona) arise from the cladogram in sequence. Acknowledgments Paul Oman has always been gracious and very helpful in his support of leafhopper sys- tematics; it is a high honor to contribute to a Fig. 119. PAUP cladogram. volume dedicated to his many contributions to our science. Merv Nielson collected with the senior author in Mexico and elsewhere and has offered assistance throughout this study. Likewise, the assistance of Bob Whit- comb and Andy Hicks has added significantly, particularly with host information and species distribution. Bob Brooks frequently furnished work space in the Snow Entomological Mu- seum. Paul Cwikla has read drafts of this and related manuscripts, and JefiFry Ediger, a stu- dent at KSU, inked the illustrations of the genitalia. 42 Great Basin Naturalist Memoirs No. 12 Literature Cited Ball, E. D , and R H Beamer 1940. A revision of the genus Athysanella and some related genera (Ho- moptera: Cicadellidae). Univ. Kansas Sci. Bull. 26: 5-82. Blocker, H. D. 1984. Morphological irregularities in the external genital structures of Athysanella (Ho- moptera: Cicadellidae: Deltocephalinae). Bull. Soc. Entomol. Suisse 57: 412. Blocker, H. D., and J. W Johnson. 1988. Subgenera of the genus Athysanella Baker (Homoptera: Ci- cadellidae: Deltocephalinae) and a proposed phy- logeny. Proc. VI Auchenorrhyncha Mtg., Torino, Italy, September 1987. Blocker, H. D., andC S. Wesley 1985. Distribution of Athysanella (Homoptera: Cicadellidae: Delto- cephalinae) in Canada and Alaska with descrip- tions of three new species. J. Kansas Entomol. Soc. 52: 377-386. Hicks, A L , R F Whitcomb, H D Blocker, and K. A Allred 1988. Why are there so many species of Athysanella? Proc. VI Auchenorrhyncha Mtg., Torino, Italy, September 1987. Swofford, D L. 1986. PAUP, phylogenetic analysis us- ing parsimony. Private publication. Wesley, C S , and H D Blocker 1985. Athysanella of Mexico including descriptions of a new subgenus and six new species (Homoptera: Cicadellidae). Entomography 3: 163-180. Whitcomb, R F , A L Hicks, D E Lynn, K A Allred, andH D Blocker. 1987. Geographic variation in host relationships of leafhoppers (Homoptera: Ci- cadellidae) in North American Grasslands. In: M. R. Wilson and L. R, Nault, eds., Proc. 2nd Int. Workshop on Leafhoppers and Planthoppers of Economic Importance. Commonwealth Inst, of Entomol., London. A NEW GENUS, ILEOPELTUS, RELATED TO CHLOROTETTIX (HOMOPTERA: CICADELLIDAE) Paul S. Cwikla' Abstract — On the basis of their unique genitahc characters, the Neotropical leafhopper species previously treated in the genus Doleranus Ball are placed in a new genus, Ileopeltus. Twelve species are treated, including six described as new. A key to the species and a cladistic hypothesis using Chlorotettix Van Duzee as the outgroup are presented. The new species of Ileopeltus include: nanocanthus (Panama), dorsalus and clavatus (Brazil and Venezuela), ventriculus and haplus (Brazil), and blockeri (Venezuela). This paper is the first in a series of revisions deahng with the New World deltocephahne genera, which have their crown completely microsculptured. These leaflioppers, com- monly called the broad-headed leaflioppers, are best known from the genus Chlorotettix Van Duzee. Species of this genus are com- monly found in low-lying, grassy habitats of North and South America. Traditionally, those leaflioppers with a completely microsculptured crown have been restricted to two genera, Chlorotettix Van Duzee and Doleranus Ball (Oman 1949). Lin- navuori (1959) mentioned that some species of Paratanus Young and Stirellus Osborn & Ball also have their crown microsculptured, al- though no degree of phylogenetic relationship was implied. More recently, I (Cwikla 1988) examined the North American species o{ Doleranus and found that they fell within an acceptable range of variation for Chlorotettix. I did not, how- ever, consider the Neotropical species of Dol- eranus congeneric with Chlorotettix because of the following synapomorphies: the asym- metrical aedeagus, the aedeagus without pro- cesses, and, with the exception of only two species, the male plates fused with the valve. Because of these unique attributes, the Neotropical species of Doleranus treated by Linnavuori (1959) and six species described as new are designated a new genus, Ileopeltus. Virtually nothing is known about the biol- ogy of Ileopeltus species. Ileopeltus tethys has been collected from sugar cane, weeds, sweet potato (Wolcott 1923), and grassy pastures (Caldwell and Martorell 1950) in Puerto Rico. Hosts for the other species of Ileopeltus are probably grasses. With the exception of/, tethys, specimens of Ileopeltus species are rare in collections. For the most part they appear restricted be- tween the Tropic of Cancer and the Tropic of Capricorn. Specimens have been collected in low- to mid-altitude areas. Twelve species are included in this new genus, of which six are described as new. In addition, this paper provides a key to the spe- cies and a discussion of the phylogenetic rela- tionships among the species. Specimens were prepared for SEM study as described in Cwikla and Freytag (1983). Names of institu- tions associated with abbreviations used in the text are in the acknowledgments section. Ileopeltus, n. gen. Diagnosis. — Ileopeltus species can be sep- arated from other deltocephaline genera by the asymmetrical aedeagus without processes and the crown completely microsculptured. Color. — Yellowish green or ochraceous. Markings occasionally present on crown and forewing. Structural features. — Small, delto- cephaline leaflioppers. Crown roundly pro- duced, median length slightly longer than length next to eye. Clypellus not constructed proximally. Forewing with cross-vein in claval area. Male genitalia. — Pygofer roundly pro- duced or truncated, process present or ab- sent. Anal tube weakly sclerotized dorsally. Valve acutely triangular, usually fused to plate posteriorly. Plate short or slightly longer than 'Department of Entomology, University of Missouri, Columbia. Missouri 65211, Present address: 14560 Josephine Street, Irwin, Pennsylvania 15642. 43 44 Great Basin Naturalist Memoirs I No. 12 Figs. 1-4. Ileopeltus tethijs Van Duzee; 1, pygofer, left lateral aspect; 2, right style, dorsal aspect; 3, aedeagus, ventral aspect; 4, aedeagus and apex of connective, left lateral aspect. pygofer, rounded apically, macrosetae unise- riate, second short row of microsetaelike structures occasionally present, lateral mar- gin sinuate, straight, or slightly convex. Stylar apex linear, preapical angle usually absent. Connective as long as or shorter than aedea- gus. Aedeagus without processes, asymmetri- cal, apex acute in posterior aspect. Gonopore on right side at base of shaft. Type species. — Chlorotettix tethijs Van Duzee 1907:71. Distribution. — Neotropical, usually re- stricted between the Tropic of Cancer and the Tropic of Capricorn. Notes. — This genus has been previously described as Doleranus by Linnavuori (1959). The Neotropical species are not considered congeneric with the Nearctic forms and have been treated as synonyms oi Chlorotettix (in- cluding the type species of Doleranus, Tham- notettix longulus Gillette & Baker) in a disser- tation by Cwikla (1988). Key to the Male Ileopeltus Species 1. Pygofer without a process (Fig. 1), or if present then e.xtremely small (Fig. 7) 2 — Pygofer with long, distinct processes (Fig. 43) 3 2(1). Pygofer without a process (Fig. 1) tethys (Van Duzee) — Pygofer with short process on posterodorsal mar- gin in lateral aspect nanocanthus , n. sp. .3(1). Stylar apex curved laterally (Fig. 52) Cyclops (Linnavuori) — Stylar apex linear (Figs. 12, 30) 4 4(3). Pygofer with process directed ventrad in lat- eral aspect ventriculus. n. sp. — Pygofer with process directed mesad or dor- sad in lateral aspect 5 5(4). Pygofer with process heavily sclerotized, in- serted on medial side in lateral aspect (Fig. 27) 8 — Pygofer with process not heavily sclerotized, not inserted on medial side in lateral aspect (Fig. 11) 6 6(5). Aedeagal shaft wide in ventral aspect (Fig. 25) hapliis, n. sp. — Aedeagal shaft narrow in ventral aspect (Fig. 22) . . .' 7 7(6). Stylar apex truncate (Fig. 12) . . dorsalus, n. sp. — Stylar apex narrow (Fig. 20) . spinosus (DeLong) 8(5). Stylar apex truncate and constricted subapi- cally (Fig. 40) clavaUis, n. sp. — Stylar apex rounded or if truncate, then not constricted subapically 9 9(8). Pygofer with process inserted on middle of ventral margin (Fig. 33) blockeri, n. sp. — Pygofer with process inserted on posteroven- tral margin (Figs. 27, 49) 10 10(9). Aedeagus wide and highly asymmetrical in 1988 CwiKLA: The Genus Ileopeltus 45 Fig. 5. Ileopeltus tethys Van Duzee, dorsal habitus. ventral aspect (Fig. 49) cuneus (DeLong & Martinson) — Aedeagus narrow in ventral aspect 11 11(10). Aedeagus with large gonopore in lateral as- pect (Fig. 32); plate fused with valve hasttilus (DeLong & Linnavuori) — Aedeagus with small gonopore in lateral as- pect (Fig. 46); plate not appearing fused with valve aberrans (Osborn) Ileopeltus tethys (Van Duzee), n. comb. Figs. 1-5, 57, 59; Map 1 Chlorotettix tethys Van Duzee 1907:71. Chlorotettix hidentatus DeLong 1923:264, Wolcott 1936: 86. Chlorotettix dilutus Oshorn 1923:73, Osborn 1935:118. Doleranus kinonanus Ball 1936:432, Linnavuori 1959: 274-275. Diagnosis. — Ileopeltus tethys is near /. nanocantus and can be separated from it and other Ileopeltus species by the lack of a pygoferal spine. Length. — Male 3.8-4.8 mm, female 4.1- 5.4 mm. Color, — Yellowish green. Eye reddish brown. Forewing yellowish subhyaline, small brown patch occasionally present on middle of wing and/or on claval area. Male genitalia. — Pygofer with posterior margin roundly produced, processes absent. Plate short, approximately half the length of pygofer, lateral margin straight or slightly concave, not fused to valve. Stylar apex stout, linear, preapical angle absent. Aedeagus asymmetrical in posterior aspect, left subapi- cal margin sinuate in posterior aspect, right subapical margin straight or somewhat convex in posterior aspect. Gonopore at base on right side. Female seventh sternum. — Posterior margin shallowly excavated, median of exca- vation with short, produced projection bear- ing two short teeth (see Linnavuori 1959: Fig. 113g). Depth of excavation varies from only slightly to one-third width of segment. Type. — A single female cotype bearing the labels "Martinique W.I., VII-26" and "Aug. Busck collector" was examined from the USNM. Distribution. — ^Many specimens from the following localities were examined: Central America: Honduras, Mexico (Chiapas, Guer- rero, Michoacan, Oaxaca, Tamaulipas, Vera- cruz), Nicaragua, Panama. West Indies: An- tigua, Cayman Islands, Dominican Republic, Guadeloupe, Haiti, Jamaica, Montserrat, Nevis, Puerto Rico, Trinidad, and Tobago. South America: Venezuela (Guarico). Specimens were collected from May through December and are deposited in the UPB, UCV, UKC, USNM, OSUC, and BMNH collections. Note. — The types of hidentatus, dilutus, and kinonanus were not available for study; consequently, previous synonymies of these names under /. tethys were not verified. Be- cause the types are females, it is doubtful that they can be correctly associated with males at this time. Ileopeltus nanocanthus, n. sp. Figs. 6-10, 58, 60; Map 2 Diagnosis. — Ileopeltus nanocanthus is near /. tethys and can be separated from it and other Ileopeltus species by the short pygoferal spine. Length. — Male 4.6-5.4 mm, female 4.8- 5.3 mm. 46 Great Basin Naturalist Memoirs No. 12 Map 1. Distribution of Ileopeltiis tethijs. Color. — Ochraceous or greenish yellow. Eye reddish brown. Crown occasionally with faint orange, median band. Pronotum with four longitudinal, faint orange bands. Fore- wing yellowish subhyaline, brown spots present at bases of anteapical cells and on claval area. Male genitalia. — Pygofer with ventral margin roundly produced into short spine on dorsoposterior margin, spine directed medi- ally in ventral aspect. Plate short, anterior margin convex. Stylar apex narrowed, di- rected posteriorly, preapical angle absent. Aedeagus linear, asymmetrical in posterior aspect, apex with opposite curve compared to /. tethys. Gonopore at base on right side. Female seventh sternum. — Posterior margin with U-shaped excavation extending half length of segment, base of excavation with small V-shaped notch, notch surrounded by light brown coloration, lateral angle rounded. Type. — Male holotype, Panama, Canal Zone, Curundu, 26-3'()-XI-1986, B. Sieber- glied collector. Four female paratypes, same data as holotype; one male paratype, Panama, Chiriqui, Rio Colorado, 1,200 m, 8°5'N, 82°43'W, 26-XII-1974, H. Wolda collector; male paratvpe, Panama, Panama, Las Cum- bres, 17-23-11-1982, H. Wolda collector. Holotype, two paratypes with same data as holotype, and one paratype from Las Cum- bres deposited in OSUC, remainder in HWC. Etymology. — Greek, nano (dwarf) and acantha (spine) refer to the small pygofer pro- Ileopeltiis dorsalus, n. sp. Fig.s. 11-14; Map 2 Diagnosis. — Ileopeltus dorsalus is near 7. ventriculus and can be separated from it by the acute, dorsally directed pygoferal spine. LENCiTH. — Male 4.4 mm, female unknown. Color. — Ochraceous, either without dark markings or crown with light brown surround- ing coronal sulcus. Eye red. Pronotum with six light brown, longitudinal bands. Scutel- lum with brown lateral angles. Forewing sub- hyaline with brown patches at base of anal 1988 CwiKLA: The Genus Ileopeltus 47 Figs. 6-10. Ileopeltus nanocanthus, n. sp.: 6, pygofer, left lateral aspect; 7, pygofer, ventral aspect; 8, right style, dorsal aspect; 9, aedeagus, ventral aspect; 10, aedeagus and apex of connective, left lateral aspect. Map 2. Distribution of Ileopeltus nanocanthus (squares), /. dorsalus (circles), and I. ventriculus (triangles) 48 Great Basin Naturalist Memoirs No. 12 Figs. 11-14. Ileopeltus dorsalus, n. sp.: 11, pygofer, left lateral aspect; 12, right style, dorsal aspect; 1.3, aedeagus, ventral aspect; 14, aedeagus and apex of connective, left lateral aspect. veins, proximal and distal ends of inner ante- apical cell, and proximal end of central ante- apical cell. Male genitalia. — Pygofer with posterior margin produced into dorsally directed spine, acute apically, not heavily sclerotized, small group of microsetaelike structures near mid- dle of posterior margin. Plate triangular, lat- eral margin straight. Stylar apex truncate, di- rected posteriorly, preapical angle small. Aedeagus only slightly asymmetrical. Type. — Male holotype, Venezuela, Aragua, El Limon, 22-11-1973, Malaise trap, C. J. Resales collector. Male paratype, Brazil, Ceara, Barbalha, V-1969, M. Alvarenga, B.M. 1971-165. Holotype deposited in OSUC, paratype in BMNH. Etymology, — Latin, dorso (back) refers to the dorsally directed pygoferal spine. Ileopeltus ventriciilus , n. sp. Figs. 1.5-18; Map 2 Diagnosis. — Ileopeltus ventriculus is near spinosus and can be separated from it and other Ileopeltus species by the pygoferal pro- cess directed ventrally. Length. — Male 4.4 mm, female unknown. Color.— Ochraceous, without distinct dark markings. Eye reddish brown. Male genitalia. — Pygofer with short, ven- trally directed process inserted on posterior margin. Plate elongate, lateral margin sinu- ate, apex curved dorsally, bluntly rounded. Style elongate, apex linear, slightly curved laterally. Aedeagus elongate, broadly curved dorsally in lateral aspect, apex acute in poste- rior aspect. Type. — Male holotype, Brazil, Ceara, Crato, May 1969, M. Alvarenga collector. Type deposited in BMNH. Etymology. — Latin, ventricul (belly) refers to the ventrally directed pygofer pro- cess. Ileopeltus spinosus (Dehong), n. comb. Figs. 19-22; Map 3 Chlorotettix spinosus DeLong 1945:10-11. Diagnosis. — Ileopeltus spiiiosus is near /. dorsalus and can be separated from it and other Ileopeltus species by the presence of a preapical angle on the style and the unique pygoferal process. Length. — Male 5.2-5.5 mm, female un- known. Color. — Greenish yellow, without dark- markings. Eye gray or greenish yellow. 1988 CwiKLA: The Genus Ileopeltus 49 Figs. 15-18. Ileopeltus ventricuhis , n. sp.: 15, pygofer, left lateral aspect; 16, right style, dorsal aspect; 17, aedeagus, ventral aspect; 18, aedeagus and apex of connective, left lateral aspect. Male genitalia. — Pygofer with posterior margin broadly rounded apically, heavily sclerotized process inserted on posteroventral margin, directed posteriorly then curved me- dially. Plate with lateral margin slightly sinu- ate, apex elongate, pointed. Stylar apex nar- rowly elongate, directed slightly laterally, preapical angle roundly triangular. Aedeagus elongate, not strongly asymmetrical, narrow in ventral aspect. Gonopore on right side near base. Type. — Male holotype bearing the labels "Buena Vista, Gro. [Guerrero, Mexico], 23-X- 1941, 3,400 ft." and "DeLong, Good, Cald- well and Plummer" was examined from OSUC. Distribution. — Besides the type, three paratypes from Mazaclan, Guerrero, Mexico, 3-X-1945, were examined from OSUC. Note. — DeLong (1945) reported this spe- cies from meadow grasses. Ileopeltus haplus, n. sp. Figs 23-26; Map 3 Diagnosis. — Ileopeltus haplus is near /. spinosus and can be separated from it and other Ileopeltus species by the unique pygofer process and the wide aedeagus in ventral as- pect. Length. — Male 5.3 mm, female unknown. Color. — Yellowish green, without dark markings. Eye yellowish green. Male genitalia. — Pygofer with posterior margin produced into dorsally directed spine, spine not heavily sclerotized. Plate triangular, lateral margin straight, apex rounded. Stylar apex linear, preapical angle absent. Aedeagus asymmetrical, left margin forming carina in posterior aspect. Gonopore on right side near base. Type. — Male holotype, [Brazil] MS [Matto Grosso du Sul], Campo Grande, l-X-1982, W. KoUer collector. Type deposited in UPB. Eri'MOLOGY. — Greek, haplo (simple) refers to the color of this species. Ileopeltus hastulus (DeLong & Linnavuori), n. comb. Figs. 27-32; Map 3 Chlorotettix hastulus DeLong & Linnavuori 1978:121- 122. Diagnosis. — This species can be separated from other members of this genus by the long pygofer process and the aedeagus with a large gonopore. Length. — Male 5.0-5.2 mm, female un- known. 50 Great Basin Naturalist Memoirs No. 12 Figs. 19-22. Ileopeltus spinosus (DeLong): 19, pygofer, left lateral aspect; 20, right style, dorsal aspect; 21, aedeagus, ventral aspect; 22, aedeagus and apex of connective, left lateral aspect. Map 3. Distribution of //eo/«'/pe is the only specimen knowTi to have the dorsum of the abdomen browTi and a slightly shorter aedeagal shaft (Fig. 25). similar to peregrina . The setal for- mulae var\- as follows: foretibia. 1.3. 1.4. or 1.5; midtibia, 4.4 or 5.4; hind femur. 2. 1. 1 or 2.1.1.1. Balclutha peregrina (Kirkaldy) Fig. 12 Nesosteles peregrina Kirkaldy, 1910: 575. Holotype 6, Hawaii (BPBM) [examined]. Balclutha peregrina (Kirkaldy): Zimmerman 1948: 86-87, Fig. 27, Namba 19.56: 108, Figs. 8a-c. Host — Cyathodes, Deschampsia (Gram- ineae), Railliardia (Compositae), Vaccinium (Ericaceae), Styphelia (Epacridaceae). Distribution — Endemic (Maui, Hawaiij. M.\TERIAL EX.^MINED — Holot>pe (5, Ha- waii: Kilauea, vi.l903, R.C.L.P. Hawaii: (nu- merous specimens from Kilaueaj; 26, 59, Stainback Highwav, 2,134 m, on Styphelia (BPBNL BMNH). ' Remarks — This species is similar to plu- tonis but is significantly smaller (6 , 2.6-3.1 mm; 9 , 2.8-3. 1 mm ), has the abdomen some- times marked with pale brown, the sides of the clypellus straight (Fig. 12), and the midtibial setal formula 4.3 or 4.4. A few speci- mens examined (Stainback Highway) are tinged with scarlet and have the abdomen and thoracic stemites marked with dark brown. Balclutha usitata Namba Fig. 27 Balclutha usitata Namba. 19.56: 104-105. Figs. 2a-b. Holot\"pe i . Kauai (BPBM) [examined]. Host. — Cyathodes (Gramineae). Distribution. — Endemic (Kauai. Oahu, Molokai. Maui. Hawaii . Material E.\.\MiNED — Holot\-pe 6 , Kauai, Kainamanu, vii.1952, 3,800' (BPBM). Nu- merous specimens from throughout its range BMNH. BPBM). Renl\RKS. — This species can be distin- guished by its small size (d , 2.6-2.9 mm, 9, 2.8-3.1 mm), its overall yellow or greenish \ellow color without brown markings on the dorsum of the abdomen, the 3.3 midtibial setal formula, and the angularly cur\-ed aedea- gal shaft (Fig. 27). Some specimens have stramineous markings similar to but paler than in volcanicola (Figs. 5, 7 . Balclutha timberlakei (Osbom) Figs. 13-15, 22 Nesosteles timberlakei Oshom. 1935: -59-60, Figs. 25a-d. Holot>pe 6. Oahu BPBM [examined]. Balclutha timberlakei iOsbom(: Zimmerman 1948: 87, Figs. 7e. 24c: Namba 19.56: 108-109, Figs. 9a-b. Host. — Eragrostis variabilis (Gramineae). Distribution — Endemic (Kauai. Oahu, 96 Great Basin Naturalist Memoirs No. 12 Molokai, Lanai, Maui, Hawaii). Material E>L\MINED. — Holotype S, Oahu (BPBM). Oahu: \S (paratype) Palolo Val. on Eragrostis variabilis (BMNH). Maui: Id, Waikapu Val. (BPBM). Hawaii: M, 29, Po- hakuloa, 1,800 m (BPBM). Remarks. — This species can be distin- guished by its small size ((5, 3.0 mm, 9 , 3.4 mm), elongate head that is distinctly wider than the pronotum (Fig. 15), general yellow color without brown markings on the dorsum of the abdomen, 3.3 midtibial setal formula, robust and slightly sinuate shaft of the aedea- gus (Fig. 22), and its elongate connective, which is longer than the style (Figs. 13, 14). Balclutha phoxocephala Namba Fig. 23 Balclutha phoxocephala Namba, 1956: 109, Fig. 10. Holotype 6, Kauai (BPBM) [examined]. Host. — Cyathodes, Eragrostis variabilis (Gramineae). Distribution. — Endemic (Kauai, Oahu, Molokai). Material e.xamined. — Holotype 6 , Kauai: Nualolo, viii.1925, on Cyathodes (BPBM). Kauai, Oahu, Molokai (96, 59, paratvpes, BPBM). Remarks. — This species can be distin- guished by its small size {6 , 2.4-2.6 mm, 9 , 2.7-3.0 mm), 3.3 midtibial setal formula, and robust aedeagus (Fig. 23). Some specimens have stramineous markings similar to but paler than in volcanicola. A series of specimens from Hawaii, Pohaku- loa, 6,300' (BPBM), may be this species but have the aedeagal shaft more robust. Balclutha incisa hospes (Kirkaldy), n. stat. Figs. 28-40 N esosteles hebe \'dr . hospes, Kirkaldy. 1910: 574; Osborn 1935: 57, Figs. 24a-b. Lectotype 9, Hawaii (BPBM), designated by Zimmerman (1948: 85) [examined]. N esosteles hospes TimberVdke, 1918: .381. Balclutha hospes (Kirkaldy): Zimmerman, 1948: 85, Fig. 24b, Namba 1956: 107, Figs. 6a-b. Host. — Chloris radiata, Panicum piirpur- ascens (Gramineae). Distribution. — Cosmopolitan and through- out the Hawaiian islands. Material examined, — Lectotype 9 ofNe- sosteles hebe var. hospes, Hawaii: Kauai (BPBM). Numerous specimens from Kauai, Oahu, Molokai, and Hawaii (BPBM, BMNH). Remarks. — ^This subspecies can be distin- guished by the head being slightly wider than the pronotum, the short vertex (Fig. 28), the ocelli being only 1.0-1.5 times their own diameter from the corresponding eye, the coronal suture visible in facial view (Fig. 29), distinctively shaped aedeagus (Figs. 39, 40), and the brown markings on the female pre- genital sternite (Fig. 36). It is similar in exter- nal appearance to B. saltuella (see Remarks under salt uella). This subspecies, which occurs in Hawaii, the Marquesas Islands, and the Americas, dif- fers from the nominate subspecies in the Old World by the more laterally directed ventral process of the aedeagus. This distinction sug- gests that the species may have reached Hawaii from the Americas rather than the Pacific. Contrary to Timberlake's data (1918), the shape of the pygofer process (Fig. 37) is similar but variable in both subspecies. Balclutha saltuella (Kirschbaum) Figs. 41-46 Jassus (Tharnnotettix) saltuellus Kirschbaum, 1868; 86. Holotype 9 , Germany (Museum Wiesbaden) [ex- amined]. Balclutha beardsleyiNamhd, 1956: 110, 112, Figs, lla-b. Holotype d, Hawaii (BPBM) [e.xamined], syn. nov. Host. — Unknown in Hawaii; elsewhere: Agrostis (Gramineae) and Gossypium (Mal- vaceae) (Knight 1987). Distribution. — Cosmopolitan and on Oahu. Material e.xamined. — Holotype 9 Jassus {Tharnnotettix) saltuellus, Germany (Mu- seum Wiesbaden). Holotvpe 6 B. beardsleyi, Oahu: Honolulu, i. 1955 (BPBM). Oahu: 16, 39, Honolulu; 19, Ewa(BPBM). Remarks. — This species is similar to B. in- cisa in external appearance but differs from this and other Hawaiian species by its reduced number of foretibial spines (1.1), its medially directed, lateral frontal sutures, and the short, broad rostrum in facial view (Fig. 44). It differs from incisa, in addition, by having the prosternum dark brown (Fig. 43), the female pregenital sternite with a medial, triangular lobe on the posterior margin, and the disc devoid of brown markings. Nesolina Osborn NesoHna Osborn, 1935: 60. Type species: Nesolina lineata Osborn, bv original designation: Zimmerman 1948: 87. 1988 Knicht, Webb: Hawaiian Endemics in Balclutha 97 46 Figs. 28-46. Balclutha species. 28-40, B. incisa hospes: 28, head and thorax, dorsal view; 29, face; 30, apex of left hind femur; 31, right forewing; 32, right hindwing; 33, apex of second valvulae, ventral view; 34, apex of left midtibia; 35, apex of left foretibia; 36, female pregenital sternite; 37, ventroposterior corner of male pygofer lobe, lateral view; 38, left second valvula, lateral view (toothed area between broken lines); 39, aedeagus, left lateral view; 40, aedeagus, posterior view. 41-46, B. saltuella: 41, aedeagus, left lateral view; 42, connective, dorsal view; 43, head and thorax, left lateral view; 44, face; 45, right style, dorsal view; 46, posteroventral corner of male pygofer lobe. Remarks. — ^This endemic Hawaiian genus can be distinguished by its long head and distinctive markings, its large aedeagus with- out processes, and the plumose macrosetae of 98 Great Basin Naturalist Memoirs No. 12 Figs. 47-53. Balclutha lucida: 47, head and thorax, dorsal view; 48, left style, dorsal view; 49, male pygofer, left lateral view; 50, left subgenital plate, ventral view; 51, aedeagus, left lateral view; 52, connective, dorsal view; 53, distribution. the legs and genitalia. The foretibia and midtibia each have one long, and one short, stout apical seta. Setal formulae: foretibia, 1.3; midtibia, 3.3; hind femur, 2.1.1. (see also Remarks under "Relationship of Balclutha to Other Hawaiian Macrostelini"). Nesolina lineata Osborn Figs. 54-66 Nesolina lineata Osborn, 1935: 60-61, Figs. 26a-c: Zim- merman 1948: 88, Figs. 7d, 8e-f, 24d. Lectotype 6 , Oahu (BPBM), here designated [examined]. 1988 Knight, Webb: Hawaiian Endemics in Balclutha 99 Figs. 54-66. Nesolina lineata: 54, head and thorax, dorsal view; 55, face; 56, apex of left foretibia; 57, right forewing; 58, right hindwing; 59, immature, head and thorax, dorsal view; 60, apex of first valvulae; 61, aedeagus, left lateral view; 62^ aedeagus, posterior view; 63, left second valvula, lateral view (toothed area between broken lines); 64, male genital segment, left lateral view; 65, apex of left style, lateral view; 66, subgenital plates, dorsal and ventral view, valve, connective, and left style, dorsal view. Host. — Eragrostis variabilis (Gramineae). Distribution. — Endemic (Oahu, Hawaii). Material examined. — Lectotype d, Oahu: Diamond Head, 18. ii. 1917, W. M. Giffard. 100 Great Basin Naturalist Memoirs No. 12 Oahu: IM, 89, 5 immatures, Diamond Head, various dates (paralectotypes) (BPBM). Hawaii: 16, Kau; 16, Pohakuloa, on bunch grass (BMNH). Balolina, n. gen. Type species; Macrosteles kilatieae Kirkaldy. Length. — 3.6-4.3 mm. Yellow to greenish yellow, variably marked with brown. Head slightly to distinctly nar- rower than pronotum. Vertex triangular, me- dial length approximately 1.5 times length next to eye, foremargin broad to narrowly parabolic. Ocelli marginal, visible dorsally, situated approximately midway between eye and midline. Face slightly wider than long, lateral margin strongly sinuate below eye. Clypeus narrow, lateral margins incurved at level of eye. Transclypeal suture indistinct. Clypellus narrow, slightly expanded to near apex, extended well beyond margin of face. Laterofrontal sutures extending one-half dis- tance to corresponding ocellus. Pronotum 1.5 times length of vertex, lateral margins short. Forewings elongate, appendix wide, extend- ing around first and second apical cells; outer subapical cell absent; inner subapical cell open basally. Hindwing with veins r and m fused distally. Forefemora with two stout api- cal setae. Foretibia setal formula 1.3. Mid- femora with one long, and one moderately long, stout apical seta. Midtibia setal formula usually 3.3. Hind femora setal formula usually 2.2.1. Base of hind tarsomere sulcate. Male. — Pygofer narrowly rounded posteri- orly; a group of submarginal macrosetae pos- teriorly. Subgenital plates gradually tapered to lightly sclerotized, fingerlike apex; a multi- seriate row of macrosetae along ventrolateral margin. Styles with apical process tapered to acute apex in ventral view, apex footlike in medial view; lateral lobe well developed; in- ner basal apophyses short. Connective Y- shaped; stem short, equal in length to arms; articulating with aedeagus. Aedeagus with shaft short and robust, cylindrical; a pair of short processes apically; gonopore apical on ventral surface; basal apodeme short. Female. — First valvulae fused. Second valvulae evenly tapered to acute apex; teeth very fine, extending over distal half of dorsal margin and onto apex of ventral margin. Remarks. — This genus is similar in appear- ance to Balchitha and Nesoclutha but can be distinguished from these genera by the ab- sence of hairy macrosetae, shorter laterofron- tal sutures, indistinct transclypeal suture, the ocelli situated more distant from the eyes, the subgenital plates with multiseriate macrose- tae, and the aedeagus with a pair of apical processes. It differs from Balchitha also by the other characters noted in the key. Balohna kilaueae (Kirkaldy), n. comb. Figs. 67-83 Macrosteles kilaueae Kirkaldy, 1910; 575. Holotype 9, Hawaii (BPBM) [examined]. Balchitha kilaueae (Kirkaldy); Zimmerman 1948; 86, Figs, 7a, Ud-f; Namba 1956; 103-104, Figs. la-b. Host. — Cihotium chamissoi and C. men- ziesii (Dicksoniaceae) (ferns). Distribution. — Endemic (Kauai, Oahu, Molokai, Maui, Hawaii). Material examined. — Holotype 9, Hawaii, Kilauea, vii.1906, RCL Perkins. Nu- merous specimens from throughout its range (BPBM, BMNH). Remarks. — The specimens examined varied in color and in the length of the vertex. Brown markings are often present on the dor- sal and ventral surfaces of the abdomen and sometimes on the thorax, the clypeus, the basal two-thirds of the ovipositor, the basal half of the corium of the forewing, and the hind legs. One specimen examined from Maunawainui Valley, Molokai, has the clav- us of the forewing pale scarlet. Setal formu- lae: foretibia, 1.3; midtibia, usually 3.3 but sometimes 4.3, 4.4, or 5.3; hind femur, usu- ally 2.2.1, sometimes 2.1.1.1, 2.2.1.1, or 2.1.2.1.1. This species is similar in external appear- ance to Balchitha nambai with which it may be confused. Acknowledgments We thank Keith Arakaki (BPBM) for the loan of material under his care. Literature Cited Blockkk, H D 1967. Classification of the Western Hemisphere Balchitha (Homoptera; Cicadelli- dae). Proceedings of the United States National Museum 122; 1-55. Fabkicius.J C. 1775. Systeme Entomologiae. Flensburgi et Lipsiae. xxviii + 832 pp. 1988 Knight. Webb: Hawaiian Endemics in Balclutha 101 Figs. 67-83. Balolina kilaueae: 67, head and thorax, dorsal view; 68, face; 69, apex of left femur; 70, right forewing; 71, right hindwing; 72, apex of left midtibia; 73, apex of left foretibia; 74, apex of hind tibia and base of first tarsus; 75, left second valvula, lateral view (toothed area between broken lines); 76, aedeagus, left lateral view; 77, right style, dorsal view; 78-79, apex of right style, ventral and medial view; 80, connective, dorsal view; 81, male genital segment, left lateral view; 82, apex of aedeagus, posterior view; 83, subgenital plates, dorsal and ventral view, valve, connective, and left style, dorsal view. FlEBER, F. X. 1866. Neue Gattungen und Arten in Ho- 497-516. moptern(CicadinaBur.). Verhandlungen der Zo- Ghauri, M. S. K 1966. Revision of the genus Orosius ologisch-Botanischen Gesellschaft in Wien 16: Distant (Homoptera; Cicadelloidea). Bulletin of 102 Great Basin Naturalist Memoirs No. 12 the British Museum (Natural History) Entomol- ogy 18: 231-252. KiRKALDY, G W 1900. Bibliographical and nomenclatu- ral notes on the Rhynchota. No. 1. Entomologist 33; 238-243. 1910. Fauna Hawaiiensis 2: 531-700. KiRSCHBAUM, C L 1868. Die Cicadinen der Gegend von Wiesbaden und Frankfurt a. M. nebst einer An- zahl neuer oder schwer zu unterscheidender Arten aus anderen Gegenden Europas. Jahrbuch des Nassauischen Vereins fur Naturkunde 21-22: 1-202. Knight, W. J. 1987. Leafhoppers of the grass-feeding genus Balclutha (Homoptera, Cicadellidae) in the Pacific region. Journal of Natural History 21: 1173-1224. LiNNAVUORl. R 1960. Cicadellidae (Homoptera, Auchen- orrhyncha) of Fiji. Acta Entomologica Fennica 15: 1-70. Namba, R. 1956. A revision of the Balclutha species found in Hawaii, with descriptions of five new species (Homoptera: Cicadellidae). Proceedings of the Hawaiian Entomological Society 16: 101-112. Oman, P W 1949. The Nearctic leafhoppers (Ho- moptera: Cicadellidae). A generic classification and check list. Memoirs of the Entomological So- ciety of Washington No. 3: 1-253. OSBORN, H 1935. Cicadellidae of Hawaii. Bulletin Ber- nice P. Bishop Museum 134: 1-62. OSSIANNILSSON, F 1983. The Auchenorrhyncha (Ho- moptera) of Fennoscandia and Denmark. Part 3: the family Cicadellidae: Deltocephalinae, cata- logue, literature and index. Fauna Entomologica Scandinavica 7: 594-981. Simon, C. 1987. Hawaiian evolutionary biology: an intro- duction. Trends in Ecology and Evolution 2: 175-229. SoUTHWOOD, T R E 1978. Ecological methods. Chap- man & Hall, London. 524 pp. TiMBERLAKE, P H 1918. Note on the non-identity of a common Hawaiian jassid with Nesosteles hebe Kirkaldy of Fiji. Proceedings of the Hawaiian En- tomological Society 3: 381. Triplehorn. B. W.. and L. R Nault 1985. Phylogenetic classification of the genus Baldulus (Homoptera: Cicadellidae), and notes on the phylogeny of the Macrostelini. Annals of the Entomological Society ofAmerica 78: 291-315. Webb, M D 1980. The Cicadellidae from Aldabra, As- tove and Cosmoledo atolls collected by the Royal Society E.xpedition 1967-68 (Hemiptera, Ho- moptera). Journal of Natural History 14: 829-863. 1986. Sugar-cane cicadellids of the genus Yanwjfo- tettix (Homoptera: Auchenorrhyncha). Journal of Natural History 20: 131-141. 1987. The endemic Macrostelini of the island of St. Helena (Homoptera, Cicadellidae). Revue de Zo- ologie Africaine 100: 453-464. Zimmerman, E C 1948. Insects of Hawaii. Vol. 4. Ho- moptera: Auchenorrhyncha. University of Hawaii Press, Honolulu. 268 pp. COLLADONUS AND RELATED GENERA OF MEXICO AND CENTRAL AMERICA WITH NEW TAXA AND SYNONYMY (HOMOPTERA: CICADELLIDAE) M. W. Nk'lson' Abstract. — A study of the genus CoUadonus and related genera of Mexico and C^entral Ameriea was based almost exclusively upon the examination of nearly the entire type series of all the Idiudomis species described by the late Dr. D. M. DeLong. All species referable to CoUadonus are redescribed, and illustrations of the male genitalia are given for the first time. A key to the species of Mexico and Central America is also included. A brief discussion of the distribution and phylogeny of the group is given. Among 35 species described by DeLong in Idiodonus , 21 belong in CoUadonus, 1 is assigned to OUarianus Ball, 1 to Paratanus Young, 2 to Bonncyana Oman, 2 are retained in Idiodonus , 5 are relegated to four proposed new genera, and the remaining 3 have uncertain generic position. One new combination in CoUadonus is proposed, and 16 names are treated as new synonyms. Eight new combinations are proposed in the treatment of related genera. New genera include: ParacoUadonus, Paracrassana, Paranurenus, and Jaacunga. The subgenus Angulanus DeLong is elevated to generic rank. The genus CoUadonus Ball was first revised by Nielson (1957), who treated primarily the Nearctic species north of Mexico and "one known single Palearctic species. After the ma- jor part of the revisionary work was finished, a number of Mexican leafhoppers described by DeLong (1946) in the genus Idiodonus Ball were studied, and several species were found referable to CoUadonus . Twelve species were listed as new combinations and treated as in- certae sedis in the appendix of my 1957 paper. Two species-level names were suppressed as new synonyms of an older name. Time did not permit a more thorough study of suspected additional synonymy and possible dissociation of the sexes between the female holotype and male allotype of several species in this group. The descriptions of nearly all of De Long's Idiodonus species were based on females, and a female specimen was designated as the holo- type in nearly all cases. Illustrations were lim- ited to a dorsal view of the head and thorax, and a ventral view of the female seventh ster- num, which with color patterns formed the basis of his classification. In a much later pa- per, DeLong (1983) described additional spe- cies of Idiodonus from Mexico and Bolivia; then he (DeLong 1984) prepared a revised key to all the known species in the New World, separating them on the basis of color patterns and configuration of the female sev- enth sternum. Although the female seventh sternum alone is a useful character for separating some spe- cies o( CoUadonus, its utility is enhanced only in combination with male genital characters. This is particularly true in certain species that show similarities in male characters that re- quire support by associated female seventh sternal characters to differentiate these spe- cies. In this study the entire type series, with the exception of a few specimens, of all species of Idiodonus described by DeLong were exam- ined with attention given to association of the female holotype and her counterjDart male al- lotype or male paratype specimen. A few cases of improper sex association were found, and these are discussed below. All species found referable to CoUadonus are re- described, and illustrations of the male geni- talia are given for the first time. A key to the species of Mexico and Central America is also included. Six new species of CoUadonus are described from that region. One new combi- nation is proposed, those treated as new com- binations in my earlier paper are confirmed, and 16 names are treated as new synonyms under their respective nominate species-level name. CoUadonus delongi described by Lin- navuori (1959) from Panama and based on an apparent mislabeled and misdetermined fe- male specimen was treated as a new synonym of CoUadonus montanus (Van Duzee) by Niel- son (1966). 'Monte L. Bean Life Science Museum, Brigham Young University, Provo, Utah 84602. 103 104 Great Basin Naturalist Memoirs No. 12 Among 35 species described in Idiodonus by DeLong, 21 belong in CoUadonus (includ- ing 14 species treated in my earlier work), 1 is assigned to Ollarianus Ball, 1 to Paratamis Young, 2 to Bonneyana Oman, 2 are retained in Idiodonus, 5 are relegated to four proposed new genera, and the remaining 3 have uncer- tain generic position. One species, /. plum- meri DeLong, is considered the only valid and correctly placed species among all those de- scribed in Idiodonus by DeLong. Idiodonus bakeri DeLong is suppressed as a new syn- onym of plummeri. DeLong treated 6 older species of Idiodonus in his 1946 paper, 5 of which were subsequently assigned to three other genera by Oman (1949). Descriptions of four new genera, ParacoUadonus, Paracras- sana, Paranurenus, and Jaacunga, are pro- vided with redescriptions and illustrations of the species assigned to their respective genus. The subgenus Angulanus DeLong is elevated to generic rank. Distribution of Colladonus . — The origin of the genus appears to have been centered in the southern highland range of the Nearctic region (Rocky Mountain subregion) in Mexico where radiation northward produced the bulk of species in United States. Only one species is known to occur as far south as Panama, and one is widespread in the Palearctic region. Fifteen species occur in Mexico proper, only one of which ranges into Arizona. Among 44 species in United States (including Alaska) and Canada, seven occur marginally in Mex- ico (Bliven 1954, Nielson 1962, 1966). Most of the species of CoUadonus occur on trees and shrubs in the mountainous regions of western United States and Canada. Genera closely allied to CoUadonus {Nigri- donus Oman, Caladonus Oman, Bonneyana Oman, and two new genera) are known only from Mexico and the extreme southwestern mountain areas of the United States, lending support to the Mexican origin of the genus CoUadonus. With respect to the New World distribution of Idiodonus, it is doubtful that the genus occurs south of Mexico. Members of the genus are more common in the United States than in Mexico. Considerations of phyloceny— Nearly all female CoUadonus of species that occur north of Mexico have a spatulate process on the caudal margin of the seventh sternum. None of the species in Mexico and Central America possess this structure, suggesting that it is a derived (apomorphic) character. The spatulate process thus far has attained its highest development in C. furculatus (Os- born) that occurs in the eastern United States and Canada. There does not appear to be any correlation between the presence of this structure and male genital structures, indicat- ing that its development was independent of the evolutionary development in male struc- tures. However, there appears to be a correla- tion between the absence of the spatulate pro- cess on the female seventh sternum and a short, male pygofer spine among the species (United States, Canada, and Palearctic re- gion) that possess these characters, viz., beUi, youngi, hrunneus, and torneeUus. This rela- tionship does not hold true for species in Mex- ico and Central America, suggesting that the northern population evolved separately from its sister population in the United States and Canada that does not exhibit this relationship. About half of the CoUadonus species have crowns that are produced, which is consid- ered a more primitive condition. Associated with this character is the relative position of the gonopore on the shaft of the aedeagus. A gonopore basad of the midlength of the aedea- gal shaft is considered more primitive, where- as a position distad of middle is more ad- vanced. About 82% of the United States and Canadian species bearing a basal gonopore have a produced crown, whereas all of those species bearing a distal gonopore have a short, rounded crown. Among the Mexican and Central American species, nearly all species with a basal gonopore have a produced crown, whereas all species with a distal gonopore have a short, rounded crown. Each of those sister groups apparently evolved indepen- dently. Key to Males oi Colladonus of Mexico and Central America 1. Pronotum with yellow or ivory, (sometimes pale) transverse band 2 — Pronotum without such band 13 2(1). Pygofer with a ver\- short, exposed spine on caudal margin, exposed length about twice as great as exposed basal width 3 — Pygofer with moderately long to very long, exposed spine on caudal margin, exposed length nearly three times to more than five times as great as exposed basal width 8 1988 NiELSON; COLLADONUS OF MEXICO AND CENTRAL AMERICA 105 14(13). 15(14). 3(2). Pygofer spine near middle of caudal margin; connective short, not reaching to apex of style; aedeagus with bifurcated processes long, reaching to midlength of shaft 4 — Pygofer spine distinctly hasad of middle of caudal margin; connective long, reaching to apex of style; aedeagus with bifurcated pro- cesses short, not reaching to midlength of shaft albocinctus (DeLong) 13(1). 4(3). Pygofer spine near middle or just below mid- dle of caudal margin 5 — Pygofer spine distinctly above middle of can- dal margin belli (Uhler) 5(4). Pygofer with caudoventral margin smooth; style with apical stylar spine 6 — Pygofer with caudoventral margin toothed or serrate; style with subapical stylar spine .... serratus, n. sp. 6(5). Aedeagus with bifurcated processes long, ex- tending beyond gonopore in lateral view .... 7 — Aedeagus with bifurcated processes short, not reaching to gonopore in lateral view fasciaticollis (Stal) 7(6). Style with apical shaft broad and lateral mar- gins parallel, stylar spine apical and project- ing laterad ultimus, n. sp. — Style with apical shaft narrowed distally, lat- eral margins not parallel, stylar spine apical and projecting laterodistad . . . tolucensis, n. sp. 8(2). Pygofer spine with exposed length not more than 4 times exposed basal width 9 — Pygofer spine with exposed length more than 5 times exposed basal width 11 9(8). Connective short, not reaching to apex of style; aedeagus in lateral view with bifurcated process short, reaching to about midlength of shaft 10 — Connective long, reaching to about apex of style; aedeagus in lateral view with bifurcated processes long, extending beyond midlength of shaft dampfi (DeLong) 10(9). Pronotum with narrow, black band on ante- rior margin and narrow, black, transverse band on middle; style with lateral margins of distal shaft smooth bicinctus (DeLong) — Pronotum with narrow, black band on ante- rior margin but without medial, black, trans- verse band; style with inner lateral margin of distal shaft toothed claustriis (DeLong) 11(8). Pygofer with caudoventral margin and pygofer spine smooth, without serrations or teeth 12 — Pygofer with caudoventral margin serrate or toothed, pygofer spine toothed basally verecundus (DeLong) 12(11). Pygofer with spine arising about middle of nearly truncate caudal margin; connective very long, extending beyond apex of style; aedeagus with bifurcated processes short, 16(14) nearly reaching to midlength of shaft titulus (DeLong) Pygofer with spine arising from apex of dis- tally produced caudoventral margin; connec- tive short, extending nearly to apex of style; aedeagus with bifurcated processes long, ex- tending beyond midlength ol shaft sirifiularius, n. sp. Pronotum never with yellow, longitudinal band; claval veins not deeply marked with yellow, if yellow, very pale or ivory 14 Pronotum with yellow, lontiituclinal band, claval veins deeply marked with yellow .... trabilis, n. sp. Pygofer spine very long, exposed length more than 5 times exposed basal width 15 Pygofer spine short, exposed length not more than 3 times exposed basal width 16 Crown with two handlebar-shaped markings on anterior margin; aedeagus with very long, reflexed, bifurcated processes in lateral view, processes parallel in dorsal view clathrus (DeLong) Crown with as many as 8 black spots on ante- rior margin, some spots sometimes con- nected, if so, never handlebar shaped; aedea- gus with short bifurcated processes nearly reaching midlength of shaft in lateral view, processes crossing over in dorsal view beameri (Ball) Pygofer with caudal margin smooth 17 Pygofer with caudal margin serrate or toothed 18 17(16). 18(16). Pygofer with very short spine arising from caudoventral margin; aedeagus with gono- pore at about middle of shaft; connective long, reaching to about apex of style albocinctus (DeLong) Pygofer with moderately long spine arising from middle of caudal margin; aedeagus with gonopore basad of middle shaft; connective short, not reaching apex of style tessellatus. n. sp. Pygofer with caudoventral margin produced distally, small, robust spine arising from cau- dodorsal angle, projecting dorsally; style with distal shaft long and narrow, serrate on inner lateral margin; aedeagus with gonopore at middle of shaft. . anademus (DeLong) Pygofer with obtusely angled caudal margin, very small spine arising from apex of angle and projecting distad; style with distal shaft robust and short, smooth on inner lateral margin; aedeagus with gonopore basad of middle of shaft incidus (DeLong) Colladonus albocinctus (DeLong) Figs. 1-5 Idiodonus albocinctus DeLong 1946;22 [Holotype 9 (OSU) (examined).] Colladonus albocinctus: Nielson 1957:51 106 Great Basin Naturalist Memoirs No. 12 Figs. 1-5. Colladonus albocinctiis (DeLong): 1, male pygofer, lateral view; 2, connective and right style, dorsal view; 3, aedeagus, lateral view; 4, apex of right style, enlarged dorsal view; 5, apex of aedeagus and distal processes, enlarged dorsal view. Idiodonus albocinctus: Metcalf 1967:1286 Idiodonus albocinctus: DeLong 1984:10 Idiodonus nigridens DeLong 1946:29 [Holotype 9 (OSU) (examined).] New synonymy Colladonus nigridens: Nielson 1957:51 Idiodonus nigridens: Metcalf 1967:1301 Idiodonus nigridens: DeLong 1984:10 Idiodonus sexpunctatus DeLong 1983:90 [Holotype 9 (OSU) (examined).] New synonymy Idiodonus sexpunctatus: DeLong 1984:10 Length. — IVIale, 4.9-5.0 mm; female, 5.0- 5.3 mm. Color variable from absence to presence of narrow, ivory, transverse band about middle of pronotum, with two black spots on anterior margin of crown and absence or presence of narrow, black, transverse band on disc of crown. Similar in general habitus to verecun- dus (DeLong) and in male genitalic characters to fasciaticoUis (Stal). Head with anterior margin produced, apex obtusely angulate. Male. — Pygofer in lateral view with very short, exposed caudoventral spine arising be- low middle of caudal margin, basal nine- tenths of spine sclerotized and fused to pygofer wall (Fig. 1); aedeagus in lateral view with bifurcated distal processes less than half as long as aedeagal shiift, not reaching gonopore (Fig. 3), crossing over in dorsal view (Fig. 5), gonopore at about midlength of shaft; 1988 NiELSON: COLLADONUS OF MEXICO AND CENTRAL AMERICA 107 connective long, extending to about apex of style (Fig. 2); style with distal shaft broad, lateral margins smooth, stylar spine broad and bluntly pointed, projecting laterodistallv (Fig. 4). Female. — Seventh sternum produced dis- tally to form broad medial lobe, with median, shallow notch on its caudal margin, configura- tion of lobe variable, sometimes with small lobe at base of notch. Distribution. — This species has been col- lected in the Mexican states of Distrito Fed- eral, Michoacan, Morelos, and Hidalgo. Host. — Some specimens were collected in pine forests at elevations of 7,500 feet from July to October (DeLong 1946). Remarks. — The configuration of the female seventh sterna of the holotypes of nigridens and sexpunctatiis is similar, with some intra- and interpopulation variation in the angle of the posteriolateral margins. The abdomen of the female holotype oi albocinctus is missing, but comparisons were made with paratype female specimens after they were associated with holotype and with DeLong's illustration of the seventh sternum. Although the male genital structures of the allotypes of albocinc- tus and nigridens are nearly identical, the latter is more deeply sclerotized than the for- mer. The sexes of both species appear to be properly associated. The female holotype of sexpunctatus is identical in general habitus with the female holotype oi nigridens, but both species differ markedly in color patterns from albocinctus. The latter species has a distinctive pronotal band that is absent in the specimens of the two former species. The male genital structures, however, are identical among these popula- tions, which suggests that two infraspecific entities may be involved. Such action is de- ferred until more material is collected and studied. Colladonus albocinctus is distinguished homfasciaticollis by the position of the pygo- fer spine, which is distinctly basad of the mid- dle of the caudal margin of the pygofer. Colladonus belli (Uhler) Figs. 6-10 Jassus belli Uhler 1877:471 [Holotype 9 (USNM) (exam- ined).] Thamnotettix belli: Van Duzee 1892:306 Colladonus belli: Ball 1936:58 Idiodonus belli: Medler 1943:18 Colladonus belli: Nielson 1957:38 Colladonus belli: Metcalf 1967: 1245 Thamnotettix semipullatus Van Duzee 1892:306 Colladonus semipullatus: Oman 1949:125 Thamnotettix fiilletti Van Duzee 1892:306 Colladonus gilk'tti: Oman 1949:125 Thamnotettix sonorae (;illette & Baker 1895:100 Colladonus sonorae: Oman 1949:125 Length. — Male, 4.0-4.4 mm; female, 4.4- 5.2 mm. A well-marked species from pale yellow to deep yellow with two black spots on anterior margin of crown and sometimes with a black, transverse band on disc of crown; pronotum with pale yellow to deep yellow, transverse band; forewings with veins pale to yellow; color intensity highly variable. Similar to fas- ciaticollis in general habitus but with distinc- tive male genitalia. Head with anterior margin not produced, apex rounded. Male. — Pygofer in lateral view with short, stout spine near or above middle of caudal margin, spine projecting dorsally (Fig. 6); aedeagus in lateral view with long, bifurcated distal processes extending beyond midlength of shaft (Fig. 8), crossing over in dorsal view (Fig. 10), gonopore about middle of shaft; con- nective short, not reaching apex of style (Fig. 7); style with moderately broad stylar shaft, inner lateral margin serrate, outer one smooth, stylar spine moderately long, pro- jecting laterally (Fig. 9). Female. — Seventh sternum with truncate caudal margin, shallow, U-shaped excision medially. Distribution. — This species is widespread in mountainous areas from Mexico north to British Columbia. In Mexico the author col- lected specimens in the state of Chihuahua. It has been reported in seven western states in the United States and in the province of British Columbia (Nielson 1957). Host. — Adults have been swept from a va- riety of trees and shrubs by the author. Sev- eral specimens were taken from Monarda sp. near citriodora Cerv. in Chihuahua state in Mexico by the author. Remarks. — Colladonus belli may repre- sent a species complex that occupies a rather narrow, longitudinal range in western North America. Specimens north of Utah and Colorado are almost uniformly yellow-green without a dark band across the crown, whereas southern populations are more 108 Great Basin Naturalist Memoirs No. 12 Figs. 6-10. CoUadomts belli (Uhler): 6, male pygofer, lateral view; 7, connective and right style, dorsal view; 8, aedeagus, lateral view; 9, apex of right style, enlarged dorsal view; 10, apex of aedeagus and distal processes, dorsal view. deeply marked with yellow and black mark- ings. The position of the pygofer spine, al- though similar in configuration, varies from distad of the middle of the caudal margin in northern populations to the middle of the cau- dal margin in southern populations. No varia- tion in the configuration of the female seventh sternum was evident, however. Much additional collecting over the entire range of this species complex is needed. Pop- ulations are not commonly collected, and when found, specimens are usually few in number. A reexamination of the type speci- mens of belli, semipullatus, ^illettei, and sonorae is recommended for any future study. From fasciaticollis, to which it is similar, belli can be distinguished by the short pygofer species has been made, nor has any type of die to upper middle portion of the caudal margin. Colladonus serratiis, n. sp. Figs. 11-14 Length. — Male, 4.8-5.0 mm; female, 5.4 mm. Color dark brown to blackish; crown yellow with two large, (juadrate, black spots on ante- rior margin, disc with broad, black, transverse band; pronotum with yellow, narrow to broad, transverse band; forewings black ex- cept for transculent area along middle of costa, veins yellow; face yellow with black markings. Head with anterior margin produced, apex obtusely angled. Male. — Pygofer in lateral view with broadly roimded caudal margin, caudoventral 1988 NiELSON: COLLADONUS OF MEXICO AND CENTRAL AMERICA 109 Figs. 11-14. CoUadonus serratus, n. sp.: 11, male pygofer, lateral view; 12, connective, right style, and aedeagus, dorsal view; 13, aedeagus, lateral view; 14, apex of right style, enlarged dorsal view. margin toothed or serrate, short spine above middle of caudal margin, directed dorsad (Fig. 11); aedeagus in lateral view narrow along major portion of shaft with moderately long, distal, bifurcated processes reaching to about middle of shaft (Fig. 13), processes broad medially, gradually tapered distally, crossing over in dorsal view (Fig. 12), gono- pore basad of middle of shaft; connective short, not reaching apex of style (Fig. 12); style with broad stylar shaft, stylar spine sub- apical, small, sharply pointed (Fig. 14). Female. — Seventh sternum with truncate caudal margin, small tooth medially. HoLOTYPE (male). — Mexico: Mex., 7 miles east of Amecameca, 9,300 ft, 31. VI. 1974, C. W. and L. O'Brien and Marshal (USNM). Allotype female, same locality as holotype, 10,000 ft, 17. VIII. 1982, C. W. and L. O'Brien and G. Wibner (USNM). Paratypes: one male, same data as female allotype (in au- thor's collection); one male, Mexico, #16, 7 miles southeast of Amecameca, 9,000 ft, 17. VIII. 1969, George W. Byers (UK). Remarks. — ^This is a rather unique species that is not related to other species oi CoUa- donus in Mexico. The combination of the sub- apical stylar spine and toothed caudovcntral margin of the pygofer will readily distinguish it from all other known species in the region. CoUadonus fasciaticollis (Stal) Figs. 15-22 J assiis fasciaticollis Stal 1864:86 [Holotype 9 (NM) (ex- amined).] Thamnotettix fasciaticollis: DeLong 1946:13 CoUadonus fasciaticollis: Nielson 1957:37 Thamnotettix fasciaticollis: Metcalf 1967:756 Idiodonus tubtdus DeLong 1946:22 [Holotype 9 (OSU) (examined).] CoUadonus tubulus: Nielson 1957:37 Idiodonus tubulus: Metcalf 1967:1303 Idiodonus tubulus: DeLong 1984:10 Idiodonus diserus DeLong 1946:24 [Holotype 9 (OSU) (examined).] CoUadonus diserus: Nielson 1957:37 Idiodonus diserus: Metcalf 1967:1297 Idiodonus diserus: DeLong 1984:11 Idiodonus pravus DeLong 1946:24 [Holotype 9 (OSU) (examined).] New synonymy Idiodonus pravus: Metcalf 1967:1301 Idiodonus pravus: DeLong 1984:11 Idiodonus dampfi DeLong 1946:20 [In part, allotype 6 and 8 paratypes (OSU)] Idiodonus dampfi: Linnavuori 1959:280 [Misdetermined specimens.] no Great Basin Naturalist Memoirs No. 12 Figs. 15-22. Colladonus fasciaticollis (Stal): 15, male pygofer of specimen from Costa Rica, lateral view; 16, male pygofer of allotype oi tubulus , lateral view; 17, male pygofer of allotype oi disertis, lateral view; 18, male pygofer of allotype of dampfi (= fasciaticollis), lateral view; 19, connective and right style, dorsal view; 20, aedeagus, lateral view; 21, apex of right style, enlarged dorsal view; 22, apex of aedeagus and distal processes, enlarged dorsal view. Length. — Male, 4.4-4.6 mm; female, 4.8-5.0 mm. A well-marked black and yellow species usually, but sometimes paler in some speci- mens. Similar in general habitus to belli and to albocinctus in male genitalic characters but distinct from both species. Head with anterior margin not produced, apex nearly rounded. Male. — Pygofer in lateral view with short, bluntly pointed spine arising below middle of caudal margin (some variation in origin of spine on caudoventral margin), basal 3/4 of spine sclerotized and fused to pygofer wall (Figs. 15, 16, 17, 18); aedeagus in lateral view with moderately long and narrow (sometimes broad), distal, bifurcated processes extending to about middle of shaft (Fig. 20), crossing over in dorsal view (Fig. 22), gonopore at about middle of shaft; connective short, apex not reaching to apex of style (Fig. 19); style with tapered stylar shaft, lateral margins smooth, stylar spine slanted laterodistally (Fig. 22). Female. — Seventh sternum with caudal margin obtusely angled with shallow, U- shaped, median emargination. Distribution. — Colladonus fasciaticollis is widely distributed from southern California to Costa Rica. Its northern range into the United States is restricted to the southern border of California. It is common in southern Mexico in the states of Distrito Federal, Veracruz, Michoacan, and Puebla. Host. — This species presumably occurs on unspecified trees or shrubs in Mexico from September to November at elevations from 1988 NiELSON:CoLLADONus OF Mexico AND Central America 111 5,000 to 10,000 feet (DeLong 1946). In Costa Rica it was taken on Lippia herlandieri in February (Nielson 1957). Remarks. — In my earlier work (Nielson 1957), tubulus and diserus were transferred to Colladonus and suppressed as synonyms of fasciaticollis . Reexamination of the female holotypes and comparison of them with au- thentically determined female specimens of fasciaticollis confirm that action. The female holotype of pravus is identical to its counteipart holotype specimens of the above species. The male allotype of dampfi from Bella Vista, Mexico (MB 366), which apparently had been misidentified and im- properly associated with the female holotype, was associated with the male holotype of pravus from Zitacuaro, Michoacan. Two of five male paratype specimens and three fe- male paratype specimens of dampfi from Orizaba, Veracruz, were also associated with pravus. From albocinctus and belli, to which it has similar genitalia, fasciaticollis can be distin- guished by the pygofer spine that arises from near the middle of the caudal margin, the shorter aedeagus, and the stouter, bifurcated distal processes. Colladonus idtimus, n. sp. Figs. 23-26 Length. — Male, 3.9-4.2 mm; female, 4.7-4.9 mm. General color dark brown to black; crown yellow with two black spots on anterior mar- gin and narrow, black, transverse band be- tween eyes; pronotum black with narrow, yel- low, transverse band at middle; forewings black with yellow veins, costal area hyaline; face yellow with black markings. Similar in general habitus to dampfi (DeLong) but with distinctive male genitalia. Head with anterior margin produced, apex rounded. Male. — Pygofer in lateral view with caudal margin broadly rounded, pygofer spine short, slightly longer than basal width, arising just below midline of pygofer and projecting cau- dodorsally (Fig. 23); aedeagus in lateral view with narrow shaft, distal, bifurcated processes long, more than 1/2 length of shaft and extend- ing beyond gonopore (Fig. 25), processes of uniform width except for tapered apex, gonopore at about midlength of shaft; connec- tive short, apex not reaching apex of style (Fig. 24); style with broad stylar shaft, nearly straight, stylar spine short, sharply pointed distally and projecting laterally (Fig. 26). Female. — Seventh sternum with caudal margin nearly truncate, with small indenta- tion medially. Holotype (male). — Panama: Chiriqui, Bambito, 1,400 m, 10. VI. 1976, Wolda and Estribi(USNM). Remarks. — Numerous specimens of this species that were collected in Costa Rica and preserved in alcohol were too badly distorted and discolored to be used in the type series. They were all taken at Cerro de la Muerte, 10,000 ft, I. VIII. 1966, S. L. Wood, and are in the M. L. Bean Museum collection. A single female specimen of this species from Yepo- capa, Guatemala, XII. 1948, H. T. Dalmat, was also examined. This species thus far represents the south- ernmost extremity of the range ofColladonus. It can be distinguished from dampfi by the shorter pygofer spine, by the gonopore, which lies midlength of the aedeagal shaft, and by the shorter connective. Colladonus tolucensis, n. sp. Figs. 27-30 Length. — Male, 4.5-4.6 mm; female, 5.0- 5.2 mm. Color tan with black markings; crown with two spots on anterior margin, black, trans- verse band on middle of crown; pronotum with broad, pale ivory, transverse band, sometimes with broken black stripe on either side of middle; forewings with pale veins, clavus sometimes dark brown, and with dark brown, longitudinal stripe above costal cell; face deep tan with black markings on clypeus. Similar in general habitus to several species possessing broad pronotal band and in male genital characters to idtimus. Head with anterior margin produced, apex obtusely angled. Male. — Pygofer in lateral view with broad, obtusely rounded caudal margin, very short spine arising from middle of margin and some- times hidden from lateral view, projecting distally (Fig. 27); aedeagus in lateral view with narrow shaft, distal, bifurcated processes long, extending slightly beyond midlength of shaft (Fig. 29), processes broad medially, pointed distally, gonopore at about midlength 112 Great Basin Naturalist Memoirs No. 12 Figs. 23-26. CoUadonus ultimus, n. sp. : 23, male pygofer, lateral view; 24, connective, right style, and aedeagus, dorsal view; 25, aedeagus, lateral view; 26, apex of right style, enlarged dorsal view. of shaft; connective short, not reaching apex of style (Fig. 28); style with tapered stylar shaft, stylar spine small, directed laterodistallv (Fig. 30). Female. — Seventh sternum with caudal margin nearly truncate, with broad median excavation. HoLOTYPE (male). — Mexico: Distrito Fed- eral, Toluca road, 26. IX. 1945, DeLong, Hershberger, and Elliot (OSU), allotype fe- male, Mexico City, D.F., 13.IX.1939, D. M. DeLong (OSU). Paratypes: one male, same data as holotype (in author's collection); one female, same data as allotype (in author's col- lection); 29 females, same data as allotvpe (OSU). Remarks. — From ultimus, to which it is similar, tolucensis can be distinguished by the unexposed pygofer spine, by the tapered sty- lar shaft, and by the color patterns on the pronotum. CoUadonus dampfi (DeLong) Figs. 31-34 kliodoniis dampfi DeLong 1946:20 [Holotype 9 (OSU) (examined).] CoUadonus dampfi: Nielson 1957:51 Idiodonus dampfi: Metcalf 1967:1297 Idiodomis dampfi: DeLong 1984:11 Length. — Male, 3.9-4.0 mm; female, 4.5 mm. A well-marked, black and yellow species with two distinct, black spots on anterior mar- gin of crown, narrow, transverse, black band on disc; pronotum with broad, black band along anterior margin, broad, yellow, trans- verse band medially, narrow, black band on posterior margin; forewings with pale yellow to deep yellow veins, cells brown to black. Similar to other pronotal banded species and to bicinctus (DeLong) in male genital charac- ters. Head with anterior margin not produced, apex obtusely rounded. Male. — Pygofer in lateral view with mod- erately long, narrow spine arising from mid- dle of caudal margin and projecting dorsally 1988 NiELSON: COLLADONUS OF MEXICO AND CkNTHAI. AMERICA 113 Figs. 27-30. Colladonus tohicensis, n. sp. : 27, male pygofer, lateral view; 28, connective, right style, and aedeagus, dorsal view; 29, aedeagus, lateral view; 30, apex of right style, enlarged dorsal view. (Fig. 31); aedeagus with distal, bifurcated pro- cesses long, extending beyond midlength of shaft (Fig. 33), crossing over in dorsal view, gonopore basad of middle of shaft; connective long, extending to about apex of style (Fig. 32); style with narrow stylar shaft, inner lat- eral margin roughly dentate, stylar spine long and projecting laterally (Fig. 34). Female. — Seventh sternum with caudal margin narrowed distally, shallow, U-shaped indentation medially. Distribution. — Colladonus dampfi is known only from the Mexican states of Veracruz and Morelos. Host. — Recorded from unspecified trees and shrubs from 3,000 to 9,000 feet during September to November (DeLong 1946). Remarks. — The type series of dampfi is mixed. The female holotype and male allotype were not properly associated; the latter is identical to pravus, which is treated as a ju- nior synonym oi fasciaticollis in this paper. A male paratype oi dampfi from Cordoba, Vera- cruz (type locality), was associated with the female holotype of dampfi and used in illus- trating the male genitalia. Five additional male paratypes, three from Cuernavaca, Morelos, and two from unspecified localities (MB 95, Mexico, and MF 8523, Mexico) were identical to the male paratype specimen asso- ciated with the female holotype. The remain- ing specimens in the type series are discussed under fasciaticollis. Colladonus darnpfi can be distinguished from bicinctus by the short and more rounded anterior margin of the head, by the longer pygofer spine, and by the longer, distal, bifur- cated aedeagal processes. Colladonus bicinctus (DeLong) Figs. 35-39 I diodomis bicinctus DeLong 1946:18 [Uo\otype 9 (OSU) (examined).] Colladonus bicinctus: Nielson 1957:51 Idiodonus bicinctus: Metcalf 1967:1287 Idiodonus bicinctus: DeLong 1984:10 Length. — Male, 4.2 mm; female, 4.5-4.8 mm. A well-marked species of deep yellow with black markings; crown with two triangular, 114 Great Basin Naturalist Memoirs No. 12 Figs. 31-34. CoUadonus dampfi (DeLong): 31, male pygofer, lateral view; 32, connective, right style, and aedeagus, dorsal view; 33, aedeagus, lateral view; 34, apex of right style, enlarged dorsal view. black spots on anterior margin and narrow, black, transverse band on middle of disc; pronotum with narrow, black band along anterior margin and narrow, black, transverse band on middle transecting broad, yellow, transverse band; forewings with veins yellow, cells black. Similar in general habitus and male genitalic characters to claustrus (De- Long). Head with anterior margin produced, apex angled. Male. — Pygofer in lateral view with mod- erately long spine on middle of caudal margin, spine directed caudodorsally (Fig. 35); aedea- gus in lateral view with stout, distal, bifur- cated processes reaching to about midlength of shaft (Fig. 37), not crossing over in dorsal view (Fig. 39), gonopore slightly basad of mid- length of shaft; connective short, not reaching apex of style (Fig. 36); style with narrow stylar shaft, lateral margins smooth, stylar spine very short and bluntly pointed (Fig. 38). Female — Seventh sternum with caudal margin shallowly and broadly concave, small indentation medially. Distribution — This species is known only from the Mexican state of Distrito Federal. Host. — According to DeLong (1946) the species was collected at 7,500 feet from un- specified shrubs or trees during a June- September period. Remarks, — ^This species was described from the female holotype and a female para- type specimen. However, a male paratype specimen (MB 352, Mexico, A. Dampf, coll.), which was without a name label and not cited in the original description, was found among the type series. The specimen was associated with the female holotype and used in illustrat- ing the genitalia. The species can be separated from claus- trus by the presence of a narrow, transverse, black band on the middle of the pronotum, by the narrower pygofer spine, by the shorter 1988 NiELSON: COLLADONUS OF MEXICO AND CENTRAL AMERICA 115 Figs. 35-39. Colladonus bicinctus (DeLong); 35, male pygofer, lateral view; 36, connective and right style, dorsal view; 37, aedeagus, lateral view; 38, ape,\ of right style, enlarged dorsal view; 39, apex of aedeagus and distal processes, enlarged dorsal view. connective, and by the smooth-margined sty- lar shaft. Colladonus claustrus (DeLong) Figs. 40-44 Idiodonus claustrus DeLong 1946:18 [Holotype 9 (OSU) (examined).] Colladonus claustrus: Nielson 1957:51 Idiodonus claustrus: Metcalf 1967:1289 Idiodonus claustrus: DeLong 1984:11 Length. — Male, 4.4 mm; female, 4.8-5.0 mm. A well-marked species with color patterns nearly identical to bicinctus except paler and without median, transverse, black band on pronotum. Similar in male genital characters to bicinctus. Head with anterior margin produced, apex angled. Male. — Pygofer in lateral aspect with mod- erately long, stout spine on middle of caudal margin, spine directed caudodorsally (Fig. 40): aedeagus with slender, bifurcated, distal processes reaching to about middle of shaft, apices not reaching gonopore (Fig. 42), cross- ing over in dorsal view (Fig. 44), gonopore basad of midlength of shaft; connective short, not reaching apex of style (Fig. 41); style with inner lateral margin serrate, outer one smooth, stylar spine short, bluntly pointed and directed laterad (Fig. 43). Female. — Seventh sternum with caudal margin shallowly and broadly concave, small 116 Great Basin Naturalist Memoirs No. 12 Figs. 40-44. Colladonus claustrus (DeLong); 40, male pygofer, lateral view; 41, connective and right style, dorsal view; 42, aedeagus, lateral view; 43, apex of right style, enlarged dorsal view; 44, apex of aedeagus and distal processes, enlarged dorsal view. indentation medially. Distribution. — This species is known only from the Mexican states of Veracruz and Chia- pas. Host. — Specimens were collected on un- specified shrubs or trees at elevations be- tween 4,000 and 7,000 feet from August to October (DeLong 1946). Remarks. — The female holotype and male allotype, although from widely separate local- ities, appear to be properly associated. From bicinctus, to which it is closely related, claus- trus can be separated by the absence of the narrow, transverse, black band on the middle of the pronotum, by the stouter pygofer spine, by the serrated, inner lateral margin of the stylar shaft, and by the more basad position of the gonopore. Colladonus verecundus (DeLong) Figs. 4.5-50 Idiodonus verecundus DeLong 1946:20 [Holotype ? (OSU) (examined).] Colladonus verecundus: Nielson 1957:51 Idiodonus verecundus: Metcalf 1967:1304 1988 NiELSON: COLLADONUS OF MEXICO AND CENTRAL AMERICA 117 Figs. 45-50. Colladonus verecundus (DeLong); 45, male pygofer (allotype oi vereciindus), lateral view; 46, male pygofer (allotype of acus), lateral view; 47, connective and right style, dorsal view; 48, aedeagus, lateral view; 49, apex of right style, enlarged dorsal view; 50, apex of aedeagus and distal processes, enlarged dorsal view. Idiodonus verecundus: DeLong 1984:11 Idiodonus acus DeLong 1946:20 [Holotype 9 (OSU) (ex- amined).] New synonymy Colladonus acus: Nielson 1957:51 Idiodonus acus: Metcalf 1967:1286 Idiodonus acus: DeLong 1984:11 Idiodonus mexicanus: DeLong 1946:27 [Holotype 9 (OSU) (examined).] New synonymy Idiodonus mexicanus: Metcalf 1967:1300 Idiodonus mexicanus: DeLong 1984:11 Length. — Male, 4.5-4.6 mm; female, 4.9- 5.0 mm. A well-marked species with two distinctive, black spots on anterior margin of crown and narrow to broad, ivory or yellow, transverse band along middle of pronotum. Similar to other pronotal banded species, but with unique pygofer spine. Head with anterior margin produced, apex angled. Male. — Pygofer in lateral view with long, dentate caudoventral spine, spine projecting caudodorsally with its exposed distal part about 1/2 as long as its entire length from its basal origin (Figs. 45, 46); aedeagus in lateral view with bifurcated distal processes about 1/2 as long as shaft, apices reaching gonopore (Fig. 48), crossing over in dorsal view (Fig. 50), gonopore at midlength of shaft; connec- tive long, extending distally beyond apex of style (Fig. 47); style with stylar shaft serrate on inner lateral margin with short stylar spine projecting laterally (Fig. 49). Female. — Seventh sternum with caudal margin slightly rounded or angled on either side of median, shallow indentation; some- times with slight protrusion laterally next to indentation. Distribution. — This species is restricted to the Mexican states of Distrito Federal, Michoacan, and Morelos. 118 Great Basin Naturalist Memoirs No. 12 53 54 Figs. 51-55. CoUadonus tituhis (DeLong): 51, male pygofer, lateral view; 52, connective and right style, dorsal view; 53, aedeagus, lateral view; 54, apex of right style, enlarged dorsal view; 55, ape.x of aedeagus and distal processes, enlarged dorsal view. Host. — Specimens were collected on pine at elevations from 9,500 to 10,300 feet during September and October (DeLong 1946). Remarks. — The female seventh sterna of the holotypes of verecundus and mexicanus are identical in configuration. In acus the lat- eral margins are angled toward the middle. I consider these traits normal variation in the population. The sexes of each species oi vere- cundus and acus appear to be properly associ- ated, although there is some variation in color between the two species. The color patterns and configuration of the seventh sternum of the holotype oi mexicanus are similar to vere- cundus. The male genitalic features of the allotypes of acus and verecundus are identi- cal. This species can be distinguished from all members of the genus CoUadonus by its unique, very long, dentate pygofer spine. CoUadonus titulus (DeLong) Figs. 51-55 Idkxiomi.s titulu.s DeLong 1946:24 [Holotype S (OSU) (examined).] CoUadonus titulus: Nielson 1957:51 Idiodonus titulus: Metcalf 1967:1302 Idiodonus titulus: DeLong 1984:11 Idiodonus goodi: DeLong 1946:27 [Holotype 9 (OSU) 1988 NiELSON: COLLADONUS OF MEXICO AND CENTRAL AMERICA 119 (examined).] New synonymy Idiodonus '^oodi: Metcalf 1967:1298 Idiodonus goodi: DeLong 1984:11 Length. — Male, 4.4-4.6 mm; female, 4.9-5.2 mm. General color light tannish to yellow with deep fuscous to black markings on crown, pronotum, and forewings; crown with two subtriangular spots on anterior margin; pronotum with anterior border black, broad, transverse, tannish band below; forewings with veins tannish to yellow, cells suffused with light fuscous to black. Similar in general habitus and certain male genital characters to claustrus, but more nearly related in male genital characters to singularius. Head with anterior margin not produced, apex slightly angled. Male. — Pygofer in lateral view with long spine arising about middle of caudal margin and directed nearly dorsad (Fig. 51); aedeagus in lateral view with short, rather stout, distal, bifurcated processes not reaching midlength of shaft (Fig. 53), crossing over in dorsal view (Fig. 55), gonopore slightly distad of middle of shaft; connective long, extending distad of apex of style (Fig. 52); style with narrow stylar shaft, stylar spine long, narrow, directed lat- erad (Fig. 54). Female. — Seventh sternum with caudal margin slightly produced along middle, with small excision at middle. Distribution. — This species is known only from the Mexican states of Distrito Federal and Veracruz. Host. — Reported from unspecified trees or shrubs above 7,500 feet during September and October by DeLong (1946). Remarks. — The male holotype and female allotype of titulus appear to be correctly asso- ciated. The general habitus and female sev- enth sternum of the holotype of goodi are identical to the female allotype of titulus. Males o( goodi are unknown. Colladonus titulus can be separated from claustrus by the much longer pygofer spine and from singularius by the shorter aedeagal processes, the more distal position of the gonopore, and the shorter stylar spine. Colladonus singularius, n. sp. Figs. 56-59 Length. — Male, 4.6 mm; female, 5.0-5.2 mm. Color tan to pale yellow with two small spots on anterior margin of crown, narrow, dark, transverse band on middle of crown in male, marking incomplete in female; prono- tum with broad, ivory, transverse band; forewings with veins pale yellow, cells translucent; face yellow with dark markings on clypeus. Similar to belli in general habitus but with distinctive male genitalia. Head with anterior margin not produced, apex rounded. Male. — Pygofer in lateral view with pro- duced caudoventral margin, pygofer spine very long, arising from apex of caudoventral margin and projecting caudodorsally (Fig. 56); aedeagus in lateral view with narrow shaft and long, distal, bifurcated processes extending beyond midlength of shaft (Fig. 58), crossing over in dorsal view (Fig. 57), gonopore basad of middle of shaft; connective moderately long, nearly reaching to apex of style (Fig. 57); style with narrow stylar shaft, curved laterally and with short, bluntly pointed stvlar spine (Fig. 59). Female. — Seventh sternum with truncate caudal margin with small, narrow, median ex- cision. HoLOT\'PE (male). — Me.xico: Guanajuato, 10 mi south San Luis de la Paz 24.X. 1981, M. W. Nielson (Nielson collection), female allo- type, same data as holotype (Nielson collec- tion). Paratype: seven females, same data as holotype (BYU, Nielson collection). Remarks. — From belli, to which it is simi- lar in general habitus, singularius can be sepa- rated by the very long pygofer spine and the longer, aedeagal, bifurcated processes. Colladonus trabilis, n. sp. Figs. 60-63 Length. — Male, 4.8 mm. Color. — Yellow and black. Crown yellow with two black spots on anterior margin, broad, black marking on either side of middle on disc; pronotum black with broad, yellow, longitudinal band on middle and yellow mark- ings on lateral angles; scutellum black with large, yellow marking medially; forewings black with claval veins yellow except for black marking on basal part of posterior portion, remaining veins pale ivory, cells fuscous ex- cept in translucent costal area, first and sec- ond apical cells, and appendix; face black ex- cept yellow above. Similar to tessellatus. 120 Great Basin Naturalist Memoirs No. 12 Figs. 56-59. Colladonus singtilariiis, n. sp. : 56, male pygofer, lateral view; 57, connective, right style, and aedeagus, dorsal view; 58, aedeagus, lateral view; 59, ape.\ of right style, dorsal view. Head with anterior margin produced, apex rounded. Male. — Pygofer in lateral view with acutely angled caudal margin, pygofer spine moderately long and arising from apex of cau- dal margin, spine curved mesad and directed caudodorsad (Fig. 60); aedeagus in lateral view with short, distal, bifurcated processes, not reaching midlength of shaft (Fig. 62), pro- cesses crossing over in dorsal view (Fig. 61), gonopore at midlength of shaft; connective long, nearly reaching to apex of style (Fig. 61); style with broad stylar shaft, curved laterad, outer lateral margin serrate, stylar spine very small, blunt distally and projecting laterally (Fig. 63). Female. — Unknown. HoLOTYPE (male). — Mexico: D.F., De- sierto de los Leones, 9,600 ft, 6. VII. 1961, George W. Byers (KU). Remarks, — This unusual, marked species can be distinguished from tessellatus by the distinctive markings on the pronotum and clavus; the broad, curved styler shaft; the shorter, distal, bifurcated processes of the aedeagus; and the gonopore at midlength of the shaft. Colladonus clathrus (DeLong) Figs. 64-68 Idiodonus clathrus DeLong 1946:28 [Holotype $ (OSU) (examined).] Colladonus clathrus: Nielson 1957:51 Idiodonus clathrus: Metcalf 1967:1288 Idiodonus clathrus: DeLong 1984:11 Idiodonus turpiter DeLong 1946:28 [Holotype 9 (OSU) (examined). ] New synonymy Colladonus turpiter: Nielson 1957:51 Idiodonus turpiter: Metcalf 1967:1303 Idiodonus turpiter: DeLong 1984:10 Length — Male, 4.7-4.9 mm; female, 5.0-5.2 mm. General color tawny with fuscous mark- ings on disc of crown and forewings of mature specimens; crown with two distinctive, black, handlebar-shaped markings on anterior mar- gin; pronotum without transverse band. Simi- lar in general habitus to anademus (DeLong) 1988 NiELSON: COLLADONUS OF MEXICO AND CENTRAL AMERICA 121 62 Figs. 60-63. CoUadonus trabilis, n. sp.; 60, male pygofer, lateral view; 61, connective, right style, and aedeagus, dorsal view; 62, aedeagus, lateral view; 63, apex of right style, enlarged dorsal view. but with distinctive male genitalia. Head with anterior margin not produced, apex rounded. Male. — Pygofer in lateral view with long, robust spine arising from middle of caudal margin and projecting caudodorsally (Fig. 64); aedeagus in lateral view with distal, bifur- cated processes more than 1/2 half as long as shaft, reflexed at distal half (Fig. 66), pro- cesses parallel in dorsal view except for distal third (Fig. 68), gonopore distad of midlength of shaft; connective long, extending beyond apex of style (Fig. 65); style with narrow stylar shaft, inner lateral margin serrate, outer one smooth, stylar spine broad, projecting later- ally. Female. — Seventh sternum with broad, V- shaped, median emargination, caudal margin on either side convex. Distribution. — This species appears to be restricted to the Mexican state of Distrito Federal. Host. — Unknown. Remarks. — The female holotype and male allotype o{ clathrus appear to be properly as- sociated. The female holotype of turpiter is identical with the female holotype of clath- nis , based on similar color patterns (handle- bar-shaped markings on anterior margin of crown) and configuration of the seventh ster- num. Although DeLong did not designate an allotype of turpiter in his description of that species, a specimen so labeled was found and examined. This specimen could not be associ- ated with the female holotype o( turpiter but is identical in color patterns and male genitalia to the male allotype of nigridens, which is treated as a junior synonym of albocinctus in this paper. From anademus, to which it is similar in general habitus, clathrus can be separated 122 Great Basin Naturalist Memoirs No. 12 66 Figs. 64-68. CoUadonus clathrus (DeLong): 64, male pygofer, lateral view; 65, connective and right style, dorsal view; 66, aedeagus, lateral view; 67, apex of right style, enlarged dorsal view; 68, ape.x of aedeagus and distal processes, enlarged dorsal view. from all known Mexican and Central Ameri- can species by the reflexed, distal, bifurcated aedeagal processes. CoUadonus beameri (Ball) Figs. 69-75 Idiodonus beameri: Ball 1937:28 [Holotype ? (USNM) (examined).] Idiodonus beameri: DeLong 1946:17 CoUadonus beameri: Oman 1949:125 CoUadonus beameri: Nielson 1957:45 CoUadonus beameri: Metcalf 1967:1244 Idiodonus beamercUus: DeLong 1983:92 [Invalid re- placement name for nonexistent species, Id- iodonus beameri: DeLong 1946:17 nee Idiodonus foeamert Ball 1937:28.] Idiodonus beamereUus: DeLong 1984:10 Idiodonus nuirginatus: DeLong 1983:90 [Holotype 6 (OSU) (examined).] New synonymy Idiodonus marginatus: DeLong 1984:10 Length. — Male, 4.2-4.6 mm; female, 5.0-5.3 mm. General color tawny throughout with black spots along anterior margin of crown, spots sometimes numbering as many as eight, four on anterior margin and two each above and below margin next to eye, spots sometimes connected. Similar in general habitus to in- cidiis (DeLong) but larger and with additional spots on crown and distinctive male genitalia. Head with anterior margin not produced, apex rounded. Male. — Pygofer in lateral view with very 1988 NiELSON: COLLADONUS OF MEXICO AND CENTRAL AMERICA 123 73 72 Figs. 69-75. CoUadonus beameri (Ball): 69, male pygofer, lateral view; 70, male pygofer (holotype ofnuirginatus), lateral view; 71, connective and right style, dorsal view; 72, aedeagus, lateral view; 73, aedeagus (holotype of tnarginatus), lateral view; 74, apex of right style, enlarged dorsal view; 75, apex of aedeagus and distal processes, enlarged dorsal view. long spine, exposed length about as long as fused basal half (Figs. 69, 70); aedeagus in lateral view with short, bifurcated distal pro- cesses, processes reaching to gonopore (Fig. 72, 73), crossing over in dorsal view (Fig. 75), gonopore distad of middle of shaft; connective long, extending to or slightly beyond apex of style (Fig. 71); style with long stylar shaft, lateral margins smooth, stylar spine very short and projecting laterally (Fig. 74). Female. — Seventh sternum with subtrun- cate caudal margin, with shallow, V-shaped emargination medially. Distribution. — This is a widespread spe- cies known from the Mexican states of Mi- choacan, Distrito Federal, Puebla, Hidalgo, Veracruz, Jalisco, and Morelos, and from the state of Arizona. Host. — Specimens were collected from pine at elevations from 5,500 to 9,800 feet during October in Mexico (DeLong 1946) Remarks. — DeLong (1983) proposed heam- erellus as a replacement name for his sup- posed "beameri Delong" treated as beameri Ball in his 1946 paper. That action was actually based on a nonexistent species. The male holotype of marginatus is identi- cal to authentically determined males of beameri. From incidus, to which it is similar in general habitus, beameri can be distin- guished by the eight spots on the anterior margin of the crown and by the very long pygofer spine. 124 Great Basin Naturalist Memoirs No. 12 Figs. 76-79. Colladonus tessellatus , n. sp. : 76, male pygofer, lateral view; 77, connective, right style, and aedeagus, dorsal view; 78, aedeagus, lateral view; 79, apex of right style, enlarged dorsal view. Colladonus tessellatus, n. sp. Figs. 76-79 Length. — Male, 4.3 mm; female, 4.7-4.9 mm. Color tan to tannish brown with dark brown markings on dorsum; crown with two large, subquadrate spots on anterior margin, small spot in ocellocular area next to lateral frontal suture, two large spots on disc separated widely at middle; pronotum with dark, mosaic markings separated by narrow, tannish line medially; forewings with dark markings in cells, veins mostly tannish to yellow tannish; face tan with dark markings. Similar to trahilis in male genitalic characters. Head with anterior margin produced, ape.x rounded. Male. — Pygofer in lateral view with caudal margin obtusely angled, pygofer spine mod- erately long, arising at apex of caudal margin above middle and slightly curved dorsad (Fig. 76); aedeagus in lateral view narrow along major portion of shaft, with moderately long, distal, bifurcated process (one process broken off in holotype) (Fig. 78) reaching to about midlength of shaft, gonopore basad of mid- length of shaft; connective short, not reaching apex of style (Fig. 77); style with narrow stylar shaft, inner lateral margin serrate, slightly ta- pered distally and curved laterad, stylar spine very small (Fig. 79). Female. — Seventh sternum with truncate caudal margin, very small, shallow excavation medially. Holotype (male). — Mexico: Mex. D.F., 5 mi west of Tlamacas, P.N. Ixtapopo, 10,000 ft, 31. V. 1974, C. W. and L. O'Brien and Mar- shall (BYU), female allotype, same data as holotype (BYU). Paratypes: one male and one female, same data as holotype (author s collec- tion); three females, same data as holotype (BYU); one female, 11 mi east of Amecameca, 10,300 ft, 31. V. 1974, C. S. and L. O'Brien and Marshall; one female, P.N., Cumbres de 1988 NiELSON: COLLADONUS OF MF.XICX) AND CENTRAL AMERICA 125 Figs. 80-84. Colladonus anademus (DeLong): 82, male pygofer, lateral view; 81, connective and right style, dorsal view; 82, aedeagus, lateral view; 83, apex of right style, enlarged dorsal view; 84, apex of aedeagus and distal processes, enlarged dorsal view. Ajusco, Llano de Cantimplora, 3,340 m, 4.IX.1982, C. W. and L. B. O'Brien and G. J. Wibmer (CAS). Remarks. — From trabilis, to which it is similar in certain male genitalic features, tes- sellatus can be separated by the gonopore, which is basad of midlength of the aedeagal shaft, by the short connective, and by the slender stylar shaft, which is serrate on the inner lateral margin. Colladonus anademus (DeLong) Figs. 80-84 Idiodonus anademus DeLong 1946;29 [Holotype 9 (OSU) (examined).] Colladonus anademus: Nielson 1957:51 Idiodonus anademus: Metcalf 1967:1286 Idiodonus anademus: DeLong 1983:90 Idiodonus anademus: DeLong 1984:10 Length. — Male, 4.8 mm; female, 4.9-5.1 mm. General color tawny throughout with two distinct, black, elongate, triangular spots on anterior margin of crown, small black mark- ings on disc of crown and pronotum. Similar in general habitus to incidus (DeLong) but with distinctive male genitalia. Head with anterior margin not produced, apex rounded. Male, — Pygofer in lateral view with cau- doventral margin produced distally, subtrun- cate and serrate (Fig. 80), pygofer spine very short, basal 3/4 sclerotized and fused to inner mesal margin of pygofer, spine projecting dor- sad (Fig. 80); aedeagus in lateral view with slender, distal, bifurcated processes reaching to midlength of shaft (Fig. 82), crossing over in dorsal view (Fig. 84), gonopore at about middle of shaft; connective long, nearly reaching to apex of style (Fig. 81); style with long, narrow stylar shaft, inner lateral margin 126 Great Basin Naturalist Memoirs No. 12 89 Figs. 85-89. Colladonus incidus: 85, male pygofer, lateral view; 86, connective and right style, dorsal view; 87, aedeagus, lateral view; 88, apex of right style, enlarged lateral view; 89, apex of aedeagus and distal processes, enlarged dorsal view. serrate, outer one smooth, stylar spine short, blunt, and directed laterad. Female. — Seventh sternum with caudal margin convex, shallow, narrow, V-shaped excision medially. Host. — Specimens were collected on pine by DeLong (1946). Distribution. — This species is known only from the Mexican state of Distrito Federal. Remarks. — The female holotype and male allotype appear to be correctly associated. This species, similar to incidus, can be distin- guished by the produced, subtruncate cau- doventral margin of the pygofer, by the short, dorsally directed pygofer spine, and by the long, narrow stylar shaft. Colladonus incidus (DeLong), n. comb. Figs. 85-89 Idiodonus incidus DeLong 1946:29 [Holotype 9 (OSU) (examined).] Idiodonus iticidus: Metcalf 1967:1299 Idiodonus incidus DeLong 1984:11 Idiodonus pallidas: DeLong 1983:90 [Holotype 6 (OSU) (examined).] New synonymy Idiodonus pallidus: DeLong 1984:10 Length. — Male, 4.8 mm; female, 4.9-5.1 General color tawny throughout with two small, black spots on anterior margin of crown. Similar in general habitus to anade- ynus but with distinctive male genitalia. Head with anterior margin produced, apex angled. Male. — Pygofer in lateral view with broadly angled, serrate caudal margin, very small spine at middle of margin, spine di- rected distad (Fig. 85); aedeagus with slender, distal, bifurcated processes extending beyond midlength of shaft (Fig. 87), crossing over in 1988 NiELSON: COLLADONUS OF MEXICO AND CENTRAL AMERICA 127 dorsal view (Fig. 89), gonopore at midlength of shaft; connective short, not reaching to apex of style (Fig. 86); style with slender stylar shaft, lateral margins smooth, stylar spine long, directed laterad (Fig. 88). Female. — Seventh sternum with broadly convex caudal margin, very small indentation medially. Distribution — This species is known only from the Mexican state of Distrito Federal. Host. — Specimens were collected from unspecified trees or shrubs at 8,500 feet dur- ing September and October in Mexico (De- Long 1946). Remarks, — No males of incidus were de- scribed in the type series. The female holo- type of incidus was associated with the male holotype of paUidus and is identical with the female allotype ofpaUidiis . The latter name is the more recent and is, therefore, suppressed as a junior synonym oi incidus. From anademus, to which it is similar, in- cidus can be easily separated by the serrated, broadly angled caudal margin of the pygofer with its very small, distally directed spine, by the shorter connective, and by the longer, distal, bifurcated processes of the aedeagus. Paracolladonus , new genus Type-species : Idiodonus insculptus De- Long 1946:25. Similar to Colladonus Ball and Dolyobius Linnavuori in general habitus but with dis- tinctive male genitalia. General color tannish brown in female to black in male with dark markings on crown, pronotum, and scutel- lum. Head slightly wider than pronotum, ante- rior margin broadly rounded and slightly pro- duced medially; pronotum with posterior margin broadly concave; scutellum as in Col- ladonus ; clypeus broad anteriorly; venation of forewing similar to Colladonus, inner an- teapical open basally. Male pygofer with large spine on caudo- ventral margin, setose caudodorsally; aedea- gus long, broadly recurved, somewhat compressed dorsoventrally, with paired, re- curved, short, distal processes, processes sometimes with secondary process medially, gonopore subapical on dorsal surface; connec- tive long and narrow; style broad at basal half, with foot-shaped apex. Paracolladonus insculptus (DeLong), n. comb. Fi^s. 90-94 Idiodonus in.sctdptiis: DeLong 1946:25 [Holotype $ (OSU) (examined).] Idiodonus insculptus: Metealf 1967:1299 Idiodonus insculptus: DeLong 1984:11 Length. — Male, 5.3 mm; female, 5.7-5.9 mm. Long, slender species with dark markings on crown, pronotum, and scutellum; prono- tum black in male, light brown in female with dark markings on anterior margin; forewing light to dark brown with yellowish veins. Male pygofer with very long, stout spine on caudoventral margin, spine projecting cau- dodorsad, numerous setae confined to cau- dodorsal area (Fig. 90), aedeagus in lateral view long, recurved, with distal processes short, broad medially (Fig. 92), sometimes with secondary process medially (Fig. 94) (if absent, basal attachment apparent in dorsal view), curved dorsad in lateral view, slightly flared laterally in dorsal view (Fig. 93), shaft somewhat flattened dorsoventrally (Fig. 94), gonopore subapical on dorsal surface; connec- tive long and narrow, reaching to or beyond apex of style (Fig. 91); style with narrow ridge or flange from inner lateral margin above preapical lobe to apex of style in dorsal view (Fig. 92). Remarks. — ^This is the only known species assigned to the genus. It is known from the states of Distrito Federal and Morelos in Mex- ico. DeLong (1946) collected specimens from pine at 10,000 feet in September and Octo- ber. A pair of specimens were collected on the Cuernavaca road in Mexico by Ball and Stone. Paranurenus, new genus Type-species: Idiodonus latidens DeLong 1946:26 Similar to Colladonus in general habitus and Nurenus in some male genital characters. General color tawny with black markings on head. Pronotum with ivory, transverse band, fore wings with ivory stripe on claval suture. Head about as wide as pronotum; crown produced, longer medially than next to eye; pronotum and scutellum as in Colladonus; forewing as in Colladonus; venation partially obscured; clypeus long; clypellus long and narrow. 128 Great Basin Naturalist Memoirs No. 12 Figs. 90-94. Paracolladonufi insculptus (DeLong); 90, male pygofer, lateral view; 91, connective and right style, dorsal view; 92, aedeagus, lateral view (allotype); 93, aedeagus, enlarged dorsal view (allotype); 94, aedeagus, dorsal view (specimen from Cuernavaca Road, Mex.). Male pygofer without caudal spine, margin obliquely truncate as in Niirenus; aedeagus small, recurved, flattened dorsoventrally with pair of short, acuminate distal processes; gonopore subapical on dorsal surface; connec- tive and style as in Colladonus. Paranurenus is represented by a single known species from Mexico. From Colladonus, to which it is similar in general habitus, Paranurenus can easily be differentiated by the flattened aedeagus and by the absence of a pygofer spine. The head characters and the flattened shaft of the aedea- gus will differentiate the genus from Nurenus. Paranurenus latidens (DeLong), n. comb. Figs. 95-98 Idiodonus latidens: DeLong 1945:26 [Holotype 9 (OSU) (examined).] Idiodonus latidens: Metcalf 1967:1300 Idiodonus latidens DeLong 1984:11 Length. — Male, 4.4-4.5 mm; female, 4.8- 5.1 mm. Color tawny, crown ivory with two black spots on anterior margin and narrow, black, transverse band on disc between eyes; prono- tum tawny with narrow, ivory, transverse band above middle; scutellum tawny; fore- wing tawny with ivory on claval suture ex- tending to pronotal band, fuscous band above costal area; face ivory to tawny with black markings. Male. — Pygofer in lateral view with obliciuely truncate caudal margin, pygofer spine absent, macrosetae on caudodorsal quadrant (Fig. 95); aedeagus in lateral view recurved, socle large, shaft slender, flattened 1988 NiELSON; COLLADONUS OF MEXICO AND CeNTKAL AMERICA 129 95 97 98 Figs. 95-98. Paranurenus latidens (DeLong): 95, male pygofer, lateral view; 96, connective and right style, dorsal view; 97, aedeagus, lateral view; 98, aedeagus, ventral view. dorsoventrally (Fig. 97) with pair of short, acuminate distal processes, processes di- rected laterad in ventral view (Fig. 98); gono- pore subapical on dorsal surface of shaft; con- nective long and narrow, extending just be- yond apex of style (Fig. 96); style with stylar shaft short and enlarged distally, stylar spine prominent, subapical, and projecting laterally (Fig. 96). Female. — Seventh sternum with short, median, spatulate process on caudal margin. Remarks. — This species has no known close relatives. The styles are typical of many species oi Colladonus, and the aedeagus, ex- cept for its flattened condition, is similar in configuration to Nurenus. The head charac- ters and general habitus are similar to species oiColladonus. The species is known from the states of Michoacan and Distrito Federal in Mexico, and occur at elevations from 5,000 to 8,000 feet in February and September on unspeci- fied shrubs or trees (DeLong 1946). Jaacunga, new genus Type-species: Idiodonus vinculus DeLong 1945:15 Similar to Paracolladonus in general habitus but more robust with slightly wider head and distinctive male genitalia. General color tawny with markings on crown, narrow band on pronotum and iridescent forewings. Head wider than pronotum, anterior mar- gin broadly rounded and slightly produced as in Paracolladonus; pronotum large, posterior margin broadly and shallowly concave; fore- wing venation as in Paracolladonus. Male pygofer without caudal spine, setose submarginally; aedeagus stout with enlarged socle, recurved at apical third with terminal 130 Great Basin Naturalist Memoirs No. 12 Figs. 99-101. Jaacunga vinculus (DeLong): 99, male pygofer, lateral view; 100, connective, right style, and aedeagus, dorsal view; 101, aedeagus, lateral view. processes, gonopore large, subapical on dor- sal surface; connective Y-shaped; style with- out apical spine. Female seventh sternum with median spatulate process. Two species, described below, are assigned to the genus. Jaacunga vinculus (DeLong), n. comb. Figs. 99-101 Idiodonus vinculus: DeLong 1946:15 [Holotype 9 (OSU) (examined).] Idiodonus vinculus: Metcalf 1967:1304 Idiodonus vinculus DeLong, 1984:10 Length. — Male, 5.8 mm; female, 6.4 mm. Long, robust species with two spots on anterior margin of crown and narrow, black, transverse, irregular band on disc in male, less marked in female; pronotum with narrow, yellow or ivory, transverse band near middle and similar, less pronounced band on poste- rior margin; forewing iridescent, veins black to tawny. Similar to spatulatus in male genital character. Male. — Pygofer without caudal spine, macrosetae on submargin from dorsal to ven- tral margin (Fig. 99); aedeagus in lateral view stout, socle enlarged, shaft tubular, curved dorsad at apical half with six terminal spines, two long and projecting anteroventrally, two moderately long and projecting anterodorsad, and two very short and projecting anteriorly (Fig. 101), gonopore large, subapical on dorsal surface; connective stout, Y-shaped, extend- ing slightly beyond apex of style (Fig. 100); style similar to CoUadonus but broader at basal two-thirds, stylar apex narrow, without distinct stylar spine, apex obliquely truncate. Female. — Seventh sternum with short, apically bifid, medial projection on caudal margin, shallowly excavated on either side of projection. 1988 NielsonColladonus OF Mexico AND Central America 131 Figs. 102-104. jaacunga spatiilatiis (DeLong): 102, male pygofer, lateral view; 103, connective, right style, and aedeagus, dorsal view; 104, aedeagus, lateral view. Remarks. — This species is known from the female holotype and male allotype specimens collected at Jacala, Hidalgo, Mexico, and Aca- pulco Road, Mexico, respectively. Other specimens that I have not seen are reported from Mexico City and Jalapa Road in Mexico by DeLong (1946). All specimens were col- lected from September to October at eleva- tions from 5,000 to 8,500 feet from unspeci- fied shrubs or trees. Jaacunga vinculus can be distinguished from spatulatus by its six terminal spines, which differ in their configuration, and by the shape of the apex of the style. Jaacunga spatulatus (DeLong), n. comb. Figs. 102-104 Idiodonus spatulatus DeLong 1946:15 [Holotype 9 (OSU) (examined).] Idiodonus spatulatus: Metcalf 1967:1302 Idiodonus spatulatus DeLong 1984:10 Idiodonus ruhellus DeLong 1946:15 [Holotype 9 (OSU) (examined).] New synonymy Idiodonus ruhellus: Metcalf 1967:1301 Idiodonus ruhellus DeLong 1984:9 Length. — Male, 5.5 mm; female, 5.5-5.8 mm. Similar in general habitus to vinculus but with distinctive male genitalia. Male pygofer without caudal spine, mac- rosetae submarginal in caudodorsal area (Fig. 102); aedeagus in lateral view stout but not as robust as in vinculus, socle enlarged, shaft similar to vinculus, with four terminal pro- cesses, longest pair projecting ventrally and broad before gradually pointed apex, second pair shorter, projecting anteriorly with tips curved ventrally (Figs. 103, 104); gonopore large, subapical on dorsal surface; connective similar to vinculus except shorter, not reach- ing apex of style (Fig. 103); style as in vinculus except apex curved mesally and rounded dis- tally. 132 Great Basin Naturalist Memoirs No. 12 Remarks — Jaacunga spatulatus is known from the type series collected from Mexico City and from the female holotype o^nibeUus, which has the same type locality as spatulatus. The specimens were collected from unspeci- fied shrubs at elevations of 7,500 to 8,500 feet in September (DeLong 1946). The species can be distinguished from vinculus by the shape of four terminal spines on the aedeagus and by the shape of the stylar apex. Paracrassana, new genus Type-species: Idiodonus nigrifrons De- Long 1983:91 Near Crassana DeLong & Heishberger but with distinctive head, facial, and genitalic fea- tures. General color tawny with black mark- ings on head and thorax, face black. Head slightly wider than pronotum, ante- rior margin slightly angled and produced me- dially, not rounded to face, clearly demar- cated but not carinate, narrow, transverse impression submarginally between ocelli, surface shiny, striate longitudinally; ocellus remote from eye; forewing as in Crassana; clypeus broad anteriorly, about as wide as long; clypellus long, lateral margins expanded distally. Male pygofer long, with deep incision basally to near middle, dorsal margin broadly excavated near middle, row of fine teeth on ventral margin; aedeagus symmetrical, long, narrow, nearly tubelike, socle basal, gono- pore terminal, sclerotized endotheca basad of aedeagal shaft; connective very short as in Crassana and Neocrassana Linnavuori; plate long, triangular, tenth segment sclerotized laterally. Paracrassana is represented by a single known species from Mexico. It can be distin- guished from Crassana by the narrow demar- cation between the crown and face, the nar- row impression submarginally on the crown, the long pygofer with basal incision and tubu- lar aedeagus. From Neocrassana, it can be distinguished by the remote position of the ocelli from eyes, the position of the socle on the aedeagus, and characters described above. Paracrassana nigrifrons (DeLong), n. comb. Figs. 105-107 Idiodonus nigrifrons DeLong 1983;91 [Holotype 6 (OSU) (examined).] Length. — Male, 5.4 mm. Long, robust species, crown with irregular, narrow, black, transverse band between ocelli; pronotum with black markings on ante- rior lateral angles; face black except for nar- row, yellow anterior margin; forewing iri- descent, veins fuscous. Male pygofer in lateral view long and nar- row, macrosetae on distal third except caudal margin, ventral margin with fine spines (Fig. 105); aedeagus long, tubular, curved dorsally at distal third, with subdistal flange laterally on dorsal margin (Fig. 106), gonopore apical, endotheca not forming a loop; connective short, broad; style narrow, with narrowed curved apices. Female. — Unknown. Remarks. — This species is known only from the male holotype from Huanchinango, Pue- bla, Mexico. The host is unknown. The nomenclature of a few species de- scribed in Idiodonus and Colladonus are treated below: Bonneyana caldwelli (DeLong), n. comb. /(/io(/onusca/duJe//i DeLong 1946:16 [Holotype 9 (OSU) (examined).] Idiodonus caldwelli: Metcalf 1967:1288 Idiodonus caldwelli DeLong, 1984:10 Idiodonus apertus DeLong 1946:14 [Holotype 9 (OSU) (examined).] New synonymy. Idiodonus apertus: Metcalf 1967:1287 Idiodonus apertus Delong 1984:10 I have compared dissected specimens of caldwelli and schwartzi (Ball) and found them to be distinct. Two remaining known species in the genus, osborni (Ball) and terminalis (Ball), have not been examined; but, based on the original descriptions, they appear to be distinct from caldwelli. Further study of the group is warranted, however, including illus- trations and descriptions of the male diagnos- tic characters. Ollarianus tripartitus DeLong Ollarianus tripartitus DeLong 1944:398 /r/tO(/oni/sfl/fo(frons DeLong 1983:89 [Holotype 9 (OSU) (examined).] New synonymy Idiodonus alhifrons DeLong 1984:10 Examination of the female holotype of /. alhifrons and comparison with specimens of several species o{ Ollarianus show that it be- longs in that genus and is identical with O. tripartitus. The genus Ollarianus has wide 1988 NiELsoN: CoLLADONus OF Mexico AND Centiul America 133 107 Figs. 105-107. Paracrassana nigrifrons (DeLong): 105, male pygofer, lateral view; 106, connective, right style, and aedeagus, dorsal view; 107, aedeagus, lateral view. distribution from the Nearctic to the Neotrop- ical region and needs revision. Paratanus costatus (DeLong), n. comb. Idiodonus costatus DeLong 1984:92 [Holotype 3 (OSU) (examined).] This species is close to Paratanus wygod- zinskyi recurvatus (Linnavuori) but differs from it in color characters. It may be a distinct subspecies of the nominate form or a separate species pending studies of the genus Para- tanus. Angulanus DeLong, n. status Idiodonus subgenus Angulanus DeLong 1946:30. Type- species, Idiodonus incisurtis DeLong 1946:30. [Holotype 9 (OSU) (examined).] Idiodonus (Angulanus): Metcalf 1967:1304 Idiodonus: DeLong 1984:9. [Inadvertent synonymy of Angulanus DeLong in the treatment of the type- species, incisurus DeLong, in a key to the species oi Idiodonus .] Examination of the genital structures of the male allotype clearly showed that Angulanus deserves generic-level rank. The aedeagus is asymmetrical, and the pygofer possesses a prominent spine on the caudal margin, char- acters that are not descriptive of the genus Idiodonus. Angulanus incisurus (DeLong), n. comb. /diodonus incisurus DeLong 1946:30 [Holotype 9 (OSU) (examined).] Idiodonus incisurus: DeLong 1946:13, 14 Idiodonus (Angidanus) incisurus: Metcalf 1967:1304 Idiodonus (Angulanus) incisurus: DeLong 1984:10 134 Great Basin Naturalist Memoirs No. 12 Idiodonus plummeri DeLong Idiodonus plummeri DeLong 1946:25 [Holotype 9 (OSU) (examined).] Idiodonus plummeri: DeLong 1984:11 Idiodonus bakeri. DeLong 1984:26 [Holotype 9 (OSU) (examined).] New synonymy Idiodonus bakeri: DeLong 1984:11 The generic placement of this species ap- pears to be correct. No males were described in the type series oi plummeri or bakeri, but a male paratype specimen labeled "Idiodonus bakeri DeLong, MB 73, Mexico, A. Dampf coll." was found and examined. The genitalic structures clearly show that it belongs in Id- iodonus . The species was associated with the female holotype of bakeri, which was found to be identical with the female holotype o( plum- meri. Species incertae sedis Three species of Idiodonus described by DeLong, edentulus, excavatus, and copidus, are treated herein as incertae sedis until males are known and associated with females of these species. Idiodonus copidus appears to be close to Jaacunga and may belong in that genus. Acknowledgments I thank Dr. Charles A. Triplehorn and Dr. Paul Cwikla, Ohio State University (OSU), for placing on loan nearly the entire type series of all the Idiodonus species from Mexico and Central America described by the late Dr. D. M. DeLong. Without this material it would not have been possible to unravel the taxonomic problems involved with the Id- iodonus and Colladonus fauna in that part of the world. I also thank Dr. James P. Kramer, U.S. National Museum, for the loan of the type of Colladonus delongi Linnavuori and material of genera related to Colladonus. My appreciation is extended to Jeanette Price and Pamela Scott for the illustrations and to Kaye Thome for identifying the host plant of a population of the Colladonus belli complex. I thank Dr. Leon Hepner for read- ing the manuscript and offering helpful sug- gestions to improve its content. References Cited Ball, E D 1937. Some new species of leafhoppers in groups recently segregated from Thamnotettix. Bull. Brooklyn Ent. Soc. 32: 26-31. Bliven, B P 1954. New Hemiptera from redwood (Miri- dae, Cicadellidae). Bull. Brooklyn Ent. Soc. 49: 119. DeLong, D M 1944. The genus Ollarianus (Homoptera: Cicadellidae) in North America, including Mex- ico. J. Wash. Acad. Sci. 34: 391-398. 1946. The Mexican species oi Idiodonus (Homop- tera-Cicadellidae). Ohio J. Sci. 46: 13-30. 1983. New species and distribution notes of Mexi- can and Bolivian Idiodonus (Homoptera: Cicadell- idae) of Mexico and Bolivia. Entomol. News 95(1): 9-15. 1984. A revised key to the Idiodonus (Homoptera: Cicadellidae) of Mexico and Bolivia. Entomol. News 95(1): 9-15. Gillette, C. P, AND C. F. Baker 1895. A preliminary list of the Hemiptera of Colorado. Colorado Agric. Expt. Sta. Bull. 31. 1.37 pp. Medler, J T 1943. The leafhoppers of Minnesota. Ho- moptera: Cicadellidae. Minnesota Agric. Expt. Sta. Tech. Bull. 155. 196 pp. LiNNAVUORL R 19.59. Revision of the Neotropical Delto- cephalinae and some related subfamilies (Ho- moptera). Suomal. Elanja Kasv. Seuran Vanamon Elaint. Julk. 20: 1-370. Nielson, M W 19.57. A revision of the genus Colladonus (Homoptera, Cicadellidae). U.S. Dept. Agric. Tech. Bull. 11,56: 1-.52, 1962. New species of leafhoppers in the genus Colladonus (Homoptera, Cicadellidae). Ann. En- tomol. Soc. Amer. .55: 143-147. 1966. A synopsis of the genus Colladonus (Ho- moptera: Cicadellidae). J. Kansas Entomol. Soc. .39: .3.33-.336. Oman, P W 1949. The Nearctic leallioppers (Ho- moptera: Cicadellidae). A generic classification and check list. Washington Entomol. Soc. Mem. 3: 1-2,53. Stal, C 1864. Hemiptera Mexicana enumeravit spe- ciesque novas descripsit. Stettiner Entomol. Zeit. 25: 49-86. Uhler, P R 1877. Report upon the insects collected by P. R. Uhler during the explorations of 1875, in- cluding monographs of the families Cynidae and Saldae and the Hemiptera collected by A. S. Pack- ard, Jr. U.S. Geol. and Geog. Survey of the Terr. Bull. 3(2): 355-475. NEW WORLD FULGORIDAE, PART I: GENERA WITH ELONGATE HEAD PROCESSES Lois B. OBi-it-n' Abstract. — Genera new to science described below include: Amerzanna, Sinuala, and Staluhra. New species include: Amerzanna peruana (Peru), Amijcle hrevis (Mexico), A. grandis (Mexico), A. mankimi (Honduras), Aphro- disias shaman (Mexico), Enchophora maculata and E. uniformis (Peru), Phrictus delicatiis (Brazil), P. dilinen.s (Colombia), Sinuala schmidti (Belize), S. stall (Honduras and El Salvador), S. tuherculata (Costa Rica), and Staluhra brunnea (Brazil and Guyana). In addition, Artacie dufourii (Signoret) is removed from synonymy with A. haemoptcra (Perty). New generic synonymy includes C/H/oi>ifl Stal(= Ecuadoria) Distant, En/iyr/na (Walker) (= UluhraStdl). New synonymy of species includes: Copidocephala merula Distant (= Coanaco melanoptera Schmidt), Copiducephala viridiguttata Stal (= Coanaco ornanda Distant), Diareusa conspersa Schmidt (= D. dahli Ossiannilsson), Enchophora nigromactdata Distant (= E. nigrolimhata Lallemand), Enchophora recurva (Olivier) (= £. hohemani Stal), Kri- chophora sanguinea Distant (= £. /Zorens Distant and E. longirostris Distant), Enchophora tuberctdatu (Ohvit'r) (= E. parvipennis Walker), Enchophora viridipennis Spinola (= E. emincnta Schmidt), Enhydria tessellata (Walker) (= £. brachialis Stal), Fulgora graciliceps Blanchard (= Laternaria orthocephahi Fonseca), Fulgora laternaria (Linnaeus) (= F. servillei Spinola), Phrictus auromactdatus Distant (= P. notatus Lellemand), Phrictus moebiusi Schmidt (= P. sordidus Caldwell). The two species of Fulgora are synonymized by Ridout. New combinations include Chilobia dichopteroides (Distant) (Ecuadoria) and Stahtbra ruftda (Lallemand) (Enhydria). Although the genera of New World Fulgori- dae were monographed in an excellent paper by Stal (1870b), 25 of the 64 genera have been described since then. Also, there have been many subsequent species described, many in the wrong genus, and there are few keys to species. Consequently, even though they are large and beautiful insects, it is very difficult to identify most species. The purpose of this paper is to provide a guide for identification with emphasis on the use of external charac- ters. Very little is known about the biology of Fulgoridae. Two species have been reported to be of economic importance (Wilson and O'Brien 1987), Phrictus diadema (L.) on cocoa trees {Theobroma cacao L.) in Brazil (Silva 1945) and Pyrops candelaria (L.) on longan and mango (Kershaw and Kirkaldy 1910). Host records for species contained in this pa- per include Amycle pinyonae Knull & Knull on pinyon pine, Pinus monophylla Voss (Knull and Knull 1947), Fulgora laternaria (L.) on Hymenaea courbaril L. or quapinol (Janzen 1983), Enchophora pallidipunctata Lalle- mand on Copaifera dourajeauni (Legumi- nosae) (label data, species not found in botani- cal references), and Rhabdocephala brunnea Van Duzee on Baccharis sarothroides Gray. Johnson and Foster (1986) explored host specificity in a tropical forest, where plant diversity might make host finding difficult, and found 71% of Enchophora longirostris Distant (= sanguinea Distant) on Simarouba amara Aubl. (Sapindales) and 84% of Phric- tus quinquepartitus Distant on Terminalia oblonga Steud. (Combretaceae). Adults have been reported to aggregate on tree trunks in ranks and to move in conjunction with each other (Johnson and Foster 1986, O'Brien and Wilson, 1985). Further host data will be given in parts to be published later. Little information on life histories is avail- able. Kershaw and Kirkaldy (1910) noted that the eggs o{ Pyrops candelaria were laid in flat clumps on tree trunks and covered with wax from the female abdominal segments; nymphs dispersed and fed on branches; and adults fed on tree trunks. Hingston (1932) reported rear- ing fulgorid nymphs from what he described as a mantidlike egg case. Other descriptions of immatures are those of Fulgora phosphorea L. (= laternaria L.) (Hagmann 1928); Pris- tiopsis serratus (F.) (= Cathedra serrata [¥.]) (da Fonseca 1931), and Itzalana subnuiculata Schmidt (Wilson and O'Brien 1986). As far as 'Entomology, Florida A & M University. Tallahassee. Florida, USA. 135 136 Great Basin Naturalist Memoirs No. 12 is known, fulgorids feed in phloem tissue and are univoltine. No accounts of other typical fulgoroid behaviors, such as accoustical com- munication and association with ants, have been reported. Fulgoridae are subject to predation by fly- catchers (D. Wechsler, personal communica- tion) and parasitization by the epipyropid moths (Lepidoptera, Epipyropidae). The col- ors of the closed wings would appear to make them inconspicuous against tree trunks, but they can be found fairly easily by the human eye if observed in profile. I assume that the red, orange, yellow, and white coloration so common at the base of the hind wings is simi- lar in function to that of the underwing moths (Noctuidae), in which the color has been shown to startle birds seeking prey. Eye spots on the hind wings of Cathedra and Fulgora would seem to serve a startle or warning func- tion as well. But for all the speculation on the bizarre shapes of the head processes, no one has reported the response of predators to the "alligator head" of the peanut bug, nor to any other species. One common name for the family Fulgori- dae, "lantern-flies," was given because Ful- gora were reported to emit light (de Merian 1771), but recent inquiries and enzyme stud- ies have failed to substantiate this phe- nomenon (Ridout 1983). Many Amerindians feared Fulgora, reporting that they flew a zig- zag path through the forest, killing everything they touched (Branner 1885). Shamans car- ried one in their amulet bags. The modern version of this myth, probably invented and surely repeated in coed entomology classes far out in the bush, is that if a human is bitten by a Fulgora, death is certain unless one is saved by the antidote, mating within 24 hours; non- sense, of course. Lallemand (1959, 1963) revised the 20 gen- era and 102 species of Fulgoridae from Africa and the 28 genera and 178 species of Asia and Australia. The New World has a much richer fauna, composed of 64 genera and 250 species as I begin this monograph. The only revisions of New World genera in which authors ex- amined types are Cyrpoptus (Kramer 1978) and Fulgora (Ridout, personal communica- tion). Other generic reviews are those on Ful- gora (da Fonseca 1932), Enchophora (Metcalf 1938), Phrictus (Caldwell 1945), and Poiocera (Gerstaecker 1860), which are either out of date or relied on published descriptions. I have examined types of all but the following species: Artacie dufourii (Signoret) (MZF?, letter not answered); A. haeriioptera (Perty) (ZSBS, not found); Cathedra serrata (Fabri- cius) (not listed by Zimsen 1964); Copidoced- phala melanoptera Schmidt, Diareusa con- spersa Schmidt, and Enchophora etninenta Schmidt (IZW); Enchophora ensifera (Ger- mar) and E. tuba (Germar) (lost in Lvov, Rus- sia); E. nigrolimbata Lallemand and Enhydria rufula Lallemand (FSAG, sent but not yet arrived); Olivier s species, £. recurva, E. tu- berculata, and F. caerulescens, and the types of Fulgora, which were studied by Ridout. Olivier's (1791) species were redescriptions with Latin binomials of Stoll's (1781, 1788) descriptions and figures in French and Dutch or Flemish, using names in those languages. Olivier's species are not in the Paris Museum (MHNP), nor in his personal collection de- posited there, which is all Coleoptera (Bour- goin, personal communication). Olivier cites "Du cabinet de M. Holthuisen" in some spe- cies; Horn and Kahle (1935) say of Holthuisen collection "in Hamburg am 3. II. 1796 u. 16. V. 1797 sowie in Stralsund 1800 verauk- tionert." If any specimens went to the Ham- burg Museum (ZMUH), they were destroyed by bombing in World War II, except for a few specimens out on loan. I have not been able to trace Olivier's or Stoll's specimens further. The first three of the species listed here were well illustrated when described; Schmidt's descriptions clearly fit the specimens I have; only Lallemands, Germar's, and Olivier's species are in doubt. Fulgoridae may be distinguished from other Fulgoroidea by the presence of numer- ous cross-veins in the hind wings, a feature lacking in all other families. Stal also noted that the carina between the frons and gena continues onto the clypeus, but this feature also occurs in some Derbidae, Dictyophari- dae, and Lophopidae. Fulgoridae share with the Dictyopharidae similar bilaterally sym- metrical, usually trilobed, inflatable male genitalia. Emelyanov (1979) transferred some Asian genera from Dictyopharidae to Ful- goridae, citing other characters, but these characters are not diagnostic for New World families. The stable characters easiest to use are color, especially of the hind wing; the shape 1988 O'BRIEN; New World Fulgoridae 137 and length of the head process when present; and the shape and carinae of the frons, vertex, and thorax. Since the proportion of the head length to the body length varies greatly with the genera and cannot be determined from the illustrations, measurements are given for the genera (unless it is impossible to get an accurate measurement because of the vari- ability of the curvature of the head process as in Enchophora). The ratio of head/pronotum is given for most species. Male genitalia are difficult to inflate consistently, but they are figured for some specimens; I state when I believe they must be used for a valid determi- nation, such as in Scolopsella. These descrip- tions are brief and diagnostic; illustrations are more helpful than words. A character matrix is being prepared and will be published when all of the genera have been studied. Part I covers those genera with head pro- cesses. These keys are artificial. When the definition of genera and placement of species is completed, an attempt will be made to pro- duce cladograms and to identify monophyletic taxa. Two species are consigned to incertae sedis. I do not recognize the genus of En- chophora ensifera (Germar) among any New World Fulgoridae or Dictyopharidae, nor Fulgora caerulescens Olivier as figured by Stoll (1788, plate 13, Fig. 65) among any New World Fulgoridae or Flatidae. Illustrations of the characters used are found in Figures 1-10 and 33. Three words that may not be commonly used elsewhere are defined as ioWows: fossette — a small, deep pit; porrect — extending forward horizontally; terete — round in cross section. The head pro- cess may have 10 carinae or fewer, which are named in Figure 4. The median vein is pecti- nate in Figure 5. The convention in Ful- goroidea is to consider the "base" of the insect the dorsal junction between head and thorax, with the base of the head and pronotum both here. The ventral part of the head has its base anterad, at the junction with the vertex, and its apex at the frontoclypeal suture, which is the base of the clypeus. To save duplication in each section, 1 have included within the final part of this paper the literature cited for all parts (although papers published subsequent to Metcalf 's catalog are cited here); a fist of localities with provinces or longitude and latitude (only the city or dis- tance for a city plus elevation where given will be included earlier), and leclotypes or neo- types will be designated. There is a list of genera and species and their distribution by country at the end of this part. Type reposi- tory abbreviations are listed in the acknowl- edgments. To complete this monograph successfully, I will need help in locating specimens, since many species are known from uni(}ues or one sex only, and distribution records are incredi- bly incomplete. For at least a year after the publication of this article, I will accept speci- mens for identification for the traditional rec- ompense, a sample of the specimens in return for identification. Perhaps your specimens will become types. The genus Chilohia is a case in point: there are four described species, 1 have 10 new ones on hand, and only one species is known from more than one speci- men. Please help if you have unidentified specimens. Key to Genera of New World Fulgoridae with Head Processes 1 . Tegmen usually transparent, rarely opaque in basal half 2 — Tegmen opaque, rarely transparent at apex . . 4 2(1). Tegmen transparent brown with small bris- tles on veins; prothorax with small, white nod- ules Enhydria — Tegmen without bristles on veins; prothorax without small, white nodules 3 3(2). Vertex longer than broad (Fig. 1) Stalubra — Vertex 3 times as broad as long in midline, emarginate medially (Figs. 51, 54, 60) or apex of vertex hidden by recurved head process (Fig. 57) Chilohia 4(1). Each hind wing with an eyespot visible from below with wings unspread; head process gib- bous or laterally serrate with narrow apex; large species, over 50 mm in length 5 — Hind wings without large, round eyespots; head process porrect or curved, but not as above 6 5(4). Head process peanut-shaped, in lateral view with markings mimicking an alligator head; eyespot on hind wing with iris or pupil Fulgora — Head process subtriangular, narrowing at apex, strongly serrate (Fig. 26); eyespot on hind wing without iris or pupil Cathedra 6(4). Head process porrect (Figs. 14-32, 46-50, 102-117) 9 — Head process curved up and back (Figs. 33-35, 40-42, 63-76) 7 7(6). Preocular horizontal flange between eye and junction of frons and vertex; head process ad- pressed to vertex, flattened, rugosely tuber- culate (Figs. 33-35) Artacie 138 Great Basin Naturalist Memoirs No. 12 Figs. 1-13. See facing page for identification. 1988 O'BRIEN; New World Fulgoridae 139 Figs. 1-13. 1-3, 5, 6: Staluhra hrevis O'Brien: 1, dorsal view, head and thorax; 2, frons and clypeus; 3, lateral view, head; 5, tegmen; 6, hind wing. Hypothetical insect: 4, diagramatic cross section of head process showing position of carinae; 7, stylized hind tibia. Male genitalia, lateral, dorsal, and ventral views: 8-10, Scolopsella mexicana O'Brien; 11-13, Scolopsella reticulata Ball. Morphological characters: a-antenna A,-lst anal vein aed— aedeagiis af-anal fold of hind wing all- anal flap (10th and 1 1th abdominal segments) an— anal area ant— anterior margin of wing ap— apical area of hind wing ax-apex b-base C— costa or costal vein c— clypeus cl-clavus clas-clasper cm— commissural margin cor-corium cs— claval suture CU— cubitus or cubital vein cx-coxa e-eye f-frons fe— femur fo-fossette g-gena hp-head process Ic^lateral carina Ih-length of head Ihp-length of head process Ip-length of pronotum M -media, medial vein m-mesonotum mac-marginal carina mc-median carina mem-membrane of tegmen nl-nodal line o-ocellus p-pronotum pc-pleural carina Pcu-postcuhital vein pf-postocular flange post-posterior area of wing pr-pronotum prf-preocular flange (see Fig. 33) py-pygofer R-radius, radial vein sa— sutural angle Sc-subcostal vein sf-supraocular flange t-tegula ta-tarsomeres ti-tibia tr-trochanters vertex of head (Continued from page 137) — Preocular horizontal flange absent; head process usually not adpressed to vertex (see E. tuber- culata, subviridis, pyrrhocrifpta. Figs. 67, 73, 75) ' 8 8(7). Pronotal median carina raised, with deep fos- settes at base; mesonotal apex depressed, stri- ate (Figs. 63-76) Enchophora — Pro- and mesonota smooth and gently rounded (Figs. 40-42) Copidocephala 9(6). Medium-sized species, 26-50 mm in length 10 — Small species, under 25 mm in length 13 10(9). Head process with apex twice as wide as nar- rowest part, dorsolateral edges of process with spines Phrictus — Head process with apex not so widened, sides of process without spines 11 11(10). Sutural angle of tegmen triangularly pro- duced; head with small, horizontal, preocular flange (Figs. 116-117) Odontoptera — Sutural angle of tegmen rounded; head with slight vertical flange on vertical carina (Figs. 27-32) 12 12(11). Head process elongate, head 1/3 length of insect, subtriangular, slightly bent dorsad at apex; hind wing clear with veins tinged with green at base, brown elsewhere .... Ainerzanna — Head process proportionally shorter, less than 1/5 length of insect, with sides subparal- lel, apex slightly enlarged, not bent dorsad; hind wing brown with either red or white spots Diareusa 13(9). Head process semicircular in cross section; with dorsal surface slightly compressed, ven- tral rounded Amycle — Head process more nearly rectangular or cir- cular in cross section 14 14(13). Head, including process, longer than thorax; tegmen with major veins straight 15 — Head, including process, shorter than thorax; tegmen with major veins irregular, slightly angulate 16 15(14). Head process subrectangular in cross section, carinae foliaceous, angulate (Figs. 24, 25) . . . Scolopsella — Head process smoothly rounded, almost cir- cular in cross section, carinae normal (Fig. 23) Rhabdocephala 16(14). Costal margin of tegmen markedly sinuate (Fig. 43); head process not transversely striate Sinuala — Costal margin of tegmen slightly convex; head process transversely striate (Fig. 49) Aphrodisias 140 Great Basin Naturalist Memoirs No. 12 Figs. 14-28. Head and thorax, dorsal view: 14, Amycle brevis O'Brien, 15, A. pinyonae KnuW & Knull; 16, A. saxatilis Van Duzee; 17, A. tumacacoriae Knull & Knull; 18, A. vernalis Manee; 19, A. g,randis O'Brien; 20, A. sodalis Stal; 21, A. manhinsi O'Brien; 22, A. amabilis (Westwood); 2.3, RhahdoccpJuila hninnea Van Dnzee; 24, Scolopsella mexicana O'Brien; 25, S. reticulata Ball; 26, Cathedra scrrata (Fahricius); 27, 28, Amcrztinna prrnana O'Brien (27, lateral view). Amerzanna, n. gen. Figs. 27, 28 Type-species: Amerzanna peruana O Brien. Medium-sized, narrow, brown insect.s, 31-41 mm long, head 1/3 length of insect. Head process porrect, terete, gradually nar- rowed to apex; apex bent upward; dorsal and ventral median and lateral carinae present on anterior third; weak hut traceable pleu- ral carina cvuving ventrally to end in slight 1988 O'BRIEN: New World Fulgoridae 141 ventral tubercle. Preocular flange absent. Pronotum smoothly rounded, traces of fos- settes and pleural carinae faintly visible. Teg- men with M pectinate; cross-veins irregular, forming diamond-shaped cells. Profemur Ion- longer than metafemur, posterior tibia with 6 lateral spines. Ninth abdominal tergite in fe- males twice as long as eighth. I name this genus Amerzanna because su- perficially it resembles the genus Zanna from Africa and Asia. The type-species is A. peru- ana, new species. Atiierzanna differs from Odontoptera, which has the head similarly shaped, by the differences in the tegmina, for it lacks the sharp sutural angle and the spot on the nodal line of Odontoptera. Amerzanna peruana, n. sp. Figs. 27, 28 Length. — Male 31-33 mm, female 35-41 mm; head 15.6 mm; ratio, head/pronotum 8.3. Ground color reddish brown. Tegmina well pigmented, transparent, but not obviously so (the hind wings may be seen through the teg- mina). Hind wings transparent, greenish cloud at base, brown cloud at apex, brown veins between. HOLOTYPE (male) and ALLOTYPE (female). — Peru: Madre de Dios: Rio Tambopata Res., 30 air km SW Pto. Maldonada, 290 m, 16-20- XI-1979, J. B. Heppner, subtropical, moist forest (USNM). Paratypes (10 female, 1 male): 3, same data (USNM); 2, 11-15-XI-1979 (USNM); 2 females, 1 male 6-10-XI-1979 (USNM, LOB); 2, Loreto, Ucayali R. Yarina Cocha, 30-XI-1954, leg. Peter Hocking, 1 head process broken (LOB, FMNH); 1, Yahuarmayo, Ape. Group (MCZ). Amycle Stal Figs. 14-22, 93-101 Amtjcle Stal 1861:148. Type species: A. amahilis (West- wood), subsequent designation by Van Duzee 1916:78 Cyrpoptus (Amyele) [sic] Stal, 1862:305. Small, brown insects, 12-23 mm long, head 1/8 to 1/4 length of insect. Head process por- rect, dorsoventrally flattened, dorsal surface flat, ventral convex; usually elongate triangu- lar, but sometimes parallel-sided, sometimes slightly enlarged at apex; dorsal lateral carinae sinuate, it and other carinae variable between species. Preocular flange horizontal, about as produced as marginal carinae. Tegmen with M not pectinate (2 main branches), cross- veins reticulate, apical margin more angulate than in most genera. Metatibia with 4-5 lat- eral spines (sometimes 3 or 7 on a single tibia). Female ninth tergite 1.5-2 times as long as eighth. This genus may be separated from other small, brown fulgorids by the porrect head being flattened dorsoventrally. It resembles Cyrpoptus in wing coloration. Diagnoses are not provided; these species may be identified by the shape of the head (Figs. 14-22) and the color and color pattern of the hind wings (Figs. 93-101). Key to the Species oi Amycle 1. Hind wing with 2 colors, brown posterior and base white, yellow, pink, or red (Fig. 93) saxatilis Van Duzee — Hind wing with 3 colors, brown posterior, translucent area (Figs. 94-101), and base white, yellow, orange, or red 2 2(1). Base of hind wing yellow, white, or pink 3 — Base of hind wing red or orange 4 3(2). Base of hind wing yellow or yellow-orange, con- tiguous with brown suffusion, apical area lightly suffused with brown, intermediate area translu- cent (Fig. 101) amahilis (WesUvood) — Base of hind wing pale pink, contiguous with white band, posterior margin brown (Fig. 97); sides of head process parallel (Fig. 18) vernalis Manee 4(2). Vertex at least twice as long as broad (Figs. 17-21) 5 — Vertex less than twice as long as broad (Figs. 14, 15) 8 5(4). Hind wing with brown apical and anal areas contiguous (Figs. 93-97) 6 — Hind wing with transparent area between brown apical and anal areas (Figs. 98-100) 7 6(5). Hind wing with transparent area restricted to narrow anterior band (Fig. 94); head process parallel-sided (Fig. 20) sodalis Stal — Hind wing with transparent area about 1/4 area of wing (Fig. 9.5); head process subtriangular (Fig. 21) mankinsi, n. sp. 7(5). Head process expanded at tip (Fig. 19); hind wing with red area separated from brown anal and apical areas by transparent band (Fig. 98) grandis, n. sp. — Head process subtriangular (Fig. 17); red area and brown anal area contiguous (Fig. 100) .... tumacacoriae Knull & Knull 8(4). Head broader than long (Fig. 14); apex of hind wing narrowly brown (Fig. 99) brevis, n. sp. — Head longer than broad (Fig. 15); apex of hind 142 Great Basin Naturalist Memoirs No. 12 wing broadly brown (Fig. 96) pinyonae Knull & Knull Amycle amabilis (Westwood) Figs. 22, 101 Fttlgora amabilis Westwood 1842:119. Type repository: UMO. Amycle amabilis (Westwood), Stal 1861: 148. Length. — Male 14 mm; head 3.4 mm, ra- tio, head pronotum 3. 1. Head process elongate, parallel-sided. Hind wing yellow to yellow-orange at base, followed by brown suffusion, apical area less darkly suffused, area between latter two hya- line. Distribution. — Holotype and paratype: Mexico. No other specimens seen. Amycle brevis, n. sp. Figs. 14, 99 Length. — Male 13 mm, female 14 mm; head 1.8 mm; ratio, head/pronotum 1.0. Head triangular, not produced into distinct process (Fig. 14). Hind wing pale orange at base, anal area pale brown, dark brown area on each side of anal fold, apical row of cells brown (Fig. 79). Tegmina in male pale orange with brown maculation apically; in female mottled brown throughout, forming a pattern with a clean, apical, diagonal line. The two specimens upon which this species is based may represent different species, but the head and hind wings are similar; examina- tion of male genitalia will be necessary to de- termine whether these are two different spe- cies. A. brevis may be separated from other species by its short head. Holotype (male). — Mexico: Nayarit, San Bias, 4-IX-1971, W. J. Hansen (LOB). Fe- male (not allotype or paratype): Guerrero: Ciudad Altamirano 20-X-1983, H. Brailov- sky, E. Barrera (LOB). Amycle grandis, n. sp. Figs. 19, 45, 98 Length. — Female 20-22 mm; head 5 mm; ratio, head/pronotum 4.5. Head elongate, slightly expanded at apex (Fig. 19). Hind wing orange in basal 1/3, brown in anal 1/3 and apical 1/4, translucent areas between (Fig. 98). Tegmina brown with most veins orange-red; two diagonal, brown lines delineating translucent half oval on costa, translucent triangle at apical angle, and mottled brown and white pattern in anal angle (Fig. 45). Holotype (female). — Mexico: Sinaloa, Ma- zatlan, 5-VIII-1971, D. W. Davis (LOB). Paratype (female): Sonora, Canyon Sapopa, 15-X-1934, Rio Mayo, H. S. Gentry (CIS). Possible male (not paratype): Jalisco, Chapala, 6 mi W Jahsco, 30-VM963, J. Doyen (LOB). Amycle rnankinsi, n. sp. Figs. 21, 95 Length. — Female 17 mm; head 3.5 mm; ratio, head/pronotum 2.9. Head slightly elongate triangular, turned up at apex (Fig. 21). Hind wing red in basal third, apical and anal thirds brown, all colored areas separated from each other by pale areas (Fig. 95). Tegmen brown with most veins red. I dedicate this species to Dr. J. V. Mankins, with whom we have spent many happy hours collecting. Holotype (female). — Honduras: Lago Yojoa, 2-IX-1977, J. V. Mankins (LOB). Fe- male paratype: same data except 6-X-1977 (LOB). Amycle pinyonae Knull & Knull Figs. 15, 96 Amycle pinyonae Knull & Knull 1947:397. Type reposi- tory: OSU. Length. — Male 8 mm, female 14-14.5 mm; head 2.6 mm, ratio of head/pronotum 1.9. Head process triangular, lateral margins sinuate. Hind wing reddish orange in basal 2/5, anal and apical angles brown, translucent area between with brown veins. Tegmen dark brown with many clear areas, these not form- ing definite pattern. Distribution. — Holotype (male): Califor- nia: Pinyon Flat, Santa Rosa Mts. on Pinus cembroides var. monophylla Voss. I have seen three other specimens collected in August from California: Pinyon Flats and Valvermo and New Mexico: Juan Tabo, 7,000 ft. Amycle saxatilis Van Duzee Figs. 16, 93 Amycle saxatilis Van Duzee 1914:33. Type repository: CAS. Length. — Male lectotype 12.5 mm; head 2.8 mm, ratio, head/pronotum 3.5. Head process elongate triangular. Hind wing pale orange at base, followed by white 1988 O'BRIEN: New World Fulgoridae 143 Figs 29-45 Head lateral view: 29, Diareusa conspersa Schmidt; 30, D. kemneri Ossiannilsson; 31, D. annularis (Olivier) 32 D imitatrix Ossiannilsson. Dorsal view: 33, Artacie haemoptera (Perty); 34, A. dufouri (Signoret). Lateral view 35 A dufouri (Signoret) (pronotal nodules shown only in lateral view); 36, Enhydria longicornuta Lallemand; 37, £ tesseilata (Walker); 38, E. cicadina Gerstaecker. Frontal view: 39, E. cicadina Gerstaecker Lateral view: 40, Copidocephala merula Distant (nodules not shown); 41, C. viridiguttata Stal; 42, C. guttata white. Tegmen: 43, Sinuala tuberculata O'Brien; 44, Odontoptera carrenoi Signoret; 45, Amycle grandis O Bnen. 144 Great Basin Naturalist Memoirs No. 12 band, posterior half light brown. Tegmen pale, only slightly darker at apex. Van Duzee said the color at the base of the hind wing varied from red to luteus in his specimens. This has been verified (N. Penny, personal communication). The color pattern, however, in both specimens has no translu- cent area. Distribution. — Lectotype, right wing re- moved and pinned beneath specimen: Cali- fornia: San Diego County. I have seen no other specimens. Amijcle sodalis Stal Figs. 20, 94 Amycle sodalis Stal 1861:148. Type repository: NRS. Length. — Male 13-14 mm, female 16.5- 18 mm; head 4.5 mm, ratio, head/pronotum 4.5. Head process elongate, sides parallel to apex. Hind wing orange at base, dark brown posteriorly, anterior 1 1/2 rows of cells behind orange base translucent (Fig. 94). Tegmina red basally, apically translucent with 2 dark apical lines (similar to Cyrpoptiis); some dark patches caudally. Distribution. — Holotype: Mexico. I have seen six other specimens collected from July through November from El Salvador: Santa Tecla, 900 m and Mexico: Atlixco, Canada de Negros, Cordoba, Huachinango. Amycle tumacacorae Knull & Knull Figs. 17, 100 Amycle tumacacorae Knull & Knull 1947:398. Type repository: OSU. Length. — Male 14 mm, female 14-15.5 mm; head 3.7 mm; ratio, head/pronotum 3.7. Head process elongate triangular, slightly widened just before apex, slightly bent dorsad in lateral view. Hind wing red-orange at basal third, dark anal and apical areas, gray mem- branous areas between with dark veins. Teg- men dark brown with few translucent apical cells. This species is very close to A. vernalis in shape and length of head process; it differs in the color and pattern in the hind wing. Distribution. — Holotype (male): Arizona: Tumacacori Mts. I have seen 12 other speci- mens from Arizona: Catalina Mts., Chiric- ahua National Monument, Molino Basin, and Pepper Sauce Canyon; Texas: Eastland County, Forestburg, 31 mi W Ozone, Tyler. Amycle vernalis Manee Figs. 18, 97 Amyele[sic] vernalis Manee 1910:117. Type repository; USNM. Length. — Female 14.5 mm; head 4 mm; ratio, head/pronotum 4.0. Head elongate, sides gradually narrowing to apex, apex slightly turned up. Hind wing pale pink in basal fourth, followed by band of white, posteriorly black. Distribution. — Lectotype: North Caro- lina: Southern Pines (USNM). I have seen one other specimen from Louisiana: Gal- braith. Aphrodisias Kirkaldy Figs. 49, 50 Compsoptera Stal 1869:236. Type species C. cacica Stal, original designation. Aphrodisias Kirkaldy 1906:248, new name for Compsop- tera Stal. Small, reddish brown insects, 18-23 mm long, head about 1/8 length of insect. Head process porrect, terete, transversely striate on all sides; dorsolateral carinae present; short, nonstriate, apical portion with dorsal median carina and longitudinal ridge on verti- cal face. Preocular flange horizontal. Tegmina with longitudinal veins irregularly angled, not straight; few cross-veins. Female ninth tergite slightly longer than eighth. This genus may be separated from all other genera by the wrinkled, porrect head process. The head process of Artacie also is textured, but it is recurved and recumbent on the ver- tex. Key to the Species of Aphrodisias 1. Head with wrinkled or striate portion of process shorter than pronotum; hind wings red at base, brown apically cacica (Stal) — Head with wrinkled portion of process longer than pronotum; hind wing red, white, and brown shaman, n. sp. Aphrodisias cacica (Stal) Fig. 49 Compsoptera cacica Stal 1869:237. Type repository: NRS. Aphrodisias cacica (Stal), Kirkaldy 1906:248. Length. — Male 18-20 mm; head 2 mm; ratio, head/pronotum 1.5. Head process siibe(|ual in length to vertex. Hind wing red at base, brown posteriorly. Tegmen brown. 1988 OBrien; New World Fulc;ohidae 145 Figs. 46-62. Head and pronotum, dorsal view: 46, Sinuala stali O'Brien; 47, S. schmidti O'Brien; 48, S. tuberculata O'Brien; 49, Aphrodisias cacica (Stal); 50, A. shaman O'Brien. Head and thorax, dorsal view and lateral view, and head, ventral view: 51-53, Chilobia silena Stal; 54-56, C. smaragadina (Walker); 57-59, C. cinxia Stal; 60-62, C. dichopteroides (Distant). 146 Great Basin Naturalist Memoirs No. 12 Distribution. — Holotype (female): Mex- ico. I have seen six specimens collected from Mexico: Alamos, 15 mi S Cuernavaca, Cuer- nonica, from September through November. Aphrodisias shaman, n. sp. Fig. 50 Length. — Female 23 mm; head 3.5 mm; ratio, head/pronotum 2.3. Head process 1.7 x longer than vertex. Hind wing pale pink at base, triangular, white area intruding into pink, apex and anal area missing, but edges of anal area brown, apical area brown with white spots. The wings of this specimen are badly dam- aged, but it may be separated from cacica on the basis of head shape. This specimen has lost its left tegmen and hind wing and part of the right wing, but it is a very distinctive species based on the head alone. Holotype (female). — Mexico: Sinaloa, 5 mi N Mazatlan, l-VII-1965, J. A. and M. A. Chemsakand E. G. and J. M. Linsley (UCB). Artacie Stal Figs. 33-35 Artacie Stal 1866:132. Type species: Flata haemoptera Perty, subsequent designation by da Costa Lima 1935:488. Brown-winged, medium-sized insects, 20- 27 mm long. Head process terete, recurved to lie against vertex (Fig. 35), texture of apex unevenly, rugosely tuberculate, ventral me- dian and marginal carina on process, lateral carinae on frons. Pre- and supraocular flanges large (Figs. 33, 34). Pronotum with white nodules. Tegmina with main veins slightly raised, surface concave between them, M 3- branched, cross- veins reticulate in corium, forming rectangular cells in membrane. Red or orange-red color at base of hind wing al- most reaching apical margin medially, divid- ing it into apical and anal areas. I synonymized the two species in this genus in 1985. After studying additional specimens, I conclude that I was in error and that the artist's illustrations of the species were correct in each case and there is a difference in the shape of the vertex of the two species. The white spots in the apical portion of the wings do vary in size, placement, and number be- tween specimens, but a general pattern for haemoptera is described below. Key to the Species oi Artacie 1. Median length of head process before preocular flange subequal to median length of vertex .... dufourii (Signoret) — Median length of head process before preocular flange half of median length of vertex haemoptera (Perty) Artacie dufourii (Signoret), new status Figs. ,34, 35 Encophora [sic] dufourii Signoret 1858:497, pi. 12, no. 2, Fig. 1. Type repository: MZF? = Artacie hemoptera (Perty), O'Brien 1985:661. Error. Length. — Male 25 mm, female 27 mm; head length 2.6 mm; ratio, head/pronotum 1.4. Length of head process in front of preocu- lar flange subequal to length of vertex. Wings orangish red at base, white spots scattered through brown apical and anal ar- eas, not forming a figure with an empty center (a square, circle, or diamond). The two known species may be separated by the proportion of the head process in front of the preocular flange to the vertex, subequal in dufourii and about half in haemoptera (Figs. 33, 34). Distribution. — Type (not seen): Cayenne. I have seen two other specimens collected in March from French Guiana: Maron River; Guyana: Wineperu. Part of Signoret's collec- tion went to the Naturhistorisches Museum, Vienna, and part to Museo Zoologico dell'Universita degli Studi di Firenze, Flo- rence, according to Horn and Kahle (1935). I did not see the type in Vienna and have not heard from Florence. Artacie haemoptera (Perty) Fig. 33 Flata haemoptera Perty 1833:176, pi. 35, Fig. 3. Type repository: ZSBS, not found. Artacie haemoptera (Perty), Stal 1866:389. Encophora [sic] dufourii Signoret, synonymized O'Brien 1985:661. Error. Length.— Male 20-22 mm, female 23-25 mm; head length 2.1 mm; ratio, head/prono- tum 1.2. Length of head process in front of preocular flange half length of vertex. Wings red at base, brown in apical and anal areas, 4-6 white spots in circle, diamond, or rectangle on api- cal area, few smaller dots also. These two species may be separated by the length of the head process and the pattern of white dots on the hind wing. 1988 O'BRIEN: New World Fulgoridae 147 Distribution. — Holotype (not seen) : Ama- zon basin [Brazil?]. I have seen nine other specimens collected in March, August, and October from Brazil: Canum, Manaus, Obidos, Rio Negro, and Teffe. The type could not be found in the Zoologische Sammlungen des Bayerischen Staates, Munich, in 1975. Cathedra Kirkaldy Fig. 26 Cathedra Kirkaldy 1903:179. Type-species: Phrictus ser- rata (F.), original designation. Pristiopsis Schmidt 1905:332. Type-species: Fulgora ser- rata F. , original description; synonymized by Dis- tant 1906:20. Large, yellowish brown insects, 52-71 mm, head 1/3 length of insect. Head process porrect, terete, gradually narrowed to apex; marginal and lateral carinae of frons produced into 11 and 8 spines, respectively; head about 1/3 length of insect. Preocular flange horizon- tal, spinelike. Tegmen with M irregularly branched (2 main branches); clavus open; cross-veins forming almost round cells anteri- orly, arched and parallel behind nodal line. Hind wings brown with ochre eye spot at apex, pale, elongate spots at base, some waxy spots and dashes throughout; emarginate on posterior border before spot. This monotypic genus has a head process like a cathedral spire, with many pairs of lat- eral spines, which distinguishes it immedi- ately from any other fulgorid genera. Cathedra serrata (Fabricius) Fig. 26 Fulgora serrata Fabricius 1781:313. Phrictus serratus (Fabricius), Schaum 1850:65. Cathedra serrata (Fabricius), Distant 1906:20. Length. — Male 52-53 mm, female 61-71 mm; head 19 mm; ratio head/pronotum 6.3. Ground color ochre mottled with brown. Distribution. — Holotype: Suriname. I have studied eight other specimens collected from January to November from Bolivia: Santa Cruz; Brazil: Manaus-Itacoatiana road, km 30; Panama: Barro Colorado Island. Zim- sen (1964) did not list the repository of this Fabrician type. Chilobia Stal Figs. 51-62 Chilobia Stal 1863:237. Type-species: C. cinxia Stal, sub- sequent designation by da Costa Lima 1935:497. Ecuadoria Distant 1906:21. Type-species: E. dichop- teroides Distant, original designation. New syn- onymy. Tegmina membranous throughout or mem- branous in apical half, basal half opaque, brown or brownish red, medium-sized in- sects, 19-28 mm long. Head process usually recurved, usually dorsoventrally compressed, usually gradually expanding in ventral view towards apex. Lateral carinae of frons parallel or diverging to near base, carinate throughout or becoming broad, fused with median carina into broad hump. Frons elevated above level of frontoclypeal suture, expanded into lobes near apex, with transverse carina extending across frons from lobe to lobe. Vertex with lateral carinae sinuate or angulate in lateral view, supraocular flanges present, apical mar- gin V-shaped, sometimes angulately emar- ginate medially in addition, sometimes hid- den under recurved process. Preocular flange absent. Pronotum with few white nodules. Tegmen with M pectinate (5-8 branches); clavus closed; cross-veins irregular and dense in basal half, few and as thick as longitudinal veins in apical half; longitudinal veins with small spines alternately angled to left or right; transition zone between membrane and opaque area, when present, is arched. Hind wings variable, from only slightly enfumed at base to bright red with sinuate, brown, con- tiguous area, only two marginal rows of cells enfumed at apex. Female ninth abdominal tergite shorter than eighth. Because I have intermediates and because the male genitalia are very similar, I have synonymized t\i^ genera Ecuadoria and En- hydria in spite of several character differ- ences. The species of Ecuadoria have a long head process and brightly colored tegmen and hind wing, whereas the Enhydria species have a short head process, transparent teg- mina, and clear hind wings. The four described species are illustrat- ed, but 1 have not provided a key because I am unable to determine the range of varia- tion within a species due to lack of available material. Chilobia dichopteroides (Distant), n. comb. Figs. 60-62 Ecuadoria dichopteroides Distant 1906:22. Type reposi- tory: BMNH. 148 Great Basin Naturalist Memoirs No. 12 Distribution, — Holotype: Ecuador (nw): Rio Durango, 350 ft. I have seen no other specimens. Chilobia cinxia Stal Figs. 57-59 Chilobia cinxia Stal 1863:238. Type repository: NHMV. Distribution. — Holotype (female): Vene- zuela. I have seen no other specimens. Chilobia silena Stal Figs. 51-53 Chilobia silena Stal 1863:238. Type repository: NRS. Distribution. — Holotype (male): Ecua- dor: Quito. I have seen no other specimens. Chilobia smaragdina (Walker) Figs. 54-56 Dichoptera smaragdina Walker 1851:304. Type reposi- tory: BMNH. Chilobia smaragdina (Walker), Gerstaecker 1895:39. Distribution. — Holotype: Venezuela. I have seen no other specimens. Copidocephala Stal Figs. 40-42 Copidocephala Stal 1869:235. Type-species: Enchophora guttata White, original designation. Coanaco Distant 1887:28. Type-species: Enchophora guttata White, original designation; synonymized by Distant 1906:23. Medium-sized, greenish or brown or red- dish brown insects, 22-32 mm long. Head process recurved at junction with head, later- ally compressed, gradually narrowing ante- rad; lateral carinae of frons fused at junction with process with medium and lateral carinae of process into one large mound; dorsolateral carina also present on process. Preocular flange absent. Pronotum with small, white, hemispherical tubercles, median carina not strongly humped. Tegmen with M usually pectinate (5-6 branches); corium with cross- veins regularly, evenly reticulate, with smooth, sunken, red spots without cross- veins; membrane with cells subsquare. Fe- male eighth and ninth abdominal tergites subequal in length. This genus has the head process curved back at the base, never extending foi-ward as in some Enchophora; it is scimitar-shaped like Enchophora and Enhydria. Copidocephala differs from Enhydria, which has membran- ous tegmina, and from Enchophora in the characters of the thorax given in that generic discussion. The close proximity of the prono- tum and eyes suggests that the postocular flange serves to protect the eye from contact with the pronotum. Key to the Species of Copidocephala 1. Hind wing black, luispotted merula Distant — Hind wing brown, spotted with red or bine, green or white 2 2(1). Large species, 28 mm or longer; hind wing brown with red spots guttata (White) — Small species, 25 mm or less; hind wing brown with blue, green, or white spots viridiguttata Stal Copidocephala guttata (White) Fig. 42. Enchophora guttata White 1846:331. Type repository: BMNH. Copidocephala guttata (White), Stal 1869:236. Length. — Female 29-32 mm, head length 1.4 mm; ratio, head/pronotum .63. Pronotum and costal margin of tegmen green, pronotum with red, transverse, sometimes broken, band; ground color of rest of insect ranging from yellow or green through red to olive green. Corium with approximately 21 round, smooth, sunken, small (1 mm or smaller), red spots; spots differing slightly in number, size, and placement among individuals. Hind wing brown with red spots in basal two-thirds. This species may be separated from inerida, which is similar in size and coloration, by the absence of red spots in the hind wing in merula. It differs from viridiguttata in size and in the color of spots in the hind wing, blue or green to white spots in viridiguttata. Distribution. — Type: South America. I have seen 20 other female specimens col- lected from March through September, November and January. Panama: Barro Colo- rado Island, Chiriqui Provence, Fort Clayton Costa Rica: 6 mi W Las Canas, La Selva Honduras: Catacamas, 15 km W La Ceiba Mexico: Tamazunchale, Temiscal, Volcan Tacana. Copidocephala merula Distant Fig. 40 Copidoccpluda iiicrula Distant 1906:23. Type repository: BMNH. Coanaco melanoptera Schmidt 1907:361. T\pe reposi- tory: IZW. 1988 O'BRIEN; New World Fulgoridae 149 Copidocephala melanoptera (Schmiclt), Metcall 1947: 176, New synonymy. Length. — Female, 34-38 mm. Ground color as in guttata except hind wing black, without red spots. The differences in coloration of the tegmen found in the two specimens of mcrula and melanoptera are within the variation found in the species guttata; so on the basis of size and the color of the hind wing, I synonymize the two species. Distribution. — Holotypes: Colombia (both types). I have seen the type oimerula but not o{ melanoptera. No other specimens seen. Copidocephala viridiguttata Stal Fig. 41 Copidocephala viridiguttata Stal 1869:236. Type reposi- tory: NHMV. Coanaco ornanda Di.stant 1887:29, pi. 4, Fig. 13. Type repository: BMNH. Copidocephala ornanda (Distant), da Costa Lima 1935: 491. New synonymy. Length. — Male 22-24 mm; head 1.1 mm; ratio of head/pronotum 0.5. Ground color olive green with black or red- black, sunken, smooth, round spots in same pattern as guttata. Hind wing brown with blue or green to white spots. This species differs from the other two in having blue, green, or white spots in the hind wing rather than red spots or none. It is inter- esting that I have 13 males of this species for study and 20 females of guttata. I will examine other collections to see if this is coincidence or unexpected sexual dimorphism. I synonymize ornanda on the basis of the shape of the head process and wing coloration. Distribution. — Type: Golombia (viridi- guttata); syntypes: Panama: Bugaba, Tole (or- nanda). I have studied 13 other specimens collected from May through September and January from Panama: Achiote Road, Barro Colorado Island, Fort Clayton; Costa Riga: Golfito, La Selva; Mexico: Los Tuxtlas, 3 mi S Palenque. Diareusa Walker Figs. 29-32 Diareusa Walker 1858:43. Type-species: Phrictus annu- laris (Olivier), by monotypy. Large, reddish brown insects, 32-49 mm, head relatively short, from .1 to .2 length of the insect. Head process porrect, terete, gradually slightly enlarging at apex; median ventral carina in about anterior third of pro- cess, ventral and dorsal lateral carinae com- plete; short dorsal median carina in anterior portion. Preocular flange vertical, well pro- duced. Tegmen with M sometimes pectinate (3-4 main branches), sometimes evenly branched; cross-veins irregular in corium, forming small squares beyond stigmal line. Tegmen brown throughout with some ante- rior cells and many spots either white or red; often some apical spots translucent. Female eighth and ninth abdominal tergites subequal in length. Diareusa are large, reddish brown insects with a comparatively short (.1 to .2 length of insect), terete, but not inflated head without transverse striae or spines; these characters will separate them from other New World Fulgoridae. The species differ in the color of spots in the hind wing and in the shape of the head process. Key to the Species of Diareusa 1. Spots on hind wing red and white conspersa Schmidt — Spots on hind wing white only 2 2(1). Head process short; vertex and head process as long as width of head, including eye kemneri Ossiannilsson — Head process longer than width of head includ- ing eye (Fig. 30) 3 3(2). Dorsal surface of head process in lateral view convex (Fig. 31) annularis (Olivier) — Dorsal surface of head process in lateral view straight or concave, not convex (Fig. 32) imitatrix Ossiannilsson Diareusa annularis (Olivier) Fig. 31 Fulgora annularis Olivier 1791:568. Type repository: un- known. Phrictus annularis (Olivier), Walker 1851:264. Diareusa annidaris (Olivier), Walker 1858:44. Length. — Male 32-38 mm, female 45 mm; head 6.2 mm; ratio of head/pronotum 2.2. Head process in lateral view convex dor- sally (Fig. 31.) Hind wings with white spots. These two characters separate it from the other species. Distribution. — Type (not seen): Suri- NAME. I have seen 10 other specimens col- lected June to August and January from PERU: Brazilian frontier; BRAZIL: Serra do Navio, Sinop; French Guiana: 67 km S Cayenne; SURINAME: Mapane; GUYANA: Shudihat River. 150 Great Basin Naturalist Memoirs No. 12 Diareusa conspersa Schmidt Diareusa conspersa Schmidt 1906:375. Type repository: IZW. Diareusa dahli Ossiannilsson 1940:44, Fig. 2. Type repository: UZIL. Tslew synonymy. Length. — Male 38 mm, female 46-49 mm; head 5.3 mm, ratio of head/pronotum 1.8. Hind wings with red spots. Head process similar to that odinitatrix. This is the most easily identified species in the genus with the spots on the hind wing red and white; in all other species they are white. It is interesting that although this is the easiest character for identification, both Schmidt and Ossiannilsson described this character among the last, causing it to be overlooked. Thus, many museums have imitatrix, the common Central American species, identified as con- spersa. I synonymize conspersa and dahli on the basis of head shape and wing color. Distribution. — Holotypes: Ecuador: Pal- mar (conspersa); Colombia: Rio San Agustin (dahli). I have seen three other specimens collected from June through August from Ecuador: Amhuagu, Rio Palenque. Diareusa imitatrix Ossiannilsson Fig. 32 Diareusa itnitatrix Ossiannilsson 1940:45, Fig. 3. Type repository: UZIL. Length. — Male 35-39 mm, female 43-48; head 5.8 mm; ratio head/pronotum 2.2. Hind wing with white spots. Head process similar to conspersa, but longer and narrower than other species with white-spotted hind wings. This is the only Central American species and often has been misidentified as con- spersa. Panamanians call this insect "totoran. Distribution. — Holotype (female): Pan- ama: Chiriqui. I have seen 50 other specimens collected all year around from Colombia: Turbo; Venezuela: Rancho Grande; Panama: Barro Colorado Island, Fortuna, Hartman's Finca, Margarita; Costa Rica: La Selva, Tur- rialba; Belize: Middlesex; Mexico: Bonampa R., 1 mi W Fortin de las Flores, Los Tuxtlas. Diareusa kemneri Ossiannilsson Fig. 30 Diareusa kemneri Ossiannilsson 1940:43, Fig. 1. Type repository: UZIL. Length. — Male 35 mm; head 4 mm; ratio. head/pronotum 1.8. Hind wings with white spots. Head process short (Fig. 30). D. kemneri may be separated from other white-spotted species by the short head process. Distribution. — Holotype (female): Peru: Perene. I have studied one other specimen collected in September from PERU: Aguaitia. Enchophora Spinola Figs. 63-76 Enchophora Spinola 1839:221. Type-species: recurva, subsequent designation by Duponchel 1840:200. Medium-sized, reddish brown, green, yel- low, or mottled green, white, and red insects, 22-32 mm long. Head process recurved, lat- erally compressed; nine carinae present on process (all except dorsal median carina). Pre- ocular flange present as vertical carina linking frons and vertex laterally (Fig. 3). Pronotum with pair of deep fossettes laterad of strongly raised median carina. Apex of mesonotum with striate depression between bifurcate me- dian carina. Tegmen with few strong, longitu- dinal veins, cross-veins variable, dense to av- erage. Female abdominal segments 8 and 9 subequal in length. Enchophora may be separated from the very similar Copidocephala by the raised me- dian carina of the pronotum with a fossette or depression on each side, the depression at the apex of the mesonotum surrounded by a high ridge, and the ventrolateral fields of the pronotum being separated from the eye by a space about the width of the last antennal segment. In Copidocephala the callus behind the eye almost touches the pronotum when viewed from above, and the pronotum and mesonotum are smoothly rounded. Both gen- era lack the preocular flange. Enhijdria, which also has a scimitar-shaped head pro- cess, may be differentiated by its pale tegmen and small spines on the tegminal veins. The species may be separated by the shape of the head process (although this structure varies slightly in curvature and compression), the color of the hind wings, and sometimes by distinctive marks or colors on the body or wings. Waxy spots on the wings may be help- ful if they have not been rubbed off; often they may be identified by a smooth, pale brown area if the wax is missing. The hind wings are usually red, yellow, orange, or white in the basal 1/4 to 2/3, measured along the anterior 1988 OBrien: New World Fulgoridae 151 margin, or 5/6 if measured from anterior to posterior. The color pattern of the tegmen may be quite variable, as in sanguinea Dis- tant, nigromaculata Distant, and reciirva (Olivier) where, in females, the tegmen may be concolorous, marked with discrete red or black spots, or marked with coalescent red spots. Although I have not seen both sexes of every species, if there is a sexual difference in color pattern, the male tends to be more evenly mottled and less variable. Three species listed as Enchophora in Met- calf's (1947) catalog have been transferred since to other genera (see list of species). Key to the Species of Enchophora 1. Hind wing with white, waxy points on red area stillifera (Stal) — Hind wing without white, waxy points on red area 2 2(1). Head process about twice length of frons be- fore process, enlarged and narrowed near tip (Figs. 63, 64) .3 — Head process less than twice length of frons before process, scimitar-shaped, curved back over vertex, or curved and then bent upward 4 3(2). Apex of head process black, broadly trilobed (Fig. 63) reciirva (Olivier) — Head process brown throughout, narrowly expanded at apex (Fig. 64) tuba (Germar) 4(2). Hind wing red in basal half; posterior half medium to dark brown; tegmen brown or reddish brown, often mottled 5 — Hind wing white, pale orange, or brown at base; either only apical 1/4 pale brown or posterior half medium brown (subviridis, tu- berculata), tegmen unmottled green or yel- low or green mottled with red 9 Apex of pronotum with transverse, black band (Fig. 66) nigromaculata Distant Apex of pronotum without transverse, black band 6 Mesonotum with black spots (Figs. 67, 76) ... 7 Mesonotum without black spots 8 Mesonotum with pair of black spots, also pair on ventrolateral fields of pronotum; head pro- cess recurved to near vertex, then bent up- ward (Fig. 67) pyrrhocrypta Walker — Mesonotum with 2 or 3 pairs of black spots; head process short, only slightly recurved (Fig. 76) maculata, n. sp. 8(6). Apical fourth of pronotum pale; base of mem- brane of tegmen with a brown spot about size of eye on each side, 3-5 other such spots usually present on membrane; ca basal half of hind wing red; abdominal tergites brown pallidiptinctata Lallemand 5(4). 6(5). 7(6). — Pronotum concolorous; tegmen not as above; ca basal 2/3 of hind wing red; abdominal ter- gites bright red sanguinea Distant 9(4). Hind wing white at base 10 — Hind wing yellow, orange, or brown at base 12 10(9). Tegmen green or yellowish 11 — Tegmen greenish white mottled with red . . rosacea Distant 11(10). Tegmen green or yellowish with white costal margin; head process large (Fig. 71) prasina Gerstaecker — Tegmen greenish yellow throughout; head process small (F'ig. 72) uniformis, n. sp. 12(9). Head process scimitar-shaped (Fig. 74); hind wing yellow at base, then white, then brown at apex viridipennis Spinola — Head process recurved and wrinkled (Figs. 73, 75); hind wing orange at base, brown at apex 13 13(12). Head process recurved, then bent upward at apex (Fig. 75) subviridis Distant — Head process recurved almost to vertex (Fig. 73) tuberculata (Olivier) Enchophora maculata, n. sp. Fig. 76 Length. — Male 24 mm, female 28 mm. Ground color brown with minute dots of red and green. Head process short, extending forward then dorsad (Fig. 76). Wings brown on apical third, red basally, red extending from anterior to posterior margin. Distinctive markings: two or three pairs of brown spots on pronotum (Fig. 76); anterior spot black, poste- rior medium brown, sometimes pale brown spot laterad of latter. Abdominal tergites brownish red. This species may be identified easily by its unusual, short, partially forward-directed head process (Fig. 76) and the spots on the pronotum. HoLOTYPE (male) and allotype (female). — Peru: Madre de Dios: Rio Tambopata Res., 30 air km SW Pto. Maldonada, 290 m, 16-20- Xl-1979, J. B. Heppner, subtropical, moist forest (USNM). Enchophora nigromaculata Distant Fig. 66 Enchophora nigromaculata Distant 1906:23. Type repos- itory: BMNH. £. nigrolimbata Lallemand 1938:350. Type repository: DEI. New synonymy. Length. — Female 24-26mm. 152 Great Basin Naturalist Memoirs No. 12 Figs. 63-76. Head and pronotum, lateral view, ofEnchophora: 63, E. recurva (Olivier); 64, E. tuba (Germar); 65, £. stillifera (Stal); 66, E. nigromaculata Distant; 67, E. pyrrhocrypta Walker; 68, £. piillidipunctata Lallemand; 69, E. sanguinea Distant; 70, E. rosacea Distant; 71, E. prasina Gerstaecker; 72, £. uiiiformis O Brien; 73, E. tuhercidata (Olivier); 74, E. viridipennis Spinola; 75, E. subviridis Distant; 76, E. maculata O'Brien. Ground color reddish brown, variable, teg- men may be clear, mottled with yellow veins, or with 10 dark spots; membrane brown or with dark spots, 5 to 7 waxy points along nodal line. Head process scimitar-shaped (Fig. 66). Wings red at base, brown posteriorly. This species may be separated from all other Enchophora by the black line along the hind margin of the pronotum. I have not seen the type o(nifi,rolimhata. Distribution. — Holotype: Bolivia (nigro- maculata), Peru (nigrolimbata). I have seen 1988 O'BRIEN: New World Fulgoridae 153 11 other specimens collected from October to January from Bolivia: Chapare; Brazil: Rio de Janeiro; Peru: Chanchamayo; 30 km SW Puerto Maldonado, 290 m; Satipo; Tingo Maria; Explorama Lodge; Ecuador: Parque Nacional St. Cecelia. Enchophora pallidipunctato Lallemand Fig. 68 Enchophora pallidipunctata Lallemand 1966:.52. Type repository; FSAG. Length. — Male 23 mm, female 24-26 mm. Ground color reddish brown, mottled. Head process scimitar-shaped (Fig. 68). Wings red on basal half, brown posterior half (not the usual 2/3 red, 1/3 brown). Seventeen waxy tufts present in apical area. Distinctive markings: pale band along apical fourth of pronotum and five round, dark spots in tegmi- nal membrane, one on each side just behind nodal line, three to five distad of these. Ab- dominal tergites brown. This species may be separated from all other Enchophora by the pale apical fourth of the pronotum, the dark spots in the membra- nous area of each tegmen, and the smaller, red portion of the hind wings. It most closely resembles sanguinea, which has the abdomi- nal tergites bright red, while they are brown in pallidipunctata. Distribution. — Holotype: Bolivia: Buena Vista. I have seen 24 other specimens col- lected from February through April, August and September from Bolivia: Buena Vista, Santa Cruz; Peru: Chanchamayo, Chontilla, Iquitos, Lima, Pileopota, 600 m, Rio San- tiago, Satipo, Tarapoto. Several specimens la- beled on Copaifera Dourajeanni (Legumi- nosae) at Tingo Maria, but I am unable to find this species listed in any botanical reference. Enchophora prasina Gerstaecker Fig. 71 Enchophora prasina Gerstaecker 1895:37. Type reposi- tory: emau. Length. — Male 24-27 mm, female, 28 mm. Ground color green, with costal margin of tegmen white and head and genitalia reddish brown. Head process scimitar-shaped, long, almost as long as mesonotum and pronotum combined. (Fig. 71). Wings white. No waxy tufts evident. This species may be separated from all other Enchophora by the green tegmen with white costal margin. Distribution. — Holotype: Colombia: Nova Granada. I have seen 12 other specimens col- lected from April through June, and October from Bolivia: Buena Vista; Peru: NE; Panama: Barro Colorado Island, Cerro Azul, Santa Rita Ridge; Costa Riga: La Selva. Enchophora pyrrhocrypta Walker Fig. 67 Enchophora pyrrhocrijpta Walker 1851:272. Type repos- itory: BMNH. Length. — Male 19-20 mm, female 23-25 mm. Ground color reddish brown, mottled. Head process recurved over pronotum, then bent upward (Fig. 67). Wings red at base, brown posteriorly. Two pairs of distinctive, black spots, one on each side of mesonotum, and one on each ventrolateral margin of pronotum (Fig. 67). No waxy tufts evident. This species may be separated from all other Enchophora by the black marks on the lateral fields of the pronotum and on the sides of the mesonotum (Fig. 67). E. subviridis hsis a similarly recurved head process. Distribution. — Holotype: Brazil: Para. I have seen seven other specimens collected from October through February from Brazil: Benjamin Constant, Rio Negro, Belem, Man- gabeira; Guyana: Kartabo, St. Laurent; Venezuela: Cerro de Neblina. Enchophora recurva (Olivier) Fig. 63 Fulgora recurva Olivier 1791:34. Type repository: un- known. Enchophora recurva (Olivier), Spinola 1839: 222, pi. 10, Figs. 1, 2 Enchophora bohemani Stal 1854:244. Type repository: NRS. New synonymy. Length. — Male 24-26 mm. Ground color reddish brown, tegmen mot- tled, often with spots composed of yellow clumped veinlets; membrane with long yel- low and brown markings. Head process elon- gate, produced in club that at apex is twice average width of process, apex trilobed, black (Fig. 63). Wings red at base, brown posteri- orly. No waxy tufts evident. This species may be separated from all other Enchophora by the enlarged, blackapex of the head process. 154 Great Basin Naturalist Memoirs No. 12 Distribution. — Holotypes: Brazil {bohe- manni) and Suriname (recurva). I have seen nine other specimens from BRAZIL: Linhares, Araxa, and Painaivas, taken from October through December. I am unable to confirm Berg's report from Argentina (Berg 1879). Enchophora rosacea Distant Fig. 70 Enchophora rosacea Distant 1887:27, pi. 4, Fig. IL Type repository: BMNH. Length. — Male 22-23 mm, female 25-26 mm. Ground color green, tegmen at base green- ish white with distinctive clumps of red dots between the green veins; membrane green with 20 white, waxy spots. Head process scim- itar-shaped (Fig. 70). Wings mostly white, dusted with red spots at base, brown posteri- orly. This species may be separated from all other Enchophora by the red and green com- bination on the tegmen. Distribution. — Holotype: Nicaragua: Chontales. I have seen 10 other specimens collected from March through August from Panama: Barro Colorado Island; Costa Riga: La Selva. Enchophora sangtiinea Distant Fig. 69 Enchophora sanguinea Distant 1887:27, pi. 4, Fig. 16. Type repository; BMNH. Enchophora florens Distant 1887:28, pi 4, Fig. 12. Type repository: BMNH. New synonymy. Enchophora longirostris Distant 1887:28. Type reposi- tory: BMNH. New synonymy. Length.— Male 22-23, female 24-25 mm. Ground color greenish to reddish, tegmen greenish or reddish, or greenish black with orange or red spots encircled with yellow veins, or the red spots may be fused into large areas or confluences, usually very mottled with clumps of yellow veins and dark and/or light spots. Head process scimitar-shaped (Fig. 69). Wings red at base, brown posteri- orly. Twenty waxy tufts present in membra- nous area of tegmen. Abdominal tergites bright red. I synonymize this species with two other species; E. florens and longirostris have the same male genitalia; E. sanguinea differs from them only in the pattern of the tegmen. When I found comparable difference in coloration in other species, I felt confident in synonymizing the three. This species may be separated from all other Enchophora with red hind wings by the shape of the head and the lack of distinc- tive identifying marks such as bands and spots on the pronotum. Distribution. — Type: Guatemala, Panama (sanguinea). Nicaragua: Chontales (florens); Costarica: Cache. Colombia (lon- girostris). I have seen 39 other specimens col- lected from July to September, with one in February. Ecuador: Palmar; Panama: Barro Colorado Island, Cana 1,200 ft; Costa Riga: 1 mi N Tucurrique, Rincon, Turrialba, Cairo. Enchophora stillifera (Stal) Fig. 65 Phrictus stillifera Stal 1862:303. Type repository: nhmv. Enchophora stillifera (Stal), Stal 1864:49. Length. — Male 25-28 mm, female 32 mm. Ground color greenish brown, tegmen of- ten with spots of clumped yellow veins, also often with broad patches of waxy exudate. Head process elongate, scimitar-shaped, about twice as long as frons below process. Wings red at base, mottled with white, waxy spots; brown posteriorly. This distinctive species is the only En- chophora with many white, waxy spots on the red portion of the hind wing. Distribution. — Holotype (male): Mexico; Guatemala: Sabo; Panama: Bugaba, 2,000- 3,000 ft. I have seen 24 other specimens col- lected from June to October from Panama Chiriqui, Lino; CoSTA RiCA: Osa Peninsula Honduras: Orina del Rio Camgre, Juticalpa Belize: Northern Road, mi 28-45; Mexico San Quintin. Enchophora subviridis Distant Fig. 75 Enchophora subviridis Distant 1887:28, pi. 4, Fig. 17. Type repository: BMNH. Enchophora subviridis vdT. a. Distant 1887:497. Enchophora subviridis distanti Metcalf 1938:358. Length. — Male 22 mm, female 23-25 mm. Ground color green, brown, or yellow. Head process reflexed, then turned up at tip. Wings orange in basal half, brown posteriorly; 8 waxy tufts in costal margin of tegmen, 20 plus in membrane. This species may be separated from all other species but pijrrhocnjpta by the re- curved and bent dorsad head process, and 1988 O'BRIEN; New World Fulcoridae 155 pyrrhocrypta has distinctive, black spots on the pro- and mesonota and red on the hind wings. Distribution. — Syntypes: Panama: Chiri- qui, Bugaba. I have seen six other specimens collected from August to October fiom Panama: Chiriqui, Bugaba, 800-1,500 ft; Costa Rica: Puntarenas Prov., Osa Penin- sula, Rincon; San Vito, Golfito. Enchophora tuba (Germar) Fig. 64 Fulgora tuba Germar 1830:46. Type repository: Lvov. (All types there are lost.) Enchophora tuba (Germar), Burmeister 1845:4. Length. — Female 30 mm. Ground color reddish brown, mottled. Head process elongate, produced in club that at apex is twice average width of process, apex trilobed, brown like rest of process (Fig. 64). Wings red at base, brown posteriorly. No waxy tufts evident. This species may be separated from all other Enchophora by the long, thin head pro- cess with a trilobed, concolorous tip. Distribution. — Type: Brazil. I have seen two other specimens collected from Brazil: Agua Preta, in December. Enchophora tuherculata (Olivier) Fig. 73 Fulgora tuherculata Olivier 1791: 569. Type repository: unknown. Enchophora tuberculata (Olivier), Burmeister 1845: [4]. Enchophora tuberculata sub. sp. fuscomaculata Lallemand 1956:6. Type repository: FSAG. Enchophora parvipennis Walker 1858:30. Type reposi- tory: BMNH. New stjnonyinij. Length. — Male 22 mm. Ground color greenish brown. Head pro- cess recurved toward vertex (Fig. 73). Wings brown, orange in anterior four cells. No waxy tufts evident. This species may be separated from all other Enchophora by the recurved head pro- cess, which differs from pyrrhocrypta and subviridis by not being turned up at the tip. The basal colored area of the hind wing is also more reduced than in any other species; in one specimen it is brown with the four cells slightly paler brown. Distribution. — Type: Suriname {tuhercu- lata), Brazil: Para {parvipennis male). I have seen two other specimens from French Guiana: St. Jean du Moroni; Venezuela: Salto de las Acadencias. Enchophora uniformis, n. sp. Fig. 72 Length. — Male 24-26 mm, female 25-26 mm. Ground color yellow to greenish yellow. Head process scimitar-shaped, shorter than frons below process. Hind wing white with apical third yellowish. Tegmen with margin darker behind stigmal line; this margin with eight spots that may be indicators of waxy tufts. Distinctive markings, brown or black: apex of supraocular flange, three or more dashes along clavus, spots along stigmal line (these are variable; one specimen lacks them all). This species has a smaller process than viridipennis and prasina, the only two other species with green or yellow tegmen. It lacks the white costal margin of prasina and the yellow-orange base of the hind wing of viri- dipennis. Distribution. — Holotype (male), allotype (female), and 1 paratype (female): Peru: Iqui- tos, F6062, H. Bassler Golln. Ace. 33591 (AMNH). Four other paratypes: PERU: Chau- chamayo [sic], from W. F. H. Rosenberg (NCU, male); Jauja Prov. Satipo, Sept. 1946, A. Mailer Coll., Frank Johnson Donor (AMNH, female).; Dept. Huanuco, Vic. Tingo Maria, Jungle, 670 m, XII- 1939, Felix Woytkowski (AMNH, male); Iquitos, Rio Itaya, Herbert Osborn Colin (OSU, female). Enchophora viridipennis Spinola Fig. 74 Enchophora viridipennis Spinola 1839:225, pi. 11, Fig. 2. Type repository: NHMV. Enchophora erninenta Schmidt 1909:187. Type reposi- tory: IZW. New synonymy. Length. — Male 22 mm. Ground color yellow brown (given as green in original description). Head process scimi- tar-shaped, shorter than most other species (Fig. 74). Hind wings yellow-orange at base, posteriorly white with ochre veins. More than 10 waxy tufts present along slightly infuscate margin of tegmen. This species may be separated from all other Enchophora but uniformis by the col- oration of the hind wings. In the specimen examined the distinctive color markings are 156 Great Basin Naturalist Memoirs No. 12 the shiny, black apex of the head process, a black spot on each side of the frons before the eye, five very small, red marks on the tegmen, red longitudinal lines on the clypeus, a diago- nal, red, marginal band on each side of the pronotum, and the browner wing margin. Schmidt (1909) described the clypeus as hav- ing blood-red longitudinal stripes, the apex of the clypeus, the pronotum and scutellum, the legs, and the five spots on the tegmen blood red. The specimen at hand has fainter col- oration. Spinola (1839) said that the spots on the tegmen are black. I did not have a speci- men to compare with either of these types when I visited their respective museums; so, I am placing them in synonymy by utilizing their descriptions and illustrations and my notes. Distribution. — Holotype: Brazil: Pebas (eminenta) , locality unknown {viridipennis). I have seen one specimen from BRAZIL: Jacarea- canga, collected in February. Enhydria Walker Figs. 36-39 Enhydria Walker 1858:44. Type-species: Dichoptera tesellata Walker, by monotypy. Ulubra Stal 1866:133. Type-species: U. hrachialis Stal, by monotypy. Synonymized by Stal 1870:286. Ulubra resurrected by O Brien 1985:662. [Error] Netc synompny. Medium-sized, brown insects with trans- parent tegmina, 19-22 mm long. Head pro- cess recurved, laterally compressed, gradu- ally narrowing to apex. All five ventral carina and two dorsolateral carina present, com- plete; dorsal median carina present at base of process, extending as far as dorsal notch. Pre- ocular flange horizontal, small. Pronotum with white nodules. Tegmen with short spines along veins on upper surface, M not pectinate; cross-veins along costal margin dense, diago- nal; other cross-veins not dense, about same diameter as longitudinal veins; most cells rectangular. Female with eighth and ninth abdominal tergites subequal. This genus may be separated by the short spines on the veins on the upper surface of the tegmen and the transparent, brown hind wing. The dorsal head measurement and head/pronotal ratio were not made in this genus because the length varies greatly with the position of the head process. I resurrected Ulubra as a valid genus on the basis of illustrations and notes taken from a specimen so identified at the British Mu- seum. Recently, I discovered a photograph of the type of Ulubra brachialis taken in Stock- holm; and it is a synonym of tessellata. The genus I resurrected as Ulubra is renamed he- lowAS Stalubra(q. v.). All species o( Enhydria have a similar color pattern: brown insects with red spots on the pronotum, black markings on the head pro- cess, etc. They may be identified to species by the length and shape of the head process. Key to the Species oi Enhydria 1. Head in lateral view with distance from fronto- clypeal suture to dorsal notch greater than length from dorsal notch to apex, thus process short (Figs. 37, 38) 2 — Head in lateral view with distance from fronto- clypeal suture to dorsal notch on process less than length from dorsal notch to apex, thus pro- cess long (Fig. 36) longicornuta Lallemand 2(1). Venter of process black; thickness of process in lateral view about 1/2 width of eye (Fig. 38) . . . cicadina Gerstaecker — Venter of process pale with brown spots, only tip black; thickness of process in lateral view sube- qual to width of eye (Fig. 37) tessellata (Walker) Enhydria cicadina Gerstaecker Figs. 38, .39 Enhydria cicadina Gerstaecker 1895:38. Type repository: EMAU. Length. — Male 19.5-21.5 mm. This species has the shortest head process (Fig. 38). Male genitalia. — The two pairs of inflat- able lobes have darkened, tuberculate tips, which the other species do not have. Distribution. — Syntypes: Brazil: Bahia. I have seen six other specimens collected in December from Brazil: Caviuna, Iguazu Falls, and Tijuco Preto. Enhydria longicornuta Lallemand Fig. 36 Enhydria brachialis i. longicorntita Lallemand 1960:106. Type repository: NRS. Enhydria longicornuta Lallemand, O Brien 1985:661. Length— Male 20-22 mm, female 20-22 mm. This species has the longest head process (Fig. 36). Male genitalia. — ^The ventral lobe in ven- tral view has a slight median projection and is 1988 O'BRIEN: New World Fulgoridae 157 about half as wide as the claspers. Distribution. — Holotype: Brazil. I have seen five other specimens collected from Brazil: Manaus, Serro do Navio, and Trin- idad. Enhydria tessellata (Walker) Fig. 37 Dichoptera tessellata Walker 185L305. Tvpe repositorv: BMNH. Enhydria tessellata (Walker), Walker 1858:44. Enchophora brachialis Stal 1862: L Type repository: NHS. New sijnonytmj. Enhydria brachialis (Stal), da Costa Lima 1935:497. Length. — Male 19-20 mm, female 21-22 mm. This species has a medium-length head pro- cess (Fig. 37). Male genitalia. — Ventral lobe of the aedeagus with median emargination, ventral lobe about as wide as claspers. Distribution. — Holotype: Brazil: Rio de Janeiro (brachialis male); Para {tessellata fe- male). I have seen 11 other specimens col- lected in January, March, June, and October from Brazil: Corcovado, Faz Morelandia, Corupa, and Para; PERU: Tingo Maria. Fulgora Linnaeus Figs. 77-92 Fulgora Linnaeus 1767:703. Type-species: F. laternaria (Linnaeus), subsequent designation by Interna- tional Commission on Zoological Nomenclature, Opinion 322 (1954:187). Large insects, 65-105 mm in length, yel- lowish brown mottled with black markings and white, waxy areas, head .25 to .28 length of insect. Head process porrect, terete, in- flated, resembling a peanut from above and an alligator head, including false eye spots, in lateral view. Preocular flange large, horizon- tal. Tegmen with M not pectinate, most cross- veins forming many-sided, almost round cells. Female with abdominal tergites eight and nine subequal in length. At the museum in Sao Paulo, 1 examined the specimens used by da Fonseca (1926) in his revision; these did not fit his key. B. V. Ridout (personal communication) examined previously used characters and several new features, including male genitalia and the re- sults of morphometric analyses, and found the best way to identify species in this genus was to match specimens to the accompanying il- lustrations of head shapes. His fllustrations are labeled lectotype except for la7npetis (holotype), crocodilia, and lucifera. I give him full credit for this section and am convinced that this is the best that can be done at this time. Any errors are mine. Ridout suggested that species with narrow heads might live in dry forests, the others in rain forests. Janzen (1983) described the be- havior of F. laternaria, including at least 100 specimens seen resting on Hymenaea cour- haril or guapinol, but questioned whether this plant is the host or only host, as it does not extend throughout the range of the insect spe- cies. Fulgora castresii Guerin-Meneville Figs. 81, 82 Fulgora castresii Guerin-MeneviWe 1837:3, pis. 173, 174, Figs. 3, 4. Distribution. — Type: Mexico. I have seen 10 specimens collected from July through September from MEXICO: Hustusco, Nuevo X-Can [sic], Tamazunchale; Peru: Tingo Maria: Panama: Barro Colorado Island. Fulgora cearensis Forseca Figs. 87, 88 Laternaria cearensis Fonseca 1932:3, Figs. 1, 2, 3h. Fulgora cearensis Fonseca, Metcalf 1947:219. Distribution. — ^Type: Brazil. I have seen two specimens from Trinidad : Curepe and Maraval, collected in September. Fulgora crocodilia Brailovsky & Beutelspacher Figs. 85, 86 Fulgora crocodilia Brailovsky & Beutelspacher 1978:176, pi. 3. Distribution. — Type: Mexico. I have seen three specimens collected from MEXICO: Chamela, in September and November. Fulgora graciliceps Blanchard Figs. 83, 84 Fulgora graciliceps Blanchard 1849:pl. 2, Fig. 1. Laternaria orthocephala Fonseca 1926:493, pi. 6, Figs. 1, 2. New synonymy, Ridout. Fulgora orthocephala Fonseca, da Costa Lima 1942:42. Distribution. — Type: Brazil (orthoceph- ala, graciliceps unknown). I have seen three specimens from Bolivia: Requena and Warnes, and PERU: Tingo Maria, collected in November. 158 Great Basin Naturalist Memoirs No. 12 14.2mm I \ nn 19. Vuloin-ii Itiuwctis Burnieister; Figs. 77-101. Head, dorsal view (odd numbers)^ ^t^J^;;;: ^8^4 T^^S Bbn'wl; SS^SG, F. crocodiUa 79-80, F.iaternam(L.); 81-82, F.casfrm.Guenn-Menevlle^ 8.3 M j^ ^^_y^ ^ riofirandensts BraiWsky & Beutelspacher; 87-88, F. -^Xvan Xzt 94 1 voi^^ 95, A. u^ankinsi OBnen; 96^A. Fonseca. Hind wings oi Arnycle:93, A. '^""f^ll"^^^^^^^^^ 99 a brevis O'Brien; 100, A. f.i»Ki«iror««e K & K, pinyonae K & K; 97, A. vernalis Manee; 98, A. grandts O Bnen, 99, A. 101, A. amabilis (Westwood). 1988 OBrien: New World Fulgoridae 159 Fulgora lampetis Burmeister Figs. 77, 78 Fulgora lampetis Burmeister 1845:3. Distribution. — Type: Brazil. I have seen 11 specimens collected throughout the year from Bolivia: La Cordillera Santa Cruz; Peru: Tingo Maria; Brazil: Obidos, Sinop; Panama: Barro Colorado Island; Costa Rica: Sarapiqui; Nicaragua: Leon; Honduras: La Ceiba. Fulgora laternaria (Linnaeus) Figs. 79, 80 Cicada laternaria Linnaeus 1758:434. Fulgora laternaria Linnaeus, Linnaeus 1767:703. Fulgora servillei Spinola 1839:214. New synonymy. Rid- out. Distribution. — Type: Tropical America {laternaria); Brazil {servillei). I have seen live specimens collected in January, and April through July in Panama: Barro Colorado Is- land; Honduras: Armenia, La Ceiba; Mex- ico: LosTuxtlas. Fulgora lucifera Germar Figs. 89, 90 Fulgora lucifera Germar 1821:100. Fulgora mitrii Burmeister 1867;xxiii, Berg 1879:178. Distribution. — ^Type: Brazil. I have seen 16 specimens collected from November through February from Argentina: Abra Grande Iran; Bolivia: Coipa, El Torno, Mora Abadolz., Montero, Santa Cruz, andWarnes. Fulgora riograndensis Fonseca Figs. 91, 92 Laternaria servillei riograndensis Fonseca 1926:486, pi. IV, Fig. 1. Laternaria riograndensis Monte 1932:22. Fulgora riograndensis Fonseca, Metcalf 1947:232. Distribution. — Type: Brazil. I have seen one specimen from Panama: Barro Colorado Island, collected in June. Odontoptera Carreno Figs. 44, 116, 117 Odontoptera Carreno 1841:275. Type species: O. spectabilis Carreno, by monotypy. Medium-sized, green or brown insects, 27-37 mm long, head 1/6 to 1/3 length of insect. Head process porrect, terete, elongate triangular; apex bent dorsad; lateral carinae of dorsum and venter and pleural carinae all visi- ble. Preocular flange small, diagonal or hori- zontal. Ventropleural carina of pronotum present, dorsopleural carina absent, repre- sented by low ridge. Tegmen with apex trun- cate; anal angle angulate; dark spot along stig- mal line; M greatly expanded, 16-19 branches at level of stigmal line; cross-veins incomplete in corium, forming rectangular cells in membrane. Female with eighth and ninth abdominal tergites subequal in length. Cathedra serrata and the two species of this genus are the only Fulgoridae with head pro- cesses which I have never seen misidentified. Key to the Species oi Odontoptera 1. Head about 1.5 times as long as pro- and mesonota combined; tegmen green with nar- row, apical, brown band; hind wing green with brown and orange markings .... carrenoi Signoret — Head almost as long as pro- and mesonota com- bined; tegmen orange-brown basally, brown be- hind pale, thin, nodal band; hind wing white at base, brown apicalUy spectabilis Carreno Odontoptera carrenoi Signoret Figs. 44, 117 Odontoptera carrenoi Signoret 1849:178, pi. 6, Fig. 4. Type repository: NHMV. Length. — Male 27-28 mm, female 30-33 mm; head length 9.7 mm; ratio, head/prono- tum5.4. Ground color green with brown apical band on tegmen, round, black spot surrounded by white line at apex of clavus on stigmal line (Fig. 44). Hind wing green with two black spots near anal angle, spots surrounded by orange area. This species is green with a more elongate head than the brown spectabilis. Distribution. — Holotype: locality un- known. 1 have seen 26 other specimens, col- lected May to October and February, from: Brazil: Sinop; French Guiana: Montague des Singes near Kourou; 67 km S Cayenne; Panama: Barro Colorado Island, Santa Rita Arriba, Parque Nacional Soberania Frijolito; Costarica: La Selva; Mexico: Los Tuxtlas. Odontoptera spectabilis Carreno Fig. 116 Odontoptera spectabilis Carreno 1841:277, pi. 5, Fig. 2. Type repository: ZMHB. Length. — Female 36-37 mm; head length 5.8 mm; ratio, head/pronotum 2.3. 160 Great Basin Naturalist Memoirs No. 12 Ground color yellow-orange on head, tho- rax, and apical part of tegmina, tegmen be- hind stigmal line brown. Head, thorax, and tegmen with lateral, longitudinal, brown band extending half length of costa. Hind wing white, black along apical margins, tinged with orange at base. This species is brown and has a shorter head than the green carrenoi. Distribution. — Type: America (?). I have seen four other specimens, collected from January to March, from Brazil: Boraceia Field Station and Corupa. Phrictus Spinola Figs. 102-115 Phrictus Spinola 1839:216. Type-specie.s: Fulgora di- adema Linnaeus by nionotypy. Large, reddish brown or yellowish brown insects, rarely yellow and green, 29-53 mm; head 1/8 to 1/4 length of insect. Head process porrect, terete, usually enlarged into trifur- cate apex (Figs. 102-115); dorsolateral carinae each with two pairs of spines, ventrolateral carinae each produced into flange near fronto- clypeal suture and into small, dull spine at junction of frons and clypeus. Preocular flange horizontal, small; supra- and postocular flanges produced into strong spines. Tegmen with M not pectinate, cross-veins reticulate in corium forming square cells behind stigmal line. Female with ninth tergite medially shorter than eighth. These beautiful insects with a name that means "horrible" or "terrible" may be identi- fied by the shape of the apex of the head process and the waxy tufts on it, the color pattern of the tegmina, and the color of the hind wings. Galdwell (1945) also used the me- dian notch in the caudal margin of the prono- tum and variation in the furcation of the me- dian carina around the notch, which I found difficult to quantify. The female first valvulae, as he illustrated, also serve to identify difficult specimens. The apex and venter of the head process are usually reddish orange or reddish brown with a white stripe down the dorsum reaching the apex of the pronotum. There are several color patterns on the tegmina which will be discussed under species descriptions. I have illustrated the head process with the insect at approximately a 45-degree angle in order to provide a good view of the apex. Because this makes the length of the head and process difficult to determine, I have included a head length and head/pronotum ratio. Key to the Species of Phrictus 1. Head process flattened apically, transversely arcuate, lacking definite apical teeth (Fig. 1 12); base of hind wing yellow with transverse bands of brown maculae auromaculatus Distant — Head process with 3 to 5 apical teeth (Figs. 106-111, 113, 114); hind wing lacking trans- verse bands of brown maculae 2 2(1). Head process with 5 apical teeth (Fig. 104) quinquepartitus Distant — Head process with 3 apical teeth 3 3(2). Hind wing with large (ca 2 mm), hyaline, apical spots, visible from below with wing unspread 4 — Hind wing without hyaline, apical spots, but small pruinose areas sometimes present, visi- ble when wings are spread 5 4(3). Hind wings yellow or orange at base; head process with 3 black spots dorsally just behind apical area (Fig. 113) ... . xanthopterus Schmidt — Hind wing red at base; head process with no black markings near apex, but diamond- shaped, red area in same position (Fig. 107) oceUatus Signoret 5(3). Basal area of hind wing yellow or yellow- orange moebiusi Schmidt — Basal area of hind wing red or reddish orange 6 6(5). Tegmen with continuous, broad, yellow, transverse fasciae present 7 — Tegmen lacking transverse fascia, or, if present, interrupted medially 8 7(6). Tegmen brown, transverse fascia yellow only diligens, n. sp. — Tegmen scarlet with green veins, anterior edge of fascia partially bordered with black tripartitus Metcalf 8(6). Transverse fascia absent, apical trident of head process 2/3 as wide as distance between supraocular spines (Fig. 108) delicatus, n. sp. — Transverse fascia interrupted medially; apical trident of head process wider (Figs. 105, 106, 109, 110) 9 9(8). Head process with apical teeth broader than head (Fig. 105); hind wing with black or fus- cous area covering apical three-lourths recalls Caldwell — Head process with apical teeth not wider than head (Figs. 106, 109, 110); hind wing with black or fuscous areas covering apical third . 10 10(9). Tegmen green in basal two-thirds, with a few small, roinid, red or orange maculae; apical third with large, brown spots hoffmannsi Schmidt 1988 OBrien: New World Fulgoridae 161 Figs. 102-117. Head and pronotum, dorsal view, drawn with head process at 45-degree angle so that the apex is horizontal; 102, Phrictus diadema L.; 103, P. tripartitus Metcalf; 104, P. quinquepartitus Distant; 105, P. regalis Caldwell; 106, P. punctatus Caldwell; 107, P. ocellatus Signoret; 108, P. delicatus O'Brien; 109, P. minutacanthis Caldwell; 110, P. hoffmansi Schmidt; 111, P. moebiusi Schmidt; 112, P. auromactdatus Distant; 113, P. xanthopterus Schmidt; 114, P. diligens O'Brien; 115a, P. diadema (L.) and 115b, P. triportitus Metcalf (head, lateral view); 116, Cathedra spectabilis Carreno; 117, O. carrenoi Signoret. 162 Great Basin Naturalist Memoirs No. 12 — Tegmen brown or reddish brown, sometimes with yellow spots 11 11(10). Tegmen some shade of reddish brown, with pink, calloused areas present, especially basally; head process toothed on ventral sur- face (Fig. 115a) diadeina (Linnaeus) — Tegmen brown, maculate with yellow; head process not toothed ventrally 12 12(1 1). Combined length of head process and head as long as pronotum; expanse of trifurcate apex equal to distance between ocular spines (Fig. 109) minutacanthis Caldwell — Combined length of head process and head longer than pronotum; expanse of apex much greater than distance between ocular spines (Fig. 106) punctatus Caldwell Phrictus auromaculatus Distant Fig. 112 Phrictus auromaculatus Distant 1905:672. Type reposi- tory: BMNH. Phrictus notatus Lallemand 1931:188. Type repository: FSAG, NRS. New synonymy. Length. — Male 29-32 mm, female 34-35 mm; head length 3.7 mm; ratio, head/prono- tum 1.0. Head process semicircular at apex, not di- vided into teeth, orange-red; three round, waxy plates on each side (Fig. 112). Tegmen evenly pale brown mottled with yellow, veins dark brown in corium, no transverse fascia. Hind wings yellow at base, brown posteriorly; two maculate, brown bands from anal area running into yellow area. This species is easily separated by the apex of the head not being divided into spinelike segments. It may be verified by the yellow base of the hind wings and the evenly colored tegmina. Distribution. — Syntypes: Bolivia (auro- maculatus), Ecuador (notatus). I have seen five other specimens collected from Bolivia: Santa Cruz; Prov. Sara [sic], 450 m; Brazil: Sinop; Peru: Jauja Prov., Satipo. Phrictus delicatus, n. sp. Fig. 108 Length. — Female 36 mm; head length 3.7 mm; ratio, head/pronotum 1.3. Head process tripartite, lateral portions di- rected anterad, not laterad, smaller than that oi minutacanthis; head pale brown (Fig. 108). Mesonotum with three pairs of dark spots (pin is placed through apex; there may or may not be one there). Tegmen evenly colored, no fascia. Hind wings red basally, posteriorly pale brown. This species may be identified by the pale brown, narrow head process, without a white dorsal stripe and red venter and apex. The pale apex of the hind wings separates this species from all but moebiusi. The evenly col- ored tegmen, broader and less rooflike than most of the species, is similar to that of auro- maculatus and ocellatus. HoLOTYPE (female). — Brazil: Amazonas, S. Paulo de Olivenca, Vn-32, A. Mailer Colin., Frank Johnson Donor (AMNH). The printed label says S. P. Olivenca/ Amazonas/ Brasil Vn-32. The handwritten label under- neath gives the more complete data. Phrictus diadema (Linnaeus) Figs. 102, 115a. Fulgora diadema Linnaeus 1767:703. Type repository: UZIU. Phrictus diadema (Linnaeus), Spinola 1839: 219. Length. — Male 38 mm, female 45 mm; head length 8 mm; ratio, head/pronotum 2.0. Apex of head process trifurcate, three or four pairs of lateral, waxy plates and one me- dian oval (Fig. 102). Ventral margin of head process in lateral view sinuate. Hind wings red at base. Tegmina mottled with red and brown in large splotches, transverse, yellow- ish fascia narrow, not bordered anteriorly in black. This species most closely resembles regalis in the mottled color pattern of the tegmen and pale, transverse fascia and the sinuate ventral margin of the head process in lateral view, but regalis has a much wider, trifurcate apex on the head process, exceeding the distance be- tween the tips of the supraocular spines (Fig. 105). In diadema the spines and apex are about the same width. Distribution. — Type: India ? [error]. I have seen two other specimens collected in November from BRAZIL: Linhares. Phrictus diligens, n. sp. Fig. 114 Length. — Female 40-43 mm; head length 6 mm; ratio, head/pronotum 1.3. Apex of head process trifurcate, narrow band of waxy plates each side (Fig. 114). Hind wings red at base. Tegmen evenly brown with unbroken clear transverse fascia. This species may be identified easily by the 1988 O'BRIEN: New World Fulgoridae 163 browii tegmen with the unbroken transverse band. I name it diligens, which means prim, because it is more conservative in color and shape of head process than the other species. It is most closely related to diadema and re- galis in the color pattern of the tegmen. HoLOTYPE (female). — Colombia: 1,500 ft, Anchicaya, 26-VII-1970, J. M. Campbell (LOB). Paratype (female): Colombia: Dept. El Valle, Anchuaya, 200-300 mi, 27-IV- 1969, Gonin-H Balazar-S. Draachsler Rec. (MNHP). Phrictus hoffmannsi Schmidt Fig. 110 Phrictus hoffmannsi Schmidt 1905:338. Type repository: IZW. Length. — Male 37 mm, female 48 mm; head length 7 mm; ratio, head/pronotum 1.0. Apex of head process trifurcate, large, squarish, waxy plate area on each side (Fig. 110). Hind wings red at base. Tegmina green mottled with yellowish orange in large splotches, clavus predominately orange, broad, transverse, yellow fascia interrupted by brown spots medially and toward commis- sure, membrane yellowish brown with brown oval spots. This species may be distinguished easily by its bright green and yellowish orange tegmen. Distribution. — Holotype (female): Peru: Chanchamayo. I have seen four other speci- mens collected in January from Peru: Tingo Maria; Ecuador: Guayaquil, Loja. Phrictus minutacanthis Caldwell Fig. 109 Phrictus jninutacanthis CaldweW 1945:180, pi. 7, Figs. 2, 16, pi. 8, Fig. 2, pi. 9. Type repository: USNM. Length. — Female 36 mm; head length 4.8 mm; ratio, head/pronotum 1.5. Apex of head process trifurcate, small; waxy plate on each side; pale red, midline near apex with reddish triangle (Fig. 109). Hind wings red at base. Tegmina light brown evenly mot- tled with yellow, transverse yellowish fascia indistinct, broken. This species may be identified by its small head. It resembles xanthopterus and punc- tatus most closely in the light brown tegmen with yellow markings, and ocellatus and punc- tatus in having the red subapical triangle on the pale dorsum of the head process. Distribution. — Holotype: Peru: Chaqui- mayo. I have seen only the type. Phrictus moebiusi Schmidt Fig. Ill Phrictus moebiusi Sclimiclt 1905:335. Type repository: ZMHB. Phrictus sordidus Caldwell 1945:180, pi. 7, Figs. 3, 20, pi. 8, Figs. 1, 13, pi. 9. Type repository: USNM. New synonymy. Length — Male 40 mm, female 43-48 mm; head length 6 mm; head/pronotal ratio 1.2. Apex of head process trifurcate, large, waxy plate on apex (Fig. 111). Hind wings yellow at base, posterior pale brown. Tegmina brown evenly mottled with yellow, transverse fascia pale, broad, interrupted by two brown spots in middle and near commissural margin. This is one of three species with yellow coloration at the base of the hind wing. P. auromaculatus has a head process without apical spines, and xanthopterus has white spots in the posterior brown area of the hind wing. The type of sordidus is a pale specimen that agrees in all external characters with moe- biusi. Distribution. — Syntype (female): Colom- bia: Antioquia (moebiusi); Ecuador (male holotype sordidus). I have seen four other specimens collected in January from "New Granada" and Ecuador: Route Mono-Los Banos, km 37, 2,385 m. Phrictus ocellatus Signoret Fig. 107 Phrictus ocellatus Signoret 1855:v. Type repository: nhmv. Length. — Male 35 mm, female 42 mm; head length 6.0; ratio, head/pronotum 1.9. Apex of head process trifurcate; large, waxy plates on each side; red diamond and median line along pale dorsum of process (Fig. 107). Hind wings red at base, white spots in brown posterior area. Tegmina evenly mottled with pale yellow and red brown, no fascia. Two species have hind wings with white spots in the brown posterior area, xan- thopterus with a yellow base and ocellatus with a red base. Distribution — Holotype (female): Co- lombia: Fusagasuga. I have seen two other specimens collected in June from VENEZUELA: Rancho Grande. 164 Great Basin Naturalist Memoirs No. 12 Phrictus punctatus Caldwell Fig. 106 Phrictus punctatus Ca\dwe\\ 1940:181, pi. 7, Figs. 12, 23, pi. 8, Fig. 6, pi. 9. Type repository: USNM. Length. — Female 40 mm, head length 6.5 mm; ratio, head/pronotum 1.4. Apex of head process trifurcate, with trans- verse, waxy plates; triangular red area on mid- line behind apex (Fig. 106). Hind wings red at base, pale brown posteriorly. Tegmina brown mottled with yellow in large splotches, clavus predominantly yellow; transverse, yellowish fascia narrow, interrupted medially; diagonal yellow line from clavus to costal margin. In coloration, the tegmen o( punctatus most closely resembles xanthoptertis, while the color of the head process resembles oceUatus and minutacanthis. Distribution. — Holotype (female): Pana- ma: Bugaba. Paratype: El Volcan. I have seen two other specimens, one without data, the other collected from CosTA RiCA: Palo Verde. Phrictus quinquepartitus Distant Fig. 104 Phrictus quinquepartitus Distant 1883:24, pi. 4, Fig. 8. Type repository: BMNH. Length. — Male 40-42 mm, female 45-48 mm; head length 9.3 mm; ratio, head/prono- tum 2.06. Apex of head process trifurcate, one median and three lateral pairs of waxy tufts in apex (Fig. 104). Hind wings red at base. Tegmina red with green veins; transverse, yellowish fascia bordered anteriorly in black. This species is closely related to tripartitus in the color of the tegmen but differs in the apex of the head process. Distribution. — Syntypes: Panama: Buga- ba, 3,000-4,000 ft; Colombia: Bogota. I have seen nine other specimens collected in Feb- ruary, June, August, October and December from Panama: Barro Colorado Island, Code, Chiriqui; CoSTA RiCA: La Selva. Phrictus regalis Caldwell Fig. 105 Phrictus regalis Ca\dwe\\ 1945:183, pi. 7, Figs. 7, 19, pi. 8, Fig. 8, pi. 10. Type repository; USNM. Length. — Male 42 mm, female 49-53 mm; head length 10; ratio, head/pronotum 2.3. Apex of head process trifurcate, three waxy tufts on each side, two larger ones medially. semicircular carinate area astride median carina (Fig. 105). Hind wings red at base. Tegmina mottled with red and brown in large splotches; transverse, yellowish fascia nar- row, not bordered anteriorly in black. This species is very similar to diadema in the color of the tegmen, but the apex of the head process is much wider in regalis (Figs. 105, 102). Distribution. — Holotype (female): French GuiANA: Maroni River, near Duserre. I have seen three other specimens collected in March from French Guiana: St. Jean du Ma- roni; Suriname: Nassaugebergte. Phrictus tripartitus Metcalf Figs. 103, 115b Phrictus tripartitus Metcalf 1938:365, pis. 20, 21. Type repository: AMNH. Length. — Male 35-40 mm, female 43-45 mm; head length 9.4 mm; ratio, head/prono- tum 1.8. Apex of head process trifurcate, three round, waxy plates and one oval on each side (Fig. 103). Head process sinuate ventrally in lateral view (Fig. 115b). Hind wings red at base. Tegmina red with green veins; trans- verse, yellowish fascia bordered anteriorly in black. This species most closely resembles quin- quepartitus in the color of the tegmen, but the species may be separated by the shape of the apex of the head process. Some specimens of tripartitus do have five spines, but the inter- mediate spines are smaller than those of quin- quepartitus (Fig. 104) and the shape of the other teeth is like that illustrated for triparti- tus (Fig. 103). Distribution. — Type: Belize. I have seen 21 other specimens collected from August through October from Belize: 24 mi SE Bel- mopan, 10-11 mi S Georgeville; Honduras: La Ceiba; MEXICO: Nuevo X-Can [sic], San Quintin, Yucatan. Phrictus xanthupterus Schmidt Fig. 113 Phrictus xanthopterus Schmidt 1910:144. Type reposi- tory: IZN. Length. — Female 39-41 nnn; head length 5 mm; ratio, head/pronotum 1.0. Head process with apex trifurcate, without waxy plates, orange-red, with three triangu- lar, black spots behind apex (Fig. 113). Hind 1988 O'BRIEN; New World Fulgoridae 165 wings yellow at base with white spots in brown posterior area. Tegmen brown with yellow markings, half of clavus predominantly yellow; transverse, yellow fascia interrupted medially. This species is one of the most distinctive with the hind wings yellow at base and with white spots in the apical brown area and the three black, subapical spots on the head pro- cess. The tegmen most closely resembles punctatus. Distribution. — Holotype (female): Ecua- dor: Canelos. I have seen six other specimens collected in June and July and December from Peru: Cusco; Sta. Isabel; Route Olmoc- Tarapoto, km 385, 1,800 m; Tingo Maria; Ecuador: Lumbagui. Rhabdocephala Van Duzee Fig. 23 Rhabdocephala Van Duzee 1929; 190. Type-species; R. brunnea Van Duzee, original designation. Small, brown, narrow insects, 12-16 mm long; head about 1/3 length of insect. Head process porrect, terete, sides parallel; apex slightly widened; dorsal lateral carinae com- plete, ventral incomplete, present only in middle of process. Preocular and supraocular flanges absent. Tegmen with M not pec- tinate, cross-veins reticulate, apical margin angled at apex of clavus, forming angulate, not smoothly rounded, apical margin. Ninth ab- dominal tergite 1.5 times length of eighth. These small insects may be separated from other small Fulgoridae by the simple cephalic process with no dorsolateral undulate carinae as in Scolopsella and the head process not dorsoventrally flattened as in Amycle. Rhabdocephala brunnea Van Duzee Fig. 23 Rhapdocephala brunnea Van Duzee 1929; 191. Type repository; CAS. Length. — Male 12-14 mm, female 14.5- 16 mm; "head 5.2 mm; ratio, head/pronotum 2.1. Tegmen brown mottled with white. Hind wing red in basal third, brown posteriorly. Distribution. — Holotype (female): USA: Arizona. I have seen 15 other specimens col- lected from May to October from USA: ARI- ZONA: Pima, Cochise, and Santa Rita counties; Mexico: Sonora, 15 mi S Hermosfllo. Two specimens were collected on Baccharis sarothroides Gray. Scolopsella Ball Figs. 8-13, 24-25 Scohypsella Ball 190.5; 118. Type-species; S. reticulata Hall by original designation. Small, brown insects, 14-21 mm long, head .23-. 28 length of insect. Head process por- rect, terete, very slightly enlarged at apex; dorsolateral carinae undulate; median and lat- eral ventral and marginal carinae complete on process and frons. Preocular flange indistinct. Tegmen with M pectinate (3 branches), com- missural margin emarginate between claval apex and wing tip. This genus of small, brown insects may be separated from other similar genera by the straight head process with dorsolateral mar- gins crenulate or undulate and without the apex greatly expanded. The head process is not as unadorned as Rhabdocephala, lacks the transverse wrinkles o( Aphrodisias , and is not flattened as in Amycle. Diareusa specimens are much larger insects with much shorter heads. Key to the Species oi Scolopsella 1. Length of anal flap of male greater than width at apical expansion (Fig. 23); apical margin of pygofer emarginate ventrally (Fig. 25) reticulata Ball — Length of anal flap of male less than width at apical expansion (Fig. 20); apical margin of pygofer smoothly convex (Fig. 22) mexicana O'Brien Scolopsella niexicana O'Brien Figs. 8-10, 24 Scolopsella niexicana O'Brien 1985:73. Type repository; LOB. Length. — Male 14-15 mm, female 17-21 mm; head length 5.6 mm; ratio, head/prono- tum 3.7. Tegmen various shades of brown, cross- veins and reticulations pale, some apical cells white. Structural characteristics common to both species given in generic description. Scolopsella reticulata and S. mexicana may be separated by the shape of the anal flap, which can be seen without dissection, the applicable characters being given in the key. Once males are identified in this way, females may often be associated with them because the golden brown reticulata is slightly larger 166 Great Basin Naturalist Memoirs No. 12 and the head process is wider, and mexicana is darker. Distribution. — Type: Mexico. I have seen seven other specimens collected from June through September from Baja Califor- nia. Scolopsella reticulata Ball Figs. 11-13, 25 Scolopsella reticulata Ball 1905:118. Type repository: CAS. Length. — Male 14-16 mm, female 17-19 mm; head length 5 mm; ratio, head/pronotum 4.1. The structural characters and coloration do not differ sufficiently from S. mexicana to de- termine the species without either using the shape of the male anal flap or comparing the two species as discussed above. Distribution. — Holotype: Arizona. I have seen other specimens collected from April to November in Arizona: Maricopa, Pima, Pinal, and Santa Cruz counties; California: Joshua Tree National Momument, Riverside or San Bernardino County. Sinuala, n. gen. Figs. 43, 46-48 Type-species: Sintiala stall O Brien. Reddish brown, medium-sized insects, 19-21 mm long, head .12 to .15 length of insect. Head process porrect, terete, en- larged at apex; vertex truncated triangle, basal angle rounded, supraocular lobes variable. Frons with lateral carinae diverging anteriorly to form process; lateral margins expanded near frontoclypeal suture, ridge from lateral carina to margin at this expanded area. Frons and clypeus about in same plane, frons not raised above clypeus; clypeus medially and marginally carinate. Preocular flange length- ened into pleural carina. Pronotum without raised median carina, lateral fossettes and two pleural carinae present; apex of mesonotum with striate triangle, delimited by carinae, not high ridge. Tegmen with M not pectinate, clavus open, longitudinal veins angulate, cross-veins variable. Fore and mid femora ex- panded. Posterior tibiae with eight lateral spines. Female abdominal tergite nine twice length of eight. This genus is similar to Aphrodisias , from which it differs by the costal margin of the tegmen sinuate and the head process not transversely ridged. Key to the Species oi Sinuala 1. Supraocular flange expanded, height equal to width of eye; tegmen with 10 strong tubercles tuberculata, n. sp. — Supraocular flange smaller than above; tegmen with tubercles evanescent 2 2(1). Vertex as long as broad (Fig. 46); hind wing red at base stali, n. sp. — Vertex 1.7 times longer than broad (Fig. 48); hind wing pink at base sclimidti, n. sp. Sinuala stali, n. sp. Fig. 46 Length. — Male 19-20 mm; head 2.3 mm; ratio, head/pronotum 1.4. Head process short, vertex as long as broad. Hind wing red at base, narrow anterior mar- gin and broad posterior area brown, apical angle brown with some white spots and crossveins. This species may be easily separated from tuberculata, which it resembles in wing color and head length, by the small, supraocular flange oi stali. I name this species in honor of Carl Stal, who provided a fine basis for generic work in this family. Holotype (male). — Honduras: Lago Yojoa., 6-VI-1976, J. V. Mankins, coll. (LOB). Paratype (male): El Salvador: Alfredo Mar- tinez Cuestas, no further data (NCSR). Sinuala schmidti, n. sp. Fig. 47 Length. — Female 21.5 mm; head 3.3 mm; ratio, head/pronotum 2.2. Head process elongate, vertex 1.7 times as long as broad. Hind wing pink at base, then white; anal margin pale brown, posterior mar- gin with cells infuscate, veins white; apical third brown with white lines following each cross- vein. This species may be separated from the others by the long head and the pale base of the hind wings. I name this species in honor of Edmund Schmidt, who had an excellent eye for species and a gift for describing the most significant characters precisely. HoLOT\'PE (female): BELIZE: Corazol, 1 mi NW Corozal, 12-VIII-1977. C. W. and L. B. O'Brien and Marshall (LOB). Sinuala tuberculata, n. sp. Figs. 43, 48 1988 O'BRIEN: New Would Fulgoridae 167 Length. — Female 21 mm; head 2.6 mm; ratio, head/pronotum 1.6. Head process short, vertex with postocular flange raised above vertex equal to width of eye, vertex as long as broad. Hind wing red in basal half, anal area brown, apical 2/5 brown with some white spots and cross-veins. Five strong tuberculations on veins in anterior half of tegmen, five smaller in apical half. This species may be separated easily from the others by the strong tubercles on the teg- men and the strong supraocular flange. Holotype (female): Costa Rica: P. R. Uh- ler Colin. (NCSR). Stalubra, n. gen. Figs. 1-3, .5, 6 Type-species: Stalubra brunnea O'Brien. Medium-sized, brown or reddish, transpar- ent-winged insects, 19-28 mm long, head . 14 to .19 length of insect. Head process terete, gradually narrowing to apex, upturned at apex. Vertex twice as long as broad, elongate medially (Fig. 1). Lateral carinae of frons par- allel or diverging toward head process, me- dian carina on process and basal 2/3 of frons; frons only very slightly raised above fronto- clypeal suture; lateral margins of frons ex- panded near apex; weak ridge from lateral carina to lateral margins. Clypeus laterally carinate. Preocular flange reduced to slight tubercle on vertical carina joining marginal carinae of frons and vertex (Fig. 3). Pronotum without median carina raised, fossettes and two pleural carinae present; mesonotal apex not depressed or striate. Tegmen with M pectinate (5-7 branches); clavus open; cross- veins undulate, parallel; no small spines on veins, only on wing margins. Posterior tibiae with six lateral spines. Female abdominal ter- gite nine shorter than eight. This genus is similar to Enhydria in the compressed, recurved head process and translucent tegmen; however, it lacks the hairs on the veins of the tegmen and the dense, diagonal cross-veins along the costal margin found in Enhydria, and the head pro- cess is less strongly curved. It differs from Chilobia in the vertex being elongate medi- ally, whereas the vertex in Chilobia is broader than long and emarginate medially (Figs. 51, 54, 60) or hidden by the recurved head pro- cess (Fig. 57). Lallemand's species Enhydria rufula (1966) belongs in this genus, but I cannot tell from his description which it is of two of the three species I have before me. The requested type has not yet arrived, and so these two species will be discussed in Part II. Stalubra brunnea, n. sp. Figs. l-,3, 5, 6 Length. — Male 19-22 mm; head 2.6 mm; ratio, head/pronotum 1.9. Head short, vertex 1.5 times as long as wide. Hind wing clear with brown veins. Teg- men translucent with brown markings and minute red dots along veins, apex pale brown with white circles plus some darker brown spots. Holotype (male). — Brasil [sic]: Mato Grosso, Sinop (12°31'S, 55°37'W), X-1975, M. Alvarenga (LOB). Paratypes (25 male): same data (LOB, MZSP); Guyana: 1 male. Upper Mazaruni River, IX-X-1938, A. S. Pinkus(AMNH). ACKNOWLE DGM ENTS It is with great pleasure that I dedicate this paper to our Latin American colleagues and to Paul Oman, among whose many accomplish- ments is a paper on a U.S. fulgorid (Oman 1936). I thank the American Philosophical Society for a grant to study at European museums (Grant No. 5771, Penrose Fund), without which this work would have been impossible. I am deeply indebted to B. V. Ridout for his work on Fulgora. I greatly appreciate the crit- ical reading of the manuscript by F. W. Mead, G. J. Wibmer, and S. W. Wilson and the sending and examining of types and paratypes of R. C. Froeschner and N. D. Penny. I am also grateful to the following museums for allowing me to examine types and borrow specimens in their collections: AMNH — American Museum of Natural History, New York; BMNH — British Museum (Natural His- tory), London, England; CAS — California Academy of Sciences, San Francisco; DEI — Institute fiir Pflanzenschutzforschung (for- merly Deutsches Entomologisches Institut), Kleinmachnow, Eberswalde, East Germany; EMAU — Zoologisches Institut und Museum, Ernst Morits Arndt Universitat, Greifswald, East Germanv; FMNH — Field Museum of 168 Great Basin Naturalist Memoirs No. 12 Natural History, Chicago, Illinois; FSAG — Faculte des Sciences Agronomiques, Gem- bloux, Belgium; IZW — Instytut Zoologii PAN, Warsaw, Poland; LOB— Lois O'Brien collection, Tallahassee, Florida; MCZ — Museum of Comparative Zoology, Harvard University, Cambridge, Massachusetts; MHNP — Museum National d'Histoire Na- turelle, Paris, France; MRSN — Museo Re- gionale di Scienze Naturali, Turin, Italy; MZF — Museo Zoologico dell' Universita de- gli Studie di Firenze, Florence, Italy; MZSP — Muzeo de Zoologia, Universidade de Sao Paulo, Brazil; NCSU— North Carolina State University, Raleigh; NHMV — Natur- historisches Museum, Vienna, Austria; NRS — Naturhistoriska Riksmuseet, Stock- holm, Sweden; OSU — Ohio State University, Columbus; UCB — University of California, Berkeley; UMO — University Museum, Ox- ford University, Oxford, England; UNAM — Instituto de Biologia and Museo de Zoologia, Universidad Nacional Autonoma de Mexico, Mexico City, D.F.; USNM— United States National Museum, Washington, D.C.; UZIL — Universitets Zoologiska Institut, Lund, Sweden; UZIU — Universitets Zoolo- giska Institut, Uppsala, Sweden; ZMHB — Zoologisches Museum, Humboldt Universi- tat. East Berlin, East Germany; ZMUC- Zoologisk Museum, Universitets Copen- hagen, Denmark; ZSBS — Zoologische Samm- lungen des Bayerischen Staates, Munich, East Germany. Literature on New World Fulgoridae Since Metcalfs Catalog (a complete bibliography will be included with the final part.) Brailovsky, a., H. and C. R. Beutelspacher B 1978. Una nueva especie de Fulgora Linneo (Ho- moptera: Fulgoridae) de Mexico. An. Inst. Biol. Univ. Nal. Auton. Mexico, Ser. Zool. 49: 175-182. Dlabola, J , AND F Zayas 1980. Eine neue Fulgoride der Gattung javiaicastes von Cuba (Homoptera, Auchenorrhyncha). Acta Faun. Entomol. Mus. Nat. Pragae 16: 83-84. Emelyanov, A. F 1979. The problem of the family dis- tinction between the Fulgoridae and the Dic- tyopharidae (Homoptera, Auchenorrhyncha). In: Phylogeny and systematics of insects. Acad. Sci. USSR, Proc. Zool. Inst. 82: 3-22. Fennah, R C , and J. C M. Carvalho. 1963. On a new genus and species of fulgorid from Brazil (Ho- moptera, Lystrini). An. Acad. Brasileira Cienc. 35: 459-461. Hemming, F 1955. Validation, under the plenary pow- ers, of the generic name: "Fulgora" Linnaeus, 1767 (Class Insecta, Order Hemiptera) and desig- nation for the genus so named of a type species in harmony with current nomenclatorial practice. Opinion 322. International Commission on Zo- ological Nomenclature. Opin. & Declar. ICZN. 9(13): 185-208. Horn. W., and I Kahle 1935. Uber entomologische Sammlungen (Ein Beitrag zur Geschichte der En- tomo-Museologie). Ent. Beih. Berlin-Dahlem 2: 1-540. Janzen. D H , andC L Hogue 1983. Fulgora late rnaria (machacha, peanut-head bug, lantern fly.) In: Costa Rican natural history. University of Chicago Press, Chicago, Illinois, xi + 816 pp. Johnson, L. K , and R B Foster 1986. Associations of large Homoptera (Fulgoridae and Cicadidae) and trees in a tropical forest. J. Kansas Entomol. Soc. 59: 41.5-422. Knull, D J . and J. N Knull 1947. Two new Amycle (Homoptera: Fulgoridae). Ann. Entomol. Soc. Amer. 60: 397-400. Kramer, J P 1978. Taxonomic study of the American planthopper genus Cyrpoptus (Homoptera: Ful- goroidea: Fulgoridae). Proc. Biol. Soc. Washing- ton 91: 303-335. Lallemand, V 1956. Contribution a I'etude des Fulgori- dae (Hemiptera) (Ire. note). Bull. Inst. R. Sci. Nat. Belgique. 32: 1-7. 1959. Description de nouvelles especes de Ful- gorides d'Asie et d'Afrique. Zool. Meded. 36: 267-272 [no African species, five New World]. 1960. De quibusdam Fulgoris. Ent. Mitt. 24: 101-107. 1963. Nouvelles especes d'Homopteres. Bull. Inst. R. Sci. Nat. Belgique. 39: 1-5. 1966. Fulgorides nouveaux du Musee Zoologique de Berlin et de ma collection. Bull. Recher. Agron. Gembloux. N. S. 1: 51-54. Metcalf, Z p. 1947 General catalogue of the Hemiptera. Ease. IV. Fulgoroidea, Part 9. Fulgoridae. Smith College, Northampton, Massachusetts. 280 pp. Nast, J 1950. New genera and species of Neotropical Fulgoridae in the collection of the British Museum (Homoptera). Ann. Mus. Zool. Polonici 14: 167- 175. 1951. Some remarks on Neotropical Fulgoridae with descriptions of new genera and species (Ho- moptera). Ann. Mus. Zool. Polonici 14: 267-279, 3 plates. O Brien, L. B 1985. New synonymies and combinations in New World Fulgoroidea (Achilidae, Del- phacidae, Flatidae, Fulgoridae: Homoptera). Ann. Entomol. Soc. Amer. 78: 657-662. 1986. Five new species of Fulgoroidea (Ho- moptera) from the western United States and Mexico. Southwestern Entomol. 11:67-74. O Brien, L B , and S W Wilson 1985. Planthopper systematics and external morphology. Pages 61-102 in L. R. Nault and J. G. Rodriguez, eds., The leafhoppers and planthoppers. Wiley and Sons, Inc. 1988 O'BRIEN: New World Ful(;oridae 169 RiDOUT. B V 1983. Structure, form and function of the lantern fly head process {Fulgora laternaria Linn.) Unpublished dissertation, Birheck C'ollege. Wilson, S W , and L B O Brikn 1986. Descriptions of nymphs of Itzalana suhinaculata Schmidt (Ho- moptera: Fulgoridae), a species new to the United States. Great Lakes Entomol. 19: 101-105. 1987. A survey of planthopper pests of economi- cally important plants (Homoptera: Fulgoroidea). Pages 343-360 in Proc. Second Annual Workshop on Leafhoppers and Planthoppers of Economic Importance, held Provo, Utah, USA, 28 July-1 August 1986. Commonwealth Inst. Entomol., London. ZiMSEN, E 1964. The type material of 1. C. Fabricius. Munksgaard, Copenhagen. 280 pp. List OF Species (Countries are listed from south to north, west before east.) imitatrix Ossiannilsson Venezuela, Panama, Costa Rica, Belize, Mexico Peru Amerzanna O Brien, new genus peruana O'Brien Peru Amycle Stal aniahilis (Westwood) Mexico brevis O'Brien Mexico grandis O Brien Mexico mankinsi O Brien Honduras pinyonae Knull & Knull USA:CA, AZ, NM saxatilis Van Duzee USA:CA sodalis Stal Mexico, El Salvador tumacacoriae Knull & Knull USA: TX, AZ vernalis Manaee USA: NC, LA Aphrodisias Kirkaldy cacica (Stal) Mexico shaman O'Brien Mexico Artacie Stal dufourii (Signoret), resurrected name Fr. Guiana, Guyana haemoptera (Perty) Brazil Cathedra Kirkaldy serrata (Fabricius) Bolivia, Peru, Brazil, Fr. Guiana, Suriname, Colombia, Panama, Costa Rica Chilobia Stal = Ecuadoria Distant, new synonymy cinxia Stal Venezuela dichopteroides (Distant), (Ecuadoria), new combination Ecuador silena Stal Ecuador sinaragdina (Walker) Venezuela Copidocephala Stal guttata (White) Panama, Costa Rica, Guatemala, Mexico merula Distant Colombia = melanoptera (Schmidt), new synonymy viridiguttata Stal Columbia, Panama, Costa Rica, Honduras, Mexico = ornanda (Distant), new synonymy Diareusa Walker annularis (Olivier) Peru, Brazil, Fr. Guiana, Suriname, Guyana conspersa Schmidt Ecuador, Colombia = dahli Ossiannilsson, new synonymy keinneri Ossiannilsson Ecuadoria, sec Chilobia Encho})hora Spinola atonuiria Brancsik, see Cornelia, teste Lallcmand 1959:88 dufourii Signoret, see Artacie, teste O'Brien 1985:661 ensifera (Germar), to inccrtae sedis maculata O'Brien Peru nigroinaculata Distant Bolivia, Peru, Ecuador, Brazil = nigrolimbata Lallcmand pallidipunctata Lallcmand Bolivia, Peru prasina Gerstaecker Bolivia, Peru, Panama, Costa Rica pyrrhocrypta Walker Brazil, Guyana, Venezuela recurva (Olivier) Suriname, Brazil = bohe7nani Stal, new synonymy rosacea Distant Panama, Costa Rica, Nicaragua sanguinea Distant Ecuador, Colombia, Panama, Costa Rica, Nicaragua, Guatemala =florens Distant, new synonymy = longirostris Distant, new synonymy stillifera (Stal) Panama, Costa Rica, Honduras, Guyana, Belize, Mexico subviridis Distant Panama, Costa Rica = subviridisv ar. distanti Metcalf tuba (Germar) Brazil tuberctdata (Olivier) Brazil, Fr. Guiana, Suriname, Venezuela = tuberculata vm. fuscoinaculata Lallcmand = parvipennis Walker, new synonymy unifonnis O Brien Peru viridipennis Spinola Brazil = eminenta Schmidt, new synonymy Enhydria Walker = Ulubra Stal, new synonymy cicadina Gerstaecker Brazil ferruginea (Walker) to Hydriena (Dictyophariadae) longicornuta Lallcmand Brazil, Trinidad tessellata (Walker) Peru, Brazil = brachialis (Stal), new synonymy Fulgora L. (New synonymies in this genus are attributed to B. V. Ridout) caerulescens Olivier, to incertae sedis castresii Guerin-Meneville Peru, Panama, Mexico cearensis (Fonseca) Brazil, Trinidad crocodilia Brailovsky & Beutelspacher Mexico graciliceps Blanchard Peru, Bolivia, Brazil = orthocephala (Fonseca), new synonymy, Ridout lampetis Burmeister Bolivia, Peru, Brazil, Panama, Costa Rica, Nicaragua, Honduras laternaria (Linne) Brazil, Panama, Honduras, Mexico = servillei Spinola, new synonymy, Ridout lucifera Germar Argentina, Bolivia, Brazil riograndensis (Fonseca) Brazil Odontoptera Carreno carrenoi Signoret Brazil, Fr. Guiana, Guyana, Panama, Costa Rica, Guatemala, Mexico spectabilis Carreno Brazil Phrictus Spinola auromaculatus Distant Bolivia, Peru, Ecuador, Brazil 170 Great Basin Naturalist Memoirs No. 12 = notatus Lallemand, new synonymy delicatus O'Brien Brazil diadema (Linnaeus) Brazil, Fr. Guiana, Suriname = diadema var. walkeri Metcalf diligens O'Brien Colombia hoffmannsi Schmidt Peru, Ecuador minutacanthis Caldwell Peru moebiusi Schmidt Colombia = sordidus Caldwell, new synonymy ocellatus Signoret Venezuela, Colombia punctatus Caldwell Panama quinquepartitus Distant Colombia, Panama, Costa Rica regalis Caldwell Fr. Guiana, Suriname tripartitus Metcalf Panama, Honduras, Belize xanthopterus Schmidt Peru, Ecuador Rhabdocephala Van Duzee brunnea Van Duzee USA; AZ, Mexico Scolopsella Ball mexicana O'Brien Mexico reticulata Ball USA: AZ, CA Sinuala O'Brien, new genus schmidti O'Brien Belize stali O'Brien Honduras, El Salvador tuberculata O'Brien Costa Rica Stalubra O'Brien, new genus brunnea O'Brien Brazil, Guyana ruftda (Lallemand), (Enhydria), new combination Brazil A NUMERICAL TAXONOMIG ANALYSIS OF INTERSPECIFIC MORPHOLOGICAL DIFFERENCES IN TWO CLOSELY RELATED SPECIES OF CICADA (HOMOPTERA, CICADIDAE) IN PORTUGAL J. A. Quartan' Abstract — Cicada orni Linnaeus is among the most common and widespread cicadas in Portugal, and, unless a critical study of the male genitalia is made, it is easily confused with the much less widely distributed C. barhara lusitanica Boulard. These species are morphologically very similar and sometimes diflkult to separate using existing keys. This study attempts to test the discriminating capabilities of numerical technicjucs commonly used for classifica- tory purposes, as well as to discover the most effective characters to distinguish between the two species. F"or these purposes, cluster analysis and principal component analysis were applied to a sample of 64 male specimens character- ized by 40 characters (33 derived from the external morphology and 7 from genitalia). In WPGMA cluster analysis, product-moment correlations gave a better separation between these species than did taxonomic distance coefficients; moreover, the analysis derived from the genital characters alone gave better separation than the analyses based on the 33 external characters. Principal component analysis yielded a clear, interspecific separation along the first axis. The best characters to discriminate between males of the two species were the lengths of the pygofer (and its dorsal spine), the tenth abdominal segment, and the appendages of the latter (which are smaller in barbara lusitanica), as well as the width of the shaft of the aedeagus (thinner in orni).' Finally, the uniformity of the general clustering pattern resulting from the two multivariate techniques suggests the presence of two distinct species, as also clearly indicated by behavioral data. Cicada orni Linnaeus is among the most common and widespread cicadas in Portugal, and, unless a critical study of the male geni- talia is made, it is easily confused with the much less widely distributed C barbara lusi- tanica Boulard (Quartan and Fonseca 1988). As live specimens, however, they are easily distinguished by the male calling songs, which are quite distinct. Oscillograms are found in Claridge et al. (1979) and Boulard (1982), respectively, for C. orni and C. bar- bara lusitanica. The two species are externally very similar and sometimes even difficult to separate by existing keys (e.g., Gomez-Menor 1957). In fact, the main distinguishing character used for their separation has been the presence in barbara of only two spots on the cross-veins of the forewings instead of four; however, some specimens o( barbara lusitanica have the full four spots as they occur in orni (Fig. 4). Boulard (1982), when describing the Por- tuguese form of C. barbara, which he origi- nally named lusitanica, provided a good diag- nosis of the genital characters of this species. However, no detailed comparison of the two species has been made, nor has any type of multiple-character analysis involving the simultaneous use of several measurements or counts been attempted. It was felt of interest, therefore, to see how far some common tech- niques of numerical taxonomy would discrim- inate between this pair of closely related spe- cies. This study was undertaken with two main objectives in mind. The first was to apply current techniques of numerical taxonomy commonly used for class ificatory purposes with the aim of testing their general discrimi- nating power with respect to these two spe- cies. The techniques chosen were a form of hierarchical cluster analysis and principal component analysis. It is known that apart from the explicit use of the former, principal component analysis can also serve as a cluster technique of great generality and can be used to distinguish pairs of putative morphs as in the classical study of Temple (1968). The sec- ond objective was to discover new characters that might help to separate C. orni from C. barbara. 'Depto. de Zoologiae Antropologia, Faculdade de Ciencias de Lisboa, 1200 Lisboa, Portugal. 171 172 Great Basin Naturalist Memoirs No. 12 Material The data on which this study is based were taken from dried male specimens (OTUs) of 32 Cicada orni and 32 C. harhara lusitanica (Table 1). These samples were mostly taken by the author in Portugal: all 32 males of orni were collected in central Portugal; an equal number of males of harhara hisitanica were taken in several areas of Algarve (the southern province of Portugal), where the species ap- pears to be particularly common, with the exception of two specimens only that were collected in Sesimbra (south of Lisbon). The localities and sample sizes are C. orni: Albu- ritel, Vila Nova de Ourem (n = 32); C. har- hara hisitanica: Carvoeiro (n = 25), Praia da Rocha (n = 4), Serra de Monchique (n = 1), and Sesimbra (n = 2). Methods Measurements and Counts Thirty-seven of the 40 characters were mea- surements; the remaining 3 were counts. Measurements were made using a Wild M3 microscope with a graduated eyepiece and were taken as described in Table 2 or as illus- trated in Figures 1-9. Of these 40 characters, 33 refer to external morphology and the re- maining 7 to male genitalia. Data Analysis Data processing was carried out on the CDC 6500 computer at the Imperial College Computer Center (University of London) us- ing two multivariate statistical programs de- veloped by Prof. R. G. Davies (Department of Pure and Applied Biology, Imperial College) for cluster analysis and ordination (Quartan and Davies 1983, 1985). In most analyses, characters were standard- ized by expressing each state as a deviation from the mean in standard deviation units. For Q-mode analysis, taxonomic distances as well as product-moment correlations were found and structured by the WPGMA method of cluster analysis (Weighted Pair Group Method with Arithmetic Averaging). Pheno- grams, expressing the phenetic relationships among the OTUs in a hierarchy of increasingly larger clusters, were thus obtained (Figs. 10-16). For R-mode analysis, character correlations Table L List of specimens (males) oi Cicada orni Lin- naeus and of C. harhara lusitanica Boulard investigated (OTUs). OTUs Locality, date of capture, and collector Cicada orni 1-15 Alburitel, 10.viii.l979, J. A. Quartau 16-32 Alburitel, vii. 1971, J. A. Quartau C. harhara hisitanica 33-34 Carvoeiro, 14.viii.l966, P. D. Rodrigues 35-38 Praia da Rocha, IT.viii. 1973, J. A. Quartau 39-40 Carvoeiro, 14. vii. 1978, J. A. Quartau 41-47 Carvoeiro, 30. vii. 1978, J. A, Quartau 48-55 Carvoeiro, 28. vii. 1978, J. A. Quartau 56-57 Carvoeiro, 9.viii-10.i.x.l980, L. Mendes 58-59 Sesimbra, 2.viii. 1980, J. A. Quartau 60 Carvoeiro, 31. vii. 1978, J. A. Quartau 61-62 Carvoeiro, 18. vii. 1978, J. A. Quartau 63 Monchique, 2.i.x.l971, F. Carvalho 64 Carvoeiro, 24.viii.1981, J. A. Quartau based on data standardized by OTUs were subjected to principal component analysis (PGA). This ordination method transforms the original characters, generally continuous, cor- related characters, into a suite of uncorre- cted, composite variables — the principal components (principal axes). In addition to being mutually independent, these compo- nents account for maximum variance as fol- lows: the variance along the first axis (i.e., the corresponding eigenvalue) is the maximum possible. The second axis describes the next largest variance orthogonal to (uncorrected with) the first. The third axis follows similarly but is independent of both first two axes, and so on, for as many axes as one wishes to extract (e.g., Gibson et al. 1984). A transposed matrix of the character loadings was post-multiplied by the standardized data matrix to yield a matrix of OTU projections in the principal component space. Two-dimensional ordina- tion diagrams of the representations of the two species, together with the character loadings (scaled eigenvectors), were thus obtained (Figs. 17-18, Table 3). Results Phenograms The seven phenograms resulting from vari- ous analyses based on all characters, on the genitalia only, or on the external characters alone are shown in Figures 10-16. The WPGMA clustering technique was followed 1988 QuARTAU: Cicada Analysis 173 Table 2. Description of characters: measurements and counts (terminology mostly follows Myers [1928]). Character No. Description 1. Overall length measured from tip of crown to apical margin of the right forewing with the latter in posi- tion of rest alongside the body (Fig. 1). 2. Length of crown measured along a medial line pass- ing through the median ocellus (Fig. I). 3. Minimum distance between the ocular sutures mea- sured along the paired ocelli (Fig. 2). 4. Medial length of frons measured dorsally as indicated (Fig. 2). 5. Medial length of pronotum measured dorsally as in- dicated (Fig. 1). 6. Medial length of mesonotum measured dorsally from anterior margin to posterior margin of cruciform ele- vation or scutellum (Fig. 1). 7. Width of pronotum measured at the level of anterior lateral margins (Fig. 1). 8. Width of pronotum measured at the level of postero- lateral margins (Fig. 1). 9. Width of crown measured at the level of median ocellus and as indicated (Fig. 2). 10. Inner distance between the paired ocelli (Fig. 2). 11. Distance between the right paired ocelli and the right ocular suture as indicated (Fig. 2). 12. Distance between the base of the left antenna and the left ocular suture as indicated (Fig. 3). 13. Inner distance between the base of antennae (Fig. 3). 14. Length of frons as illustrated (Fig. 3). 15. Length of clypeus as illustrated (Fig. 3). 16. Length of the exposed part of beak. 17. Length of dorsal margin of the left fore femur as illustrated (Fig. 5). 18. Length of ventral margin of the left fore femur as illustrated (Fig. 5). 19. Length of basal spine in ventral margin of left fore femur as illustrated (Fig. 5). 20. Length of apical spine in ventral margin of left fore femur as illustrated (Fig. 5). 21. Distance between tips of the apical and basal spines in the ventral margin of the left fore femur as illus- trated (Fig. 5). 22. Distance from anterior right corner to posterior left corner of left operculum as illustrated (Fig. 6). 23. Distance from anterior left corner to posterior right corner of left operculum as illustrated (Fig. 6). 24. Length of right forewing as illustrated (Fig. 1). 25. Greatest width of right forewing as illustrated (Fig. 4). 26. Length of subcostal cell ("gancho" cell of G6mez- Menor 1957) in right forewing (Fig. 4). 27. Length of anterior margin of basal cell in right forewing (Fig. 4). 28. Length of posterior margin of basal cell in right forewing (Fig. 4). 29. Maximum width of basal cell in right forewing. 30. Minimum width of basal cell in right forewing. 31. Number of apical cells in right forewing. 32. Number of cells other than apicals of right forewing. 33. Number of spots in cross-veins of right forewing. 34. Length of pygofer in lateral view as indicated (Fig. 7). 35. Overall length of tenth abdominal segment as indi- cated (Fig. 7). 36. Overall length of appendages of tenth abdominal segment as indicated (Fig. 7). 37. Distance in basal curvature of shaft of aedeagus as indicated (Fig. 9). 38. Width of shaft of aedeagus as indicated (Fig. 9). 39. Width of shaft of aedeagus in area of curvature as illustrated (Fig. 9). 40. Medial length of eighth sternite or hypandrium (Fig. in all; and Pearson's product-moment coeffi- cient and the taxonomic distance coefficient were used as measures of taxonomic proxim- ity. (a) Genital analyses Figures 10 and 11 illustrate a correlation and a distance phenogram, respectively, both based on the seven standardized variables of the male genitalia. Both analyses, notwith- standing their being based on a small number of variables, resulted in two main clusters, one with C. orni and the other with C. bar- bara lusitanica. However, in the former phe- nogram, the cluster oi barbara lusitanica in- cludes one specimen of orni (No. 13). (b) External characters These analyses resulted in the production of the phenograms depicted in Figures 12 and 13. Both were based on standardized data, but only the correlation coefficient succeeded in giving an almost complete separation of the two species of cicadas. In fact, OTUs were grouped into two main clusters as in the geni- tal analyses, but specimen No. 19 belonging to C. orni appeared misplaced within C. bar- bara lusitanica (Fig. 12). On the contrary, the distance phenogram provided much less satis- factory results than the previous analysis, since each of the two major clusters incorpo- rates elements of both species of cicadas (Fig. 13). (c) Combined characters The phenograms of this group of analyses, involving all 40 characters combined, are il- lustrated in Figures 14-16. Considering the phenograms based on standardized data (Figs. 14, 16), it is clear that the correlation phenogram gave a much bet- ter distinction between the two species than the distance analysis. In fact, the latter (Fig. 16) clustered six specimens of C. orni with C. 174 Great Basin Naturalist Memoirs No. 12 1 10 'i i ' 1 1 9 ' 1 1 ' 1 : n 1 : Figs 1-6. Diagrams of a male Cicada orni Linnaeus illustrating most of the measurements taken; 1, body, dorsal view; 2, head, dorsal view; 3, head, anterior view; 4, right forewing, dorsal view; .5, left foreleg, mner view; 6, lett operculum, ventral view. 1988 QuARTAU: Cicada Analysis 175 Figs. 7-9. As in Figures 1-6: 7, pygofer and tenth abdominal segment, ventral view; 8, eighth stemite or hypandrium, ventral view; 9, aedeagus, lateral view. barbara lusitanica. Even when the data were unstandardized, correlations gave a good pic- ture of the relationships between these two species (Fig. 15). Principal Component Analysis This analysis involved all 40 characters and was computed from a between-character cor- relation matrix based on data standardized by OTUs. As in similar analyses carried out with leafhoppers of the genus Batracomorphus (Quartan 1983), slightly more than half (54%) of the total variation in the study was ex- plained by the first three axes. The first component accounted for 38.90% of the variation in the data and is interpreted as a contrast between the lengths of the pygofer, tenth abdominal segment, or ap- pendages of the latter and the width of the shaft of the aedeagus. It does not represent overall size as commonly is the case, since many of the characters (Table 3) are not posi- tively correlated with it (e.g., Jolicoeur and Mosimann 1960, Blackith and Reyment 1971, Baker 1980, Gibson et al. 1984, Shea 1985). In fact, it must represent both size and shape as has been pointed out by several authors (Mosimann 1970, Oxnard 1978, Humphries et al. 1981). A complete separation of C. orm and C. barbara lusitanica was given by the dis- crimination afforded by this axis, which is probably close to the orientation of the opti- mum discriminant function. The characters loading most heavily on this component (Table 3) are therefore of considerable taxo- nomic interest, since they are diagnostic for this pair of species. The highest negative scores, in decreasing order, were for charac- ters numbered 35 (length of tenth abdominal segment), 34 (length of pygofer), and 36 (length of appendages of tenth abdominal seg- ment). The highest positive score was for character numbered 38 (width of shaft of aedeagus). The second principal component accounted for 8.51% of the total variation and was inter- preted as a contrast between the number of spots in cross-veins of the wings and the width of the crown. It was most heavily loaded, neg- atively and positively, on characters num- bered 33 and 9, respectively. The third principal component accounted for 6.61% of the total variation and was inter- preted as a factor resulting from the lengths of 176 Great Basin Naturalist Memoirs No. 12 Table 3. Eigenvector matrix (character loadings) in a principal component analysis of the matrix of correlations among the 40 variables (data standardized by OTUs.) Variables Scaled eigenvectors I II III 1 0.296 -0.1.58 0.441 2 0.404 0.123 0. 155 3 0.531 0.557 -0.174 4 0.843 -0.160 0.126 5 0.492 0.300 -0.119 6 0.037 -0.096 0.280 7 0.567 0.280 -0.069 8 0.314 0.365 -0.131 9 0.336 0.633 -0.1,33 10 0.738 -0.237 -0.000 11 0.687 0.155 -0.200 12 0.859 -0.127 -0.142 13 0.313 0.423 0.414 14 0.117 0.422 0.306 15 -0.604 -0.196 -0.120 16 -0.769 0.104 -0.192 17 -0.316 0.338 0,524 18 -0.667 0.174 0.,375 19 0.565 0.2.52 -0.091 20 0.531 -0.066 0.111 21 -0.132 0.019 0.257 22 0.784 0.203 0.160 23 0.847 0.112 0.000 24 0.524 -0,340 0.060 25 0.683 -0.104 0.154 26 0.187 -0.207 -0.412 27 -0.334 0.126 -0,561 28 0. 133 0.117 -0.,549 29 -0.011 -0.207 0.471 30 0.129 -0.287 0.511 31 0.818 -0.490 -0.049 32 0.818 -0.479 -0.059 33 0.266 -0.745 -0.163 34 -0.952 -0.011 -0.040 35 -0.958 -0.153 -O.OOI 36 -0.944 -0.180 0.043 37 -0.869 -0.250 0.028 38 0.942 -0.065 -0.023 39 0.892 -0.249 0.024 40 -0.858 -0.125 -0.090 Latent roots 15.559 3.406 2.644 Percentage of component 38.898 8.514 6.609 variation cumulative 38.898 47.412 ,54.021 the anterior and posterior margins of the basal cell of the wings. It was most heavily loaded (negative scores) on characters numbered 27 and 28. Neither the second nor the third axis leads to a separation of the two species of cicadas. Figures 17 and 18 are two-dimensional views of the relationships among specimens of both species in the space determined by compo- nent I combined with component II and by component I with component III, respec- tively. It is clear that these two plots gave a good separation between C. orni and C. bar- bara lusitanica. The plot combining axes II and III did not, however, succeed in giving a correct assignment of the two species and therefore was not illustrated here. Conclusions Cluster analysis and principal component analysis are two of the methods most com- monly used for recognition of group structure in numerical ta.xonomy (e.g., Quartau 1987). This study suggests that the two general methods are also very useful for discriminat- ing between pairs of closely related species. In fact, because of the uniformity of the gen- eral clustering pattern that resulted from the application of both methods, it is clear that two distinct species exist, a fact also indicated by behavioral data. However, concerning cluster analysis, it is worthwhile noting that the hierarchical struc- ture within each of the two major groups of OTUs differs a good deal from one particular method to another. Moreover, it appeared that correlations were more effective than tax- onomic distances in describing relationships between the two cicadas, a result in keeping with Boyce (1964) or Cheetam (1968), for in- stance. It is interesting to note, in this regard, that such finding is in disagreement with a similar study carried out with leallioppers (Quartau and Davies 1983) or with results based on other groups (e.g.. Smith 1972). Fi- nally, the failure of the cluster analysis using distances with nongenital characters shows that the use of such techniques needs to be undertaken with care and that it might be best to use a consensus of several clustering tech- niques when applying numerical methods to a novel taxonomic situation. Principal component analysis succeeded in giving a good distinction of the two species along axis I, in spite of its accounting for only .38.90% of the total variation. This analysis also showed that the main distinguishing charac- ters between C. orni and C. barbara lusitan- ica are the following: lengths of the pygofer, the tenth abdominal segment, and the ap- pendages of the latter (which are smaller in C. barbara lusitanica. Figs. 19, 22), as well as the width of the shaft of the aedeagus (which is 1988 QuARTAU; Cicada Analysis 177 -0.8 Correlations Data standardized Genital characters D C. orni • C. barbara lusitanica 10 it Distances Data standardized Genital characters DC. orni • C. barbara lusitanica 11 Figs. 10-11. 10, correlation phenogram based on the seven genital characters with standardized data; 11, distance phenogram based on the seven genital characters with standardized data. thinner in orni. Figs. 20, 23). Moreover, de- tailed examination of the male genitalia also showed that the dorsal spine of the pygofer is smaller in barbara lusitanica than in orni (Figs. 19," 22). Acknowledgments 1 thank Prof. R. G. Davies, Department of Pure and Applied Biology, Imperial College, for having made his computer programs avail- able for my use as well as for his critical com- ments on the manuscript. Prof. R. M. Ander- son is also acknowledged for having granted working facilities at the Imperial College (University of London). 1 am also grateful to my colleague Dr. M. L. Mathias for her help with the preparation of most phenograms, and to Miss A. R. Cruz for help with other illustra- tions and the measurements. For enabling me to publish this study in honor of Prof. P. Oman (Oregon State University, USA), who has been a source of encouragement since the beginning of my taxonomic studies on leafhoppers, I also acknowledge Dr. M. W. Nielson (Brigham Young University, USA) and Dr. H. D. Blocker (Kansas State Univer- sity, USA). 178 Great Basin Naturalist Memoirs No. 12 Correlations Data standardized Cxt. Morphology characters D C. orni • F. barbara lusitanica Figs. 12-13. 12, correlation phenogram based on the 33 external morphological characters with standardized data; 13, distance phenogram based on the 33 external morphological characters with standardized data. This paper was completed at Imperial Col- lege (University of London) under an INIC (Ministerio da Educagao, Lisbon) sabbatical fellowship. Literature Cited Baker, A. J 1980. Morphometric differentiation in New Zealand populations of the house sparrow (Passer domesticus). Evolution 34: 638-653. BLACKrrH, R E., AND R. a Reyment 1971. Multivariate morphometries. Academic Press, London. BOULARD, M. 1982. Les cigales du Portugal, contribution a leur etude (Homoptera, Cicadidae). Annales de la Societe entomologique de France (N.S.) 18(2): 181-198. BoYCE, A J. 1964. The value of some methods of numeri- cal taxonomy with reference to hominoid classifi- cation. Pages 47-65 in V. H. Heywood and J. McNeill, eds., Phenetic and phylogenetic classification. Systematic Association Publication No. 6, London. Cheetam, A H 1968. Morphology and systematics of the bryozoan genus Metrarabdotiis. Smithsonian Miscellaneous Collections 153(1); 1-121. Claridge. M F . M R Wilson, and] S Singhrao 1979. The songs and calling sites of two European ci- cadas. Ecological Entomology 4: 225-229. Gibson, A R , A J Baker, and A Moeed 1984. Morpho- metric variation in introduced populations of the common myna (Acridotheres tristis): an applica- tion of the jackknife to principal component analy- sis. Systematic Zoology 33(4): 408-421. Gomez-Menor, O J 1957. MonografiadeCicadidos (Ho- moptera) de Esparia. Academia de Madrid, Madrid. 1988 QUARTAU: CiCADA ANALYSIS 179 0.0 0.993 0.998 Correlat ions Data standardized All characters DC. orni o C. barbara lusitanica 14 T 31 Correlat ions Data unstandardized All ctiaracters n £. orni o C. barbara lusitanica Figs. 14-15. 14, correlation phenogram based on all 40 combined characters with standardized data; 15, correlation phenogram based on all 40 combined characters with unstandardized data. Humphries, J M , F L Bookstein, B Chernoff, G R Smith, R L Elder, and S G Poss 1981. Multi- variate discrimination by shape in relation to size. Systematic Zoology 30: 291-308. JOLICOEUR, P , AND J E MosiMANN 1960. Size and shape variation in the painted turtle. A principal compo- nent analysis. Growth 24: 339-354. MosiMANN, J E 1970. Size allometry: size and shape variables with characterizations of the lognormal and generalized gamma distributions. Journal of the American Statistical Association 65: 930-945. Myers, J G 1928. The morphology of the Cicadidae (Homoptera). Proceedings of the Zoological Soci- ety of London 1928: 265-472. OXNARD, C E 1978. One biologist's view of morphomet- ries. Annual Review of Ecology and Systematics 9: 219-241. QuARTAU, J A 1983. A comparison of different numerical methods applied to the taxonomy of Batracomor- phus Lewis (Insecta, Homoptera, Cicadellidae, lassinae): ordination techniques. Journal of the Entomological Society of Southern Africa 46(1): 9-35. 1987. Ordination vs. cluster analysis in the numer- ical taxonomy of some leafhoppers (Homoptera, Cicadellidae). Pages 207-215 in M. R. Wilson and L. R. Nault, eds.. Proceedings of the 2nd Interna- tional Workshop on Leafhoppers and Planthop- pers of Economic Importance, CLE., London. QuARTAU, J A , AND R G Davies 1983. A comparison of different numerical methods applied to the taxon- omy of Batracomorphus Lewis (Insecta, Ho- moptera, Cicadellidae, lassinae): cluster analysis. Revista de Biologia 12: 550-596. 180 Great Basin Naturalist Memoirs No. 12 Dis tnnrcs Dat a slanda diziMl All characters DC. or. larbara usil.iruc-.-l • D • D ; • ••-.= •. • . • • • D - orni barbara lusicanica 17 D D D * • □ a barbara lusiiamca 18 Figs. 16-18. 16, distance phenogram based on all 40 combined characters with standardized data; 17, a two-dimen- sional view of the relationships among the 64 OTUs in a space determined by component I on ,x-axis (38.90%) and component II on y-axis (8.51%) of a principal component analysis of the matri.x of correlations among all 40 characters with standardized data; 18, as in Fig. 17 but referred to component I on x-axis and III on y-axis (6.61%). 1985. Character selection by information content in the numerical taxonomy of some male Batraco- morphus (Homoptera: Cicadellidae). Zeitschrift fiir zoologische Systematik und Evolutionsfor- schung 23; 100-115. QUARTAU, J a . AND P. J. FONSECA. 1988. An annotated check-list of the species of cicadas known to occur in Portugal (Homoptera: Cicadoidea). Pages 367- 375 in C. Vidano and A. Arzone, eds. Proceedings of the 6th Auchenorrhyncha Meeting, Turin. Shea, B T 1985. Bivariate and multivariate growth allometry: statistical and biological considerations. Journal of Zoology 206: 367-390. Smith, J D 1972. Systematics of the chiropteran family Marmoopidae. University of Kansas, Publications of the Museum of Natural History 56: 1-132. Temple, J T 1968. The Lower Llandovery (Silurian) bra- chiopods from Keisley, Westmorland. Palaeon- tographical Society 521: 1-58. 1988 QuARTAU: Cicada Analysis 181 Figs. 19-24. Diagrams of the male genitalia oiCicada orni (Figs. 19-21) and of C. barbara lusitanica (Figs. 22-24): 19, 22, pygofer and tenth abdominal segment, lateral view; 20, 23, aedeagus, lateral view; 21, 24, eighth sternite or hypandrium, ventral view (scale = 0.5 mm). REVISION OF THE NIRVANINAE (HOMOPTERA: CICADELLIDAE) OF THE INDIAN SUBCONTINENT C. A. Viraktamath' andC. S. Wesley^ Abstract — ^Three tribes of Nirvaninae, namely, Balbillini (two genera and three species), Nirvanini (five genera and 26 species), and Occinirvanini (one genus and one species), occur on the Indian subcontinent. The tribes, genera, and species are redescribed, illustrated, and keyed. New ta.xa recognized are BalbiUus indicus n. sp. (India; Kerala), Stenotortor stibhimalaya n. sp. (India: West Bengal), Kana bispinosa n. sp. (India: Tamil Nadu), K. nigropicta n. sp. (India: Kerala), Sophonia bakeri n. sp. (India: Karnataka, Kerala, Tamil Nadu, Uttar Pradesh; Nepal), S. bifida n. sp. (India: West Bengal), S. complexa n. sp. (India; Meghalaya), S. compUcata n. sp. (India: Mizoram, Meghalaya), S. keralica n. sp. (India; Kerala), Nirvana peculiarisn. sp. (India: Mizoram, Meghalaya), and N. striata n. sp. (India; West Bengal, Uttar Pradesh, Himachal Pradesh). The genus Qitercinirvana Ahmed & Mahmood (type species; Q. longi- cephala Ahmed & Mahmood) is treated as a junior synonym oi Sophonia Walker. Kana signata Distant and Nirvana greeni Distant are suppressed as junior synonyms of K. illuminata and N. Hnealis , respectively, and are transferred to Sophonia . Three other species transferred to Sophonia are Kana modesta Distant, Quercinirvana longicephala Ahmed & Mahood, and Nirvana insignis Distant. The relationships among various genera are discussed briefly. The genera Crispina Distant and Mohiinia Distant are excluded from Nirvaninae. The Nirvaninae, one of the smaller sub- families of the leafhopper family Cicadellidae, include a predominantly tropical group of leafhoppers. They are fragile, often brightly colored, depressed leaflioppers and common on lush green vegetation. They are often mis- taken for Typhlocybinae, but the structure of the head and the more depressed form and truncate basitarsus of the hind leg distinguish Nirvaninae. The subfamily includes 111 spe- cies (excluding Mukariinae and other non- Nirvaninae genera, which were traditionally included in the Nirvaninae) distributed in the Oriental (56), Afrotropical (24), Neotropical (7), Palearctic (7), Australian (11), and Pacific island (6) regions. The fauna of Afrotropical (Linnavuori 1979), Australian (Evans 1966), Papua New Guinea (Evans 1973), Neotropical (Kramer 1964), and Palearctic (Kuoh and Kuoh 1983) regions have recently been re- vised. The tribe Mukariini earlier included in the subfamily by Distant (1908g) and Metcalf (1963) is now considered a subfamily distinct from Nirvaninae (Linnavuori 1979). Information on host plants of Nirvaninae is very meagre. Nirvana pallida Melichar and N. suturalis Melichar are destructive to sugar- cane and grasses in Taiwan (Schumacher 1915a). They also feed on rice, mulberry, and camphor tree (Esaki and Ito 1954a). Baker (1923a) found Stenotortor inocarpi Baker on Otaheite chestnut, Inocarpus edulis, in Sin- gapore. In Australia, Nirvaninae feed on /. edulis, casuarina, and croton (Evans 19411). Teco7naria capensis is recorded as the host of Narecho tecomariae Theron in South Africa (Theron 1970). Quercinirvana longicephala Ahmed & Mahmood feeds on Aesculus indica, Quercus dilatata, and Vihurmmi nervosum (Ahmed and Mahmood 1970). Chudania delect a Distant breeds on Ficus carica (Ahmed and Mallik 1972). In South America, Tahura fowleri Kramer is found on Passiflora (Kramer 1964). Kuoh and Kuoh (1983) recorded Camellia sinensis. Acacia confusa. Citrus reticulata, Pterocarpus indicus, Orijza sativa, Psidium guajava, and Chimonanthus praecox as host plants of species oi Pseudonir- vana Baker { = Sophonia Walker) in China. Nirvana pallida and N. greeni Distant breed on black gram, green gram, cowpea, field bean, pigeon pea, and soybean in India (Ra- makrishna 1980). The Nirvaninae of the Indian subcontinent consist of eight genera and 30 species grouped in three tribes, namely, Balbillini, Nirvanini, and Occinirvanini. The genera Mohunia Dis- tant and Crispina Distant are excluded from the study, as they probably belong to the sub- family Deltocephalinae. Department of Entomology, University of Agricultnral Sciences. CKVK Campus, Bangalore 560 065, India. ^Department of Ecology and Evolution, S.U.N.Y., Stony Brook, New York 11794, US.\. 182 1988 VlRAKTAMATII, WESLEY: REVISION OF INDIAN NiKVANINAE 183 Material and Abbreviations This study was based mainly on specimens collected over the years and deposited in the insect collection of the Department of Ento- mology, University of Agricultural Sciences by the senior author and his students. Addi- tional specimens were borrowed from various institutions for the study. Abbreviations used for these institutions and for the depositories of the types of new taxa are as follows: BMNH - British Museum (Natural History), London. lARI -Indian Agricultural Re.search Institute, New Delhi. IRSNB - Institute Royal des Sciences, Naturelles de Bel- gique, Bru.xelles. JU - Department of Biosciences, Jammu Univer- sity, Jammu. PAU - Punjab Agricultural University, Ludhiana. UAS - University of Agricultural Sciences, Bangalore. USNM -U.S. National Museum (Natural History), Washington, D.C. ZMHU -Museum fiir Naturkunde der Humboldt, Urii- versitat zu Berlin, Berlin. ZSI -Zoological Survey of India, Calcutta. Distant (1908g, 1918b) did not mention the number of specimens (syntypes) of each new species he described, except when a single specimen was involved, viz., Chundania de- lecta Distant (1908g; 268) In this study lecto- type designations were made even though a single specimen of the type series (syntypes?) was present in the BMNH collection. Unless otherwise stated they were considered as part of the syntype series. The terminology used for describing hind leg spinulation follows Davis (1975). Description of Taxa Nirvaninae Delicate, fragile, depressed, small to mod- erately large (4.0-9.0 mm) leafhoppers. Usu- ally yellow or white, often marked with red, black, orange, or brown fascia. Head as wide as or wider than pronotum. Vertex usually flattened, strongly produced, and with rugae or sculpturing on lateral and anterior regions. Ocelli on crown along lateral margin in front of eyes. Frontoclypeus and clypellus either flat or tumid, in the latter case often provided with a median keel. Lora often small, extreme anterior margin of genae bluntly prominent, extending beyond apex of clypellus. Antennal pits deep, antennal ledges more or less strong, antennae long. Lower margin of eyes sinuate ventrally. Anterior tentorial branches L-shaped. Lateral margins of pronotum cari- nate (except in Omaranus Distant), moder- ately long. Forewing venation reduced, with- out cross-veins at base (except Balbillini), longitudinal veins represented by a series of paired pits basally, appendix either narrow (Nirvanini) or broad (BcdhiUus Distant). Hind- wing with three or lour apical cells, veins lA and 2A fused basally. Fore and middle tibiae cylindrical or flattened (Balbillini), hind femoral spinulation 2+1 + 1 (Nirvanini), 2+1 + 0 (Balbillini and Occinirvanini), or 2+0+0 (Balbilhni). Male pygofer either cylindrical or de- pressed, with or without anal collar process, but may be armed with ventral process, heav- ily macrosetose. Valve small, fused with pygofer laterally. Plates usually parallel- sided. Style variable. Connective Y-shaped (Nirvanini) or platelike (Balbillini). Aedeagus with single shaft and often provided with pro- cesses. Linnavuori (1979) suggested that the sub- family Nirvaninae is a derivative of the Aphro- dinae-Cicadellinae stock and that Nirvanini is the most advanced tribe. Key to Tribes 1. Fore and middle tibiae flattened; head notched in front of eyes so that the scape is visible in dorsal aspect Balbillini — Fore and middle tibiae rounded; scape not visi- ble from above 2 2(1). Ocelli nearer to apex of head than to eyes; forewing appendix wide and extending around apical wing margin; lora large; vertex about half as long as wide, obliquely produced in front of eyes, with hind femoral spinulation 2+1 + 0; hind basitarsus with six platellae . . . Occinirvanini — Ocelli nearer to eyes than to apex of head; forewing appendix narrow and does not extend around apex of wing; lora small; vertex as long as wide or longer, not obliquely produced in front of eyes; hind femoral spinulation 2+1 + 1; hind basitarsus with two or three platellae . . Nirvanini Tribe Balbillini This tribe has been well characterized by Linnavuori (1979). The following additional characters are noted. Forewing with supernu- merary cross-veins along costal margin and with two m-cu cross-veins, four to five apical cells, two anteapical cells, and appendix (ab- sent in Stenotortor Baker). Hindwing with four apical cells. Hind femoral spinulation 184 Great Basin Naturalist Memoirs No. 12 2 + 2 + 0 or 2 + 0+0. Male connective long and incrassate or lamellate and fusiform. Aedea- gus simple with large gonopore. Key to Genera of Balbillini 1. Forewing with a well-developed appendix; hind femoral spinulation 2+1+0 Balhillus Distant — Forewing without an appendix; hind femoral spin- ulation 2 + 0 + 0 Stenotortor Baker Genus Balhillus Distant Balhillus Distant 1908g: 287. Type species; Balhillus granulosus Distant, by original designation and monotypy. Head narrower than pronotum. Crown bluntly, angularly produced, convex, de- clivous anteriorly, with median and submar- ginal, anterior carinae between ocelli. Face horizontal, flat. Clypellus twice as wide at base as at apex; lora small. Lateral margins of head in front of eyes notched, exposing scape of antenna. Pronotum declivous laterally, with two lateral, carinate margins; mese- pisternum attaining lateral position in front of base of wing and strongly ridged. Fore and middle tibiae angular; hind femoral spinula- tion 2+1 + 0, inner spine very slender and minute. Apex of hind basitarsus with three platellae. Hind tibial spinulation Rj 12 ± 1, R, 19±1, R3 25±l, R4 25±l. Pygofer much shorter than its height, with- out processes. Anal tube short. Genital plate parallel-sided, 3.4 times as long as wide with uniseriate, submarginal setae and few hairlike setae. Apophysis of style simple, without preapical lobe. Connective lamellate and fusi- form. Aedeagus articulate with connective, simple and with large gonopore. Remarks. — As pointed out bv Linnavuori (1979) and Baker (1923), Balhillus and Steno- tortor are closely related. They can be sepa- rated by the characters used in the key. The bodies of both genera remain appressed to the leaf surface, which explains the flat nature of the tibiae and the undersurface of the body. The Indian species, B. granulosa and B. indica, have hind femoral spinulation 2+1 + 0 and three platellae at the apex of the hind basitarsus. The male plate of B. indica is paral- lel-sided. The connective and aedeagus are articulated rather than fused. The connective is spindle-shaped, lamellate rather than long and incrassate, and the preapical lobe of the style is wanting. These characters strongly suggest that the Afrotropical species of Balhil- lus, namely, B. ahas Linnavuori and B. tri- maculatus Linnavuori, may not belong to Bal- hillus. Key to Species of Balhillus 1. Vertex and scutellum with dark brown spots .... granulosus Distant — Vertex and scutellum immaculate . . . indicus, n. sp. Balhillus granulosus Distant Figs. 1-5 Balhillus granulosus Distant 1908g: 288. Holotype 9 , Sri Lanka (BMNH, examined). Yellow. Vertex with a median spot, a spot in apical half on either side divided by carina, and a spot near inner margin of eye dark brown. Pronotum with a median, short stripe on anterior margin fuscous. Basal triangles on scutum and apex of scutellum dark brown; blackish, median line on scutellum not reach- ing apex of scutellum. Longitudinal veins and claval veins lemon yellow, entire surface of forewing punctate, punctations along radial vein and on outer apical cell and a spot at base of appendix fuscous. Vertex bluntly conical, 1.24 times as wide between eyes as its median length, median carina prominent in its apical half, apical 0.75 of disc sculptured, basal 0.25 polished. Face about as long as wide. Pronotum widened posteriorly. Scutellum longer than pronotum, transversely, finely rugulose beyond sulcus. Forewing 3.4 times as long as wide, with one m-cu cross-vein abut 0.33 distance from base and another near apex limiting the short, in- ner anteapical cell, outer anteapical cell open behind, numerous veins arising from apical half of outer anteapical cell and reaching costa, few of them forked at costal margin, third apical cell divided by cross-vein. Hind- wing with four apical cells. Female genitalia. — Seventh sternum about twice as long as sixth, its hind margin slightly concave with a strong, U-shaped, me- dian excavation, very finely, transversely rugulose. Ovipositor exceeding pygofers. Measurements. — Female 7.10 mm long, head 1.85 mm wide, pronotum 1.87 mm wide. Spegimen e.xamined. — Holotype 9 la- beled /Type/ /Balhillus granulosus Dist., tvpe/ /Ceylon (Green)/ /Distant Coll., 1911-383/ (BMNH). 1988 ViRAKTAMATH, WeSLEY: REVISION OF INDIAN NiRVANINAE 185 /6 17 Figs. 1-17. Species oi Balbillus. B. granulosus Distant; 1, head and thorax; 2, same, profile; 3, face and part of prothorax;4, forewing; 5, ovipositor. B. indicus, n. sp.: 6, general habitus; 7, head and thorax; 8, face; 9, head and thorax, profile; 10, forewing; 11, pygofer; 12, male plate; 13, connective; 14, style; 15, 16, 17, aedeagus, lateral, cephalic, and caudal views. Remarks. — This species is closely related to B. indicus, from which it differs by its distinctive coloration. Balbillus indicus, n. sp. Figs. 6-17 Uniformly pale yellow. Apex of clavus. 186 Great Basin Naturalist Memoirs No. 12 second and third apical cells of forewing suf- fused with brown. Disc of vertex polished, medially, longitu- dinally grooved for 0.66 distance and ridged beyond. Forewing 2.5 times as long as wide, punctate along veins of corium, in cells of clavus, and along the costal margin, m-cu cross-veins two (as in B. granulosus) but not easily seen, fewer oblique veins reaching costal margin from outer anteapical cell. Male genitalia. — Pygofer simple, three times as long as its height with a few stout setae on caudoventral area, anal tube short. Plate parallel-sided, with single row of sub- marginal, stout, short setae, and hairlike setae along outer margin. Style without preapical lobe, apophysis long, its apex pointed and directed ventrad. Aedeagus with preatrium elongate, dorsal apodeme unpaired, short, shaft tubular for greater distance caudad, then flattened and deflected anteriorly, apex pointed; gonopore apical, large. Measurements. — Male 5.50 mm long, head 1.28 mm wide, pronotum 1.53 mm wide. Specimen examined. — Holotype S, India: Kerala: Thekkadi, 27.iii.1977, s' Viraktamath CoU. (UAS). Remarks. — In coloration and size it is simi- lar to B. alhellus Baker from the Philippines, but differs in forewing venation in having five apical cells rather than four as in the latter species. Genus Stenotortor Baker Stenotortor Baker 1923a: 375, 377. Type species: Steno- tortor inocarpi Baker, by original designation and monotypy. Stucture similar to Balhillus but differs in the following respects. Orange with brick red or reddish brown markings. Median carina of vertex more prominent. Forewing without appendix, with one m-cu cross-vein, both an- teapical cells open behind. Hind femoral spin- ulation 2+0+0. Hind tibial spinulation Rj 20, R2 13, R3 10. Apex of hind basitarsus with three platellae. Male pygofer simple, its caudoventral area thickly setose. Anal tube short. Male plate parallel-sided without macrosetae or long, hairlike setae. Style with well-developed, preapical lobe. Connective triangular, articu- lated with simple aedeagus. Gonopore large on ventral margin. Remarks. — Baker (1923a) differentiated Stenotortor from Balhillus by its strongly de- pressed body, tectiform tegmina with ex- panded costal area, strongly curved outer margin, and obscure venation. However, more important differences in Stenotortor ap- pear to be the absence of an appendix on the forewing, the presence of one m-cu cross- vein, and hind femoral spinulation that is 2+0+0. Stenotortor subhiinalaija, n. sp. Figs. 18-27 Orange yellow. Vertex with oblique band on either side of median line, oblique line laterally; pronotum with lateral, oblique band, transverse band on scutellum dark red- dish brown. Forewing with submarginal band on costal area, a stripe on clavus along com- missure at basal 0.33 and then obliquely ex- tending to inner claval margin, a median stripe connecting both claval stripe basally and inward extension of submarginal stripe both medially and caudally, dark reddish brown. Vertex bluntly conical, slightly longer than its width between eyes. Face about as wide as long. Clypellus strongly narrowed apically. Lateral margin of pronotum strongly diverg- ing, 2.7 times as wide as its median length. Forewing 2.4 times as long as its width. Hind- wing with four apical cells. Male genitalia. — Pygofer twice as long as height, caudoventral margin with short, stout setae. Anal tube short. Male plate parallel- sided, about four times as long as wide, caudal apex rounded. Style with well-developed, stout, preapical lobe, apophysis fingerlike. Connective triangular. Aedeagus with elon- gate preatrium, without dorsal apodeme, shaft tubular in basal 0.33, then spatulate with large gonopore on its ventral margin occupy- ing 0.66 of its length. Measurements. — Male 4.90 mm long; head 1.18 mm wide, pronotum 1.55 mm wide. Specimens e.xamined. — Holotvpe 6, In- dia: West Bengal: Sukna, 3.xi.l981, C. A. Vi- raktamath Coll. no. 277 (UAS). Remarks. — This species differs from S. in- ocarpi in coloration of the vertex, pronotum, and wing pattern, in its larger size, and in its relatively longer aedeagus. 1988 ViRAKTAMATH, WeSLEY: REVISION OF INDIAN NiRVANINAE 187 Figs. 18-27. Stenotortor stibhimalaya, n. sp.: 18, habitus; 19, face; 20, head and thorax, profile; 21, forewing; 22, pygofer; 23, male plate; 24, connective; 25,26, aedeagus, lateral and caudal views; 27, style. Tribe Nirvanini Pale yellow with dark or red markings. Body long, narrow, more or less depressed. Head as long as or longer than its width, disc of vertex longitudinally rugose in anterior re- gion. Ocelli near lateral margins in front of eyes. Face with anterior, median carina and usually with lateral, oblique ridges on fronto- clypeus. Gena narrow, lora small. Antennal pits rather deep, near anterior margin of eyes not visible from above; antennae long, reach- ing at least hind margin of pronotum. Forewings narrow, with reduced appendix and venation indistinct basally, second api- cal cell not widened apically. Hindwing with three or four apical cells. Fore and mid- dle tibiae rounded. Hind femoral spinulation 2+1 + 1. Hind basitarsus terminated by two platellae. Pygofer with well-developed, lateral lobes. Anal tube elongate, often with apical or cau- doventral process. Male plate elongate, Ion- longer than pygofer and apically rounded. Style variable, usually with slender apophysis that is curved, hooked, or avicephaliform. Connective free, Y-shaped. Aedeagus weakly sclerotized, usually with appendages. Remarks. — This group is most abundant among Nirvaninae and appears highly evolved compared to other tribes, consider- ing the reduced wing venation, the more streamlined body form, and the specialized male genitalia. 188 Great Basin Naturalist Memoirs No. 12 Key to Genera of Nirvanini 1. Hindwing with four apical cells (Fig. 59) 2 — Hindwing with three apical cells (Fig. 100) .... 3 2(1). Most parts of head, pronotum, and scutellum black dorsally; frontoclypeus and clypellus tu- mid (Figs. 85, 86); male connective rather T- shaped (Fig. 95) Chudania Distant — Dorsum of head, pronotum, and scutellum at most with black, narrow stripe terminated by round spot (Figs. 50, 60); frontoclypeus and clypellus flat (Figs. 29, 45, 49); male connective Y-shaped (Figs. 32, 75) Kana Distant 3(1). Crown of head depressed in middle, margined by carina around its margin (Figs. 268, 269); male plate with angulate projection on its lateral margin near apex (Fig. 272); male style short, stout, appearing like loosely closed fist (Fig. 273) Ophiuchus Distant — Crown of head either flat or convex; male plate either smoothly rounded near apex or with spine; style variable (Figs. Ill, 148, 160, 182), slender and with slender apophysis that is curved, hooked, or avicephaliform 4 4(3). Dorsal apodeme of aedeagus robust, bearing processes at least at base of shaft (Figs. 118, 139, 150, 171, 180, 206); male plate about four times as long as its width at its midlength (Figs. 114, 149, 170, 196); frontoclypeus with prominent, lateral ridges Sophonia Walker — Dorsal apodeme of aedeagus slender, U- shaped, without prominent processes (Figs. 225, 240, 254, 267); male plate five times or more as long as its width at its midlength (Figs. 227, 264); frontoclypeus with weak, lateral ridges Nirvana Kirkaldy Genus Kana Distant Kana Distant 1908g: 285. Type species: Kana thoracica Distant, by original designation. Pale yellow, often with bright-colored patches and stripes. Crown slightly (less than 1.5 times) longer than its width. Ocelli placed laterad of submarginal carina. Frontoclypeus with anterior ridge, rather flattened, with lat- eral, very prominent, ridgelike folds. Clypel- lus large, broad at base. Lora small. Face slightly longer than wide between eyes. Sec- ond apical cell of forewing parallel-sided. Hindwing with four apical cells. Hind tibial spinulation R^ 20, R. 12 ± 1, R, 23. Hind ba- sitarsus with two platellae. Pygofer rounded caudally, with one or two rows of submarginal macrosetae; ventral pro- cess robust at base, narrowed apically. Male plate elongate, narrow, with outer margin sin- uate. Connective Y-shaped, with arms joining broadly; apophysis robust, short, with slender apical extension. Aedeagus partially sclero- tized, often depressed, with lateral, lamellate margin, gonopore surrounded by membra- nous tube with or without process, dorsal apodeme reduced. Remarks. — Some species of Kana, espe- cially K. bispinosa and K. nigropicta, resem- ble externally the species of Sophonia. The four closed apical cells in the hindwing, how- ever, distinguish this genus from Sophonia and Nirvana. The aedeagus of Kana is charac- teristic in that it is bent in its apical half and bears appendages surrounding the gonopore; the second apical cell on the forewing is not narrowed apically as is the case in both Sopho- nia and Nirvana. Kana is closely related to Yaoundea Linnavuori and even may prove to be synonymous with it. Both genera are re- lated to Chudania, from which they differ in having a flatter frontoclypeus. Key to Species of Kana 1. Vertex, pronotum, and scutellum traversed by a median, longitudinal, black line (Figs. 50, 60); third apical cell of forewing with large, round, black spot (Figs. 58, 61) 2 — Vertex or pronotum or both either immaculate or with yellow, reddish, or orange stripes (Figs. 42, 44); forewing with brown or reddish orange, oblique, or transverse bands, black spot on third apical cell, if present verv small (Figs. 36, 45, 47, 79) ; 3 2(1). Vertex without black spot near apex (Fig. 50); aedeagal shaft with pair of pronglike processes (Fig. 55) bispinosa, n. sp. — Vertex with black spot near apex (Fig. 60); aedeagal shaft without processes (Fig. 67) .... nigropicta, n. sp. 3(1). Forewing with transverse bands onlv (Figs. 36, 79) ' 4 — Forewing with both oblique and transverse bands (Figs. 43, 47) 7 4(3). Forewing with single, broad, mottled brown, transverse band, area beyond apical cross-vein fuscous; vertex and pronotum marked with white stripes; male genitalia as in Figs. 74-78 illahorata Distant — Forewing with two bands either both yellowish, orange, or red, or one orange red (more basal) and the other fuscous (median); apical cells at most with a few fuscous spots 5 5(4). Pronotum without colored stripes; vertex with clear yellow, longitudinal stripe on each lateral side; forewing with two transverse, yellow fas- ciae decora (Melichar) — Pronotiun with orange stripes; vertex either im- maculate or with orange-yellow, submarginal, apically converging stripes; forewing with basal, orange-yellow or red, and median, fuscous, transverse stripe 6 1988 ViRAKTAMATH, WeSLEY: REVISION OF INDIAN NiRVANINAE 189 Figs. 28-36. Kana ordinate Distant; 28, head and thorax; 29, same, profile; 30, ventral pygofer process; 31, male plate; 32, connective; 33, style; 34, connective and aedeagus, lateral view; 35, aedeagus, caudal view; 36, forewing. 6(5). Vertex with pair of anteriorly converging, or- ange-yellow stripes continued on pronotum . . . fasciata Pruthi — Vertex without stripes; pronotum with four or- ange stripes; male genitalia as in Figs. 30-35 . . ordinata Distant 7(3). Pronotum with two median, red stripes and two lateral, anterior, lemon yellow spots (Figs. 44); forewing with two red, transverse bands on corium (Fig. 47) thoracica Distant — Pronotum with anterior margin and two spots on posterior margin orange-yellow (Fig. 42); corium of forewing with three transverse, red bands (Fig. 43); male genitalia as in Figs. 37-41 ramificata Distant Kana ordinata Distant Figs. 28-36 Kana ordinata Distant 1908g: 287. Lectotype 6, Sri Lanka (BMNH, examined). Yellow. Pronotum with two inner and two outer orange stripes joined posteriorly in male. Scutellum with brownish basal trian- gles. Forewing with reddish orange band across basal area, fuscous speckled band about midlength and series of fuscous spots at apices of anteapical cell. Head slightly narrower than pronotum. Vertex of head slightly longer than its width between eyes (112:123), apex bluntly conical. Face with four prominent, lateral ridges. Male genitalia. — Pygofer rather squarish with caudodorsal area angulate, and with two rows of stout, long setae; ventral process with triangular, dorsal lobe and median, dorsal hump. Plate with rounded, caudal apex, with row of stout setae in caudal 0.66 and with 190 Great Basin Naturalist Memoirs No. 12 marginal row of long, hairlike setae. Style with well-developed, preapical lobe, apophy- sis short, stout, its apical extension slender, avicephaliform. Connective Y-shaped with stem as long as the length of arms. Aedeagus tubular with unsclerotized, poorly developed dorsal apodeme. Shaft strongly bent and di- rected ventrad and cephalad in apical 0.33, appearing rather like a compressed, inverted J, cephalic part of shaft laterally expanded and constricted in middle in cephalic view; caudal part with two long, curved processes that cross over and are longer than those in K. ramificata. Measurements. — Male body 3.00 mm long, forewing 4.10 mm long, head 1.17 mm wide, pronotum 1.25 mm wide. Female 6.40 mm long, head 1.30 mm wide, pronotum 1.32 mm wide. Specimens examined. — Lectotype S la- beled /Bogawantalawa, Ceylon, April/ /Dis- tant Coll., 1911-383/ /1625/ /Nirvana ordi- nata Dist., type/ here designated (BMNH). Paralectotype 9 labeled /Maskeliya, Ceylon, 8-05/ /Distant Coll., 1911-383/ here desig- nated (BMNH). Remarks. — Kana ordinata is closely re- lated to K. ramificata and differs in coloration and structure of the pygofer process. Kana ordinata has a shorter caudal part of the aedeagus and longer processes that cross over. Kana ramificata Distant Figs. 37-43 Kana ramificata Distant 1908g: 286. Lectotype 6, Sri Lanka (BMNH, examined). Pale yellow, with submarginal band on ver- tex, anterior margin of pronotum and two spots on posterior margin orange. Scutellum golden yellow. Forewing with basal band con- nected by similar but more oblique band from costa to clavus and then continued on clavus but not reaching claval tip, orange; another oblique band with well-defined, orange, ante- rior margin and cross-veins limiting apical cells, reddish; spot at apex of clavus and an- other on third apical cell fuscous. Head slightly narrower than pronotum (46:49). Vertex slightly longer than width be- tween eyes, bluntly conical. Lateral ridges on face four in number, not prominent. Scutel- lum longer than pronotum. Male genitalia. — Pygofer rather squar- ish, with caudal tufts of long, stout setae; ven- tral process stout with dorsal, triangular pro- cess near caudal apex. Anal tube stout. Male plate with oblique row of stout setae in caudal 0.66, with long, hairlike setae, apex bluntly rounded. Style and connective as in K. ordi- nata. Aedeagus as in K. ordinata, but caudal bent part 0.66 as long as tubular cephalic part, apical processes shorter. Measurements. — Male 5.20 mm long, head 1.15 mm wide, pronotum 1.22 mm wide. Specimen examined. — Lectotype S la- beled/Green, Ceylon, 95-221/ /Ntroana ram- ificata Dist., type/ here designated (BMNH). Remarks. — Kana ordinata and K. ramifi- cata share a similar basic pattern of male geni- talia; however, they differ in the relative size of the aedeagal processes and the shape of the pygofer process. Both hindwings and the right side of the forewing of the lectotype are dam- aged and were placed in a microvial. Kana thoracica Distant Figs. 44-48 Kana thoracica Distant 1908g; 285. Lectotype 9 , Sri Lanka (BMNH, examined). Pale yellow. Two anteriorly converging, submarginal stripes on vertex, two stripes one on either side of median line red, two lateral spots on anterior margin of pronotum lemon yellow. Forewing with red bands (Fig. 47) along anal margin, obliquely crossing over to clavus; short band on corium near base, an- other longer, oblique band about midlength from costal margin to clavus. Series of spots on either side of cross-veins separating apical cells fuscous. Head narrower than pronotum (46:50). Vertex longer than its width between eyes (37:26). Face with three lateral, oblique ridges; longer than width including eyes. Pronotum shorter than both vertex (24:37) and scutellum (24:28). Female genitalia. — Hind margin of sev- enth sternum slightly concave with a median protuberance. Measurements. — Female 5.90 mm long, head 1.15 mm wide, pronotum 1.25 mm wide. Specimen examined. — Lectotype 9 la- beled /Pundaloya, Ceylon/ /Ceylon, Green 1988 ViRAKTAMATH, WeSLEY: REVISION OF INDIAN NiRVANINAE 191 Figs. 37-43. Kana ramificata Distant: 37, male pygofer; 38, connective and aedeagus, lateral view; 39, same, caudal view; 40, male plate; 41, style; 42, head and thorax; 43, forewing. Coll. 90-115/ /Nirvana thoracica Dist., type/ here designated (BMNH). Remarks — This species appears to be re- lated to K. ramificata and K. ordinata, from which it differs in coloration. Kana bispinosa, n. sp. Figs. 49-59 Pale yellow. Vertex with median, longi- tudinal, black line from before apex and ex- tending to the tip of scutellum, gradually widening posteriorly, lateral margin orange. Pronotum reddish. Forewings with suffused, red stripe adjoining thin, light brown stripe terminating on claval suture at 0.66 distance from base on anal margin, corium pinkish distally, claval apex and apical wing mar- gin brown, third apical cell with large, black spot, surrounding area light brown, costal margin with three reddish fascia. Head narrower than pronotum. Vertex slightly longer than width between eyes, flat, slightly depressed about center, apex bluntly conical, granulose. Apical half of face con vex, basal half depressed, with three oblique ridges. Male genitalia. — Pygofer rounded, with stout, long setae along caudal margin; ventral process broad basally, abruptly narrowed and curved dorsally near apex. Male plate about four times as long as wide, distal 0.66 with an oblique row of stout spines and with marginal, hairlike setae. Style with stout, preapical lobe and short, stout apophysis, apical extension slender, beaklike. Connective Y-shaped with arms as long as stem. Aedeagus with short, well-developed preatrium; shaft with ridged 192 Great Basin Naturalist Memoirs No. 12 Figs. 44-48. Kana thoracica Distant: 44, head and thorax; 45, head, profile; 46, face; 47, forewing; 48, female seventh sternum. lateral margins, somewhat rectangular, me- dian ridge on caudal aspect running from base to apex, gonopore surrounded by membra- nous, tubelike structure bearing pair of ven- trally directed, pronglike processes. Measurements. — Male 4.50 (4.50-4.60) mm long, head 0.93(0.90-0.95) mm wide, pronotum 0.99 (0.98-1.00) mm wide. Specimens examined. — Holotype d, In- dia: Tamil Nadu: Naduvattam, 6.vi.l977, C. A. Viraktamath Coll. (UAS). Paratypes 2 6, data as in holotype but collected by S. Virakta- math (BMNH, UAS). Remarks. — This species is closely related to Kana nigropicta but only distantly related to K. illaborata and K. ordinata. It differs from the latter two species in coloration of the head, thorax, and forewing and in the struc- ture of male genitalia. Kana nigropicta, n. sp. Figs. 60-68 Uniformly orange-yellow. Vertex with round, black spot near apex, contiguous with median, longitudinal, black line extending to tip of scutellum. Lateral margins of vertex and pronotum orange. Forewing with irregular, hyaline patches, claval apex brown, third api- cal cell with large, black spot, apical margin reddish, costal margin with three red, oblique fasciae, black spot in fourth apical cell. Head as wide as pronotum. Vertex about 1.5 times as long as wide, anterior half convex, faintly granulose. Apical 0.25 of face slightly bulged with four lateral, oblique ridges. Pro- notum transversely, finely rugulose. Scutel- lum longer than pronotum. Male genitalia. — Pygofer roundish with macrosetae along caudal margin; ventral pro- cess broad basally, abruptly narrowed near apex to mesally directed, pointed process. Anal tube moderately long. Male plate more than four times as long as wide, single row of macrosetae on caudal 0.75. Style with small, preapical lobe, apophysis stout with short, relatively stout apical extention. Connective Y-shaped with stem longer than arm. Aedea- gus flattened, elongate with lateral, marginal, poorly sclerotized ridge, dorsal apodeme very short, gonopore large, on caudal margin, sur- rounded bv membranous extension caudallv. 1988 ViRAKTAMATH, WeSLEY: REVISION OF INDIAN NiRVANINAE 193 Figs. 49-68. Species ofKana: K. bispinosa, n. sp.: 49, head and thorax, profile; 50, same, dorsal view; 51, face; 52, male pygofer; 53, style; 54, connective; 55, 56, aedeagus, lateral and ventral views; 57, male plate; 58, forewing; 59, hindwing. K. nigropicta, n. sp.: 60, head and thorax; 61, forewing; 62, male pygofer; 63, apex of ventral process of pygofer in two views; 64, style; 65, connective; 66, male plate; 67, 68, aedeagus, lateral and caudal views. 194 Great Basin Naturalist Memoirs No. 12 Female genitalia. — Hind margin of sev- enth sternum almost straight. Ovipositor ex- ceeding pygofers. Measurements. — Male 4.35 (4.20-4.50) mm long, head 0.92 (0.90-0.95) mm wide, pronotum 0.95 (0.90-0.95) mm wide. Female 5.13 (5.00-5.20) mm long, head 1.03 (1.00-1.08) mm wide, pronotum 1.11 (1.08-1. 15) mm wide. Specimens examined. — Holotype 6, In- dia: Kerala: Thekkadi, 26.iii.1977, C. A. Vi- raktamath Coll. (UAS). Paratypes: 5 d, 2 9, data as in holotype but collected on 27. iii. 1977 by C. A. Viraktamath (2 d, 1 9) (BMNH, USNM, UAS). Remarks. — This species can be easily rec- ognized by the round, black spot on the vertex and the absence of processes on the membra- nous tube surrounding the gonopore. Kana illaborata Distant Figs. 69-79 Kana illaborata Distant 1908g: 287. Lectotype 3 , Burma (BMNH, examined). Yellow. Vertex with median, white stripe. Pronotum with narrow, median, irregular, lateral spot whitish. Forewing with broad, median, mottled, brown band on apical 0.33. Head about as wide as pronotum (46:47). Vertex about as long as wide in male (29:28) but longer than its width in female (38:34). Pronotum shorter than both vertex and scutellum. Face with four lateral, oblique ridges and an apical, short, median ridge. Male genitalia. — Pygofer with lobelike, membranous process, armed with tooth mesally; caudal margin with series of hairlike setae. Male plate with oblique row of stout setae in caudal half, and with marginal, hair- like setae. Preapical lobe of style stout, apoph- ysis relatively slender, apical extension slen- der. Connective Y-shaped, with stem as long as arm. Aedeagus laterally compressed, mem- branous for the most part, with pair of slender ventral processes and sclerotized tooth ven- tral to gonopore, which is elongate; pair of lamellate processes located in apical half of shaft. Measurements. — Male 5.70 mm long, head 1.15 mm wide, pronotum 1.17 mm wide. Female 6.30 mm long, vertex 1.37 mm wide, pronotum 1.42 mm wide. Specimens examined. — Lectotype 6 la- beled /Myitta (Doherty)/ /Distant Coll. 1911-383/ /Kana illaborata Dist., det. M. Webb, 1980/ here designated (BMNH). Paralectotype 9 labeled /Tenass. Vail. Myitta (Doherty)/ /Distant Coll. 1911-383/]Vtrt;ana illaborata Dist. type/ here designated (BMNH). Remarks. — ^This species is very distinctive both in coloration and male genitalic charac- ters and does not appear closely related to other species of Kana. Kana decora (Melichar) Figs. 80-81 Nirvana decora Melichar 1903b: 166. Holotype 9 , Sri Lanka (ZMHU, examined). The holotype female is a teneral specimen, and the diagnostic coloration is not very well developed. Melichar (1903b) has given a good description of this species and hence it is not redescribed here. Specimen examined — Holotype 9, la- beled /Type/ /6178/ /Ceylon, nictus/ /Nirvana/ /decora m. det. Melichar/ /Nirvana decora Melichar/ (ZMHU). Kana fasciata Pruthi Kanafasciata Pruthi 1930a; 22. Syntypes d , 9 , Sri Lanka (ZSI, not examined). The types of this species were not available for study at the time the senior author visited ZSI. Pruthi's (1930a) description and illustra- tions suggest this to be a species with very distinctive coloration. Genus Chudania Distant Chudania Distant 1908g: 268. Type species Chudania delecta Distant, by original designation and mono- typy. Yellow, with prominent broad stripe run- ning on head, thorax, and along inner margin of clavus, extending laterally to occupy entire apex of forewing, black. Vertex either as long as or slightly shorter than width, disc convex without median groove. Anterior area of disc rugose. Ocelli in front of and mesad of eyes. Frontoclypeus convex, with four to six transverse, prominent rugae evanescent medially on dorsal aspect and with median ridge in upper 0.75. Lora prominent. Clypellus large, slightly narrowed apically. Pronotum twice as wide as long, slightly longer than scutellum, polished and transversely, finely rugulose. Forewing with 1988 ViRAKTAMATH, WeSLEY: REVISION OF INDIAN NiRVANINAE 195 tho'Sfe^l^trf '^pS.^r -^'^■' '': -r' ^™^''^^ ''- --' ^-- ^2' head and 79, .a,e forewing'. .. ^.^^ (tuchT^^^^^^^^^^ -^-^-> lateral and caudal views; 196 Great Basin Naturalist Memoirs No. 12 appendix wanting, apical cells four and an- teapical cells two. Hindwing with four apical cells. Hind tibial spinulation Rj 18 ±2, Ro 12±1, R3l2±l. Male pygofer with rounded, caudal margin, length shorter than height, strongly sclero- tized with single, submarginal row of long setae and ventral process. Anal tube stout and large, about 0.8 times as high as height of pygofer. Male plate with uniseriate macrose- tae in apical 0.66 and with long, hairlike setae. Style with prominent preapical lobe, apophy- sis slender, apex avicephaliform. Style rather T-shaped. Aedeagus membranous for most part, bent in midlength, laterally expanded, with ventrally directed, subapical, pronglike, sclerotized processes surrounding gonopore. Remarks. — This genus is related to Kana and Afrokana Heller with which it shares hindwing venation; with Afrokana it shares the structure of the male pygofer. However, they differ in the shape of the head. Lin- navuori (1979) stated that A/roniruana Evans, Chudania, and Afrokana are very close, sharing, for instance, the same unique genital structure. He believes the recorded differ- ences are too slight for separating them as valid genera. Afrokana is closer to Kana than to Chudania considering the peculiar shape of the aedeagus they share. It is likely that the study of more material of these genera, espe- cially from Malaysia, Indonesia, and the Philippines, may prove that these taxa are congeneric with Chudania. Chudania delecta Distant Figs. 82-96 Chudania delecta Distant 1908g: 268. Holotype 9 , India (BMNH, examined). Male. Slender, darker than female. Vertex, pronotum, scutellum, upper part of face at level with lower margins of eyes, chocolate brown. Ocelli white, eyes dark brown. Forewing yellow, with posteriorly widening, black stripe along inner margin, occupying entire apical 0.33 of forewing (Fig. 82); two white, hyaline, triangular areas on costal area near apex and some cross-veins enclosing api- cal cells paler. Lower half of face, sterna, and legs yellow. Pygofer and fifth to eighth terga blackish. Female. Similar to male but paler. Black markings of male, chocolate brown. Entire face, lateral margins of vertex in front of eyes, and pronotum yellow. Male genitalia. — Pygofer rounded cau- dally, with submarginal row of macrosetae, ventral process broad caudodorsally with finely serrated, dorsal margin. Male plate more or less of uniform width, caudally rounded, with outer, marginal, long, hairlike setae. Anal tube large, about as long as length of pygofer. Style with anterior part larger than posterior part to the point of articulation with connective. Stem of connective thrice as long as arms, apex expanded. Aedeagus membra- nous for most part, processes sclerotized, bent about its midlength, apical half robust, preatrium with short ventral and longer lat- eral processes; two pairs of processes arising where shaft bends, dorsal pair elongate, curved at midlength, the ventral pair shorter, three to five branched. Female genitalia. — Hind margin of sev- enth sternum straight. Ovipositor exceeding length of pygofer. Distal half of pygofer black. Measurements. — Male 4.62 (4.60-4.70) mm long, head 1.07 mm wide, pronotum 1.11 (1.07-1.12) mm wide. Female 5.38 (5.20- 5.40) mm long, head 1.20 mm wide, prono- tum 1.22 (1.20-1.25) mm wide. Specimens examined. — Holotype 9 la- beled /Type, H.T./ /Chudania delecta Dist., type/ /Kurseong/ Distant Coll. 1911-383/ (BMNH). India: West Bengal: 8 d, 7 9, Kurseong, 1,483 m, 22.x. 1981, C. A. & S. Viraktamath; 1 9, 15km E Kahmpong, 1,780 m, 27.x. 1981, C. A. v.; 4 c?, 11 9, Kalimpong, 1,768 m, 29.x. 1981, S. V. ; 9 c? , 19 9 , 8 km E of Kalimpong, 1,968 m, 29.x. 1981, C. A. V. Mi- zoram: 11 d, 11 9, Aizawl, 18.xi.l981, C. S. Wesley; 11 c?, 14 9 , Lungleh, 20-25. xi. 1981, C.S.W. (UAS). Remarks. — Chudania delecta and C. afri- cana Heller are closely related but differ in coloration, shape of the ventral pygofer pro- cess, and the processes on the aedeagus. Also, the dorsal half of the aedeagus is more robust in C. delecta than in C. africana. The male genitalia of the third species, C. exposita Ja- cobi, is not known. Genus Sophonia Walker Sophonia Walker 1870b: .327. Type species: Sophonia rufitehim Walker by monotypy. Pseudonirvana Baker 1923a: 386. Type species: Pseudonirvana sandakanensis Baker, by original designation. 1988 ViRAKTAMATH, WESLEY; REVISION OF INDIAN NiRVANINAE 197 Figs 82-96 Chudania delecta Distant: 82, habitus, male; 83, head and thorax, male; 84, face, male; 85, head and thorax, profile, male; 86, same, female; 87, forewing; 88, hindwing; 89, ovipositor; 90, male pygofer; 91, ventral process of pygofer; 92, style; 93, 94, aedeagus, lateral and dorsal views; 95, connective; 96, male plate. 198 Great Basin Naturalist Memoirs No. 12 Quercinirvana Ahmed & Mahmood 1970: 260. Type spe- cies: Quercinirvana longicephala Ahmed & Mah- mood, by original designation. New synonymy. Head as wide as or wider than pronotum. Vertex as long as or longer than width be- tween eyes, anterior half rugulose or granu- lar, disc either flattened or slightly convex. Ocelli lateral in front of eyes. Frontoclypeus depressed, with prominent, lateral ridges and with median, apical ridge; face longer than width including eyes. Forewing with second apical cell narrowed apically. Hindwing with three apical cells. Hind tibial spinulation R, 20±2, R2l2±l, R3+ 1, R4l8±2. Pygofer rounded or obtusely angled cau- dally, with or without caudal or ventral pro- cess; macrosetae confined to caudal 0.33 area. Anal tube prominent. Male plate broader than in Nirvana, three to four times as long as its width and rather parallel-sided with oblique row of stout spines in caudal 0.66 area and hairlike setae. Connective Y-shaped, its stem usually more than 1.5 times as long as each arm. Aedeagus with dorsal apodeme well developed, often with processes; preatrium variable, with or without processes; shaft tubular or lamellate, with or without process. Remarks. — Evans (1947a) suppressed Pseudonirvana as a junior synonym of S apho- nia. This genus includes brightly colored spe- cies, often with red or orange streaks. It is closely related to Kana and Nirvana. It can be diffierentiated mainly by its shorter male plates and complex aedeagus that has pro- cesses on the dorsal apodeme and apex of the shaft. The holotype oi Sophonia rnfitelum is a female (Figs. 97-102). The relationships dis- cussed here are based on species assigned to this genus from the Indian subcontinent. Key to Species o( Sophonia 1. Vertex as long as wide between eyes (Figs. 186, 198) 2 — Vertex longer than wide between eyes (Figs. 112, 208) ' 3 2(1). Male pygofer with long, ventral and short, dor- sal process (Fig. 203); preatrium without pro- cesses bifida, n. sp. — Male pygofer without processes; preatrium with a pair of processes (Fig. 191) modesta (Distant) 3(1). Vertex and pronotum with either single or paired, black, longitudinal stripe(s) usually connected to black spot or spots near apex of vertex (Figs. 102, 1 12, 122) 4 — Vertex and pronotum without black, median stripe or spot near apex; vertex sometimes with median, bright crimson streak (Figs. 164, 177) or orange stripe (Fig. 154) 8 4(3). Vertex with two rather oval, contiguous spots, each connected to longitudinal, black line (Figs. 103, 104); aedeagal shaft bifid (Figs. 109, 110) linealis (Distant) — Vertex with a large, round, black spot at apex either connected to or short distance away from one or paired longitudinal line(s) (Figs. 112, 122); aedeagal shaft not bifid 5 5(4). Male pygofer without ventral or caudal process (Fig. 125); aedeagus as in Fig. 129; median, black lines on vertex not connected to its api- cal, round spot (Fig. 122) longitudinalis (Distant) — Male pygofer with ventral or caudal process; aedeagus variable but not as above (Figs. 118, 150); median, black stripe(s) of vertex con- nected to spot at apex (Figs. 112, 132) 6 6(5). Vertex with single, median, black stripe (Fig. 142); male pygofer process short and narrow, apophysis of style hooklike; apex of each arm of dorsal apodeme with two long and short, me- dian processes (Fig. 150) keralica, n. sp. — Vertex with two median, black stripes (Fig. 112); male pygofer process broad and long; apo- physis of style beaklike; each arm of dorsal apodeme with a process on caudal margin (Fig. 118) 7 7(6). Male pygofer process with serrated, dorsal margin; process of dorsal apodeme exceeding width of shaft; shaft with pair of long, caudally directed processes (Fig. 118) longicephala (Ahmed & Mahmood) — Male pygofer process not serrate apically; pro- cess of dorsal apodeme not exceeding width of shaft; processes of shaft short with basal, short, ventral tubercle (Figs. 139, 141) . . bakeri, n. sp. 8(3). Male pygofer without process; preatrium of aedeagus with process (Figs. 171, 180) 9 — Male pygofer with process (Figs. 159, 214); preatrium of aedeagus without process 10 9(8). Preatrium longer than broad in lateral aspect; longer process of dorsal apodeme not divided (Fig. 171); vertex with short, median, crimson streak in apical 0.66 but not reaching apex . . . insignis (Distant) — Preatrium broader than long, longer process of dorsal apodeme bifid (Fig. 180); vertex uni- formly whitish yellow with median, crimson streak bifid near apex (Fig. 175) . complcxa, n. sp. 10(8). Vertex with median, broad, orange stripe con- tinued on pronotinn, anteriorly narrowed; pos- terior half on pronotum broadened and black- i.sh, scutellum blackish (Figs. 154, 155); pygofer process short, bladelike, and pointed at apex (Fig. 159); aedeagus shaft tubular with apical, ventrally directed, platelike process (Figs. 162, 163) illuminata (Distant) 1988 ViRAKTAMATH, WESLEY: REVISION OF INDIAN NiRVANINAE 199 Figs. 97-102. Sophonia rufitehnn Walker: 97, head and thorax; 98, face; 99, head and thorax, profile; 100, hindwing; 101, basal half of clavus; 102, ovipositor. — Vertex, pronotuni, and scutellum whitish yel- low; pygofer process long, slender, and el- bowed (Fig. 214); aedeagus not as above (Fig. 218) complicata, n. sp. Sophonia linealis (Distant) Figs. 103-111 Nirvana linealis Distant 1908g: 282. Holotype 9 , India (BMNH, examined). Nirvana greeni Distant 1908g: 283. Holotype 9 , Sri Lanka (BMNH, examined). New synonymy. Yellow. Vertex with two piceous, apical, elongate, fused spots from which longitudi- nal, piceous lines traverse posteriorly meet- ing posterior margin of vertex, lateral margin often with orange-yellow stripe confined to ocelli or extending anteriorly. Pronotum and scutellum with median, longitudinal, piceous line often interrupted before apex of scutel- lum. Claval margin of forewing piceous with piceous stripe bent obliquely near claval apex and reaching claval suture; spot on second apical cell and two oblique lines from costa in apical half piceous. Head about as wide as pronotum. Vertex more than 1.5 times as long as wide in male, nearly twice as long as wide in female, lateral margin raised. Scutellum longer than prono- tum. Second apical cell of forewing narrowed apically. Male genitalia. — Pygofer ovate, without ventral process. Plate slightly more than 3.5 times as long as median width, slightly nar- rowed caudally. Style with broad, preapical lobe and beaklike extension of apophysis rela- tively short. Stem of connective flared cau- dally, about twice as long as each arm. Dorsal apodeme of aedeagus robust, each arm with caudally directed, elongate process that ex- ceeds width of shaft; shaft tubular, curved, hooked apically where it appears bifid. 200 Great Basin Naturalist Memoirs No. 12 Figs. 103-121. Sophonialinealis {Distant): 103, head and thora.x of holotype female Niruaria greeni Dist.; 104, head and thorax of holotvpc of Nirvana linealis Dist.; 105, profile; 106, ovipositor; 107, male pygofer; 108, forewing; 109 110, aedeagus, lateral and dorsal views; HI, connective and style. S. km^icephala {Ahmed in Mdhmood): 112, head and thorax; 113, face; 114, male plate; 115. male pygofer; 116, ventral process of pygofer; 117, style; 118, 119, aedeagus, lateral and caudal views; 120, connective; 121, forewing. 1988 ViRAKTAMATH, WeSLEY: REVISION OF INDIAN NiRVANINAE 201 Female genitalia. — Hind margin of sev- enth sternum straight with median protuber- ance. Measurements. — Male 4.06 (4.00-4.10) mm long, head 0.83 (0.80-0.85) mm wide, pronotum 0.88 (0.85-0.93) mm wide. Female 4.82 (4.50-5.00) mm long, head 0.93 (0.88-0.95) mm wide, pronotum 0.98 (0.93-1.03) mm wide. Specimens examined. — Holotype 9, la- beled/Type, H.T./ZPeradeniya, 2.08, Ceylon/ /Distant Coll. 1911-383/ /Nirvana greeni Dist., type/ (BMNH). Holotype 9, labeled /Type, H.T./ /Calcutta, 6.6.07/ /Distant Coll. 1911-383/ /Nirvana linealis Dist., type/ (BMNH). India: Karnataka: 6 9, Dharwar, 26.iii.1972; 1 9 29. iv. 1969; 1 9, — .i.l970; 1 9, 14. iv. 1972; 1 9, — .x.l979 C. A. Virakta- math Coll.; 3 d, 3 9, Chamarajanagar, 12.viii.l977; 4 9, Hunsur, 16.i.l978; 1 9, Bangalore, 11. vi. 1977; 1 9, Arsikere, 23.vii.1978; 1 d, 1 9, 16.i.l979; 1 9, Sulik- ere, 30.xii.l976, all collected by C. A. Virak- tamath (UAS). Kerala: 3 c? , 6 9 , Kayangulum, — .viii. 1983, ex coconut (UAS). Remarks. — The holotypes of linealis and greeni show color differences, especially the extension of the median, dark fuscous stripe of the pronotum along the inner margin of the clavus in greeni, which is absent in linealis. The female seventh sterna are identical; hence the species are considered as syn- onyms. This species is unique among Sopho- nia in having a bifid aedeagal shaft. Sophonia longicephala (Ahmed & Mahmood), n. comb. Figs. 112-121 Quercinirvana longicephala Ahmed & Mahmood 1970: 263. Holotype <5, Pakistan (University of Karachi, not examined). Pale yellow, elongate, black spot near apex of vertex contiguous with two longitudinal, black lines that fuse at base of vertex and continue as single line to near tip of scutellum. Forewing with broad, brown stripe along claval commissure reaching claval apex and obliquely crossing corium, apical margin suf- fused with brown, costal margin with ante- rior, oblique and two posterior, transverse, black fasciae; second apical cell with black spot. Head nearly as wide as pronotum. Vertex about 0.33 times longer than wide, disc de- pressed, faintly granulose. Second apical cell narrowed caudally. Male c;enitalia. — Pygofer trapezoidal, with long, caudally direct(^d ventral process dorsally serrated. Plate about four times as long as wide, distal 0.66 with a single row of macrosetae and hairlike setae. Style with large, preapical lobe, beaklike extension of apophysis long. Aedeagus with dorsal apodeine well-developed, caudally directed, apically pointed processes that exceed width of shaft, shaft tubular, dorsoanteriorly curved with subapical, caudally directed, slender, elongate processes. Measurements. — Male 4.50 (4.30-4.70) mm long, head 0.91 (0.90-0.93) mm wide, pronotum 0.96 (0.93-1.00) mm wide. Specimens examined. — India: Haryana: 1 6, Kalka, 29.i.l979, C. A. Viraktamath; Hi- machal Pradesh: 1 6, Kulu, l.iv.l978, I. Dworakowska (UAS). Remarks. — This species is related to and resembles externally S. bakeri. The spot on the vertex in this species is rather oval (as in bakeri), and it differs in the details of the male pygofer process and the aedeagus. Sophonia longitudinalis (Distant) Figs. 122-1.31 Nirvana longitudinalis Distant 1908g: 283. Lectotype 9 , Burma (BMNH, examined). Vertex pale yellow with large, subapical, black spot and two median, longitudinal lines, not reaching black spot and fused near hind margin of vertex, continued on pronotum and scutellum as median stripe and on clavus as irregular stripe. Forewing with large, round, black spot in second apical cell extending partly to first apical cell, apical margin suf- fused with brown. Head as wide as pronotum or slightly nar- rower. Vertex about 1.5 times as long as wide, apex subangular, disc mildly convex, granu- lose. Second apical cell narrowed caudally. Male genitalia. — Pygofer ovate, ventral process wanting, macrosetae limited to distal 0.33. Plate about 3.5 times as long as width, with short tooth at the outer apical margin. Style with short, preapical lobe, apophysis with long, beaklike extension. Aedeagus with robust, dorsal apodeme, each of its arms bear- ing an elongate, caudally directed process, preatrium wanting, shaft lamellate with lat- eral margin caudally curved, widest at apical 202 Great Basin Naturalist Memoirs No. 12 Figs. 122-141. Sophonialongitudinalis (Vistsint): 122, head and thorax; 123, face; 124, ovipositor; 125, male pygofer; 126, 127, male plate; 128, connective and stvle; 129, 130, aedeagus, lateral and caudal views; 131, forewing. S. I^^ken, n. sp.: 132, head and thorax; 133, face; 134, forewing; 135, male pygofer; 136, pygofer process; 137, male plate; 138, connective; 139-141, aedeagus, lateral, caudal, and dorsal views. 1988 ViRAKTAMATH, WeSLEY: REVISION OF INDIAN NiRVANINAE 203 half with long, slender, arcuate, apically pointed, ventrally curved process that bears a short tooth on its outer margin at base. Female genitalia. — Hind margin of sev- enth sternum slightly concave. Ovipositor slightly exceeding pygofer. Measurements— Male 4.43 (3.80-5.40) mm long, head 0.96 (0.80-1.43) mm wide, pronotum 0.96 (0.85-1. 13) mm wide. Female 4.90 (4.60-5.30) mm long, head 1.02 (0.98-1.10) mm wide, pronotum 1.05 (1.00-1. 10) mm wide. Specimens examined. — Lectotype 9 la- beled /Myitta (Doherty)/ /Distant Coll. 1911-383/ /Nirvana longitudinalis Dist., type/ here designated (BMNH). India: Karna- taka: 6 (5, Mudigere, 4-6. iv. 1980, C. A. Vi- raktamath; 1 9, 7.iv.l975, C.A.V.; 1 9, Kodlipet, 6.iv.l975; 1 S, Hunsur, 10. i. 1978; 1 9, Bannerghatta, 12. ix. 1976, B. Mallik; Kerala: 1 9, Thekkadi, 27.iii.1977, C.A.V. Nepal: Nagarkot, 1 (5, 1 9, 15.ix.l979,- I. Dworakowska (UAS). Burma: 1 9, Amberst Dist., Kawkareik, 19-20.xi.l911, F.H.G. (ZSI). Remarks. — This species can be distin- guished from all other species of Sophonia by its lamellate aedeagal shaft and tooth on the apex of the male plate. Sophonia bakeri, n. sp. Figs. 132-141 Pale yellow to whitish yellow. Vertex with black spot near apex, joined by two black, longitudinal lines that fuse basally and extend to scutellum. Pronotum and scutellum some- times with reddish spots, scutellum in male entirely reddish. Forewing with brown stripe close to or along anal margin, marked with yellow along inner margin, apex of clavus and apical margin brown, costal margin with ante- rior, oblique and posterior, transverse, brown fasciae, second apical cell with large, black spot. Vertex about 1.5 times as long as wide, apex subtriangular, disc depressed. Posterior half of pronotum transversely rugulose. Second apical cell narrowed at apex. Male genitalia. — Pygofer with ventrocau- dal angle produced to process with attenuated apex. Plate 3.5 times as long as wide. Apophy- sis of style with long, beaklike extension. Stem of connective 1.5 times as long as each arm. Aedeagus with each arm of dorsal apo- deme robust, bearing triangular process on caudal margin; preatrium short; shaft tubular, caudodorsally curved, apex strongly incurved with ventrally directed, short, pronglike pro- cess on caudal margin subapically. Female genitalia. — Hind margin of sev- enth sternum concavely emarginate. Oviposi- tor exceeding length of pygofer. Measurements,— Male 4.48 (4.30-5.00) mm long, head 0.90 (0.85-1.00) mm wide, pronotum 0.88 (0.78-1.03) mm wide. Female 5.08 (4.70-5.30) mm long, head 1.04 (0.93- 1.23) mm wide, pronotum 1.05 (1.00-1.10) mm wide. Specimens examined. — Holotype 6, In- dia: Karnataka: Mudigere, at light, 4-6.iv.l980, C. A. Viraktamath (UAS). Paratypes: India: Karnataka: 2 6, Dharwar, — .i.l970, C. A. Viraktamath; 1 9, Kodlipet, 6.iv.l975, C.A.V. ; 1 9, Bangalore, 9.iii.l975, C.A.V.; 1 9, Kalhalla, 18.i.l978, C.A.V., 1 9, Mudigere, 23. ix. 1973, C.A.V.; 1 (?, 1 9, Kemmanagundi, 8.iv.l975, K. D. Ghorpade; Kerala: 1 c?, Thekkadi, 27.iii.1977, S. Virakta- math; Maharashtra: 1 9 , Matheran, 915 m, 23.xi.1977, C.A.V.; Tamil Nadu: 1 6, Ko- daikanal, 2,250 m, 10. vi. 1980, C. S. Wesley; Uttar Pradesh: 1 6, 1 9, Dehra Dun, 28. iv. 1975, C.A.V.; Nepal: 2 9, Gokarna, 14. ix. 1979, I. Dworakowska; 1 9, Sarankot, 17. ix. 1979, exQiiercus sp., I.D. (1 6 and 1 9 in BMNH, lARI, USNM, and rest in UAS). Remarks. — Sophonia bakeri is related to S. longitudinalis and S. longicephala as they share similar external coloration and male genitalia. They form a closely knit group. S. bakeri differs from the other two by its shorter process of the dorsal apodeme and aedeagal process. Sophonia keralica, n. sp. Figs. 142-153 Uniformly yellow. Vertex with round, black spot near apex connected by single, median, longitudinal, black line, discontinuous for short distance near black spot and then con- tinued to apex of scutellum. Lateral margin of vertex orange basally. Forewing with claval suture and punctures along cubital vein brown, second and part of third apical cells occupied by large, black spot; costal margin with anterior, oblique and posterior, trans- verse, black fasciae, apex of clavus and apical margin brownish. 204 Great Basin Naturalist Memoirs No. 12 Figs. 142-153. Sophonia keralica, n. sp.: 142, head and thorax; 143, face; 144, head and thorax, profile; 145, male pygofer; 146, pygofer process; 147, connective; 148, style; 149, male plate; 150, 151, aedeagus, lateral and cephalic views; 152, forewing; 153, hindwing. Head nearly as wide as pronotum. Vertex slightly longer than wide, disc flat, depressed near apex. Scutellum slightly longer than pronotum. Second apical cell of forewing nar- rowed slightly near apex. Male genitalia. — Pygofer caudally rounded, its ventrocaudal angle produced to sharp, thin process, five macrosetae along caudal margin in straight line, black, rest hya- line. Anal tube as long as pygofer. Plate four times as long as wide. Style with strongly hooked apophysis. Stem of connective 1.5 times as long as each arm. Dorsal apodeme of aedeagus slender, each arm with ventral, dor- sal, and short, median process; shaft tubular with pair of laterally directed processes, gonopore subapical. Measurements. — Male 3.90 mm long, head 0.90 mm wide, pronotum 0.93 mm wide. Specimen examined. — Holotvpe 6, India: Kerala: Thekkadi, 27.iii.1977, s! Viraktamath Coll. (UAS). Remarks. — Sophonio keralica is related to S. iUuminata. Both have similar male geni- talia. They differ, however, in coloration and detailed structure of the aedeagus. Sophonia iUuminata (Distant), n. comb. Figs. 154-163 Kana iUuminata Distant 1918b: 35. Lectotype d, India (BMNH, examined). Kana sigruiffl Distant 1918b: 34. Lectotype 9 , India (BMNH, examined). New synonymy. Female. Vertex and pronotum creamy white with yellowish lateral areas, median stripe narrowing to apex of vertex san- guineous on vertex and anterior half of prono- tum, widened in posterior half where it is black. Scutelhun black except for two lateral, basal spots. Clavus along claval suture and apical half piceous except for subapical, elon- gate, orange spot. Corium along its inner mar- gin piceous which widens near apex of cla- vus and changes to orange; appendix fuscous, two large, round spots in second apical cell. 1988 ViRAKTAMATH, WESLEY: REVISION OF INDIAN NiRVANINAE 205 Figs. 154-163. Sophonia illuminata (Distant): 154, head and thorax of lectotype male Kana illuminata; 155, head and thorax of lectotype female Kana signata Distant; 156, forewing, female; 157, ovipositor; 158, apex of male pygofer; 159, pygofer process; 160, style; 161, connective; 162, 163, aedeagus, lateral and caudal views. oblique line at apical 0.33 of costa and more distal, transverse band piceous, rest of wing yellow and areas anterior to piceous spots whitish. Male. Coloration less extensive compared to female. Stripe on vertex continuing on pronotum and scutellum, gradually widening and sanguineous throughout. Forewing col- oration comparatively less developed. Head narrower than pronotum, median length of vertex slightly longer than width between eyes. Face longer than wide. Male genitalia. — Pygofer lobe gradually narrowed caudally and terminated by sharp, triangular process and with series of stout, long spines. Plate parallel-sided, with six stout setae in oblique row commencing at midlength, third to fifth setae black in color. Style with stout, preapical lobe and hooklike, rather slender apophysis with attenuated apex. Connective with stem 1.5 times as long as each arm. Aedeagus with short, dorsal pair of blunt processes and ventral, fingerlike pro- cess; shaft robust at base, narrowed apically with dorsal, large gonopore surrounded by two lateral, pronglike processes and ventrally directed, lobelike plate. Female genitalia. — Hind margin of sev- enth sternum concave. Ovipositor exceeding length of pygofer. 206 Great Basin Naturalist Memoirs No. 12 Measurements. — Male 4.40 mm long, head 1.00 mm wide, pronotum 1.05 mm wide. Female 5.1 mm long, head 1.12 mm wide, pronotum 1.12 mm wide. Specimens examined. — 3 syntype 6 of Kana illuminata mounted on single card la- beled /S. India, Madras, Kodaikanal, T. V. Campbell, K.K.4.16/. One of these was re- moved, remounted, and relabeled by the se- nior author, /Kodaikanal, S. India, T. V. Campbell/ and is here designated as lecto- type; others designated paralectotypes. One paralectotype 6 labeled /Kodaikanal, India, T. V. Campbell/ /S. India, E. A. Butler, 1915-60/ /Kana illuminata Dist., type/ (BMNH). Lectotype 9 labeled /Kodaikanal, S. India, T. V. Campbell, K.K.5.11//S. India, E. A. Butler, 1915-60/ /54,9/ /Kana signata Dist., type/ here designated (BMNH). India: Tamil Nadu: 1 9, Trichinopoly, J. Dubre- nil(IRSNB); 1 9 Kadaikanal, 2,030-2,150 m, — .viii.1922, S. Kemp(ZSI). Remarks. — Sophonia illuminata and S. sig- nata are the male and female, respectively, of the same species based on the similarity in external coloration and structure. This species is closely related to S. keralica. Both have a similar type of male genitalia but differ in coloration and structure of the pygofer pro- cess and aedeagus. Sophonia insignis (Distant), n. comb. Figs. 164-174 Nirvana insignis Distant 1918b: 33. Lectotype 6, India (BMNH, examined). Whitish yellow. Median groove on vertex crimson from about center to just short of apex, lateral marginal area orange. Forewing hyaline, light brown stripe along anal margin rarely reaching apex of clavus, if so occupying entire quarter of clavus, or stripe wanting, apex of clavus brown; costal margin with two long, oblique fasciae, third transverse and fourth short, broad, oblique fasciae in the dis- tal half brown; apical margin brown; second apical cell with black spot. Head narrower than pronotum. Vertex about 1.5 times as long as wide between eyes; disc flat or slightly depressed toward base, apex granulose. Second apical cell of forewing slightly narrowed distally. Male genitalia, — Caudal lobe of pygofer rounded, without processes. Plate about 3.5 times as long as wide. Preapical lobe of style robust, apophysis slender, with apical exten- sion forming short, beaklike extension. Aedeagus with preatrium well developed, with ventrocaudally directed process, about as long as preatrium; dorsal apodeme broad, platelike, with two processes (one short, pronglike on cephalic margin, other long, slender, and inwardly curved in middle) and ventral, short process; shaft tubular, curved anteriorly then ventrally near apex. Female genitalia. — Hind margin of sev- enth sternum straight. Ovipositor slightly ex- tending beyond pygofer. Measurements— Male 5.00 (4.80-5.20) mm long, head 0.97 (0.93-1.00) mm wide, pronotum 1.04 (1.03-1.07) mm wide. Female 5.62 (5.00-6.10) mm long, head 1.06 (0.98- 1.13) mm wide, pronotum 1.14 (1.03-1.23) mm wide. Specimens examined. — Lectotype S la- beled /Tenmalai, W. Ghat (W. side). Travan- core, 22.xi.08, Annandale/ /Distant Coll. 1911-383/ here designated (BMNH). Paralec- totype 9 , with same data as in holotype, here designated (ZSI). India: Kerala: 11 (5, 6 9, Thekkadi, 26-27. iii. 1977, C. A. Viraktamath, S. Viraktamath, and B. Mallik Coll.; Kar- nataka: 2 9, Mudigere, 7.iv.l980, C.A.V. (UAS). Remarks. — This species is closely related to S. complexa and can be differentiated by the structure of male genitalia and coloration. Sophonia complexa, n. sp. Figs. 175-185 Similar to S. insignis but more elongate. Vertex with median, crimson-colored line oc- cupying about 0.66 area and apically bifid, short, slightly oblique, sublateral stripe on apical half of vertex orange. Forewing with fuscous spots (Fig. 178). Apical 0.75 of vertex elongately rugulose, disc flat, margined by carina. Male genitalia. — Pygofer elongate, heav- ily setose, caudally rounded. Anal tube stout, long. Plate parallel-sided, about 3.5 times as long as wide, macrosetae restricted to apical 0.66. Apophysis of style short, apex avicephal- iform. Connective with stem about three times as long as each arm. Aedeagus complex, each arm of dorsal apodeme with pair of ven- trally directed processes, preatrium with cau- dally directed process, shaft tubular, slightly curved dorsally near apical 0.33, slightly flared. 1988 ViRAKTAMATH, WESLEY: REVISION OF INDIAN NiRVANINAE 207 173^ 170 174 Figs. 164-174. Sophonia insignis (Distant): 164, head and thorax; 165, face; 166, ovipositor; 167, male pygofer; 168, style; 169, connective; 170, male plate; 171-173, aedeagus, lateral, cephalic, and caudal views; 174, forewing. Female genitalia. — Hind margin of sev- enth sternum slightly concave. Measurements.— Male 5.32 (5.20-5.50) mm long, head 1.00 (0.97-1.02) mm wide, pronotum 1.06 (1.05-1.07) mm wide. Female 5.92 (5.80-6.00) mm long, head 1.10 (1.05- 1.12) mm wide, pronotum 1.16 (1.15-1.17) mm wide. Specimens examined. — Holotype 6, In- dia: Meghalaya: Nongpoh, 762 m, 4.xi.l981, C. A. Viraktamath (UAS). Paratypes: 13 d, 4 9 , data as in holotype but 7 d , 2 9 S. Virakta- math Coll. (1 d and 1 9 in BMNH, lARI, USNM, and rest in UAS). Remarks. — This species is related to and resembles S. insignis as they share coloration and similar aedeagi. However, S. cornplexa differs from S . insignis in having the process of the dorsal apodeme bifid ventrally, a longer stem on the connective, and the crimson line of the vertex being forked apically. Sophonia rnodesta (Distant), n. comb. Figs. 186-197 Kana rnodesta Distant 1918b: 36. Lectotype 6, India (BMNH, examined). Pale yellow. Vertex of female creamy white, eyes lemon yellow. Forewing hyaline with brown stripe close to anal margin often reaching claval apex, apex of corium lemon yellow; costal margin with an anterior, long, oblique, median and a slightly oblique, poste- rior, transverse, brown fasciae; second apical cell with black spot, apical margin brownish. Head slightly broader than pronotum. Ver- tex as long as wide between eyes, disc convex, apex subangular, granulose. 208 Great Basin Naturalist Memoirs No. 12 Figs. 175-185. Sophonia complexa, n. sp.: 175, head and thorax; 176, same, profile; 177, face; 178, forewing; 179, pygofer; 180, 181, aedeagus, lateral and caudal views; 182, style; 183, male plate; 184, connective; 185, ovipositor. Male genitalia. — Caudal lobe of pygofer rounded, without process. Anal collar well developed. Plate four times as long as wide. Preapical lobe of style small, rounded, apoph- ysis setose on lateral margin, apical extension beaklike. Stem of connective four times as long as each arm. Aedeagus with preatrium short, pair of ventrocaudally directed pro- cesses, dorsal apodeme robust, each arm rounded; shaft tubular, slender, dorsally 1988 ViRAKTAMATH, WESLEY. REVISION OF INDIAN NiRVANINAE 209 Figs. 186-197. Sophonia modesta (Distant); 186, head and thorax; 187, same, profile; 188, face; 189, ovipositor; 190, pygofer; 191, aedeagus and part of connective, lateral view; 192, 193, aedeagus, dorsal and cephalic views; 194, style; 195, connective; 196, male plate; 197, forewing. 210 Great Basin Naturalist Memoirs No. 12 strongly curved anteriorly, apex ventrally di- rected with pair of elongate, basal processes that are directed dorsoanteriorly then later- ally. Female genitalia, — Hind margin of sev- enth sternum slightly convex. Ovipositor fair- ly exceeding pygofer. Measurements. — Male 4.43 (4.40-4.50) mm long, head 1.08 (1.07-1.10) mm wide, pronotum 1.05 (1.03-1.08) mm wide. Female 5.20 (5.00-5.40) mm long, 1.24 (1.20-1.28) mm wide, pronotum 1.19 (1.15-1.25) mm wide. Specimens examined. — Lectotype S la- beled /South India, Madras, Coonoor, T. V. Campbell/ here designated (BMNH). India: Tamil Nadu: 3 (5 , 1 9 , Kotagiri, 23.x. 1975, C. A. Viraktamath; 1 9, Coonoor, ll.viii.l979, S. Viraktamath; 1 6, Wellington, 12.viii. 1979, S.V.; 5 9, Ootacamund, 2,350 m, 13. viii.1979, S.V. (UAS). Remarks. — Sophonia modesta externally resembles S. bifida in having a wider vertex, but they are only distantly related. It is re- lated to S. insignis and S. complexa and differs from them in having paired preatrial pro- cesses and lacking processes on the dorsal apodeme. In these three species the stem of the connective is more than three times as long as each arm. Sophonia bifida, n. sp. Figs. 198-207 Bright lemon yellow. Two spots on inner claval margin, one at midlength, another at apex, small costal patch about midlength, an outer, oblique and posterior, transverse fascia on costal area fuscous, large, round spot on second apical cell black. Hind tibial apical pecten black. Head slightly narrower than pronotum. Disc of vertex about as long as wide between eyes, broadly rounded with apical area finely rugulose, posterior half polished. Fronto- clypeus rather tumid. Second apical cell of forewing slightly narrowed caudally. Male genitalia. — Caudal margin of pygofer rounded with caudodorsal, short, stout, strongly hooked process and caudoven- tral, elongate, sharply pointed process; macrosetae forming single row. Anal tube large. Preapical lobe of style short, rounded, apophysis slender, rather hooked, apex beak- like. Arms of connective about as long as stem. Aedeagus with elongate preatrium; each arm of dorsal apodeme rounded; shaft tubular with basal pair of elongate processes extending to about 0.75 of length and with subapical pair of denticles on caudal margin. Measurements. — Male 4.50 mm long, head 1.07 mm wide, pronotum 1.10 mm wide. Female 5.30 mm long, head 1.25 mm wide, pronotum 1.25 mm wide. Specimens examined. — Holotype 6, In- dia: West Bengal: Darjeeling, 2,176 m, 11. xi. 1981, C. S. Wesley (UAS). Paratype 9, India: West Bengal: Kalimpong, 1,370 m, 29.x. 1981, S. Viraktamath (UAS). Remarks. — Although the shape of the head is similar to that in S. modesta, S. bifida occu- pies an isolated place in Sophonia. The pecu- liar pygofer process and aedeagus differenti- ate it from all other species. Sophonia complicata, n. sp. Figs. 208-220 Whitish yellow. Area surrounding ocelli, lateral-most area of pronotum lemon yellow. Second apical cell with round, black spot. Costa with outer, oblique fascia, posterior, transverse fascia near apex, and apical margin of forewing fuscous. In a few specimens, me- dian area of clavus pale fuscous, which meets fuscous stripe on claval suture and continues angularly across wing where it is joined by an oblique, fuscous fascia from costa. Hind tibial pecten fuscous. Vertex more conically produced, disc con- vex with median carina in apical half and api- cal area rugulose, rugae crisscrossing. Vertex of female distinctly longer than male. Head wider than pronotum. Male genitalia. — Pygofer with caudodor- sal angle roundly produced, with excavation below which long, slender process arises, di- rected caudally, upcurved at right angles at midlength, apex slightly hooked, bearing spinelike branch at base. Anal tube robust, slightly exceeding pygofer lobe, but not pygofer process. Plate 3.75 times as long as wide. Style with elongate, slender, preapical lobe, apophysis slender avicephaliform. Stem of connective more than twice as long as each arm. Aedeagus complex, two arms of dorsal apodemes well separated, winglike, com- pletely covering entire length of shaft later- ally, each arm with dorsomedian, caudal, and caudoventral, long processes; dorsomedian 1988 ViRAKTAMATH, WESLEY; REVISION OF INDIAN NiRVANINAE 211 204 206 207 Figs. 198-207. Sophonia bifida, n. sp.: 198, head and thorax; 199, same, profile; 200, face; 201, forewing; 202, male pygofer; 203, pygofer process; 204, style; 205, connective; 206, 207, aedeagus, lateral and caudal views. process curves caudally, caudal process di- rected anteriorly and in contact with dorso- median process, caudoventral process crosses over caudad of shaft, thus enclosing it; shaft depressed, strongly curved anteriorly, ex- panded, and hoodlike. Measurements. — Male 4.40 (4.30-4.50) mm long, head 0.92 (0.90-0.95) mm wide, pronotum 0.94 (0.90-0.95) mm wide. Female 4.88 (4.60-5.10) mm long, head 1.01 (0.97- 1.05) mm wide, pronotum 1.02 (1.00-1.05) mm wide. Specimens examined. — Holotype 6, In- dia: Mizoram: Lungleh, 23. xi. 1981, C. S. Wesley (UAS). Paratypes: 15 (5, 16 9, India: Mizoram: Aizawl, 18. xi. 1981, C. S. Wesley; 3 (5, 1 9, Lungleh, 22-24.xi.l981, C. S. Wesley; Meghalaya: 2 9, Cherrapunji, 1,299 m, 3.xi.l981, C. A. Viraktamath; 1 9, Shillong, 1,961 m, 3..xi.l981, S. Viraktamath (1 d and 1 9 paratypes in BMNH, lARI, USNM, ZSI, and rest in UAS). Remarks. — The complex nature of the aedeagus and the slender, long pygofer pro- cess distinguish this species from other spe- cies oi Sophonia. Genus Nirvana Kirkaldy Nirvana Kirkaldy 1900d: 293. Type species: Nirvana pseudommatos Kirkaldy, by original designation and monotypy. Vertex longer than width between eyes, disc depressed apically, with median ridge. Ocelli situated laterally mesad of submarginal carina. Face slightly depressed basally, fron- toclypeus with short, apical, median ridge and lateral, oblique rugae weakly developed. Sec- ond apical cell of forewing narrowed apically. Hindwing with three apical, closed cells. 212 Great Basin Naturalist Memoirs No. 12 Figs. 208-220. Sophonia complicatd. n. sp.: 208, head and thorax; 209, same, profile; 210, face; 211, forewing; 212, ovipositor; 213, male pygofer; 214, pygofer process; 215, style; 216, male plate; 217, connective; 218-220, aedeagus, lateral, caudal, and dorsal views. Hind tibial spinulation R, 20 ± 2. R. 12 ± 1, R, 12 ±2. Male genitalia. — Pygofer lobe caudally, bluntly conical or rounded, with ventral, .short or long proccs.s, caudal area with numer- ous macrosetae. Plate parallel-sided, elon- gate, more than five times as long as wide near middle, with oblique row of macrosetae in apical 0.66 and hairlike setae. Style with well- developed preapical lobe; apophysis short, apical extension more than twice length, beaklike. Aedeagal shaft tubular, with or without processes, dorsal apodcme slender in cephalic view, U-shaped; gonoporc apical. Remarks. — Nirvana and So})honia are closely related and are ditficult to separate by external characters. Nirvana has a simple dorsal apodeme, usually without processes, whereas in Soplionia it is complex with more elaborate processes; the male plate in Nirvana is narrow and elongate compared to Sopho- nia. Specimens oiN. psetido)n))iato.s were not available for the study, and hence the genus is only tentatively defined here. Key to Species oiNirvana I. Vertex with median, white stripe and lateral, orange or lemon yellow, suhmarginal stripe; aedeagal shaft with two slender, lateral, long, ventral processes directed ventrally (Figs. 225, 226) pallida Melichar — Vertex with black line or lines or with orange, suhmarginal stripes traversing on pronotum and 1988 ViRAKTAMATH, WeSLEY: REVISION OF INDIAN NiRVANINAE 213 scutellum and forewing; aedeagus without pro- cesses (male of pseudommatos is not known) ... 2 2(1). Vertex, pronotum, scutellum traversed by pair of orange stripes (Fig. 256); male pygofer cau- dally produced into an acute, dorsally curved process (Fig. 261); anal tube with two stout spines (Fig. 262) pecuhahs. n. sp. — Vertex, pronotum, and scutelhun traversed by black lines or stripes; male pygofer and anal tube unarmed (not known in pseudommatos ) 3 3(2). Vertex, pronotum, scutellum, and anal margin of forewing traversed by broad, black stripe .... 4 — Vertex of head and pronotum traversed by three black stripes; scutellum by a median, black line; apex of vertex with blackish brown spot con- nected by the stripes .... pseudommatos Kirkaldy 4(3). Median stripe with irregular margins (Figs. 233, 234); spot on second apical cell of forewing small (Fig. 243); dorsal apodeme of aedeagus simple (Fig. 240) suturalis Melichar — Median stripe with an even margin (Fig. 244); spot on second apical cell of forewing large (Fig. 248) extending to first apical cell; dorsal apodeme of aedeagus with a basal, short process (Fig. 254) striata, n. sp. Nirvana pallida Melichar Figs. 221-232 Nirvana pallida Melichar 1903b: 166. Syntype cJ, 9, Sri Lanka (not examined). Quercinirvana bengalensis Ahmed & Mahmood 1970: 263. Holotype d, Bangladesh (University of Karachi, not examined). New synonymy. Coloration as described by Melichar (1903b). The median, white stripe of vertex may be obscured in some specimens. Head as wide as or slightly narrower than pronotum. Vertex 1.5 to 2.0 times as long as width between eyes, disc slightly depressed. Clypellus convex. Male genitalia. — Pygofer lobe caudally rounded with marginal macrosetae, ventral process long, dorsally directed with rounded apex. Plate narrowed caudally with bluntly conical apex. Preapical lobe of style rounded, stout. Stem of connective longer than arms. Aedeagus with dorsal apodeme well devel- oped, consisting of two arms and forming a U, shaft tubular, curved slightly with short, spinelike, laterally directed, terminal pro- cesses and long, unpaired, ventrally directed, subapical process. Female genitalia. — Hind margin of sev- enth sternum rather straight. Ovipositor slightly exceeding pygofer. Measurements.— Male 4.80 (4.60-5.00) mm long, head 0.94 (0.90-0.98) mm wide. pronotum 1.02 (0.95-1.08) mm wide. Female 5.65 (5.5-5.80) mm long, h(>ad 1.06 (1.05- 1.07) mm wide, pronotum 0.93 (0.88-0.98) mm wide. Specimens examined. — India: 203 d, 274 9 , collected from the states of Andhra Pradesh, Haryana, Karnataka, Kerala, Maha- rashtra, Meghalaya, Mizoram, Sikkim, Tamil Nadu, and West Bengal from January to De- cember, from sea level (Cochin in Kerala) to 2,250 m (Kodaikanal in Tamil Nadu) above sea level. Nepal: 1 9, Sarankot, 17.xi.l979, I. Dworakowska. Sri Lanka: 1 9 , Peradeniya, 23.viii.1979, I. Dworakowska (UAS). Remarks. — This species is the most com- mon and most frequently encountered Nir- vaninae on the Indian subcontinent. It breeds on newly emerged leaves of Tecoma stans, Duranta sp., Vitis vinifera, beans, pigeon pea, etc. The species is identified based on the description and illustrations provided by Melichar (1903b). No other species of Nirvan- inae has the color and elongate male plate with two short, lateral, and caudal processes on the aedeagal shaft. The synonymy of Q. bengalensis is based on the illustrations of male genitalia of the species provided by Ahmed and Mahmood (1970) and on the spec- imens collected from northeastern India. None of the specimens collected, however, has "a pair of median stripes blackish pale to pale brown, running from anterior tip of crown to posterior margin of pronotum." Based on the illustrations, Pseudonirvana ru- brolimbata Kuoh & Kuoh may also prove to be a synonym of this species. Nirvana suturalis Melichar Figs. 233-243 Nirvana suturalis Melichar 1903b; 166. Holotype 9, Sri Lanka (ZMHU, examined). Yellow. Median, longitudinal, black stripe traversing vertex, pronotum, scutellum and continued along inner margin of clavus of forewing, with uneven, lateral margins and extending angularly on corium beyond clavus (stripe constricted a short distance from apex of vertex in Indian specimens but not in holo- type female) (Figs. 233, 234). First, second, and outer margin of fourth apical cells of forewing dark brown, costal margin with piceous, oblique fascia about 0.25 distance from apex; second apical cell with black spot. Ventral surface of body creamy white. 214 Great Basin Naturalist Memoirs No. 12 225 226 Figs. 221-232. Nirvana pallida Melichar: 221, head and thora.x; 222, same, profile; 223, face; 224, male pygofer; 225, 226, aedeagus, caudal and lateral views; 227, male plate; 228, connective; 229, style; 230, forewing; 231, hindwing; 232, ovipositor. Head narrower than pronotnm. Verte.x about 1.5 times as long as width between eyes, with median, longitudinal groove not reaching apex, rugulose in anterior half, pos- terior half polished. Male genitalia. — Pygofer bluntly conical caudally, with ventral, short notch, macrose- tae confined to apical 0.33 area. Plate more than six times as long as wide. Preapical lobe of style rounded, relatively slender. Stem of connective twice as long as length of each arm. Aedeagus with short preatrium, shaft tubular, slightly curved, each arm of dorsal apodeme wider in middle. Female genitalia. — Hind margin of sev- enth sternum concave with median, short protuberance. Ovipositor exceeding pygofer. Measurements.— Male 4.20 (4.00-4.30) mm long, head 0.85 (0.83-0.88) mm wide, pronotnm 0.88 (0.85-0.90) mm wide. Female 4.74 (4.40-5.00) mm long, head 0.90 (0.80- 0.95) mm wide, pronotum 0.93 (0.88-0.98) mm wide. Specimens e.xamined. — Holotvpe 9 la- beled /Type/ /7042/ /Ceylon, Nietr./ /sii- titralis, det. Melichar/ /Nirvana suturalis Melichar/ (ZMHU). India: 48 cJ, 66 9, col- lected from the states of Karnataka, Kerala, 1988 ViRAKTAMATH, WESLEY; REVISION OF INDIAN NiRVANINAE 215 Figs. 233-243. Nirvana suturalis Melichar: 233, head and thorax of holotype female; 234, female from India; 235, face; 236, ovipositor; 237, male pygofer; 238, connective; 239, 240, aedeagus, lateral and cephalic views; 241, style; 242, male plate; 243, forewing. 216 Great Basin Naturalist Memoirs No. 12 and Tamil Nadu from January to December, at an altitudinal range of 534 m (Jog Falls in Karnataka) to 2,350 m (Ootacamund in Tamil Nadu) above mean sea level. Remarks. — This is the second most com- monly collected Nirvaninae in southern In- dia. Nirvana suturalis and N. striata are very closely related. Both differ from other species of Nirvana and Sophonia in having a broad, black, median stripe running from the apex of the vertex to the corium, but the stripe is not as broad as in species of Chudania. Nirvana suturalis has a serrated lateral margin to the median stripe compared to the smoother lat- eral margin in N. striata. Nirvana striata, n. sp. Figs, 244-255 Coloration as in N. suturalis, median stripe more or less of uniform width and even mar- gined, black spot on second apical cell of forewing, larger than in N. suturalis, extend- ing to first apical cell. Head narrower than pronotum. Vertex slightly longer than width between eyes, anterior half sparsely rugulose. Pronotum slightly more than 1.5 times as wide as long, longer than scutellum. Male genitalia. — Pygofer bluntly conical with macrosetae confined to caudal 0.33 area, ventral margin with short, spinelike process. Plate caudally rounded, about seven times as long as wide. Style with rounded preapical lobe, apical extension of apophysis beaklike and long. Stem of connective twice as long as each arm. Aedeagus with shaft strongly curved (more than in N. suturalis), apex abruptly directed cephalad and with membra- nous area; each arm of dorsal apodeme with basal, short process. Female genitalia. — Hind margin of sev- enth sternum straight. Ovipositor exceeding pygofer. Measurements.— Male 4.90 (4.80-5.00) mm long, head 0.98 (0.95-1.00) mm wide, pronotum 1.06 (1.05-1.08) mm wide. Female 5.46 (5.30-5.60) mm long, head 1.08 (1.05- 1.10) mm wide, pronotum 1.12 (1.08-1.13) mm wide. Spegimens examined. — Holotype 9, In- dia: West Bengal: Kurseong, 1,483 m, 22.x. 1981, S. Viraktamath (UAS). Paratypes: India: Uttar Pradesh: 1 9, Mussoorie, 2,005 m, 27. iv. 1975, C. A. Viraktamath; Himachal Pradesh: 3 9, Simla, 2,133 m, 14.x. 1979, C. A. Viraktamath; West Bengal: 3 d, 3 9, Kurseong, 1,483 m, 22.x. 1981, S. Virakta- math (1 (5 , 1 9 ) and C. A. Viraktamath (2 c? , 2 9); 1 9, 15 km E of Kalimpong, 1,780 m, 27.x. 1981, S. Viraktamath; 1 <5 , 2 9 , 8 km E of Kalimpong, 1,768 m, 29.x. 1981, C. A. Virak- tamath (1 6 and 1 9 paratypes in BMNH, USNM, and rest in UAS). Remarks. — See N. suturalis. Nirvana peculiaris, n. sp. Figs. 256-267 Yellow. Small, black spot at apex of vertex visible both dorsally and ventrally (absent in female from Shillong). Fuscous, median stripe extending posteriorly to 0.25 length (absent in some females). Lateral, orange stripe on vertex mesad of ocelli, continued on pronotum, scutellum and then on to inner claval margin where they converge and con- tinue to apex of clavus; outer margin of stripe on clavus often fuscous. Transverse spot at apex of clavus, small spot at apex of inner, anteapical cell, and anterior, oblique fascia from costa fuscous. Black spot on each cau- dodorsal angle of male pygofer. Head as wide as pronotum or slightly nar- rower. Disc of vertex depressed with rolled- up, sharp margin, basal half with median sul- cus, apical half with a carina, apical half of disc very finely pitted, posterior half with minute rugulae on polished surface. Male genitalia. — Pygofer elongate, nar- rowed caudally to acute, dorsally curved pro- cess. Anal collar elongate, anterior margin armed with two pronglike processes on each side. Setae on pygofer confined to caudodor- sal area where it articulates with anal tube. Male plate elongate, 5.4 times as long as me- dian width, macrosetae and hairlike setae con- fined to apical half Connective robust, stem broad, and arms as long as stem. Apophysis of style fairly long. Aedeagus with dorsal apo- deme poorly developed, shaft tubular, nar- rowed caudally, strongly curved dorsally near apex and with elongate, caudoventrally di- rected process and two lateral, anteriorly directed processes surrounding small gono- pore. Measurements.— Male 5.80 (5.70-5.90) mm long, head 1.23 (1.22-1.25) mm wide, pronotum 1.24 (1.22-1.25) mm wide. Female 7.40 (7.00-7.80) mm long, head 1.50 (1.45- 1988 ViRAKTAMATH, WeSLEY: REVISION OF INDIAN NiRVANINAE 217 Figs. 244-255. Nirvana striata, n. sp.: 244, head and thorax; 245, face; 246, profile; 247, ovipositor; 248, forewing; 249, hindwing; 250, male pygofer; 251, male plate; 252, style; 253, 254, aedeagus, lateral and cephalic views; 255, connective. 1.55) mm wide, pronotum 1.54 (1.50-1.60) mm wide. Specimens examined. — Holotype 6, In- dia: Mizoram: Aizawl, 18. xi. 1981, C. S. Wesley (UAS). Paratypes: India: Mizoram: 2 S,4 9, Aizawl, 18.xi.l981, C. S. Wesley; 1 d, 2 $, Lungleh, 23.xi.1981, C. S. Wesley; Meghalaya: 1 9, Shillong, 1,961 m, 3.xi.l981, S. Viraktamath (1 9 and 1 9 paratypes in BMNH; 1 9 paratype in lARI, USNM, ZSI, and rest in UAS). Remarks. — This species is unique among Nirvanini in having an armed tenth segment. In coloration it resembles species of Kana, especially K. fasciata, but it differs from those species in having three apical cells in the hind- wing and unique male genitalia. Ophiuchus Distant Ophiuchus Distant 1918b: 33. Type species: Ophiuchus princeps Distant, by original designation and monotypy. Pale yellow with bright, contrasting col- oration of black and red or pink. Head about as wide as pronotum, spatulate, disc of vertex depressed in middle, margined by carina, lat- eral margin of vertex straight in front of eyes to ocelli where it is widened and then slightly narrowed to apex. Face with five lateral ridges, concave medially, with short, median carina near apex. Pronotum twice as wide as median length, disc minutely, transversely rugose. Hindwing with three closed apical cells. R2 of hind tibia with 12 spines. 218 Great Basin Naturalist Memoirs No. 12 Figs. 256-267. Nirvana peculiaris, n. sp.: 256, head and thorax; 257, same, profile; 258, face; 259, female seventh sternum; 260, forewing; 261, male pygofer; 262, male tenth segment; 263, style; 264, male plate; 265, connective; 266, 267, aedeagus, lateral and cephalic views. 1988 ViRAKTAMATH, WeSLEY: REVISION OF INDIAN NiRVANINAE 219 Male pygofer elongate, heavily macrose- tose and with candal, hooklike process. Valve semicircular. Plate rounded apically, macro- setae scattered and confined to caudal 0.40. Style appearing like partially closed fist. Aedeagus large, poorly sclerotized, with ven- tral, platelike process, shaft short, cylindrical with large gonopore. Remarks. — This genus appears related to Sophonia and Nirvana with which it shares general facies of the head. The peculiarly shaped style and scattered macosetae on the male plate distinguish this genus. Ophiuchus princeps Distant Figs. 268-276 Ophiuchus princeps Distant 1918b: 34. Lectotype 6 , India (BMNH, examined). Pale yellow. Vertex laterally margined by black line that does not reach apex, black, median, longitudinal line does not reach apex. Eyes centrally fuscous surrounded by red. Ocelli red. Face ochraceous with lateral in- foldings black. Pronotum sanguineous with submarginal, anterior, transverse, whitish spot. Scutellum anteriorly marked with fus- cous band with backward, lateral projections. Base of forewing fuscous, apical 0.33 with an irregular, pink spot margined by fuscous, rest yellowish. Head shorter than combined length of pro- notum and scutellum (36:44). Pronotum shorter than scutellum (21:23). Face longer than wide. Male genitalia. — Pygofer elongate, lobe covered with numerous stout setae and termi- nated by curved, hooklike process. Valve semicircular. Plate elongate, with angulate projection on lateral margin near apex. Style flat, preapical lobe with few setae, apophysis with two angular projections and apically pointed. Aedeagus with very short, dorsal apodeme, elongate, platelike, ventral pro- cess, armed with median ridge and two cau- dal, spinelike processes on its laterocaudal angles, shaft short, cylindrical, with large, round gonopore and with pair of lateral pro- cesses about its midlength. Measurements. — Male 4.60 mm long, head 1.07 mm wide, pronotum 1.05 mm wide. Specimen examined. — Lectotype <5 la- beled /Travancore/ /Distant Coll. 1911-383/ /Ophiuchus princeps Dist., type/ here desig- nated (BMNH). Remarks. — This species has a very distinc- tive coloration not possessed by any other species of Nirvaninae known from the Indian subcontinent nor by any other species of OpJiiuchus known from the Oriental region and Australia. The male genitalia of Ophi- uchus species except O. distanti Evans (1973: Fig. 5B) are not sufficiently known to suggest any relationships among them. Judging from the illustrations of O. distanti, it does not appear to be related to O. princeps. Tribe Occinirvanini Body elongate, slightly depressed. Head produced anteriorly, spatulate. Face horizon- tal, frontoclypeus and clypellus either flat {Occinirvana Evans) or convex; facial sulci deep, prominent; face longer than wide, genae laterally emarginate below eyes, lora fairly prominent. Antennae arising close to anterior margin of head. Vertex elongate, with median sulcus and transverse ridge or carina across ocelli, which are situated on disc of vertex closer to anterior than to posterior margin. Lateral margin of pronotum rounded, without carina, rather parallel-sided. Forewing with com- plete and prominent venation; at least three cross-veins reaching vein R from costal mar- gin; anteapical cells three, outer anteapical cell half as long as median, inner anteapical cell open basally; apical cells four; appendix distinct, often extending around apical margin as in Omaranus Distant. Hindwing with three apical cells. Hind femoral spinulation 2+1+0. Hind basitarsus with six platellae (in Omar- anus). Male genitalia is of deltocephaline type. Pygofer laterally not fused with valve, lobe covered by stout setae. Valve triangular. Plates triangular with stout macrosetae. Style deltocephahne type with well-developed, pre- apical lobe and fingerlike apophysis. Connec- tive Y-shaped with diverging arms. Aedeagus simple, with large gonopore, articulated with connective. Remarks. — The tribe, judged from the structural features, is not closely related to either Nirvanini or Balbillini. It has several deltocephaline features, especially the male genitalia. 220 Great Basin Naturalist Memoirs No. 12 Figs. 268-276. Ophiuchus princeps Distant: 268, habitus; 269, profile; 270, male pygofer; 271, male valve; 272, male plate; 273, style; 274, aedeagus, lateral view; 275, aedeagus and connective, ventral view; 276, aedeagus, cephalic 1988 ViRAKTAMATH, WESLEY: REVISION OF INDIAN NiRVANINAE 221 Genus Omaranus Distant Omaranus Distant 1918b: 5. Type species: Omaranus typictis Distant, by original designation and inoiio- typy. Didius Distant 1918b: 36. Type species: Didius sexualis Distant, by original designation and nionotypy. Sexually dimorphic. Head narrower than pronotum in male, wider than pronotum in female. Vertex with median carina in apical half, sulcus in basal half. Face longer than broad, frontoclypeus tumid. Clypellus longer than broad; frontoclypeus, lora, genae, and clypellus with scattered, short setae. Prono- tum gradually widened posteriorly in male, parallel-sided and with faint, median ridge near posterior half in female; posterior margin strongly inwardly concave; disc llattish in fe- male, slightly raised posteromedially in male, lateral margins not carinate. Male forewing exceeds abdomen, appendix runs around api- cal margin, with four apical and three anteapi- cal cells; hindwing well developed with three apical cells. Female forewings cover only three basal, visible, abdominal terga, while hindwings cover two basal, visible terga; ap- pendix wanting. Hind tibial spinulation Rj 16 ± 2, R, 10 ± 1, R3 3. Hind basitarsus shorter than combined length of second and third tarsi and terminated by six platellae. Male pygofer without processes. Tenth seg- ment stout, rather short. Ovipositor not ex- ceeding pygofer. Remarks. — This genus and the Austrahan Occinirvana are closely related. The shape of the head differentiates the two genera. Omaranus typicus Distant Figs. 277-289 Omaranus typicus Distant 1918b; 6. Lectotype S , India (BMNH, examined). Didius sexualis Distant 1918b: 36. Lectotype 9, India (BMNH, examined). Male. — Ochraceous (probably green in life). Anterior margin of vertex narrowly black margined, with median, black spot, two smaller, somewhat elongate spots near poste- rior margin, ocelli black. Dorsal-most area of frontoclypeus and face ventrad of antennal bases, blackish brown, marginal band dorsad of antennal base and longitudinal stripe on proepimeron blackish brown. Apical 0.25 of forewing and stripe along inner margin fus- cous. Abdomen with four black stripes; sterna marked with brown stripes. Apical half of hind femora ventrally streaked with brown. A ven- tral spot near apex of fore femora black. Genitalia. — Pygofer lobe caudoventrally produced, caudodorsal area beset with stout, short setae. Plate caudally rounded with scat- tered, stout setae. Valve straplike with caudal margin roundly and medially produced. Apo- physis of style curved laterally with series of teeth on lateral margin. Stem of connective 0. 75 as long as each arm . Aedeagus with short, dorsal apodeme, shaft stout at base, narrowed caudally in lateral aspect with a large gono- pore on dorsal aspect. Female. — Ochraceous. Head with median spot on anterior margin, ocelli, and two spots near base of vertex black; median stripe, a transverse band across ocelli, hind margin, except medially, and two spots in front of eyes reddish. Dorsal margin of face black from which two apically divergent stripes traverse face. Proepisternum traversed by longitudi- nal, piceous stripe, much longer on pro- episternum; mesopleura and mesocoxae with piceous spot. Pronotum with scattered, dark brown spots, with obscure stripes, two lateral and two median reddish. Scutellum except for two lateral, basal angles in basal half black, apical half blackish brown. Forewing veins marked with red. Fore tibiae and tarsi red- dish; fore- and mesothoracic femora black, mesocoxae with piceous spot. Abdominal col- oration as in male. Genitalia. — Hind margin of seventh ster- num with median, U-shaped excavation and translucent in middle. Anal segment con- cealed by pygofer. Measurements. — Male 4.30 mm long, head 1.00 mm wide, pronotum 1.10 mm wide. Female 7.90 mm long, head 1.00 mm wide, pronotum 1. 10 mm wide. Specimens examined. — Lectotype 6 la- beled / Calcutta, 28.viii.07, Mus. Coll./ / Omaranus typicus Dist., type/ here desig- nated. Paralectotype S labeled / Calcutta, 23.viii.07/ /Omaranus typicus Dist., type/ here designated. Paralectotype has the abdo- men and left fore and hindwings missing. Lec- totype 9 labeled / Calcutta/ /Distant Coll. 1911-384/ /Didius sexualis Dist. type/ here designated (BMNH). India: Punjab: 1 c?, 1 9 nymph, Ludhiana, 13. ix. 1966, A. S. Sohi (PAU). India: Madhya Pradesh: 3 6 , 18 9 , Nebudda Survey, Sta. No. 31, 37, 58, 21.ii-9.iii.l927, H.S. Pruthi (ZSI). 222 Great Basin Naturalist Memoirs No. 12 Figs. 277-289. Omamnus ti/picus Distant: 277, head and thorax of lectotype male; 278, same of lectotype female, Didius sexiialis Distant; 279, ovipositor; 280, 281, fore and hindwings of male; 282, male pygofer; 283, male plate; 284, male valve; 285, connective; 286, style; 287, 288, aedeagus, lateral and cephalic views. 1988 ViRAKTAMATH, WeSLEY: REVISION OF INDIAN NiRVANINAE 223 Acknowledgments The authors express thanks to Dr. W. J. Knight, Mr. M. D. Webb (BMNH), Dr. U. Gollner-Scheiding (ZMHU), Dr. H. Synave (IRSNB), Dr. A. S. Sohi (PAU), and Dr. Baldev Sharma (JU) for lending specimens for study, including primary types under their care. Also, the director of the Zoological Sur- vey of India allowed the senior author to ex- amine Dr. H. S. Pruthi's type material during his visit to ZSI. The authors are also grateful to Dr. P. H. Freytag, University of Kentucky, Lexington, for critically reviewing the manu- script. Literature Cited The references cited in the text prior to 1956 are found in Metcalf (1964), wherein Pruthi s works have been cited under the name Singh-Pruthi. Ahmed, M., and S. H Mahmood 1970. A new genus and two new species of Nirvaninae (CicadeUidae: Ho- moptera) from Pakistan. Pakistan Journal of Scien- tific and Industrial Research 12: 260-263. Ahmed, M , and K F Mallik 1972. On the true identifi- cation of genus Chudania Distant collected on fig {Ficiis carica ) in forests of Azad Kashmir and Kap- tai (Pakistan). Pakistan Journal of Forestry 22; 103-108. Davis, R. B. 1975. Classification of selected higher cate- gories of Auchenorrhynchous homoptera (Cica- deUidae and Actalionidae). Technical Bulletin No. 1494, Agricultural Research Service, United States Department of Agriculture. .52 pp. EvAN.s. J W 1966. The lealhoppers and froghoppers of Australia and New Zealand (Homoptera: Cicadel- loidea and Cercopoidea). The Australian Museum Memoir 12: 1-347. 1973. Some new genera and species of Cicadel- loidea from Australia and New Guinea (Ho- moptera). Pacific Insects 15: 185-197. KUOH. C L , AND J L KUOH 1983. New species of Pseudonirvana (Homoptera: Nirvanidae). Acta Entomologica Sinica 26: 316-.325 [In Chinese]. Kramer, J P 1964. A review of the Neotropical Nirvan- inae (Homoptera: CicadeUidae). Entomological News 75: 113-127. LiNNAVUORl, R 1979. Revision ofthe African CicadeUidae (Homoptera Auchenorrhyncha), Part II. Revue de Zoologie Africaine 93: 929-1010. Metcalf, Z P 1963. General Catalogue of the Ho- moptera, Fascicle VI. Cicadelloidea, Part 7 Nir- vanidae. Agricultural Research Service, United States Department of Agriculture. 35 pp. 1964. General Catalogue of Homoptera, Fa.scicle VI. Cicadelloidea, Bibliography of the Cicadel- loidea (Homoptera: Auchenorrhyncha). Agricul- tural Research Service, United States Depart- ment of Agriculture., 349 pp. Ramakrishna, B V. 1980. Leaihopper fauna of pulse crops and biology of Empoasca (Distantasca) ter- minalis Distant (Homoptera: CicadeUidae). Un- published thesis. University of Agricultural Sci- ences, Bangalore. 197 pp. Theron, J G 1970. A new species of Narecho (Hemiptera, Cicadelloidea: Nirvanidae) from Africa. Novos Taxa Entomologicos No. 83, 1-7. GENUS FLEXAMIA: NEW SPECIES, PHYLOGENY, AND ECOLOGY Robert F. Whitcomb' and Andrew L. Hicks' Abstract — Eight new species of Flexamia DeLong are described, and two Mexican species previously syn- onymized are reinstated. Species are assigned to 13 species groups; keys are presented for the groups and for the 44 recognized species. Host data, in many cases with ohgophagy coefficients, are presented for 37 of the species. Many species speciaUze on native, dominant, perennial, choridoid or panicoid grasses; some are monophagous. Seven sister species pairs specialize on the same (or a closely related) grass species; in addition, four closely related species appear to be restricted to Bouteloua curtipendtila. Few if any specialists colonize their host throughout its entire range. Ecological factors such as phenology and/or host patchiness strongly influence geographic distribution. An intuitive phylogeny is proposed and is used as a basis for constructing 18 character transformations. The genus Flexamia probably originated in Mexico by division of an ancestral lineage from which the modern genus Spartopyge also diverged. The first specimen of the genus Flexa7nia DeLong was apparently collected at Jack- sonville, Florida, in the height of the Lin- naean explosion; it was deposited in the British Museum (Natural History) and named Jassus productus by Walker (1851). In their studies of Iowa prairies in the latter part of the nineteenth century, Osborn and Ball encoun- tered other species, which they referred to the genus Deltocephalus. DeLong (1926) erected a subgenus Flexamia DeLong within Deltocephalus Burmeister. DeLong and Sleesman (1929) recognized that Delto- cephalus was an unworkably large assemblage of distantly related species and divided the genus into several genera (e.g., Laevi- cephalus DeLong & Sleesman, Pohjamia De- Long & Sleesman, and Flexamia). The genus concept for Flexamia was refined by Oman (1949) in his revision of North American ci- cadellid genera, but the modern generic con- cept was established by Young and Beirne (1958). These authors segregated Spartopyge Young & Beirne, synonymized Secopennis Oman, described genitalic characters of males and females (the first use of genitalic charac- ters for Flexamia females), and presented a species key. The 30 species they described included all of the widely distributed and commonly collected representatives of the genus, and their key proved to be accurate and easy to use. Young and Beirne also de- scribed possible relationships among species and, without specifying an explicit phylogeny, proposed broad outlines of hypothetical evo- lutionary events in the genus. The genus was incorrectly synonymized with Acurhinus Os- born by Linnavuori (1959). Comparison of the specimens (all females) of Acurhinus ( =Dorydium) maculatum Osborn in the Ohio State University collection led Linnavuori and DeLong (1978) to synonymize Acurhinus with the African stirelline genus Hododoecus Jacoli. The genus Flexamia as defined herein is an entirely North American taxon related to Spartopyge, to Alapus DeLong & Sleesman, and, perhaps distantly, to Aflexia Oman. The closest Old World relative may be Enantio- cephalus Haupt. Although the taxonomic status of Flexamia has been generally adequate since 1958, host data have been difficult to obtain. Earlier col- lectors were not knowledgeable agrostologists and often reported only that deltocephaline species, including Flexamia, resided on "grasses. " Astute collectors such as Osborn and Ball recognized the importance of biologi- cal data, and their notes contain some host records. In 1967, H. H. Ross encouraged the senior author to ascertain the hosts of Flex- amia species, in large part to test his intuitive belief that host relationships, especially host transfers, could be invoked to explain specia- tion of phytophagous insects. Accordingly, we sought to determine hosts of the species de- scribed by Young and Beirne. Much to our 'insect Pathology' Laboratory, AEriciiltural Research Service, Beltsville, Maryland 20705. 224 1988 Whitcomb, Hicks: Flexamia Revision 225 Fig. 1. Flexamia serrata. surprise, in the course of this search, unde- scribed species emerged, particularly when we examined dominant, perennial Muhlen- bergia or Bouteloua species (Whitcomb et al. 1986, 1987). Under these circumstances, it seems appropriate to review the species con- cept and to emend the generic concept, which, although basically intact in its essential aspects, must accommodate the new species. Genus Flexamia DeLong Deltocephalus subgenus Flexamia DeLong 1926:20, 22. Type species: Deltocephalus reflextis Osborn & Ball 1897, by original designation. Flexatnius, DeLong and Sleesman 1929:82. Flexamius subgenus Secopennis DeLong & Sleesman 1929:85. Type species: Deltocephalus slossoni Ball. Acurhinus Oshorn, Linnavuori 1959:89. Flexamia DeLong, Young and Beirne 1958; Linnavuori and DeLong 1978:208. Small deltocephaline leafhoppers. Length of male 2.4-5.0 mm, of female 2.6-5.0 mm. Head produced, median length of crown usu- ally greater than interocular width; clypellus broader at base than at apex; forewing with reflexed veinlets to costal margin in vicinity of small, outer anteapical cell. Dried specimens yellowish brown or gray, with ferruginous to black markings; live in- sects in some cases with green pigmentation changing to orange or stramineous when dried. Apical area of crown partly or com- pletely surrounded by darker markings, in latter case with clearly defined, apical areolet. Crown often with pair of oblique, dark mark- ings at base; pair of transverse, dark markings at about midlength; pair of darker markings parallel to lateral margins near apex. Face color varying from pale to black, often useful in species discrimination. Pronotum of some species with conspicuous, longitudinal stripes, but many species with faint, wider stripes. Veins of forewings usually dark-mar- gined; distinct, dark-margined areole fre- quently near center of corium. Male, — Plates usually shorter than pygo- fer, frequently fused along mesal margins; pygofer usually with pronounced posterior lobe in lateral aspect; style with longitudinally grooved, preapical lobe, which is usually not pronounced; connective linear, arms contigu- ous basally, sometimes fused to aedeagus. Connective with upper portion less heavily sclerotized than lower portion, often thinner and extended dorsad in thin keel that varies in width interspecifically; dorsal aedeagal apo- deme with pair of conspicuous appendages directed caudoventrad or caudad. Aedeagus interspecifically variable, of great value in species discrimination. Female. — Sternum VII nearly always longer than VI, often 2X or more; hind margin with 2-4 teeth and usually small, median notch. Pygofer with well-dispersed setae; ovipositor barely reaching pygofer apex. Flexamia Species Groups In the following discussion, we describe 13 species groups and provide keys to species. We describe 8 new species and reinstate 2 species previously synonymized. We also re- view important characteristics of previously described species, emphasizing diagnostic morphology, ecology, and biogeography. Our brief descriptions of previously described spe- cies complement the descriptions of Young and Beirne (1958). Key to Flexamia Species Groups 1 . Dorsum with pair of stripes extending from hind portion of disc of crown across pronotum and scutellum (Figs. 2A-D) 2 — Dorsum without such stripes (Figs. 2E,F) 4 2(1). Individuals large, at least 4.5 mm IV. grammica group — Individuals smaller 3 3(2). Forewing broadly truncate (Fig. 2D); Arizona VIII. ritana group — Forewing obliquely truncate (Fig. 2B) or not truncate (Fig. 2A) (prairie); Florida VII. alhida group 4(1). Face entirely black, genae at most with oblique, pale streak; crown without midlength transverse lines 5 226 Great Basin Naturalist Memoirs No. 12 Fig. 2. Habitus ofFlexamia species: A, alhida; B, shssonae; C, grammica; D, ritana; E, pijrops: F, areolata. 1988 Whitcomb, Hicks: Flexamia Revision 227 Fig. 3. Faces of Flexamia species: A, serrata- B, stylata; C, prairiana- D, prairiana; E, picta; F, decora; G,flexulosa; H, sandersi. — Face black basally, pale apically, or brownish in part; at least genae pale, crown usually with midlength transverse lines 6 5(4). Forewing with conspicuous black spot (Fig. 2F) in corium; crown with median length more than one-half greater than interocular width XII. areolata group — Forewing without conspicuous black spot in corium; crown with median length less than one-half greater than interocular width XI. impiitans group 6(4). Face pale with several narrow, parallel, fus- cous, interocular lines interrupted at middle (Fig. 3A); northwestern (Fig. 30) XI. serrata group — Face with (Figs. 3B,D-G) or without (Figs. 3C,H) interocular lines but not as above 7 7(6). Either with crown more than twice interocu- lar width (Fig. 2E) {pyrops Crumb), or with conspicuous, dark, apical wing blotch (Fig. 4C) and sordid yellow face with black interoc- ular band {picta [Osborn]); viewed laterally, male pygofer with conspicuous, acutely angu- late, ventral lobe (Figs. 5A,B); aedeagus (Fig. 6D) with pair of anteapical processes and api- cal, flangelike processes VI. picta group Habitus not as above; male pygofer with pos- terior lobe rounded ventrally; aedeagus not as above 8 8(7). Males 9 — Females 14 9(8). Aedeagus and connective fused (Figs. 6F,G) XIII. prairiana group — Aedeagus and connective distinctlv articu- lated (Figs. 6A-C, E) ' 10 10(9). Aedeagus without apical processes (Fig. 6C) 11 — Aedeagus with (at least minute) apical processes 12 11(10). Male plates broad, not tapered (Figs. IIA-I) I. pectinata group — Male plates more elongate and tapered (Figs. 9C,F) III. zacate group 12(10). Aedeagus (Figs. 7F,G) with 2 pairs of pro- cesses V. curvata group — Aedeagus not so 13 13(12). Aedeagus with 2 processes on shaft (Fig. 7H) II. abbreviata group — Aedeagus with 3 processes on shaft (Figs. 6E, 31A,D-M) X. flextdosa group 14(8). Face white, ivory, or very pale yellow, sharply contrasting with black interocular line (Figs. 3B,F,G) X. flexulosa group (in part) — Face with lower part not white or pale (Fig. 3y), or if pale, not contrasting sharply with interocular band, or interocular band brown, not black (Fig. 3D) 15 15(14). Body color stramineous 16 — Body color not stramineous 17 16(15). Hind margin of sternum VII with medial pro- jection (Fig. lOO) X. flexulosa group (inflata subgroup) — Hind margin of sternum VII without projec- tion 20 228 Great Basin Naturalist Memoirs No. 12 Fig. 4. Wings of Flexamia species: A, inflata; B, cur- vata; C, picta. 17(15). Sternum VII (Fig. lOE) with median, trape- zoidal projection, with a medial notch, usu- ally with smaller, shallower notches on either side II. abbreviata group — Sternum VII not as above 18 18(17). Sternum VII produced medially in a distinct lobe or projection (Figs. 10S,T) XIII. prairiana group — Sternum VII not produced medially in dis- tinct lobe, or if produced, with deep, median incision 19 19(18). Hind margin of sternum VII incised to form pattern of four teeth; hind margin usually heavily infuscated on hind margin in vicinity of teeth (Figs. 14A-H) I. pectinata group — Hind margin of sternum VII not so 20 20(19). Hind margin of sternum VII slightly concave on each side of median excision; infuscated notches usually contrasting sharply with stramineous color of sternum (Figs. 10J,K) V. curvata group — Hind margin of sternum VII not so (Figs. 101, L) III. zacate group I. The pectinata Group The greatest proposed changes in Flexamia species concepts are in the pectinata species group to which we add five new species and reinstate two others. All species of the group have aedeagal shafts without apical processes. Although aedeagal characters are not as useful for species discrimination in this group as in other groups, the male plates and female ster- num VII provide useful apomorphies that de- fine the group. The plates (except in collo- rurn) are broad and basally parallel-sided. The hind margins of the female sternum VII of all species possess four medial teeth with sur- rounding infuscation (but note the extensive modification in mescalero [Fig. 21H]). The size and shape of the teeth vary interspecifi- cally. The species are largely southwestern and/or Mexican, and many appear to have small ranges. The nominate species F. pecti- nata, however, has a wide geographic range. As defined by Young and Beirne (1958), this species ranges from western Mexico {zamora DeLong & Hershberger) through the eastern highlands (Valles and Monterrey [7ninima DeLong & Hershberger]), the Davis Moun- tains of west Texas, Otero and Guadalupe counties of New Mexico north to North Da- kota, and as far east as Akron, Ohio, in the prairie peninsula. However, our studies indi- cate that zamora and minima should be rein- stated. Recognition herein of the new species batidarita, gila, coUorum, jacala, and mesca- lero makes it clear that this group is diverse and has speciated frequently. The host of pectinata is side-oats grama (Bonteloua cur- tipendula), a widespread prairie species. In addition to pectinata, there are three south- western side-oats grama specialists of the pectinata group. Flexamia doeringae and F. gila reside in the southeastern Arizona moun- tains and Mogollon Rim region and extended conifer-oak savanna of Mexico, respectively. Flexamia bandarita, as far as known, is con- fined to the Chisos Mountains and Marathon Basin of Trans-Pecos Texas. We suspect that side-oats grama, which also has a wide range in Mexico, will prove to be the host of some Mexican species of the pectinata group. The most distinctive species of the group {mesca- lero), however, appears to specialize on Muh- lenhergia pauciflora. Several factors may account for speciation in the pectinata group. Side-oats grama is it- self a complex of ecotypes that vary greatly in structure and seasonality (Gould 1979). This 1988 Whitcomb, Hicks: Flexamia Revision 229 Fig. 5. Pygofers oi Flexamia species: A, picta; B, pyrops; C, stylata; D, producta. Redrawn from Young and Beirne (1958). Fig. 6. Aedeagus and connective of Flexamia species: A, curvata; B, abbreviata; C, canyonensis; D, picta; E, flexulosa; F, prairiana; G, graminea. Redrawn from Young and Beirne (1958). Aspects; A-F, caudoventral; G, lateral. chloridoid grass crosses many biome lines and is a dominant in mesic Sonoran and Chi- huahuan grasslands and in montane systems within these semiarid grasslands. It is com- mon through much of the prairie and occurs eastward on nonforested slopes in the Ap- palachian mountains. Like other major domi- nants of North American grasslands, there- fore, side-oats grama has a much wider geographic range than any of the cicadellid species that exploit it. The Chihuahuan desert system presents a major barrier to geographical continuity of prairie grasses, and side-oats grama is no ex- ception. Many prairie cicadellid taxa occur west of this desert system, but, depending upon the case, the taxonomic results vary. For example, populations related to the buffalo- grass specialist Athysanella (Gladionura) emarginata occur in disjunct stands of Buchloe in New Mexico's Hidalgo County. This population, until very recently, had been regarded as a subspecies (sobrina) but will be elevated to a species in a forthcoming revision oi Gladionura (H. D. Blocker, personal com- munication). On the other hand, forms of A. (Gladionura) argenteola (a Botiteloua gracilis specialist) in southeastern Arizona are not dis- tinguishable from short-grass prairie forms. The connecting link between the relatively mesic, temperate grasslands of the northern Mexican highlands and the prairie is tenuous at best, consisting of a small set of "stepping stone" montane islands within the Chi- huahuan desert of Coahuila and Chihuahua, or of the relatively narrow grassland periph- ery surrounding the desert on the east and west and meeting at the south in the state of San Luis Potosi. Also, the mesquite-acacia savanna and Trans-Pecos shrub savanna of Texas are further barriers to geographic conti- nuity of B. curtipendula. During full glacia- tions, these arid regions had climates that were much more moist than they are today (Van Devender and Burgess 1985). One of the new species, collorum, from the Edwards Plateau of central Texas, is appar- ently a specialist on Nealley grama, Bouteloua 230 Great Basin Naturalist Memoirs No. 12 LT CA CV LT Fig. 7. Aedeagal apices oi Flexamia species; A, scrrata- B, alhida; C, slossonac- D, litana- E, ^rammica; F, curvata; G, sitrcuhi; H, abbreviatii; I, zflcafc, J, canijonensis; K, /;icf«, L, ptjrop.s. Aspects: AD, anterodorsal; CA, caudal; CV, caudoventral; DS, dorsal; LT, lateral. Redrawn from Young and Beirne (1958). uniflora. Flexamia coUonim may be related to another new species, jacala, whose type lo- cahty in Mexico is close to the intersection of eastern and western montane Mexican grass- lands. The latter species is represented by a single male specimen from the USNM, so nothing is known about its biology. Finally, another new species, niescalero, discovered in a single location in southeastern New Mexico, presumably arose by host trans- fer to New Mexican muhly, Muhlenbergia paucijlora. Description of the pectinata Group Medium-sized to small deltocephaline leaf- hoppers. Length of 6 2.7-3.8 mm, of 9 2.8-4.2 mm. Species with general facies of genus; brown or in dark specimens, almost black. Crown variably produced. \6] Plates (Figs. IIA-I) elongate, broad, parallel-sided basally (except coUorum). Pygofer with dis- tinct, posterior lobe, strongly constricted in lateral aspect, both dorsally and ventrally. Aedeagus and connective distinctly articu- lated. Aedeagus (Figs. 12A-I) with no apical processes, or at most slightly capitate (mescalero), often appearing flared in dorsal aspect. [Note: Caution must be exercised when using the aedeagal apex as a diagnostic character, since this structure is membranous and is subject to tearing and breaking during copulation or preparation of the specimen.] Styles variable interspecifically (Figs. 13A-I). Gonopore subapical on caudoventral surface. [ 9 ] Sternum VII (Figs. 14A-H) with four me- dial teeth on hind margin; teeth varying inter- specifically in length and shape, surrounded by infuscated region. 1988 Whitcomb, Hicks; Flexamia Revision 231 Fig. 8. Male plates and pygolers oi Flexamia species: A, alhida: B, slossonae: C, serrata, D,ritana; E, abbreviata; F, grammica. Fig. 9. Male plates and pygofers oi Flexamia species: A, picta; B, pyrops; C, canyunensis; D, curvata; E, surcula; F, zacate. 232 Great Basin Naturalist Memoirs No. 12 Fig. 10. Female sternum VII of Flexarnia species: A, albida^ B. slossonac, C, ^errata^D, t"7;,L" o'llXa- P grammica; G, picta; H, pyrops; I, canyonensis;], surcula; K, curvata; L, zacate; M,flexulosa; N, decora, O, mjlata, F, imputans; Q, areolata; R, sfyiata; S, prainana; T, sandersi; U, fitammea. 1988 Whitcomb, Hicks: Flexamia Revision 233 Fig. 11. Male plates and pygofers oiFlexamia pectinata group, ventral aspect: A, mminui; B D, handarita- E, gila; F, doeringae; G, collorurn; HJacala; I, mescalero. zamora; C, pectinata; 234 Great Basin Naturalist Memoirs No. 12 A B u u t . - ^ Fig. 12. Aedeagal apices of Flexamia pectinata group, caudoventral aspect: A, minmui; B, zainora; C, pectinata; D, bandarita; E, gila; F, doeringae; G, coUorum; H,jacala; I, mescalero. Key to Males of the pectinata Group 1. Plates long and parallel-sided at bases 2 — Plates short, not parallel-sided collorum Whitcomb & Hicks 2(1). Plates subrectangular (Figs. IIC.F) 3 — Plates with apices rounded 4 3(2). Aedeagus in ventral aspect (Fig. 12F) with pro- tuberance close to tip; face pale; Arizona doeringae Beamer & Tuthill — Aedeagus in ventral aspect with protuberance considerably removed from tip (Fig. 12C); face brown; prairie pectinata (Osborn & Ball) 4(2). Plates (Fig. 11 A) with apices acute; Mexico . . . minima DeLong & Hershberger — Plates with apices not acute 5 5(4). Plates almost as long as pygofer (Fig. HE) .... gila, n. sp. — Plates no more than 3/4 length of pygofer 6 6(5). Aedeagus expanded apically in ventral aspect, appearing bluntly saggitate (Figs. 121, 22) .... mescalero, n. sp. — Aedeagus not expanded apically 7 7(6). Style apex (Fig. 13H) avicephaliform in lateral aspect jacala, n. sp. — Style apex not avicephaliform 8 8(7). Style apex (Fig. 13D) ventrally directed, chelate bandarita, n. sp. — Style apex (Fig. 13B) not chelate zamora DeLong & Hershberger Key to Females of the pectinata Group 1. Sternum VII with 4 teeth of approximately equal length and 3 relatively shallow incisions 2 — Sternum VII with deep incisions and/or teeth of different length 4 2(1). Sternum VII with infuscations on either side of middle incision (Fig. 14G); Texas, host Butite- loua uniflora coUorum, n. sp. — Sternum VII not as above 3 3(2). Sternum VII with heavy infuscation around teeth on hind margin (Fig. 14C); USA, prairie pectinata (Osborn & Ball) — Sternum VII with light infuscation on hind mar- gin (Fig. 14A); Mexico minima DeLong & Hershberger 4(1). Sternum VII with outer teeth longer than inner teeth 5 — Sternum VII with inner teeth longer than outer teeth 6 5(4). Sternum VII with outer teeth sharply pointed, median incision shallow (Fig. 14B); Mexico . . . zamora DeLong & Hershberger — Sternum VII with outer teeth blunt, median incision deep (Fig. 14H) mescalero, n. sp. 6(4). Inner teeth of sternum VII together forming blunt, median projection (Fig. 14F) doeringae Beamer &"Tuthill — Inner teeth of sternum VII separated, acute at tip, separated by tapered incision 7 7(6). Sternum VII with three incisions of equal depth (Fig. 14E) gila, n. sp. — Sternum VII with median incision deeper than lateral incisions (Fig. 14D) bandarita, n. sp. 1. Flexamia minima DeLong & Hershberger, n. stat. Flexamia minima DeLong & Hershberger 1947: 138. Flexamia pectinata. Young & Beirne 1958: 46. (In part.) Length of c? 2.9 (2.8-2.9) mm, 9 3.0 (2.7-3.2) mm; head width of 6 0.84 mm, 9 0.92 mm. Crown produced; median length of crown 0.74 x head width and L45 x interocu- lar width (c? n = 6; 9 n = 3). Crown hght brown to pale yellow with dark, circular spot around apex, pair of broken, transverse lines at midlength, two obli(|ue markings at rear, appearing continuous with the medial pair of six pale brown, pronotal stripes. Face with broad, dark brown, interocular band, fading to pale yellowish. Brown basally, disc and 1988 Whitcomb, Hicks: Flexamia Rf.vision 235 blX Fig. 13. Styles and style tips oiFlexamia pectinata group: A, minima; B, zamora; C, pectinate; D, bandarita; E, gila; F, doeringae; G, collorum; H,jacala; I, mescalero. apex of forewings usually with irregular, dark spot. Male. — Pygofer much like that o( pectinata but shorter; plates (Fig. 11 A) barely exceed- ing ventral lobe in lateral view or 2/3 length of pygofer, unevenly tapering to blunt apices. Genitalia similar to those of pectinata and zamora, but little more than half their size. In ventral aspect, apex of style not exceeding plane of well-developed, preapical lobe, angle formed between preapical lobe and style apex more acute than in pectinata. Connective similar to that of pectinata, but with a nar- rower, deeper, U-shaped incision at the joint with the aedeagus. Other characters as in pectinata. Female. — Sternum VII with posterior margin broadly, shallowly emarginate, hind margin with four teeth of approximately equal length (Fig. 14A), lightly infuscated. Oviposi- tor with bases of first valvulae as in zamora. Types. — Holotype i: Valles, San Luis Po- tosi, Mexico, 1 December 1938, J. S. Cald- well. Remarks. — An additional series o{ minima, collected by E. D. Ball at Monterrey, Nuevo Leon (5d, 39, 11 August 1936, USNM), has been examined. On the basis of characters of the male plates, the styles, and the distinc- tiveness of the female sternum VII from that of zamora, we reinstate minima. Species problems in this group are discussed under "Species Concept." This species, like zamora, may be entirely Mexican in distribution and is represented by the two collections discussed above (Fig. 15). The host or hosts are un- known, but the closely related pectinata spe- cializes on Bouteloua curtipendula. The grasslands of San Luis Potosi are extensive, and many other warm season grasses, includ- ing other Bouteloua spp., are present (Rze- dowski 1966); further fieldwork is required to clarify the biology oiininima. 2. Flexamia zamora DeLong & Hershberger, n. stat. Flexamin zamora DeLong and Hershberger 1947: 137. Flexamia pectinata. Young and Beirne 1958: 46. (In part.) Length of c? 3.6 (3.5-3.7) mm, 9 3.9 (3.7-4.2) mm; head width of S 1.08 mm, 9 1.15 mm. Crown not strongly produced; me- dian length of crown 0.61 x head width and 1.27 X interocular width ((5 n = 7; 9 n = 3). 236 Great Basin Naturalist Memoirs No. 12 Fig. 14. Female sternum VII ofFlexamia pectinata group: A, minima: B, zamora: C, pectinata: D, bandarita; E, gila; ", doeringae; G, collorum; H, mescalero. Crown light brown with dark, circular spot around apex, pale brown, transverse line on either side at middle. Pronotum light brown, usually with traces of four wide longitudinal stripes. Elytra subhyaline, dark brown spot on disc; costal veinlets heavily margined with dark brown. Face black above, pale brown on lower portion. Male. — Pygofer as in pectinata. Plates (Fig. IIB) extending to approximately 3/4 length of pygofer, with apices less produced than in pectinata, bluntly rounded. Style apices exceeding the plane of the moderately developed, preapical lobe, forming an angle with it smaller than that in pectinata. Other characters as in pectinata. Female. — Sternum VII (Fig. 14B) with posterior margin shallowly emarginate; four produced teeth at middle; outer teeth longer, acute, curved inward. Ovipositor with basal processes of first valvulae similar to those of pectinata, but recurved portion not separated along the middle. Types. — Holotype 6: Zamora, Michoacan, Mexico, 2 October 1941, Plummer, Good, Caldwell and DeLong (OSU). Paratvpes: 5 6 , 3 $ , same collection data (OSU and USNM). Remarks. — Although the male genitalia of zamora are similar to those o{ pectinata, the styles oi zamora differ in having a more acute angle between the apex and the less well- developed preapical lobe. Also, the plates are more evenly rounded than those of pectinata and longer than those of minima. The female sternum VII of zamora differs from that of pectinata in that the outer teeth of the hind margin are much longer than the inner teeth; in pectinata and minima the teeth are of equal length. This species, almost certainly entirely Mexican in distribution, is known only from its type locality (Fig. 15). The host or hosts are unknown, but other members of the complex feed on Bouteloua curtipendula. 3. Flexamia pectinata (Osborn & Ball) Dehocephahis pectinatus Osborn & Ball 1897: 205. Deltucephalus (Flexamia) pectinatus DeLong 1926: 32. Flexamius pectinatus. DeLong and Sleesman 1929: 83. Flexamius zatnora. Young and Beirne 1958: 46. Incorrect synonymy. Flexamia minima, Yoimg and Beirne 19.58: 46. Incorrect synonymy. Important CHARACTERS. — Length of S 3.4 (2.9-3.8), of 9 3.6 (3.1-4.0). Head slightly produced (median crown length 1.40 x inter- ocular width; 0.62 X head width) (c? n = 37; 1988 WHITCOMB, HiCKS: FLEXAMIA REVISION 237 Fig. 15. Geographic distribution of Flexaniia pectinata {i miniirui (O), And jacala (ir) with Botiteloua curfipendtila. ), doeringae (■), bandarita (A), gila (D), zatnora (A), 9 n = 29). [6] Pygofer (Fig. 62F) strongly constricted both ventrally and dorsally; plates (Fig. IIC) wide and parallel-sided for much of their length, almost subrectangular, but with rounded tips. Aedeagus (Fig. 12C) symmetri- cal, without lateral processes; gonopore sub- apical on caudoventral surface. [9] Sternum VII (Fig. 14C) with four teeth of approxi- mately equal length. Ovipositor with bases of first valvulae recurved and calipterate (Fig. 63G). Geographic distribution. — This species is a specialist of side-oats grama {Bouteloiia curtipendula) in prairie and mesic grasslands; it occurs (Fig. 15) at higher elevations of the desert plains of eastern New Mexico east to Akron, Ohio, the eastern extension of the prairie peninsula (Transeau 1935). Biology. — Like other Flexamia species, pectinata is probably multivoltine and is present throughout the growing season. In New Mexico it is often absent from its host (Botiteloua curtipendula, side-oats grama) at lower elevations where host growth may be unpredictable because of sporadic drought. In Chihuahuan and Sonoran highlands it is re- placed by bandarita and doeringae, respec- tively, and in the intervening mountains, by gila. Oligophagy coefficients. — Gramineae 1.000; Chloridoideae 0.973; Bouteloua 0.926; Bouteloua curtipendula 0.825 (n = 57). Remarks and diagnosis. — Flexamia pecti- nata can be recognized by a combination of its 238 Great Basin Naturalist Memoirs No. 12 Fig. 16. Flexamia gila, n. sp. : A, aedeagus and connective, lateral aspect; B, aedeagus and connective, dorsal aspect; C, apex of aedeagus, caudoventral aspect; D, male plates and pygofers, ventral aspect; E, male pygofer, lateral aspect; F, right style, dorsal aspect; G, ape.x of right style, lateral aspect; H, female sternum VII; I, bases of first valvulae of female, dorsal aspect. external genitalic characters and its geo- graphic range. Other species (doeringae, ja- cala) of the pectinata group with somewhat similar male plates do not occur with pecti- nata in its range. Of the nine members of the group, only mescalero (host: Muhlenhergia pauciflora) and colloruin (host: Bouteloua uniflora ) have ranges that overlap the range of pectinata. The male plates (Fig. Ill) and fe- male sternum VII (Fig. 14H) of mescalero distinguish it from pec^inaf a (Figs. IIC, 14C). The male plates oi collorum (Fig. IIG) differ from those of pectinata in general aspect and in the absence of the sclerotized ridge on the inner plate surface that characterizes coUo- rum. The female sternum VII of collorum is similar to that ofpectinata. Although the char- acters discussed above permit tentative recognition of the commonly encountered pectinata, definitive identification requires examination of the genitalia. 4. Flexamia gila, n. sp. Length of 6 3.4 (3.1-3.7) mm, 9 3.8 (3.6-4.0) mm; head width of d 1.03 mm, 9 1.08 mm. Crown not strongly produced; median length of crown approximately 0. 63 x head width and 1 . 32 x interocular width (d n = 6; 9 n = 4). Crown light brown with dark, circular spot around apex, pale brown, transverse line on either side at middle. Pronotum light brown, usually with traces of four wide, longitudinal stripes. Elytra subhyaline, dark brown spot on disc; costal veinlets heavily margined with dark brown. Face black above, pale brown on lower portion. Male. — Pygofer (Fig. 16E) much as in ban- darita; posterior margin slightly more pro- duced. Plates (Fig. 16D) broad, gradually rounded from base to apex, extending to ap- proximately 4/5 length of pygofer. Connective in lateral view with dorsal keels narrow (or rarely absent), approximately 1/4 height of dorsal apodeme; apodemal processes as in bandarita; styles (Fig. 16F) long, diverging apically in ventral aspect, preapical lobe rounded, produced ventrad and laterad; style 1988 Whitcomb. Hicks; Flexamia Revision 239 portion distad from preapical lobe gradually narrowing to embrowned, strongly divergent, acute apex in ventral aspect, or, in lateral aspect (Fig. 16G), appearing bladelike, pro- duced ventrally and flattened, bearing minute teeth along the irregularly curved ventral margin of style ending in a truncate apex that is slightly curved ventrad. Aedeagus (Figs. 16B,C) symmetrical, short, slender. Apical margin flared and convex, much as in pecti- nata. Distance between gonopore and apex less than in pectinata. Gonopore subapical on caudoventral surface. Female.— Sternum VII (Fig. 16H) with posterior margin broadly but shallowly emar- ginate, with four produced teeth; middle teeth broad. Ovipositor with basal processes of first valvulae (Fig. 161) recurved; recurved portion constricted above proximal margin, expanded slightly mesad and distad into dif- ferentially sclerotized lobes. Types. — Holotype 6 : New Mexico, Sierra Co., Kingston, 17 August 1985, R. F. Whit- comb (6,300 ft, IPL 001875, Bouteloua cur- tipendula). Paratypes: 3 6 and 1 9, same collection data; 2 6 and 2 9 , Mexico, 5 km S Casas Grandes, Chihuahua (5,600 ft, Boute- loua curtipendula, 3 September 1987, A. L. Hicks, IPL 001774); 2 c? and 1 9 , New Mex- ico, Socorro Co., Magdalena Mtns., Water Canyon, 7,000 ft., 16 August 1987, IPL 001756, A. L. Hicks. Deposited BARC, CNC, KSU, KU, OSU, and USNM. Remarks. — The aedeagal characters of gi/a are intermediate between those of doeringae and pectinata. The styles in gila are less pro- duced ventrally than in doeringae. The longer, bladelike styles of gila differentiate males oi gila from those of pectinata; females can be readily differentiated on the basis of the morphology of the female sternum VII and the bases of the first valvulae. The host of gila is B. curtipendula. The name gila, a noun in apposition, denotes the New Mexican mountains where the holotype was collected. 5. Flexamia bandarita, n. sp. Length of 6 3.4 (3.3-3.5) mm, 9 3.7 (3.4-4.0) mm; head width of 6 1.09 mm, 9 1.04 mm; head not strongly produced; median length of crown 0.64 x head width and 1.33 x interocular width (n = 8 d , 3 9 ). Color variable, ranging from very dark gray to pale, with irregular, dark markings on dor- sum and forewings; face very dark with pale arcs on upper half; variable in lower half. Head stramineous in lighter specimens with paired, transverse lines at midpoint of crown and obli(}ue markings at rear; venter and legs with irregular, fuscous markings. Male. — Pygofer strongly produced ven- trally in rounded lobe; plates (Fig. 17D) contiguous on basal half, diverging apically into narrow V; extending to approximately 3/4 length of pygofer; style (Fig. 17F) heavily sclerotized, sinuate in ventral aspect, extend- ing almost to the apex of the aedeagus, gradu- ally narrowing to acute, ventrally directed, chelate apex (Fig. 17G) that is closely associ- ated with sclerotized ridge on inner surface of each plate; connective in lateral aspect with dorsal keels narrow; aedeagus (Figs. 17B,C) symmetrical with slender shaft, straight in ventral aspect, without apical processes. Apex flared in caudoventral aspect, but varying from concave to convex. Dorsal apodemal process small, not attaining the plane of the shaft of the aedeagus, appearing suboval in caudodorsal aspect, with lobelike, converg- ing, ventral apices. Gonopore minute, sub- apical on caudoventral surface. Female. — Sternum VII (Fig. 17H) with in- ner teeth of the hind margin longer than outer pair. Ovipositor with basal processes of first valvulae (Fig. 171) recurved. Types. — Holotype 6: Brewster County, Texas, Chisos Mountains, Big Bend National Park, Texas, 7 August 1987, R. F. Whitcomb (5,200 ft, Bouteloua curtipendula, IPL 003262). Deposited in USNM. Paratypes: 2 (5 , 1 9 , same collection data; 1 S , Texas, Marathon Basin, 8 August 1984, deposited USNM, BARC, KSU, KU; 5 c?, 1 9, Chisos Mountains, D. J. and J. N. KnuU, 17 July 1946, deposited OSU. Remarks. — Flexainia bandarita resembles pectinata, gila, and doeringae, but the sinuate styles of bandarita separate it from these spe- cies. The middle teeth of the female sternum VII are longer than the outer pair, suggesting an intermediate stage in development of the median process that is present in doeringae. Like pectinata and doeringae, bandarita is a specialist of side-oats grama and, so far as known, is restricted to that host. Its known geographic range (Fig. 15) includes the higher, more mesic grasslands of the Chisos Mountains in Big Bend National Park and the 240 Great Basin Naturalist Memoirs No. 12 Fig. 17. Flexamia bandarita, n. sp.; A, aedeagus and connective, lateral aspect; B, aedeagus and connective, dorsal aspect; C, ape.x of aedeagus, caudoventral aspect; D, male plates and pygofers, ventral aspect; E, male pygofer, lateral aspect; F, right style, dorsal aspect; G, apex of right style, lateral aspect; H, female sternum VII; I, bases of first valvulae of female, dorsal aspect. Marathon Basin north of the park. This region is rich in endemism presumably derived by insularization of high grasslands by desertifi- cation in the surrounding true Chihuahuan desert. All insects in our recent series, taken August 1987, are extremely dark; the season had been one of record rainfall, and vegetative growth of all plants in the Chisos Mountains was very lush. The name bandarita, a noun in apposition, is one of the common names for side-oats grama in Mexico. 6. Flexamia doeringae Beamer & Tuthill Flexamia doeringae Beamer & Tuthill 1934: 3. Important characters. — Length of 6 3.5 (3.2-3.7) mm, of 9 3.7 (3.3-4.1) mm. Head produced (median crown length 1.40 x inter- ocular width; 0.70 X head width) (d n = 20; 9 n = 20). Face ivory with broad, black, basal, interocular line, occasional dark markings be- low. [6] Plates (Fig. IIF) broad through most of their length, then narrowing sharply to rounded apices. Style with preapical lobe strongly developed, flattened and produced posteriorly; aedeagus (Fig. 12F) with protu- berance near shaft apex close to tip in ventral aspect. [ 9 ] Sternum VII (Fig. 14F) with inner teeth strongly produced and closely associ- ated to form median process; ovipositor with each first valvula (Fig. 63A) recurved at base; recurved portion with transverse lobe cepha- lad of basal extremity. Geographic distribution. — This species is known primarily from the mountains (Babo- quivari, Catalina, Chiricahua, Huachuca, and Santa Rita ranges) of southeastern Arizona, but it also occurs in the Mexican highlands of Sonora and Chihuahua (Fig. 15). There is one record from Yavapai Co., Arizona (1 6 , Gran- ite Dell, 10 July 1933, R. H. Beamer). Biology. — This species is apparently a 1988 Whitcomb, Hicks: Flexamia Revision 241 specialist on side-oats grama, Bouteloua ciir- tipendula. Because this grass is a dominant in the mountains of southeastern Arizona and because many collectors have visited this re- gion, doeringae is a common component of large Flexamia collections (Appendix I). The climate of the southeastern Arizona moun- tains is similar to that of the Mexican high- lands but shows a definite Sonoran influence. Flexamia doeringae may therefore be a Sono- ran vicariant that separated from the Chi- huahuan bandarita and the prairie pectinata, which are also specialists of B. curtipendula. Oligophagy coefficients. — Gramineae 1.000; Chloridoideae 1.000; Bouteloua 1.000; Bouteloua curtipendula 0.943 (n = 35). Remarks and diagnosis. — The range oi do- eringae does not overlap that of any other member of the pectinata species group. This species can be readily distinguished by its ivory face, the pair of transverse lines on a pale, well-produced crown, and the structure of the external genitalia. The female sternum VII (Fig. 14F) is particularly distinct in having a medially produced structure consisting of a pair of long, central lobes and much shorter lobes on each side. The middle pair of teeth, in their size and degree of fusion, are maxi- mally developed in this species. 7. Flexamia collorum, n. sp. Length of c? 2.9 (2.7-3.0) mm, 9 3.0 (2.8-3.2) mm; head width of 6 0.87 mm, 9 0.89 mm. Crown not strongly produced; me- dian length of crown 1.53 x head width and 0.68 xinterocular width (c? n = 11; 9 n = 20). Crown stramineous to light brown, prono- tum with four faint, wide, brown stripes, forewings gray with irregular, dark markings. Face color and markings variable, without in- terocular line, but usually black in interocular area and lower area. Usually at least lorae pale. Crown with pair of oblique markings at rear, pair of transverse lines at midlength. Venter usually predominantly dark. Male. — Pygofer produced ventrally into an angulate lobe; caudoventral margin sloping dorsally to a broadly rounded, caudal lobe. Plates (Fig. 18D) short, extending to 1/2 length of pygofer, fused basally, contiguous medially for 2/3 length, acute apically. Plates produced on inner surface at about midlength into well-developed, sclerotized ridge that interfaces with style apices. Connective (Fig. 18A) in lateral view with dorsal keels narrow, approximately 1/3 height of dorsal apodeme; apodemal processes as in ban- darita; style (Fig. 18F) short, abruptly nar- rowing beyond preapical lobe to avicephali- form, mesoventrally directed apices (Fig. 18G). Aedeagus (Figs. 18B,C) symmetrical, shaft slender, without processes, slightly flared apically in caudoventral aspect. Gonopore subapical on caudoventral surface. Female.— Sternum VII (Fig. 18H) with four teeth on hind margin; outer pair not well developed, inner pair distinct, but short and rounded; ovipositor with basal processes of first valvulae (Fig. 181) recurved. Types. — Holotype curvata [s] Aedeagus with two pairs of pro- cesses, one apical, one anteapical (Figs.7F,G). [au] Apical processes long and not in bilaterally symmetrical plane (Fig. 7F). Gonopore an elongate channel. host = Buchloe dactyloides zacate —* canyonensis [s] Aedeagus curved, acute api- cally, without aedeagal processes host - (Figs. 7I,J). Muhlenbergia porteri [au] Gonopore at midlength of aedeagal shaft. ininiiTui —* zamora fs] Style apices in lateral aspect bluntly produced (Figs. 13A,B). [au] Outer teeth of female sternum VII incurved, longer than inner teeth (Fig. 14B). host unknown bandarita —* gila host = Bouteloua curtipendtila albida —* slossunae habitat specialists picta —* pyrops host = Aristida spp. [s] Female sternum VII with two produced, median teeth (Figs. 14D,E). Male plates broad, blunt- ly rounded (Figs. 11D,E). [au] Male plates nearly as long as pygofer (Fig. HE). [s] Posterior lobe of male pygofer undifferentiated (Figs. 62Q,T). [au] Male plates exceed pygofers (Fig. 8B): female sternum VII trilobed (Fig. lOB). Aedeagus and connective partially fused. [s] Aedeagal tip capitate, toothed, with pair of subapical processes (Figs. 7K,L); pygofers (Figs. 5A, B) angulate with ventral process. [au] Vertex extensively produced, anteapical aedeagal processes (Fig. 7L) more remote from apex, male plates (Fig. 9B) very short, rounded. 1988 Whitcomb, Hicks; Flexamia Revision 295 arizonensis —* Jlexulosa hosts = Aristidal Bouteloua g,racilis youngi —* decora host = Muhlerihergia richardsonis celata —* stylata hosts = Redfieldia/ Muhlenhergia spp. heameri —> texana hosts unknown [s] Male plates fFi«s. 33A,Bl short, divergent apically. [an] Apical aedeagal processes short (Fig. 3 IK). [s] Unpaired ventral aedeagal pro- cesses (Figs. 31F,G) symmetrical and much longer than paired pro- cesses. [an] Unpaired process (Fig. 31G) without median groove, aedeagal shaft curved. [s] Face (Fig. 3B) with discrete lines forming interocular band; three aedeagal processes angu- larly divergent from shaft (Figs. 31H,I). Pygofer with distinct pos- terior process (Figs. 6200,PP). [au] Apices of male plates sub- rectangular; aedeagus (Fig. 31H) asymmetrical. [s] Aedeagal processes about equal in length, but ventral pro- cesses not in sagittal plane (Figs. 31 A, D). [au] Aedeagal processes twisted apically (Fig. 31A). imputans — » areolata [s] Unpaired process lateral, with hosts ^ Muhlenhergia cuspidatal Eragrostis spectahilis ventral groove (Figs. 31B,C). [au] Aedeagus and connective fused, unique habitus (Fig. 2F). })rairiana — » rejlexa hosts andropogonoid grasses sandersi — » dehmgi hosts andropogonoid grasses satilla —* claiji hosts andropogonoid grasses [s] Gonopore dorsiJ (Figs. 44D,G). [au] Groove absent on unpaired process (Fig. 44D). [s] Lateral process appressed to aedeagal shaft (Figs. 44B,C). [au] unpaired process (Fig. 44C) at less acute angle with aedeagal shaft, [s] P'lared, elongate gonopore (Figs. 44H,J). [au] Gonopore broad basally (Fig. 44H). B. Designation ofclades. The next step consisted of appending other species to the sister sets to define clades: mtntnia —* zamora —* pectinata host probably = Bouteloua curtipendula handarita —* gila —* doeringae host = Bouteloua curtipendida graminea -> satilla — clayi hosts = andropogonoid grasses [s] Style with apical lobe not con- stricted (Figs. IIA-C); [au] Male plates (Fig. IIC) sub- rectangular; style tip (Fig. 13C) not produced in lateral aspect. [s] Middle teeth (Figs. 14D-F) of female sternum VII longer than outer teeth. [au] Middle teeth long, associated to form process (Fig. 14F). [s] Spiral gonopore (Figs. 44F,H,J) [au] Flared gonopore (Fig. 44H) C. Establishment of species groups. Clades were next assembled into species groups. These are the groups (I-XIII) described herein. Synapomorphies defining clades and species groups are given in the descriptions to the groups and in the legends to Figures 56-59. D. Establishment of polarity between Spar- topy ge and Flexamia. The dorsal and facial habitus of grammica (Fig. 2C), albida (Fig. 2A), and Spartopyge (Fig. 54) are clearly ho- mologous. On the other hand, few characters of grammica are clearly homologous to those of other Flexamia species. The morphology of the male aedeagus and connective places this species in the genus, but the reduced number of aedeagal apices and their apparent phyletic twisting make it difficult to assign homology. The male pygofer and female sternum VII suggest retention of plesiomorphic characters of an ancestor allied to Spartopyge . We there- fore hypothesize the polarity Spartopyge — ► grammica . Derivation of the similarly striped albida group, of course, could have occurred only after development of paired aedeagal processes and fusion of the anteapical pair into a single, unpaired process (Fig. 56). Revers- ing the polarity between grammica (or albida ) and Spartopyge would require generation of a wide array of apomorphies present in Spar- topyge, including a brush of setae on the inte- rior of the pygofer and the complex structures of the male genitalia (Fig. 55C), accompanied by complete loss of all apical processes. Also, the unique sternum VII (Figs. 54D,E) of S. mexicana and S. miranda would have to arise suddenly. Finally, if Spartopyge were derived from the Flexamia lineage, it would be necessary to postulate a significant apo- morphic increase in size. This would consti- tute a reversal of the overall trend for size reduction and simplification in the Delto- cephalinae in general (Whitcomb et al. 1986) and Flexamia in particular. We therefore 296 Great Basin Naturalist Memoirs No. 12 =2 5 II III 7 8 10 IV 15 19 VI 16 VII 14 12 13 Fig. 56. See facing page. 20 VIII 18 23 XI XII 30 31 2< XIII 32 25 IX 21 22 28 26 24 hypothesize (with Young and Beirne, and Ross) that Spartopyge is an outgroup of the Flexamia hneage. We beheve, with these workers, that a primitive Flexamia ancestor arose from the Spartopyge hneage and later gave rise to a hneage with paired, aedeagal processes. This hneage in some way gave rise to grammica but, ultimately, to all Flexamia groups IV-XIII. We assume that the dorsal location of the gonopore in grammica and prairiana is a result of independent evolution- ary events, since these species share few other significant characters. Whether the minute, paired processes that occur midlength on the Spartopyge aedeagus are homologous with the Flexamia aedeagal processes is conjec- tural. Phylogeny of Flexamia groups and spe- cies. The postulated relationships and polari- ties permit a cladogram to be drawn relating Flexamia groups and key species to the out- groups Alapus and Spartopyge (Fig. 56). Sev- eral significant possible variations of the pro- posed tree are given in Figure 57. Finally, the phylogeny of Flexamia species was inferred on the basis of the characters discussed above. We present cladograms for groups I-IX (Fig. 58), groups VII-XI (Fig. 59), and groups XI-XIII (Fig. 60). Synapomorphies used to define clades within each group are given in the legends to the figures. Character transformations. — Our intu- itive phylogeny provided general guidelines concerning character transformation in Flex- amia. From these concepts we constructed branched transformations for the PHYSYS program of Mickevich and Farris (1984). These transformations, which are described 1988 Whitcomb, Hicks Flexamia Revision 297 Fig. 56. Proposed phylogeny of Flexamia groiip.s and key species. Synapoinorphies [s] or, for monobasic groups, autapomorphies [an] are indicated. We hypothesize an ancestral form (1) that possessed dorsal stripes (Fig. 54 A) and a pale face with a contrasting black interoeiilar band (Fig. 54B) similar to those of Figs. 3A and 3B, relatively wide male plates (Fig. 54C); an aedeagus distinctly jointed with the connective, without apical processes, and with an anteapical gonopore on the caudoventral surface. Divergence of this lineage led (3) to Alapus (Figs. 55F,(v), represented in this diagram by elongatus, and to Spartopy^e, represented (4) in this diagram by miranda and mexicana, in which [s] the dorsal aedeagal apodeme is large and hoodlike (Fig. 55C). The transformations between Flexamia and its outgroups are tenuous (see te.xt). In Flexamia [s] the dorsal aedeagal apodeme (Fig. 6) has a pair of conspicuous appendages directed caudoventrad or caudad (5). In the first lineage (6) to diverge, the plesiomorphic aedeagal apex, which lacked processes, was retained, as were the caudoventral position of the gonopore and broad, ancestral plates; the contrasting face was retained in a minority of species, but the striped habitus was completely lost. Synapoinorphies defining this lineage include [s] the curved aedeagus lacking processes (Figs. 71, J; 16A, 17A, 2()A) and the male pygofers, which possess both dorsal and ventral constrictions (Figs. 62A-I). This lineage diverged into the pectinata (7) and the abhreviata-zacate lineages (8). In the pectinata group the plates (Fig. 11) are distinctively parallel-sided basally (except in collorum). In the abhreviata-zacate lineage the gonopore was at first elongate (Fig. 24) and [s] the posterior lobe of the male pygofer strongly produced (Figs. 62H,I). In the ahbreviata group (II) the plates (Fig. 8E) are reduced (9), and [au] a distinctive pair of minute apical processes (Fig. 7H) developed. The small aedeagal processes of the abbreviata group are not clearly homologous with other aedeagal processes in Flexamia. In the zacate group (III) the gonopore (Fig. 24) was at first elongate (10) but in canyonensis moved to the aedeagal midpoint. At this point a significant apomorphy appeared. An ancestral form (11) is hypothesized in which two pairs of aedeagal processes were present. It is unclear whether this form had fully developed before divergence of grammica (IV), in which the plesiomorphic feature of dorsal stripes was retained (12), but in which [au] the aedeagal processes were distinctively modified (Fig. 7E) and the gonopore had moved to the dorsal surface (12). A possible apomorphy linking g,rammica with the curvata group is the foot-shaped style apex (see illustrations of Young and Beirne 1958). In the cttrvata-picta lineage [s] both pairs of aedeagal processes are present (14), but many habitus features, including stripes and contrasting face, have been lost. In the curvata group (V) two pairs of processes are present (15) and [s] unmodified (Figs. 7F,G). In the picta group (VI) the apical processes are [s] modified (Figs. 7K,L) as flangelike, toothed appendages (16). Remaining species are defined by an apomorphy that developed with the presumed fusion (17) of the anteapical processes into a single, unpaired process that was plesiomorphically located ventrally in the sagittal plane (Figs. 31, 61A). A second apomorphy characterizing this divergence is the bifurcation (Figs. 7A-C) of the apical aedeagal processes, a feature that was subsequently lost (22). The plesiomorphic state of dorsal stripes and contrasting face was preserved (18) during the fusion of subapical processes and is present (Figs. 2A,B) in the albida group (VII). This lineage, which can be defined [s] in terms of the absence of dorsal or ventral constrictions in the posterior lobe of the male pygofer (Figs. 62Q,T), is represented today by the distantly related sister species albida (19) and slossonae (20); for autapomorphies defining these species, see text. In serrata (VIII) [au] additional dorsal aedeagal processes (Fig. 7A) developed (21), and, while the contrasting face was retained, dorsal stripes were lost (Fig. 1). In ritana (IX) the dorsal stripes were retained (Fig. 2D) but greatly reduced. This monobasic group (23) is defined [au] by the unique, pointed aedeagal apex (Fig. 7D). In one lineage of the^^xw/osa group (25), the unpaired aedeagal process is, plesiomorphically, ventral in the sagittal plane, permitting retention of aedeagal symmetry (Figs. 31G,I,J,K,L). The plesiomorphic pale face with black interocular band (Figs. 3B,F,G) was also retained. This clade is defined [s] by the triangulate plates (Figs. 33A-F, 34D) with conspicuous, apical divergence. (But in stylata [Fig. 34E], the plates [au] are truncated.) In the remaining species (26) the unpaired process underwent phyletic twisting (Figs. 61B-H) and the contrasting pale face, which had persisted through many nodes, was finally lost. In the inflata subgroup of group X, [s] the displacement of the unpaired process (Fig. 61 B) was relatively small (27), but [s] in the imputans-areolata (XI-XII) lineage the process had become lateral (Figs. 31B,C) and bore the gonopore as an elongate slit (29). Before divergence of groups XI-XIII, the connective and aedeagus had remained articulated (but see discussion of group VI in text). We hypothesize that after divergence of the imputans-areolata lineage, a fusion of aedeagus and connective occurred (31) in areolata . The imputans-areolata lineage is very specialized in the location of the unpaired process, in the black face, and the greenish dorsal pigmentation. Of the two species, areolata is the more highly specialized (Figs. 2F, lOQ); in fact, it may be the most highly specialized of all Flexamia species. It is most unlikely that the widely distributed and variable prairiana (see text) and the entire prairiana group (XIII) were derived from the specialist areolata lineage. Instead, we propose that divergence of a second lineage in which aedeagus and connective fused (32) led to the prairiana group. This fusion, the traces of which are evident (Fig. 6F) in prairiana , is designated [s] as a synapomorphy defining group XIII, and the fusion in areolata is defined as an autapomorphy defining that species. Details of species phylogenies are presented in detail in Figures 58-60. in Appendix II, Part III, involved the follow- ness (lateral aspect) of the aedeagus; (7) length ing characters: (1) relationship between ae- ofthe aedeagus; (8) form of the aedeagal shaft; deagus and connective; (2) morphology ofthe (9) form of margins of paired, aedeagal pro- aedeagal apex; (3) morphology of anteapical cesses; (10) form and position ofthe gonopore; aedeagal processes; (4) phyletic twisting ofthe (11) morphology of apices ofthe apodemal unpaired ventral process (Fig. 61); (5) curva- arms; (12) position of apodemal arms; (13) ture (lateral aspect) ofthe aedeagus; (6) thick- width of dorsal keels of the connective; (14) 298 Great Basin Naturalist Memoirs No. 12 ■§ 5 c o .2 I III V VI 1 II 4 1^ 2 1 A VIII X^ Xb IX 2 3 1 VII B XI XII 3 2 1 c XIII Fig. 57. Alternative resolutions for Flexamia group phylogenies. A. In this resolution (I) two groups (I and III) are defined by [s] curved aedeagus without processes (Figs. 71, J, 16A, 2IA). The alternative lineage (2) is defined by an apomorphic elongation of the gonopore. This was followed by derivation of a further lineage (3) in which aedeagal processes are present. At first these are [s] minute (II abbreviata. Fig. 7H) but later become (4) fully developed (see Figure 56). This resolution avoids hypothesizing independent origins for the aedeagal processes in abbreviata and curvata but hypothesizes additional nodes between Spartopyge and the Flexamia species with very similar habitus, implying repeated, independent losses of the complex pattern of striping in Spartopyge, grainmica, and albida. B. In this resolution, it is hypothesized (1) that the serrata group and decora subgroup of group X belong to a common lineage, and that the inflata subgroup of group X is more closely related to ritana. This construction has considerable merit and takes into account the similarities between the serrata male plates (Fig. 8C) and those of the decora subgroup (Figs. 33, 34D, E), but deemphasizes many similarities between members of the inflata and decora lineages, and would augur for elevation of these lineages to groups. C. It is possible to hypothesize a single (2) fusion of aedeagus and connective, rather than the two fusions hypothesized in Figure 56. This construction, in our view, gives insufficient weight to several synapomorphies that link areolata and imputans, which would, in fact, have been designated as a close sister species set had they not differed in the crucial relationship of aedeagus to connective. See discussion in legend to Figure 56, and text. morphology of the male pygofer (Fig. 62); (15) morphology of the male plates; (16) morphol- ogy of the female sternum VII; (17) presence or absence of dorsal stripes; (18) coloration of the face. The above transformations, as presented in Appendix II, are suitable for entry into the PHYSYS program and should serve as a useful starting point for more refined analysis of Flexamia phylogeny. Summary of phylogenetic model. — In summary, after numerous iterations, we pro- pose a formal phylogeny for Flexamia. The model is rooted in earlier informal work of Young and Beirne (1958) and Ross (1970, and unpublished notes). Our model, in addition to its formality, considers 14 species not avail- able to earlier workers and proposes species groupings based in large part on synapomor- phies. In particular, the addition of seven spe- cies to the pectinata group (I) and four species to the flexulosa group (X) adds new phyloge- netic perspectives. We have accepted the hypothesis of Young and Beirne of an ancestral form with two pairs of aedeagal processes. We also endorse the proposal of Young and Beirne (and Ross) that the anteapical pair of processes fused to form a single, unpaired process (Fig. 52). Further, we accept the proposal of Ross (Fig. 53) that the fusion of aedeagus and connective is an important synapomorphy that defines the prairiami group. The informal proposal of Ross differed from that of Young and Beirne in one major way (in addition to its presentation as a tree). Ross believed that the lack of aedeagal processes in pectimita and canyonensis was plesiomorphic and related it to the similar lack of processes in his outgroups, Spartopyge and Aflexia. In our 1988 Whitcomb, Hicks: Flexamia Revision 299 proposal we have sided with Ross. Unless dif- ferent outgroups were chosen, we feel such an assignment is inevitable. Also, transforma- tions within Flexamia are best explained by the chosen polarity. For example, viewing the pectinata group as the terminus of a simplifi- cation of the aedeagus would require the transformation of male plates from narrow to broad, in direct contrast to the general ten- dency for streamlining of the plates that runs throughout the remainder of the Flexamia tree. We have assumed, with Ross, that the minute processes in abbreviata are not ho- mologous with other Flexaryiia aedeagal pro- cesses. This assumption, which permits many other reasonable character transformations to be postulated, appeared in our model only after numerous iterations failed to find a bet- ter resolution, considering all transforma- tions. Such issues are best addressed by close examination of the structures themselves, preferably by scanning electron microscopy, for evidence that would bear on the question of homology. It is possible that a computer- ized search for a parsimonious tree would turn up a cladogram that avoids the pitfalls we encountered in our intuitive search. In any event, we present in Figure 57A an alterna- tive cladogram that, whatever its other prob- lems, assumes homology between the aedea- gal processes of abbreviata, grammica, and curvata. Young and Beirne (1958), although uncer- tain about the status oidakota, atlantica, and grajnmica, nevertheless placed them tenta- tively in their lineage IIIB (Fig. 52). We con- cur with Ross in his placement of dakota and atlantica. However, we disagree with both Young and Beirne and Ross in the placement oi grammica. Whereas the dorsal location of the gonopore and the presence of two serrate, aedeagal processes of this species is clearly highly apomorphic, we see nothing else in grammica that links it to group XIII, in which the dorsal location of the gonopore is a signifi- cant apomorphy. For example, the male plates and pygofers, the female sternum VII, the clear articulation of aedeagus and connec- tive, the striped habitus and pale face, and the overall large size oi grammica are derivable as transformations from the outgroup Spar- topyge, but they would produce chaos if in- serted into transformations in group XIII (or any other group or groups between VII and XII). In fact, the ties between Spartopyge and grammica are sufficiently reasonable that we have couched our argument for the polarity Spartopyge — ► Flexaitiia in terms oi grammica (see earlier section). The model we present solves many of the problems that we encountered. Resolution of many (juestions was made possible by propos- ing (1) that aedeagal processes arose in the course of Flexamia evolution and were not inherited from an unknown ancestor; (2) that aedeagal symmetry was plesiomorphic (but note the apomorphic symmetry in dakota). Further character transformations have been proposed (Appendix II, Figs. 61, 62) that lead to consistent hypotheses for transformation of other characters (e.g. , bases of the first valvu- lae, which are presented in Fig. 63 in terms of our proposal but were not used to derive it). The proposed resolution avoids hypothesizing independent acquisitions of the unique striped habitus present in Spartopyge, and in grammica, albida, and slossonae. It further avoids hypothesizing independent deriva- tions of the unique facial habitus of the out- group, which was retained through a consid- erable segment oi Flexamia evolution. We do hypothesize several independent losses of these distinctive sets of habitus characters. Finally, the proposed model follows the fol- lowing general polarities: (1) tropical grass- lands — ► subtropical grasslands — ► temperate grasslands — ► [semiarid grasslands, prairie]; (2) large — ► small size; (3) grass generalist — ► chloridoid specialist — ► mixed chloridoid-pan- icoid specialist — > andropogonoid specialist; (4) striped habitus —* [light habitus (semiarid), brown (prairie)]; and (5) aedeagal symmetry and caudoventral gonopore — ► highly special- ized aedeagal configurations and gonopore lo- cations. But not all the problems have been solved. We regard the cladogram for groups I-III to be tentative. The position of mescalero as a member of group I is unclear; this unique species -might have been more appropriately considered to be a separate group linking groups I and III. In fact, the general question of polarity of the pectinata group should be critically addressed. We are not certain how to interpret evidence of considerable recent spe- ciation in this group (i.e., bandarital gila , min- imal zamoral pectinata). It is likely, given our 300 Great Basin Naturalist Memoirs No. 12 1 1 e e Q Q Q S k. •S ■§ •£ .§ c o Q S Q o '^ ■* Cfc ■O ea. N o 9 10 12 15 13 2 I 17 18 II 19 III 16 II 24 25 I\ 27 23 21 28 24J S S Q -§ s ■§ .& 8C XI XII 8 13 14 16 18 19 12 17 15 11 10 9 6 XIII Fig. 60. Phylogeny of species of groups XI-XIII. Synapomorphies [s] and autapomorphies [au] are identified. In the plesiomorphic state (1) there may have been moderate aedeagal asymmetry, as in inflata (Fig. 61B). In the imputans- areolata hneage (2) [s] the unpaired processes are lateral, bearing a slitlike gonopore (Figs. 31B,C). In imputans (3) the ancestral articulation between aedeagus and connective was retained. This unique species is defined by a suite of apomorphies, including its black face, greenish dorsal pigmentation, and lack of dorsal markings, but lacks an autapomorphy. In areolata (Fig. 61C) [au] the aedeagus and connective have fused, and a conspicuous, black spot (Fig. 2F) is present in the corium (4). Because the imputans and areolata lineage is defined by several synapomorphies (see text), we have designated them as sister species, despite the important difference in the state of articulation between the aedeagus and connective. We hypothesize a second, independent fusion of aedeagus and connective that serves as a synapomorphy [s] defining group XIII (4). In the prairiana-reflexa lineage [s] the gonopore has become dorsal (Figs. 61D,E), as has the unpaired process (6). In prairiana (7) the traces of this fusion [au] are present as a transverse line (Fig. 6F) on the surface of the aedeagal shaft. The aedeagal configuration [au] is also unique (Fig. 61D). In reflexa (8) the gonopore [au] has become associated with the unpaired process (Fig. 61E). In atlantica one process has been lost, and [au] the aedeagus (Fig. 44A) has become highly asymmetrical (9). In producta [au] the gonopore has moved (10) to a lateral position (Fig, 61G), whereas in dakota (11) the gonopore remained dorsal (Fig. 441), but the unpaired process has been lost. The symmetry of the dakota aedeagus is therefore apomorphic. In the sandersi-delongi lineage (12) the unpaired process has been retained but is present [s] as a lateral process that is closely appressed to the aedeagal shaft (Fig. 61F). In all three species of the final lineage (15), the unpaired process has essentially disappeared; instead, [s] the gonopore has become spiral, with [au] a narrow, basal portion (Fig. 44F) as in graminea (16), or [s] with a wider base (17) as in clayi and satilla (Figs. 44H, J). For discussion of the minor morphological divergences that define the sister species of the sandersi-delongi and clayi-satilla sets, see text. 304 Great Basin Naturalist Memoirs No. 12 Fig. 61. Postulated character transformation: phyletic twisting and modification of the Flexamia aedeagus. In decora (A) a plesiomorphic state exists in which the unpaired process is ventral and in the sagittal plane; the aedeagus is therefore symmetrical. In inflata (B) there is a small displacement of the unpaired process. In areolata (C) the unpaired process is lateral and bears the gonopore. In prairiana (D) the process is dorsal, as is the gonopore. In reflexa (E) the gonopore has become associated with the dorsal unpaired process. In sandersi (F) the unpaired process is reduced and appressed to the shaft. In producta (G) the process is reduced or absent and the gonopore is lateral. In ^raininea (H) the process has disappeared entirely, the apical processes are reduced, and the gonopore is spiral. Aspects; AD, anterodorsal; CV; caudoventral; DS, dorsal; LT, lateral; VN, ventral. Aedeagal structures redrawn from Young and Beirne(1958). adapted to northeastern {minima) and west central (zamora) Mexican grasslands, to Chi- huahuan (bandarita ) and Sonoran {doeringae) montane grasslands of the Southwest, to the interdesert montane region of the Gila Moun- tains igila), and to the tall-grass and mixed- grass prairie of the United States (pectinata). Also derived from the side-oats lineage were jacala (Mexico: host unknown) and collorum (Edwards plateau; host: Bouteloua uniflora), a close relative of side-oats grama). Clearly, most or all of the nodes in the pectinata lin- eage are Mexican or southwestern; none rep- resent events that assuredly took place north of the Mexican-U.S. border. A second lineage of the pectinata subtree resulted in the divergence of mescalero, a distinctive species that represents a host transfer to New Mexican muhly, Midilenher- gia pauciflora. Another branch of this lineage transferred to temperate Bouteloua grass- lands of Mexico. One of the branches (abbre- viata group) inhabited the mixed Bouteloua grasslands of Mexico and, by gradual adapta- tion, to similar grasslands of the United States. Another branch adapted to semiarid grasslands by colonizing bush muhlenbergia {Muhlenbergia porteri)., divergence of this lineage led to the contemporary canijonensis (Sonoran) and zacate (Chihuahuan). Consider next the surcula-curvata lineage. Both species possess the four aedeagal pro- cesses that Young and Beirne proposed as ple- siomorphic; both species are Buchloe special- ists. We propose that an ancient Mexican an- cestral line adapted to B. dactyloides, which led to the present-day surcula and, later, to curvata. This divergence (Fig. 64) may have been mediated by climate. The grasslands of northeastern Mexico, where surcula may have arisen, have escaped major disturbance since the Cretaceous (Rosen 1978). This re- gion has an extremely warm, temperate or subtropical climate and a Chihuahuan precip- itation pattern, both of which differ im- mensely from the dry, temperate grasslands where curvata occurs today. Another branch of the curvata subtree di- verged, perhaps in Mexico or south Texas, leading to the picta-pyrops sister set. These species are Aristida specialists. One of the major events in Flexamia phy- logeny was marked by the fusion of the an- teapical pair of aedeagal processes into a sin- gle, mesal process. The most primitive species in which this feature is retained are members of the albida, serrata, and ritana groups. Young and Beirne argued (and Ross concurred) that slossonae, albida, and serrata were related. We agree but consider their divergence to be an ancient one, given their entirely different climatic reciuirements (sub- tropical, prairie, and northern grasslands, re- spectively), and their retention of the Spar- topyge habitus type. Perhaps the divergence of the albida lineage into four species, each with imiciue climatic adaptations, may have occupied a distinct stage in Flexamia evolu- tion. 1988 Whitcomb, Hicks: Flexamia Revision 305 Fig. 62. Character transformation of the Flexamia pygofer. Diagrammed Hneage was based in part on phylogeny of groups (see Figure 56). A, collorum; B, mescalero; C, handarita; DJacala; E, gila; F, pectinata, G, doeringae; H, ahhreviata- I, canyonensis; J, pijrops- K, curvata, L, surcula- M, picta; N. Spartopijge miranda; O, grammica; P, serrata,- Q, a/foida; R, A/apus elongatus; S, Spartopyge iijexicana; T, .s/ossoriae; U, ritana; V, impt^fans; W, prainana; X re/Zexa- Y af/anfica, Z, areo/ata, AA, producta; BB, ffa/coto; CC, sanrfersi; DD, c/ayi, EE, graminea, ¥¥ decora; GG, inflata; HH, texaria, II, foeani^ri, JJ, modica, KK, yowngj, LL, arenicola; MM. ari^onenm; NN,/^cxu/osa; OO, sfy/ato; PP, celata . Some pygofers were redrawn from Young and Beirne (1958) and Lowry and Blocker (1987). 306 Great Basin Naturalist Memoirs No. 12 Fig. 63. Bases of first valvulae of Flexamia species. The depicted tree represents phylogeny presented in Figures 56-60. A, doeringae; B, gila; C, handarita; D, colloruni; E, mescalero; F, curvata; G, pectinata; H, canyonensis; I, surcuhi; J, pictoi K, zacate; L, Alapus elongatiis; M, abbreviata; N, Spartopyge mexicana- O, albida, P, serrata; Q, decora; R, modica; S, arenicola; T, inflata; U, slossonae; V, ritana; W, ijoungu X, celata; Y, arizonensis; Z, beameri; AA, stylata; hB, flexulosa; CC, texana ,DD, atlantica; EE, producta; FF, sandersi; GG, c/ai/i, HH, reflexa; II, imputans; JJ, prairiana, KK, dakota; LL, graminea. Interestingly, there are present-day clues to the feeding strategies of this primitive com- plex. We suspect that slossonae and alhida are habitat specialists, whereas serrata special- izes on Muhlenbergia richardsonis . Ancestral Flexamia species in Mexican grasslands may have been general grass feeders; this strategy has been partially retained by some lineages, including those that retained the plesiomor- phic striped habitus. Perhaps the striped habitus is optimal for habitat specialists. Loss of the habitus oc- curred homoplastically several times; in at least one case (serrata), it accompanied host specialization in northern, temperate grass- lands (on Muhlenbergia richardsonis). This event could actually have taken place far south of the current range oi serrata during a glacial maximum in the early Pleistocene. Muhlen- bergia richardsonis not only occurs as far north as Wood Buffalo National Park in Canada, but also occurs in Mexico, pre- sumably as a relict. Throughout much of the northern part of its range, this warm-season grass is an "island" surrounded by a sea of cool-season grasses. Such a condition is ideal for evolution of a specialist lineage from a more general ancestor; taxonomic distinctive- ness has been recognized as an important force in determining insect species richness on plant hosts (Lawton and Schroder 1977). In any event, morphological evidence indicates that a subtree o( Flexamia evolved on Muhlen- bergia. The first event (Fig. 66) appears to have been the divergence of the serrata and decora lineages. The decora subgroup is 1988 Whitcomb, Hicks: Flexamia Revision 307 Fig. 64. Geographic interpretation of Flexamia phylogeny. Early evolution is postulated to be Mexican. The Spartopyge lineage diverged into mexicana and miranda. From this lineage Flexamia diverged. The first major node involved the divergence of the pectinata group, perhaps as a result of specialization on Boutelotia curtipendula. This divergence is hypothesized to have occurred in the grasslands of northeastern Mexico, a region thought to have remained stable over recent geologic history (Rosen 1978). Subsequent divergences, which occurred in the same general region of Mexico, were those of the abbreviata-zacate and curvata-picta lineages. The abbreviata lineage adapted to Boutelona spp., the curvata group to Buchloe dactyloides, and the picta group to Aristida spp. The divergence of the albida lineage, characterized by a single, plesiomorphically ventral aedeagal process, may have occurred somewhat north of the Sierra Madre Orientale region, perhaps in Texan grasslands. Many conteinporary species emerged from this lineage. The most plesiomorphic of these (members of the albida group) are relatively uncommon today; albida and slossonae do not appear to be strict host specialists, a feeding habit that we hypothesize to be plesiomorphic for the genus. Actual divergences in temperate zones may have occurred farther south than diagrammed, if they occurred during periods of glacial advance. presumed to have divided by vicariance dur- ing glacial episodes into a Rocky Mountain- Canadian {decora) lineage, and a Great Basin lineage (yoiingi), both specialists on M. richardsonis . Alternatively, a second lineage developed, leading to a New Mexican mon- tane lineage (modica) that transferred from Muhlenhergia richardsonis to Muhlenbergia repens. Further divergence led to arenicola (host: Muhlenbergia pungens) and to a set of sister species (celata-stylata) that specialized on Muhlenhergia or Redfieldia (and possibly other glasses of sandy grasslands), respec- tively. Finally, this line appears to have trans- ferred from Muhlenbergia to the vast resource of Bouteloua gracilis in the plains region of New Mexico. This resource has been shared with abbreviata, which arrived in the New Mexican plains by an entirely different geo- graphic route (i.e., Mexican mixed Bouteloua 308 Great Basin Naturalist Memoirs No. 12 Fig. 65. Geographic interpretation of phylogeny of the pectmata group. A focal origin in grasslands of eastern Mexico is hypothesized, A western lineage comprised gila, handarita, and doeringae. This lineage speciated exclusively on Bouteloua curtipendtda. Morphological data suggest that mescalcro, which apparently transferred to Muhlenbergia pauciflora, belongs on a deep branch. The subtree consisting of za»iora, minima, and pectinata is closely related to the doeringae subtree; like all members of that subtree, pectinata specializes on B. ctitiipendiila (side-oats grama). F. coUorum may have been derived from a lineage on B. curiipendula, since its host is B. uniflora, which is closely related to side-oats grama. The host ofjacala is unknown. grasslands). (Interestingly, in contrast to Athysanella species [Hicks et al. 1988], both Flexamia blue grama specialists colonize this host throughout most of its range in New Mex- ico). A sister species, arizonensis, transferred to another dominant southwestern grass, Aristida purpurea. We emphasize that con- siderable climatic diversity exists today in the Southwest and no doubt has existed through- out the Pleistocene and Holocene as the result of elevational gradients (Axelrod and Raven 1985, Cole 1986, Cronquist 1978, Thompson and Mead 1982, VanDevender and Spaulding 1979). Therefore, evolutionary processes that might take place only over wide geographic distances in regions of relatively constant ele- vation may be focused in small geographic areas in the Southwest. Another lineage, possibly more closely al- lied to the ritana than serrata lineage, pre- sumably diverged in grasslands of the south- ern plains. This is the inflata subgroup of the flexulosa group. One contemporary represen- tative (inflata) exhibits one of the widest host ranges in the genus; largest populations occur on Muhlcnhcrgki aspcrifolia in wetland areas. If this apparent preference indicates an ances- tral Muhlenbergia host, it would be necessary to postulate a monumental subsequent widening of host range to pooid grasses or 1988 Whitcomb, Hicks: Flexamia Revision 309 Fig. 66. Geographic interpretation of phylogeny of the serrata, ritana, flexulosa, imptitans, and areolata groups. Texan grasslands may have been a focal area for initial divergence of lineages. F. serrata appears to be a Muhlenbergia specialist. The ritana group appears to have diverged in the Southwest. The contemporary distributions of the albida, serrata, and ritana groups suggest that they may have speciated by vicariant mechanisms or, in the case oi serrata, by a combination of vicariance and host transfer. An eastern lineage consists oiinflata and beameri. From a biogeographic perspective, it is most reasonable to propose an independent origin oitexana and beameri, each from inflate or a closely related lineage. The present-day distribution oiinflata suggests that it diverged in cool, temperate grasslands; Texas, of course, was cool during glacial maxima. In general, events postulated as northern in these reconstructions could have taken place much farther south during glaciations. The imputans-areolata lineage appears to have diverged in the prairie, on Muhlenbergia cuspidata ind Eragrostis spectabilis , respectively. The western lineage of the/7ex«/osa group may be related (see Fig. 59) to the serrata group; if this association can be confirmed, the derivation of the decora set of sister species from the serrata lineage would make an attractive hypothesis from a biogeographic standpoint, since all of the species (including youngi) specialize on Muhlenbergia richardsonis. Derivation oi modica (host: Muhlenbergia ripens) and arenicola (host; Muhlenbergia pungens) represents further evolution that occurred by host transfers within Muhlenbergia. The derivation of flexulosa and arizonensis by host transfer may have been a consequence of the regional abundance (in New Mexico and eastern Arizona) of their hosts, Bouteloua gracilis and Aristida purpurea, respectively; presumably, the region was colonized from the north. Finally, derivation of celata (host: Redfieldia flexuosa and perhaps other sandhill grasses) and stylata (hosts: Muhlenbergia spp.) is hypothesized to have followed a high-plains route around the eastern edge of the high Rocky Mountains. 310 Great Basin Naturalist Memoirs No. 12 Fig. 67. Geographic interpretation of the phylogeny of the prairiana group. Although present-day distributions and host preferences do not offer obvious hypotheses for speciation mechanisms, an origin for the prairiana group in Texan grasslands is hypothesized. From this region prairiana presumably moved northward in postglacial episodes, where it adapted to little bluestem (Schizachyrium scoparium ) or other andropogonoid grasses. In the tall-grass prairie, reflexa diverged from this lineage; this species often seems to be associated with big bluestem (Andropogon gerardii), particularly in glades of the eastern United States. F. atlantica may have transferred in postglacial times from chlorodoid grasses, on which it predominates in the Southwest, to switchgrass, Panicum virgattim. This host has been a conduit for colonization by atlantica of the eastern United States, including the Atlantic coastal prairies, where its host achieves high importance. Derivation oi sandersi may have occurred by transfer to Andropogon virginicus, the first colonist in early fire succession in the Southeast. From this lineage, presumably post-Wisconsinan, delongi may have diverged by colonization of little bluestem in the glaciated area northwest of Illinois. Specialization on Schizachyrium scopariuin and subsequent divergence led to dakota in the dry prairies of the plains, to graminea in Texas and northward through the tall-grass prairie, and to a lineage of eastern species. These species, also residents of andropogonoid grasses, are residents of the Gulf Coast (satilla), or the Appalachian mountains to Maine and cool, temperate savannas of Illinois and Ohio (clayi). Juncus. It may be more likely that the cur- rently observed host range breadth repre- sents simply a retention of plesiomorphic gen- eralism. In any event, this lineage has spawned two additional species: texana (south central Texas) and beameri (west slope of the Adirondack Mountains, New York). The cir- cumstances surrounding these divergences are unclear, since we have no data on their host relationships. After divergence ofinflata, another lineage diverged in the prairie, leading to imputans (probable host: Muhlenbergia cuspidata) and areolata (host: Erag^rostis spectabilis) on one hand, and to the prairiana group on the other. All speciation events of the prairiana group (Fig. 67) occurred either in prairie, savanna, or southeastern grasslands. We hypothesize 1988 Whitcomb, HickS: Flexamia Revision 311 an ancestor for this group that was adapted to both chloridoid and panicoid grasses. From such an ancestor, prairiana (liost: Schiza- chyrium or Bothriochloa) and reflexa (host: Andropogon gerardii) were derived, as were atlantica (hosts: chloridoid grasses and/or Panicum virgatum), and prodiicta (hosts chloridoid grasses, or [possibly] a habitat spe- cialist). Finally, specialization on Schiza- chyrium scoparium and/or Andropogon vir- ginicus led (Fig. 67) to lineages that adapted to the most xeric of Andropogon habitats (da- kota, a Schizachyriiim specialist of the west- ern Great Plains), graminea (a Schizachyriiim specialist of the tall-grass prairie), and the sa7idersi-delongi and satilla-clayi sister pairs of eastern andropogonoid grasslands. Stages of Speciation Mayr (1942) believed that at any given time all stages of evolution should be observable. We believe that this is indeed the case in the genus Flexamia. We recognize at least 10 lev- els in the process of speciation in Flexamia: 1. Nonisolated ephemeral populations: (i) Many populations of Flexamia inflata appear to be ephemeral. These populations often oc- cur on pooid grasses such as Poa or Festuca; presence of immature insects confirms ovipo- sition; (ii) F. abbreviata colonizes Miihlenber- gia richardsonis in the Sacramento Moun- tains of New Mexico and M. reverchonii in the Edwards Plateau of Texas, although in the latter case we have not confirmed oviposition. 2. Nonisolated stable populations: (i) F. in- flata consistently colonizes path rush (Juncus tenuis) in Illinois and Maryland; populations, including immatures, are found regularly on that host. This is an especially interesting case, since it may represent an interfamily transfer by a species that has limited oligophagy. (ii) Flexamia atlantica regularly colonizes large plantings of weeping love grass (Eragrostis spectahilis) and bermudagrass {Cynodon dactylon) in Oklahoma and Texas. (iii) Similarly, prodiicta colonizes bermuda- grass and Zoysia japonica in Maryland. These are essentially "natural experiments " that may mimic to a large degree, but in a shorter geo- logic time period, the effect of radical changes in the dominance hierarchy of grasses induced by cyclic climatic change. 3. Disjunct populations that have not diverged morphologically: In the following cases we have found no morphological varia- tion associated with disjunct populations, (i) A disjunct population oi dakota, a specialist on Schizachyriiim scoparium in the western Great Plains, occurs on that host in the Loess Hills of western Iowa, (ii) Flexamia reflexa occurs (always, it seems, in association with big bluestem, Andropogon gerardii) in iso- lated grassland islands in the eastern decidu- ous forest as far east as the Soldier's Delight serpentine barrens in Baltimore County, Maryland. 4. Isolated stable populations with minor morphological divergence: In each of the fol- lowing cases the populations may or may not represent biological species, (i) Two disjunct populations (Nebraska Sand Hills and Four Corners or Anasazi populations) of arenicola have been discovered; the unpaired, aedeagal processes of the Anasazi population always break, whereas the processes of the Sand Hills population rarely break. Other characters have not been found to separate these popula- tions, each of which specializes on Muhlen- bergia pungens. The host population is also separated into disjunct populations, (ii) The Mammoth Lakes population oiyoungi, prob- ably isolated in eastern California in a region rich in endemics, has diverged somewhat from the population in the eastern Great Basin, (iii) Populations of prairiana on Both- riochloa in the Southwest have begun to di- verge from the prairie Schizachyriiim popula- tions, perhaps as the result of phenological isolation. In this case divergence of female rather than male structures may signal the emergence of a new species. 5. Recent (Holocene) speciations: (i) The separation of delongi from the sandersi lin- eage may represent a postglacial transfer from Andropogon virginicus to Schizachyrium sco- parium in the glaciated region of the northern prairie, (ii) The divergence ofclayi and satilla may reflect phenological isolation between Gulf Coast and Appalachian populations. 6. Pleistocene speciations: Biological and biogeographic circumstances strongly suggest that decora and youngi represent vicariant populations of a common lineage that di- verged during Pleistocene climatic cycles, re- taining Miihlenbergia richardsonis. 7. Vicariance and semiisolation: In the case of Mexican and Southwestern grasslands, spe- ciation was perhaps achieved by vicariance 312 Great Basin Naturalist Memoirs No. 12 during long periods of semiisolation. It is pos- sible, in the case of certain species, that there may have been a punctuated infusion of genes into semiisolated populations as the climate became more mesic (Milstead 1960) and the distance between host stands of side-oats grama decreased. In any event the emergence of gila, doeringae, bandarita, zamora, min- ima, and pectinata, closely related species that appear to partion the range of side-oats grama, must have entailed different specia- tion mechanisms than those described above and are clearly enforced today by phenologi- cal isolation. 8. Distinct species with obvious sisters: We consider the sister pairs of picta-pyrops, ciir- vata-surcida, canyonensis-zaccite , and prairi- ana-reflexa to be obvious sisters that occur in ambiguous circumstances that suggest no par- ticular age for their divergence. In some cases (surcula-curvata [tropical/temperate] and za- cate-canyonensis [Chihuahuan/Sonoran]) the biogeographic circumstances for the diver- gences are clear. 9. Species with distant sisters: We consider the albida-slossonae sister pair to represent a historically remote divergence. Flexamia slos- sonae is a subtropical species that has many autapomorphies separating it from other Flex- amia species. However, common characters of habitus and male genitalia leave no doubt that it is a sister to the tall-grass prairie species albida. 10. Orphans: The characteristics of gram- mica are sufficiently unique that it must be considered an orphan within the genus. In the absence of a contemporary link, the relation- ships to other extant species of the genus are obscure. From the above examples we feel that our experience with Flexamia elegantly confirms Mayr's hypothesis. Conclusions The initial choice of Flexamia as a model for the study of the role of host plants in insect evolution was a fortuitous one. Whereas some speciation events in Flexamia must have oc- curred by host transfer, others surely oc- curred by vicariance. Further, there is a ten- dency of individuals representing advanced lineages to be reduced in size and complexity, and to be more highly specialized in their host choice than their ancestors. There is therefore an immense stake in arriving at a well-sub- stantiated phylogenetic hypothesis for the genus that is supported by independent lines of evidence. In particular, if advances in the study of molecular evolution prove to be adaptable to the study of small insects, we believe that Flexamia would provide an excel- lent model for study. Because the knowledge of Flexamia hosts is now adequate, obtaining live material of most species for such a study would be an achievable task. The rewards of further detailed studies would be many but would include, especially, important insights into the evolution of life-history strategies of sap-sucking insects. When we began our studies of Flexamia, whose species number had remained rela- tively constant for a number of years, we did not dream that there were many remaining species awaiting discovery. It is difficult for us to guess, given the results reported here, how far we have fallen short, even now, of a com- plete inventory. Our discovery of the role of climate in inducing and reinforcing cicadellid speciation makes it clear that the biogeo- graphically rich areas of the Southwest may well have many interesting stories that still await us. In particular there may well be Mexican species awaiting discovery. For example, our phylogenetic hypothesis suggests that there may have been at one time a species with the plesiomorphic dorsal stripes of Spartopyge, grammica, and albida, but which possessed a full complement of four aedeagal processes. Because of the compelling logic of all phyloge- netic schemes for its existence, such a species seems so real to us that we have jokingly re- ferred to it as ancestris. Is it possible that this species, or a closely related descendant of it, may actually exist in an unexplored Mexi- can grassland? It is questions such as these that have driven us to return repeatedly to the grasslands of the Southwest, which prove to be every bit as full of enigmas as they are of answers. Acknowledgments There is a Japanese proverb that advises those who drink water from a well today to remember the ancestors who dug the well. The Flexamia well has been dug with many 1988 WniTcoMB, HiCKS; Flexamia Revision 313 spades. The early biological work of E. D. Ball, H. Osborn, D. M. DeLong, and R. H. Beamer is implicit in our work. H. H. Ross introduced us to the genus, framed many of the questions that we discuss, and left much unpublished material from his grasslands (GL) project. D. A. Young, whose 1958 mono- graph with B. P. Beirne continues to be an important tool for students of Flexamia, kindly permitted us to redraw many struc- tures from drawings in his monograph. We are particularly pleased to acknowledge the ad- vice and encouragement of P. W. Oman, given especially to the senior author during his early years at Beltsville. Further, a large fraction of USNM Flexamia had been col- lected and correctly determined by Oman; this material was extremely valuable in our study. Earlier studies on Flexamia at Beltsville were assisted by M. E. Coan, J. P. Kramer, and, in later years, J. E. Lowry, whose illustration oLserrata is presented here as Figure 1. All other drawings were done by Debi Roney. Robert Almond and Edward Clark gave advice and assistance on the graph- ics. Host records were retrieved from a com- puterized index developed by D. E. Lynn. T. Mogel assisted with the scanning electron microscopy. Assistance with phylogenetic programs was provided by Mary Mickevich. We acknowledge further the assistance of G. W. Byers, P. S. Cwikla and C. P. Triple- horn, M. W. Nielson, and T. J. Henry and R. C. Froeschner, who made available the Flexamia collections of Kansas University, Ohio State University, Monte Bean Life Sci- ence Museum (Brigham Young University), and the United States National Museum, re- spectively. K. G. A. Hamilton made available the Flexamia material from the Canadian Na- tional Collection and provided a wealth of stimulating comments on the manuscript. M. SchaufF made many useful comments on the manuscript. So many friends have helped in our field studies that all cannot be acknowl- edged. We are especially indebted, however, to Kelly Allred of New Mexico State Univer- sity for continuing lessons on New Mexico agrostology. Charles T. Mason, Jr., assisted with grasses of Arizona. R. Chambers of the U.S. Soil Conservation Service drove us to the type locality of collortwi and called our attention to the host. Mike Fleming of Big Bend National Park and Jim Walters of Guadalupe National Park assisted field stud- ies there, and Barton Warnock identified many plant specimens from Big Bend Na- tional Park and the Davis Mountains. Riley Nelson of BYU provided collections from the Colorado Plateau. P. Cubbage assisted with fieldwork. Judith Leach, wife of the senior author, assisted during many field trips and also made valuable editorial comments on the manuscript. The editorial staff of The Great Basin Naturalist helped in many unusual ways to assure timely completion of the paper. Finally, H. D. Blocker suggested that the time for this paper had come, sorted more than 1,000 unidentified Flexamia from the KU cicadellid collection, provided Flexamia ma- terial and records from the KSU collection, read various drafts of the manuscript, and en- couraged us in countless other ways. To all these workers, and many that remain un- named, we offer our sincerest gratitude. Literature Cited AXELROD. D I , AND P H Raven 1985. Origins of the Cordelleran flora. J. Biogeogr. 12: 21-47. Ball, E D 1899. Some new species of Deltocephalus. Canadian Entomol. 31: 188-192. 1900. Some new Jassidae from the Southwest. Canadian Entomol. 32: 200-205. 1905. Some new Homoptera from the South and Southwest. Biol. Soc. Washington, Proc. 18: 117-120. Beamer, R H 1936. New leafhoppers from the western United States (Homoptera-Cicadellidae). Cana- dian Entomol. 68: 252-257. Beamer, R. H , and L D Tuthill 1934. Some new spe- cies and a new genus of deltocephaloid leafliop- pers (Homoptera, Cicadellidae). J. 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Specimens examined\ Species Holotype Paratype(s) barc CNC KU OSU USNM minima OSU(h) none 0 0 0 1 8 zamora OSU(h) OSU(p) 0 0 0 9 0 pectinata USNM(c) USNM(c) 57 5 150 45 75 gila USNM(h) P^ 9 0 0 0 0 handarita USNM(h) P^ 5 0 0 6 0 doeringae KU(h) USNM(p) 35 0 68 24 91 collorum USNM(h) P^ 81 0 0 0 0 jacala USNM(h) none 0 0 0 0 1 mescalero USNM(h) P^ 26 0 0 0 0 abbreviata USNM(c) USNM(c) 765 141 661 74 409 zacate USNM(h) P^ 96 0 0 0 0 canyonensis USNM(h) USNM(p) 9 0 0 0 2 surcula osu OSU 72 0 10 57 37 curvata osu osu 560 81 209 77 107 grammica USNM(l) KU(p) 47 64 164 48 109 picta OSU(h) USNM(c) 285 107 202 135 112 pyrops KU(1) KU(c) 56 39 56 83 162 albida USNM(c) USNM(c) 46 1 7 17 19 slossonae USNM(h) none 12 0 50 0 18 serrata KU(h) KU(p) 56 98 20 0 14 ritana KU(h) USNM(p) 0 0 17 0 5 arizonensis USNM USNM(p) 177 11 114 20 94 flexulosa USNM(l) USNM(c) 380 211 254 51 353 arenicola USNM(h) USNM(p) 91 0 0 0 5 celata USNM(h) USNM(p) 49 0 4 0 5 stylata USNM(l) USNM(c) 83 67 66 9 70 decora KU USNM(p) 95 246 177 3 28 youngi USNM P^ 30 0 5 0 0 modica KU USNM(p) 377 0 110 3 8 inflata USNM(c) USNM(c) 665 103 257 146 155 beameri KU USNM(p) 0 0 7 0 0 texana USNM USNM(p) 0 0 0 0 10 imputans OSU(c) OSU(c) 4 0 10 11 19 areolata USNM(l) USNM(c) 37 51 42 51 88 prairiana OSU OSU 612 44 193 23 172 reflexa USNM USNM(c) 25 1 1 8 26 atlantica OSU OSU(p) 378 17 8 12 44 dakota USNM(h) USNM(p) 209 3 1 0 2 producta (BNM) USNM(c') 4 5 136 117 267 sandersi OSU(h) OSU(h^) 45 3 60 31 126 delongi (INHS) none 0 26 1 39 13 graminea OSU(h) none 176 28 105 10 190 clayi USNM(h) USNM(p) 148 9 115 47 3 satilla CNC(h) CNC(p) 0 2 2 0 0 Numbers reflect content of collection before redistribution at end of study. Type distribution occurred after this enumeration (c ~ cotype. h holotype; 1 ~ lectotype; p - paratype). Parentheses - specimen not examined. See text for distribution of paratypes. Cotype o( vise nda(' producta). Hohtype o( hidentata (~ sandersi). 1988 Whitcomb, Hicks: F'lexamia Revision Appendix II Part I. Character matrix for PHYSYS. 317 DISTIN SIMPLE ABSENT? 7 SINUATC THICK 1 ELONGA SHORTL TAPER ABSNT FLAREDG APODNOX APOPRIM 2 ELONGA CONNAR FORCEPY PRIMPL DEEPST ABSTR BRFACE 3 ELONGA DISTIN SIMPLE ABSENTP 7 LEFTC THICK 1 MEXICAN SHORTL TAPER ABSNT APICALG APODEXP APOPRIM 2 MEXICAN CONNAR PRIMPY PRIMPL PARALST PRESTR CONBLF 3 MEXICAN DISTIN SIMPLE ABSENTP 7 LEFTC THIN 1 MIRAND SHORTL TAPER ABSNT APICALG APODEXP APOPRIM 2 MI RAND CONNAR CNSTRPY PRIMPL EXCVST PRESTR CONBLF 3 MIRAND DISTIN FLARED ABSENTP 7 LEFTC THIN 1 MINIMA SHORTL TAPER ABSNT FLAREDG APODEXP APOPARL 2 MINIMA CONNAR BULBPY ACUTPL TOOTHST ABSTR BRFACE 3 MINIMA DISTIN FLARED ABSENTP 7 LEFTC THIN 1 ZAMORA SHORTL TAPER ABSNT FLAREDG APODEXP APOPARL 2 ZAMORA CONNAR BULBPY LNGBRPL OUTRST ABSTR BRFACE 3 ZAMORA DISTIN FLARED ABSENTP 7 LEFTC THIN 1 PECTIN SHORTL TAPER ABSNT FLAREDG APODEXP APOPARL 2 PECTIN CONNAR BULBPY RECTPL TOOTHST ABSTR BRFACE 3 PECTIN DISTIN FLARED ABSENTP 7 LEFTC THIN 1 BANDAR SHORTL TAPER ABSNT FLAREDG APODEXP APORECU 2 BANDAR CONNAR BULBPY LNGBRPL INNRST ABSTR BRFACE 3 BANDAR DISTIN FLARED ABSENTP 7 LEFTC THIN 1 GILA SHORTL TAPER ABSNT FLAREDG APODEXP APORECU 2 GILA CONNAR BULBPY LNGBRPL INNRST ABSTR BRFACE 3 GILA DISTIN FLARED ABSENTP 7 LEFTC THIN 1 DOERIN SHORTL TAPER ABSNT FLAREDG APODEXP APOPARL 2 DOERIN CONNAR BULBPY RECTPL PROCST ABSTR CONBLF 3 DOERIN DISTIN FLARED ABSENTP 7 LEFTC THIN 1 COLLOR SHORTL TAPER ABSNT FLAREDG APODEXP APORECU 2 COLLOR CONNAR BUDPY SHORTPL TOOTHST ABSTR BRFACE 3 COLLOR DISTIN FLARED ABSENTP 7 LEFTC THIN 1 JACALA SHORTL TAPER ABSNT FLAREDG APODNOX APORECU 2 JACALA CONNAR BULBPY LNGBRPL 7 ABSTR BRFACE 3 JACALA DISTIN CAPITAT ABSENTP 7 LEFTC THIN 1 MESCAL SHORTL TAPER ABSNT FLAREDG APODNOX APOCLOS 2 MESCAL CONNAR BULBPY LNGROPL DEEPST ABSTR BRFACE 3 MESCAL DISTIN SHORTPR ABSENTP 7 STRAIT LESTHIC 1 ABBREV LONGERL UNTAPER ABSNT SLITG APODEXP APOPARL 2 ABBREV CONNAR PRODPY LOBATPL INCISST ABSTR BRFACE 3 ABBREV DISTIN FLARED ABSENTP 7 LEFTC THICK 1 CANYON SHORTL TAPER ABSNT MIDLENG APODNOX APORECU 2 CANYON CONNAR PRODPY LOBATPL LONGST ABSTR BRFACE 3 CANYON DISTIN FLARED ABSENTP 7 LEFTC THIN 1 ZACATE SHORTL TAPER ABSNT SLITG APODNOX APORECU 2 ZACATE CONNAR PRODPY LOBATPL PROJST ABSTR BRFACE 3 ZACATE DISTIN ASYMPR ABSENTP 7 LEFTC THICK 1 GRAMMI SHORTL TAPER SERRATE BIPROVG APODNOX APOWING 2 GRAMMI CONNAR NARRPY SQUATPL ROUNDST PRESTR CONBLF 3 GRAMMI DISTIN LONGPR SUBAPP 7 STRAIT LESTHIC 1 SURCUL LONGERL UNTAPER ENTIRE BIPROVG APODNOX APOFORK 2 SURCUL CONWIDE SHORTPY SHORTPL BOXST ABSTR BRFACE 3 SURCUL DISTIN LONGPR DISTALP 7 STRAIT LESTHIC 1 CURVAT LONGERL UNTAPER ENTIRE BIPRSLG APODEXP APOFORK 2 CURVAT CONNAR SHORTPY SHORTPL WIDINST ABSTR BRFACE 3 CURVAT INDIST FLANGED SUBAPP 7 STRAIT THIN 1 PICTA LONGSTL , UNTAPER SERRATE SUBAPIG APODNOX APOFORK 2 PICT A CONNAR ANGLPY TAPERPL SERRST ABSTR CONBLF 3 PICTA INDIST FLANGED APPOSP ? SINUATC THIN 1 PYROPS LONGSTL , UNTAPER SERRATE SUBAPIG APODNOX APOFORK 2 PYROPS CONNAR ANGLPY REDUCPL EXTHST ABSTR TOOTHF 3 PYROPS 318 Great Basin Naturalist Memoirs Appendix II. Part I continued. No. 12 DISTIN BIFURC FUSEDP SAGGITV LESSC THICK 1 ALB I DA SHORTL TAPER SERRATE SUBVNPG APODEXP APOCLUB 2 ALBIDA CONWIDE WIDEPY TAPERPL TTHEXST PRESTR CONBLF 3 ALBIDA PARTLY BIFURC FUSEDP SAGGITV RIGHTC THICK 1 SLOSSO SHORTL UNTAPER ENTIRE SYMSLTG APODNOX APOWING 2 SLOSSO CONPAFU UNDIFPY ACUTPL TRILBST PRESTR CONBLF 3 SLOSSO DISTIN BIFURC FUSEDP SAGGITV STRAIT THICK 1 SERRAT SHORTL UNTAPER ENTIRE SUBVNPG APODEXP APOCLUB 2 SERRAT CONWIDE IRREGPY CLEAVPL LOBEDST ABSTR CONBLF 3 SERRAT DISTIN LYRIFO FUSEDP SAGGITV LESSC THICK 1 R I TANA SHORTL UNTAPER ENTIRE SUBVNPG APODNOX APOCLAW 2 RITANA CONNAR OVATEPY TAPERPL ANGLST REDUSTR CONBLF 3 RITANA DISTIN SYMMET SHORTV SAGGITV LESSC LESTHIC 1 ARENIC LONGERL UNTAPER ENTIRE SUBAPIG APODNOX APOAWAY 2 ARENIC CONNAR ROUNDPY INCISPL NOTCHST ABSTR CONBLF 3 ARENIC DISTIN SYMMET SHORTV SAGGITV STRAIT LESTHIC 1 CELATA LONGERL UNTAPER ENTIRE SUBAPIG APODNOX APOPARL 2 CELATA CONWIDE THSTYPY INCISPL NOTCHST ABSTR CONBLF 3 CELATA DISTIN ASYMME TWISTV DISPLCV RIGHTC LESTHIC 1 STYLAT LONGERL UNTAPER ENTIRE SUBAPIG APODNOX APOTHIC 2 STYLAT CONBROD STYLPY SQUARPL BINOTST ABSTR CONBLF 3 STYLAT DISTIN SYMMET SHORTV SAGGITV STRAIT LESTHIC 1 ARIZON LONGERL UNTAPER ENTIRE SUBAPIG APODNOX APOFORK 2 ARIZON CONNAR VNPRPY MINUTPL SQUARST ABSTR CONBLF 3 ARIZON DISTIN SYMMET SHORTV SAGGITV LEFTC LESTHIC 1 FLEXUL LONGERL UNTAPER SERRATE SUBAPIG APODNOX APOFORK 2 FLEXUL CONNAR CAUDPY MINUTPL MEDNST ABSTR CONBLF 3 FLEXUL DISTIN SYMMET FUSEDP SAGGITV LEFTC THICK 1 DECORA SHORTL TAPER SERRATE APPRG APODNOX APOAWAY 2 DECORA CONBROD LOBEDPY INCISPL BINOTST ABSTR CONBLF 3 DECORA DISTIN ASYMME RECURVP DISPLCV LESSC LESTHIC 1 YOUNG I LONGERL TAPER SERRATE APPRG APODNOX APOTHIC 2 YOUNG I CONBROD BROADPY INCISPL BINOTST ABSTR CONBLF 3 YOUNGI DISTIN ASYMME DECURVP DISPLCV LEFTC THICK 1 MODICA SHORTL TAPER SERRATE SUBAPIG APODNOX APOAWAY 2 MODICA CONBROD OVATEPY DEEPL NOTCHST ABSTR CONBLF 3 MODICA DISTIN ASYMME DECURVP DISPLCV RIGHTC THIN 1 INFLAT LONGSTL UNTAPER ENTIRE SUBAPIG APODNOX APOFORK 2 INFLAT CONWIDE BOATPY TAPERPL BINOTST ABSTR CONBRF 3 INFLAT DISTIN TWISTED RECURVP DISPLCV RIGHTC THIN 1 BEAMER LONGSTL UNTAPER ENTIRE APPRG APODNOX APOTHIC 2 BEAMER CONBROD SHIPPY SHORTPL BINOTST ABSTR CONBRF 3 BEAMER DISTIN TWISTED TWISTV DISPLCV STRAIT THIN 1 TEXANA LONGSTL UNTAPER ENTIRE APPRG APODNOX APOREDU 2 TEXANA CONWIDE SHIPPY SHORTPL BINOTST ABSTR CONBRF 3 TEXANA DISTIN TWISTED ? LATERLV STRAIT THIN 1 IMPUTA LONGSTL UNTAPER SERRATE LATSLTG APODNOX APOHOOK 2 IMPUTA CONWIDE LGOVPY TAPERPL LOBEDST ABSTR BLFACE 3 IMPUTA FUSED TWISTED ? LATERLV LEFTC LESTHIC 1 AREOLA LONGERL UNTAPER SERRATE LATSLTG APODNOX APOAWAY 2 AREOLA CONWIDE BIZARPY TAPERPL PROCST ABSTR BLFACE 3 AREOLA BARELY INVERT 7 DORSALV STRAIT LESTHIC 1 PRAIRI LONGERL UNTAPER SERRATE DORSALG APODNOX APOPEN 2 PRAIRI CONFUSE ELLIPY NARRPL NAROWST ABSTR VARCONF 3 PRAIRI FUSED INVERT 7 BASFUSV STRAIT LESTHIC 1 REFLEX LONGERL UNTAPER SERRATE PROFUSG APODNOX APOCLOS 2 REFLEX CONFUSE UPTNPY NARRPL NAROWST ABSTR VARCONF 3 REFLEX FUSED WINDSW 7 FUSEDV STRAIT LESTHIC 1 ATLANT LONGERL UNTAPER SERRATE LATG APODNOX APOWIDE 2 ATLANT CONFUSE THINPY NOTCHPL NAROWST ABSTR VARCONF 3 ATLANT 1988 Whitcomb, HiCKS; Flexamia Revision Appendix II. Part I continued. 319 FUSED INVTWO 7 TINYV STRAIT THICK 1 PRODUC FUSEDL UNTAPER SERRATE FUSEDG APODNOX APOPARL 2 PRODUC CONFUSE VNTLBPY BOTTLPL NAROWST ABSTR VARCONF 3 PRODUC FUSED INVTWO ? FUSEDV STRAIT THICK 1 DAKOTA FUSEDL UNTAPER SERRATE DORFUSG APODNOX APOCLOS 2 DAKOTA CONFUSE POSTPY BOTTLPL NAROWST ABSTR BRFACE 3 DAKOTA FUSED FUSPR 7 REDUCV STRAIT THICK 1 SANDER FUSEDL UNTAPER SERRATE APPRSDG APODNOX APOWIDE 2 SANDER CONFUSE HIGHPY NOTCHPL NAROWST ABSTR BRFACE 3 SANDER FUSED FUSPR ? REDUCV STRAIT THICK 1 DELONG FUSEDL UNTAPER SERRATE APPRSDG APODNOX APOWIDE 2 DELONG CONFUSE HIGHPY NOTCHPL NAROWST ABSTR BRFACE 3 DELONG FUSED ASKEW ? FUSEDV STRAIT THICK 1 GRAMIN FUSEDL SPIRALT SERRATE SPIRALG APODNOX APOPARL 2 GRAMIN CONFUSE THICKPY RECURPL NAROWST ABSTR BRFACE 3 GRAMIN FUSED ASKEW ? FUSEDV LEFTC THICK 1 CLAY I FUSEDL SPIRALT SERRATE WIDSPIG APODNOX APOCLOS 2 CLAYI CONFUSE UPLOBPY RECURPL NAROWST ABSTR BRFACE 3 CLAYI FUSED ASKEW ? FUSEDV LEFTC THICK 1 SATILL FUSEDL SPIRALT SERRATE WIDSPIG APODNOX APOCLOS 2 SATILL CONFUSE UPLOBPY RECURPL NAROWST ABSTR BRFACE 3 SATILL Appendix II. Part II. Hypothesized character transformation in Flexamia. Data for PHYSYS. aedconn;distin-indist-partly-barely-fused; aedapex;simple-flared-capitat,shortpr-[l6ngpr-flanged, bifurc, lyrifo, symmet-asymme-twisted- [ invert- windsw, [fuspr- askew] , invtwo] ] ; ANTEPR;ABSENTP-SUBAPP-DISTALP-APP0SP-[FUSEDP-DECURVP, RECURVP, [LONGV-SHORTV-TWISTV] ] ; VENTRALP;SAGGITV-DISPLCV-LATERLV-D0RSALV-BASFUSV-REDUCV TINYV-FUSEDV; AEDCURV;LEFTC-LESSC-STRAIT-SINUATC-RIGHTC; AEDTHIC ; THICK-LESTHIC-THIN ; AEDLENG ; SHORTL-LONGERL-LONGSTL-FUSEDL ; AEDTAPER; TAPER-UNTAPER- SPIRALT ; AEDPRMG; ENTIRE-SERRATE; GONOLOC;APICALG-FLARED-[SLITG-MIDLENG] , [BIPROVG- [ SUBVNPG- SYMSLTG] ,BIPRSLG, [ SUBAPIG-APPRG, LATSLTG-DORSALG- [PROFUSG- [APPRSDG-SPIRALG-WIDSPIG] , [FUSEDG-DORFUSG] , LATG] ] ] ; APOAPEX ; APODEXP- APODNOX ; APOARM;APOPRIM- [APOPARL- [APOCLAW-APOHOOK- APOPEN-APOCLOS- APOWIDE] , [APOAWAY-APOTHIC-APOREDU] ,APOFORK, [APOCLUB- APOWING ] , APORECU ] ; CONKEEL;CONNAR-CONWIDE-CONBROD-CONPAFU-CONFUSE; PYGMAL ; PRIMPY-FORCEPY- [ CNSTRPY- [ NARRPY- IRREGPY-WIDEPY- UNDIFPY], [BULBPY-BUDPY,PRODPY, [ SHORTPY-ANGLPY, [OVATEPY- [LOBEDPY-[BOATPY-SHIPPY] ,BROADPY] , [ROUNDPY- [ VNPRPY-CAUDPY] , [STYLPY-THSTPY] ] , [ LGOVPY-BIZARPY, [ELLIPY- VNTLBPY, [UPTNPY- POSTPY,THINPY, [HIGHPY-UPLOBPY-THICKPY] ]]]]]]]; PLATES ; PRIMPL- [ LNGBRPL-RECTPL , SHORTPI^ ] , SQUATPL , [ LNGROPL- LOBATPL-REDUCPL] , [TAPERPL- [NARRPL-BOTTLPL-NOTCHPL-RECURPL] , ACUTPL, [CLEAVPL-[INCISPL-SQUARPL, [DEEPL-MINUTPL] ] ] ] ; STERNUM; PAEIALST-EXCVST-ROUNDST, [TOOTHST- [OUTRST-DEEPST] , [INNRST-PROCST] , [ INCISST-LONGST-PROJST] , [BOXST-WIDINST, [SERRST-EXTHST] ] ] , [TTHEXST-TRILBST, LOBEDST- [ ANGLST- [NOTCHST-BINOTST] , [ PROCST-NAROWST ] , [ SQUARST-MEDNST ] ] ] ; DORSVIT; PEIESTR-REDUSTR- ABSTR; FACE;CONBLF-TOOTHF, [CONBRF-BRFACE] , [ VARCONF -BLF ACE] ; 1 AEDCONN 1 AEDAPEX 2 AEDAPEX 3 AEDAPEX 1 ANTEPRO 2 ANTEPRO 1 VENTRLP 2 VENTRLP 1 AEDCURV 1 AEDTHIC 1 AEDLENG 1 AEDTAPER 1 AEDPRMG 1 GONOLOC 2 GONOLOC 3 GONOLOC 1 APOAPEX 1 APOARM 2 APOARM 3 APOARM 1 CONKEEL 1 PYGMAL 2 PYGMAL 3 PYGMAL 4 PYGMAL 5 PYGMAL 1 PLATES 2 PLATES 3 PLATES 1 STERNUM 2 STERNUM 3 STERNUM 4 STERNUM 1 DORSVIT 1 FACE 320 Great Basin Naturalist Memoirs No. 12 Appendix II. Part III Character Codes of Flexomia Species: Code Identification* P = plesiomorphic; A ^ apomorphic 1. AEDCONN: Articulation (P) or fusion (A) of aedeagus and connective: DISTIN (P) (Figs. 6A,B,C,E); INDIST (Fig. 6D); BARE- LY (Fig. 6F); PARTLY (Y&B Fig. 9B) FUSED (Fig. 60). 2. AEDAPEX: Morphology of aedeagal apex which may be simple, without processes or special structures (P), or with paired pro- cesses (A): SIMPLE (P) (Figs. 55C,F); FLARED (Figs. 12A-H); CAPITAT (Figs. 121,22); SHORTPR (Fig. 7H); ASYMPR (Fig. 7E); LONGPR (Figs. 7F,G); FLANGED (Figs. 7K,L); BIFURC (Figs. 7A-C); LYRIFO (Fig. 7D); SYMMET (Figs. 31G,K); ASYMME (Figs. 31E,H,M); TWISTED (Fig. 31A); INVERT (Figs. 44D,G); WINDSW (Fig. 44A); INVTWO (Fig. 441); FUSPR (Fig. 44B); ASKEW (Figs. 44F,H). 3. ANTEPR: Anteapical processes, which are absent (P), present as paired processes, or fused (A) into a single process: ABSENTP (Figs. 55C,F); SUBAPP (Fig. 7G); DISTALP (Fig. 7F); APPOSP (Fig. 7L); FUSEDP (Figs. 7A-D); LONGV (Fig. 31G); RECURVP (Fig. 38A); DECURVP (Fig. 31E); SHORTY (Figs. 31I-L); TWISTY (Fig. 31A). 4. YENTRLP: Phyletic twisting of the un- paired ventral process (see 3. ANTEPR), which was (P) symmetrically situated on the ventral surface of the aedeagus, but moved to the dorsum, and was (A) ultimately lost (Fig. 61), presumably by fusion: SAGGITY (Fig. 61A); DISPLCY (Fig. 61B); LATERLY (Fig. 61C); DORSALY(Fig. 61D); BASFUSY (Fig. 61E); REDUCY (Fig. 61F); TINYY (Fig. 61G); FUSEDY(Fig. 61H). 5. AEDCURY: Curvature (lateral aspect) of the aedeagus, which may be curved sinistrad (P), without curvature, sinuate, or curved dextrad (A): LEFTC (Figs. 71, J); LESSC (Y&B Figs. 10B,11A); STRAIT (Fig. 7H); SINUATC (Y&B Fig. 2B); RIGHTC (Fig. 4()A). 6. AEDTHIC: Thickness (lateral aspect) of the aedeagus, which may be thick (P), less thick, or thin (A); THICK (Fig. 7J); LES- ♦Characters are illustrated herein, in Vonnj; and Beirne (1958) (V&B); Lowry and Blocker (1987) (L&B); or Hamilton and Koss (1975) (H&R). THICK (Fig. 38A); THIN (Figs. 16A, 17A, ISA). 7. AEDLENG: Length of the aedeagus, which may be short (P), longer, even longer, or, finally (A), fused with the connec- tive: SHORTL (Figs. 16A,7I,J); LONGERL (Fig. 6B); LONGSTL (Fig. 6D); FUSEDL (Fig. 6G). 8. AEDTAPR: Form of the aedeagal shaft which may be tapered (P), untapered, or, ulti- matelv (A), tapered in association with a spiral gonopore: TAPER (Figs. 7I,J); UNTAPER (Fig. 7H); SPIRALT(Fig. 6G). 9. AEDPRMG: Form of the margins of the paired, apical, aedeagal processes which, if present, may have entire (P) or serrate mar- gins (A): ABSNT (absent); ENTIRE (Figs. 7F,G); SERRATE (Figs. 44B-D). 10. GONOLOC: Form and position of the gonopore. In the plesiomorphic condition, this structure was apical and subcircular or ovate. In early stages in transformation the gonopore was subapical on the ventral sur- face. In the flexulosa group the gonopore may have reversed its distad movement from the aedeagal apex and returned to the apex, or nearly so; but in imputans and areolata the gonopore became associated with the un- paired process, which had moved to a lateral position on the aedeagal shaft. In prairiana the gradual twisting (see 4. YENTRLP) of the processes had stabilized with the gonopore in a dorsal, anteapical position. Transformation from this condition involved association with the unpaired process and movement with the process as it became appressed to the aedea- gal shaft and, eventually, fused with the shaft. Ultimately, the fusion led to an elongate structure that winds around the shaft in a helical configuration: APICALG (Y&B Fig. 32B); FLAREDG (Figs. 12A-H); SLITG (Fig. 7H); MIDLENG (Fig. 6C); BIPROYG (Fig. 7G); SUBYNPG (Fig. 7B); SYMSLTG (Fig. 7C); BIPRSLG (Fig. 7F); SUBAPIC (Figs. 31E,J,M); APPRO (Fig. 31A); LAT SLTO (Fig. 610) DORSALO (Fig. 440) PROFUSO (Fig. 44D); APPRSDO (Fig. 44B) SPIRALG (Fig. 44F); WIDSPIG (Figs 44H,J); FUSEDO (Fig. 44E); DORFUSC (Fig. 441); LATO (Fig. 44A). 11. APOAPEX: Apices of the apodema arms mav be expanded (P) or not expandet (A): APODEXP (Fig. 6B); APODNOX (Figs 6A,D,E). 1988 WHITCOMB, HICKS: Fl.EXAMIA RH\ ISION 321 12. APOARM: In dorsal aspect the apode- mal arms in the plesioniorphic condition were parallel to the aedeagal shaft. Transformations from this state (A) involved several lines of development, some of which involved reduc- tion of the arms, or direction of the apices toward or awav from the sagittal plane: APO- PRIM (Y&B Fig. 32B); APOPARL (Fig. 6B); APOCLAW (Y&B Fig. UB); APOHOOK (Y&B Fig. 21B); APOPEN (Fig. 6F); APOC- LOS (Y&B Fig. 28C); APOWIDE (R&C Fig. 2); APOAWAY (Y&B Fig. 14B); APOTHIC (Fig. 40B); APOREDU (Y&B Fig. 16B); APO- FORK (Figs. 6A,D,E); APOCLUB (Fig. 8B); APOWING (Y&B Fig. 27B); APORECU (Figs. 6C, 16B, 17B, 19B, 20B). 13. CONKEEL: In lateral aspect the dorsal keels of the connective may be narrow (P), widened, or very much widened. Alterna- tively (A), the connective fused with the aedeagus, a process that resulted in the loss of an independent identity for the keels: CON- NAR (Figs. 16A, 17A, 18A); CONWIDE (Y&B Fig. 12A); CONBROD (Fig. 40A); CONPAFU (Y&B Fig. 9B); CONFUSE (Fig. 6G). 14. PYGMAL: The male pygofer, ple- siomorphically, was apparently undifferenti- ated. From this state a defined posterior lobe differentiated, as in Spartopyge miranda. Several lines of development (A) are postu- lated (Fig. 62): PRIMPY (Fig. 62S); FOR- CEPY (Fig. 55G); UNDIFPY (Fig. 62T); CN- STRPY (Fig. 62N); BULBPY (Figs. 62C-G) BUDPY (Fig. 62A); PRODPY (Figs. 62H,I) NARRPY (Fig. 620); IRREGPY (Fig. 62P) SHORTPY (Figs. 62K,L); ANGLPY (Fig 62M); WIDEPY (Fig. 62Q); OVATEPY (Fig 62U); ROUNDPY (Fig. 62LL); STYLPY (Fig 6200); THSTYPY (Fig. 62PP); LOBEDPY (Fig. 62FF); BROADPY (Fig. 62KK); BOATPY (Fig. 62GG); SHIPPY (Figs. 62HH,II); LGOVPY (Fig. 62V); BIZARPY (Fig. 62Z); ELLIPY (Fig. 62W); VNTLBPY (Fig. 62AA); UPTNPY (Fig. 62X); POSTPY (Fig. 62BB) THINPY (Fig. 62Y); HIGHPY (Fig. 62CC); UPLOBPY (Fig. 62DD); THICKPY (Fig. 62EE) CAUDPY (Fig. 62NN); VNPRPY (Fig. 62MM). . 15. PLATES: The male plates, plesiomor- phically, were apparently long and relatively wide. The broad aspect of the plates was re- tained in the pectinata group, but in subse- quent lineages tended to be modified to be- come tapered. In the flexulosa group the ta- pered plates became notched to accommo- date dorsally produced pygofers; in arizonen- si.s and flexulosa this tendency is maximized. In tlu> praihana group the tapered design was modified by narrowing of the apical third of the plates, by the development of small apical notches, and bv the plates ultimately becom- ing recurved: PRIMPL (Fig. 54C); SQUATPL (Fig. 8F); LNGROPL (Fig. Ill); LNGBRPL (Fig. UB); RECTPL (Figs. 11C,F); SHORTPL (Fig. IIG); LOBATPL (Fig. 8E); REDUCPL (Fig. 9B); TAPERPL (Fig. 8A); NARRPL (Fig. 45A); BOTTLPL (Fig. 45E); NOTCHPL (Fig. 45C); RECURPL (Fig. 45H); ACUTPL (Fig. 8B); CLEAVPL (Fig. 8C); MINUTPL (Figs. 33A,B); INGISPL (Figs. 33C,D,E); SQUARPL (Fig. 34E); DEEPL(Fig. 33F). 16. STERNUM: The sternum VII was ap- parently plesiomorphically excavated. From this condition several lines of development (A) ensued. Many of these lines involved two pairs of teeth separated by an incision on the posterior margin of the sternum. In the pecti- nata group the development of the inner or outer sets of teeth relative to one another is an important character in species discrimination. In the flexulosa and prairiana groups, at a time that aedeagal evolution was proceeding rapidly, the sternum VII became a relatively stable apomorphv that underwent minimal transformation: PARALST (Y&B Fig. 32E); EXCVST (Y&B Fig. 33F); ROUNDST (Fig. lOF); TOOTHST (Figs. 14A,C,G); OUTRST (Fig. 14B); DEEPST (Fig. 14H); INNRST (Figs. 14D,E);PROCST(Fig. 14F); INCISST (Fig. lOE); LONGST (Fig. 101); PROJST (Fig. 10L);BOXST(Fig. 10J);WIDINST(Fig. lOK); SERRST (Fig. lOG); EXTHST (Fig. lOH); TTHEXST (Fig. lOA); TRILBST (Fig. lOB); LOBEDST (Fig. IOC); ANGLST (Fig. lOD); NOTCHST (L&B Figs. IF, 2F); BINOTST (Fig. lOR); PROCST (Fig. lOQ); NAROWST (Fig. lOS); SQUARST (Y&B Fig. 17F);MEDNST(Fig. lOM). 17. DORSVIT: Species of the genus Spar- topijge possess conspicuous dorsal stripes (P) that appear to be homologous with the stripes in albida, slossonae, ritana, and grammica. The loss of stripes (A) is presumed to be homo- plastic: PRESTR (Figs. 2A-C, 54A); RE- DUSTR (Fig. 2D); ABSTR (Figs. 2E,F). 322 Great Basin Naturalist Memoirs No. 12 18. FACE: The plesiomorphic face of Flex- amia (and its ancestor Spartopyge) is pre- sumed to be pale, with a conspicuously con- trasting, black interocular band. In later lineages this habitus aspect was modified, as the interocular band became brown rather than black, as the distinction between the line and the remainder of the face became blurred, and, finally (A), as the face became essentially brown. In some transitional spe- cies, such as prairiana, the face varies widely fi-om pale with a contrasting, brown interocu- lar line to brown to (occasionally) nearly en- tirely black. We presume that loss of the ple- siomorphic face condition, as with dorsal stripes, probably occurred in several lines: CONBLF (Figs. 3A,B,E,F,G); TOOTHF {pyrops: see text); BRFACE (Fig. 3H); CON- BRF (contrasting brown face) VARCONF (variable contrast); BLFACE (Fig. 3C). Appendix 111 Notes Plant names. — Andropogon gerardii Vit- man, big bluestem; Andropogon virginicus L., broomsedge; Aristida dichotoina Michx. var. curtisii Gray; Aristida purpurea Nutt.; Bouteloua curtipendula (Michx.) Torr., side- oats grama; Bouteloua eriopoda (Torr.) Torr., black grama; Bouteloua gracilis (Willd. ex H. B. K.) Lag. ex Griffiths, blue grama; Bouteloua hirsuta Lag., hairy grama; Boute- loua pectinata Featherley, tall grama; Boute- loua uniflora Vasey, Nealley grama; Buchlo'e dactyloides (Nutt.) Engelm., buffalograss; Calamovilfa gigantea (Nutt). Scribn. & Merr., big sandreed; Calamovilfa longifolia (Hook.) Scribner., prairie sandreed; Cynodon dactylon (L.) Pers., bermudagrass; Distichlis spicata (L.) Greene, salt grass; Eragrostis curvula (Schrad.) Nees, weeping lovegrass; Eragrostis pectinacea (Michx.) Nees, tufted lovegrass; Eragrostis spectahilis (Pursh) Steud., purple lovegrass; Juncus tenuis Willd., poverty rush; Leptoloma cognatum (Shult) Chase, fall witchgrass; Muhlenbergia asperifolia (Nees & Mey.) Paradi, scratch- grass; Muhlenbergia cuspidata (Torr. in Hook.) Rydb., plains muhly; Midilenhergia pauciflora Buckley, New Mexican muhly; Muhlenbergia porteri Scribn. ex Beal, bush muhly; Muhlenbergia pungens Thurb., sand muhly; Muhlenbergia repens (Presl.) A. S. Hitchc, creeping muhly; Muhlenbergia reverchonii Vasey & Scribn., seep muhly; Panicum capillare L., witchgrass; Panicum virgatum L., switchgrass; Poa pratensis L., Kentucky bluegrass; Redfieldia flexuosa (Thurb.) Vasey, sand blowout grass; Schiz- achyrium scoparium (Michx.) Nash, little bluestem; Sporobolus virginicus (L.) Kunth., seashore dropseed; Zoysia japonica Steudl., zoysia grass. Plant ranges. — Plant ranges were mapped using published range maps (Pohl 1968; Hitchcock 1971; McGregor 1977; Stubben- dieck et al. 1986) and/or various floras (Gould 1975; Harrington 1954; Jepson 1925; Johnston 1943, 1963; Kearny and Peebles I960; Kiich- ler 1964; Lesueur 1945; Lonard and Judd 1980; Muller 1947; Rzedowski 1966, 1973, 1975, 1983); Shreve 1939, 1942; Steyermark 1963; Pohl 1966; and Welsh et al. 1987). Insect ranges. — Occurrence of specimens was mapped on maps of the United States or North America (Goode Base Map, Depart- ment of Geography, University of Chicago, was used as a base). Specimens examined from the collections listed below were mapped. Canadian and Mexican records are mapped by locality; U.S. records were mapped by county to avoid extensive overlap of symbols. Some records from the collections of the Illinois State Natural History Survey (as reported in DeLong 1948), and the Kansas State University Collection were also mapped. Other Records. — Not all "records" were mapped. We refrained from mapping speci- mens that in our judgment represented possi- ble or probable errors. These records in- cluded the following record for clayi: "Colfax, Gal., 8-7-30," from the OSU collection. Since clayi is a common Ohio species, but appar- ently does not occur west of the Mississippi River, this record is probably in error. Three records for picta seem questionable. Perhaps "Coronado N.F., Pena Blanca Resort, Ari- zona, 19 Aug. 1970, Harris and Harris, GL2019" (CNC) is valid. However, "Panama Citv, C. Z., Julv 23, 1920" (OSU), and "Mina, Mineral Co. Nevada, Aug. 20, 1955, GL274, Galloway" (CNC) seem less likely to be cor- rect. For other unusual records, see discus- sion under decora, areolata, and graminea. 1988 Whitcomb, Hicks: Flexamia Revision 323 In all studies, some specimens (usually fe- males) remain unidentified at the end of the study. One record of this kind is intriguing. The label "Cowan, Manitoba, 8 Aug., 1946, R. H. Beamer" (KU) is affixed to a single female that we cannot name. See field notes (later in this appendix) concerning this record. Collections utilized. — We examined the Flexamia collections of the United States Na- tional Museum (USNM), Ohio State Univer- sity (OSU), University of Kansas (KU), and the Canadian National Collection (CNC). Our own collection (Beltsville Agricultural Re- search Center [BARC]) contains, in addition to mounted specimens, a large inventory of unmounted but accessioned (Lynn and Whit- comb 1987) specimens. The inventories given in Appendix I are summations of cleared and uncleared specimens. We feel that the risk of misdeterminations of uncleared specimens is very low, with some important caveats. Fe- males of the prairiana and flexiilosa groups must be cleared for determination, and males of the prairiana group should be cleared. In many cases large series were available, in which case we cleared several specimens and tentatively identified others by inference. For these reasons the numbers given probably are an accurate reflection of the relative abun- dance of each species. We attempted to curate the collections examined and to supplement them, where possible, with additional mate- rial. IPL ACCESSIONS. — The BARC collection is largely computerized (Lynn and Whitcomb 1987); collection records are given in terms of the accession number in this system. Collec- tion methods have been described (Whit- comb etal. 1986, 1987). Original Beamer field notes. — Relevant excerpts from field notes written by Mrs. Lucy D. Beamer on: ritana. "08-18-35. Santa Rita Mts., Ariz. (E. side) 11:30-5 PM. mild, fair to cloudy ... Dr. Ball, RHB (R. H. Beamer) and Jack climbed the trail up the mountain. Just as they were abandoning the search RHB took a new spe- cies of Gladioniira or Athysanella which Dr. Ball is naming. Worked for a good while, fi- nally had 30 or 40 of them. Before this only 3 specimens had been taken." There are no ex- plicit references to Flexamia, or to the exact collecting location. The Athysanella species mentioned was presumably A. (Gladionura) furculata. beameri. "07-26-46. Otter Lake, New York (5 miles west). Collected just inside Adiron- dack Park on Highway 365 at one of the first little lakes. R. [R. H. Beamer] swept tiny sedges along the shore — took 80 or so of a tiny fulgorid he thought was Bakerella sp. Later not so sure. " Again, there was no explicit men- tion of Flexamia. Unidentifiable female. "08-07-37. Cowan, Manitoba (3 miles north). R. swept in the tall green grass growing up through the heavy mat of fallen grass which surrounds a lake for fifty or a hundred yards in all directions. Found interesting cicadellid — long wings — proba- bly belonging to the genus Polyamia. Their wings were ruffed even when he took them." We have been unable to find other specimens from this collection in the KU collection. AFRONISIA, A NEW AFRICAN GENUS OF MEENOFLIDAE (HOMOPTERA: FULGOROIDEA) Michael R. Wilson' Abstract — The new genus Afronisia is founded with Eponisia albovittata Fennah as the type-species. Eponisia briinnescens Synave, £. flavescens Synave, E. pallida Linnavuori, and Nisia muiri Metcalf are transferred to this genus, and E. albinervosa Muir is placed in synonymy with Kcnnesia albinervosa Muir. Two new species are described: Afronisia bredoi, from Zaire, and A. gembtiensis, from Nigeria. Keys are provided for separation of the African genera of Kermesiinae and for the seven species oi Afronisia that are described and figured. The Old World planthopper family Meeno- plidae is one of the smaller fulgoroid families, consisting of slightly more than 100 described species. Until recently all species were placed in eight genera in two subfamilies, the Meenoplinae and the Kermesiinae ( = Nisii- nae). However, Emeljanov (1984) and Tsaur et al. (1986) described new genera for existing species in the Oriental region and named sev- eral new species. Over 50 species are known from Africa through the work of Muir (1927, 1934) and other early workers and, more re- cently, by Fennah ('l955, 1957, 1958), Lin- navuori (1973), and Synave (1957a, 1957b, 1971). It is now apparent that only a small proportion of the species has been described so far, as has been found for other African fulgoroid groups (e.g., Derbidae, Wilson 1987). Meenoplids are small fulgoroids with ten- tiform forewings and one or both claval veins with sensory pits ("granulations" in older liter- ature) on either side of the vein. Females possess wa.x-producing plates on abdominal segments 6-8. The family belongs to a group that includes Achilixiidae, Kinnaridae, and Achilidae. Meenoplidae are divided into two subfamilies, the Meenoplinae and the Kerme- siinae. In the Meenoplinae the first claval vein has a single row of sensory pits, and the sec- ond claval vein is more or less covered with them. In the Kermesiinae the claval veins are fused near the apex of the clavus (e.g. , Fig. 4). The first claval vein has a row of sensory pits on either side of the vein. Three African gen- era are placed in this subfamily: Kermcsia, Nisia, and Afronisia. Kermesiinae was first used by Kirkaldy (1906:427) as a subfamily of Derbidae to in- clude Nisia, Phaconcnra, Suva, and several derbid genera. Although the subfamily was named Kermesiinae, the genus Kermesia was not treated at that time but was included by implication in comments on Nisia by Kirkaldy (1906:427), who stated that "[Nisia] seems hardly differentiated from Kermesia." How- ever, in thefollowingyear Kirkaldy (1907:163) stated that two subfamilies of the Derbidae may be recognized, including the Nisiinae represented by Nisia. The name Nisiinae was accepted by most subsequent workers, in- cluding Metcalf (1945). Recentlv, Emeljanov (1984) and Tsaur et al. (1987) used Kermesi- inae as the subfamily name in preference to Nisiinae; this name has been followed here. The African meenoplid species have been placed in the genera Nisia, Kermesia, Epon- isia (Kermesiinae), Anifinis, and Meenoplus (Meenoplinae). Because these genera are all poorly defined, it is necessary for several new genera to be erected to accommodate some existing species and some undescribed ones. The five African species ciurently placed in Eponisia have little in common with the re- maining Asiatic species, which include the type-species, E. giittula Matsumiua, de- scribed from Taiwan (Tsaur et al. 1986). This has also been noted by Emeljanov (1984). The present paper transfers four of them to Afro- nisia and the fifth to synonymy in Kermesia. Abbreviations of Depositories BMNH British Museum (Natural History), London, UK IRSNB Institut Royal des Sciences Naturelles de Bel- gi(liie, Brussels, Belgium 'cab International Institute of Entoniolosy, i/o DepailMu-iit ol KnlonioloyN , Hntisli Museum (Natural Hist()r\), Cromwell hu.id, London. SVV7.5BD. UK. 324 1988 WiLsoN: Genus Afhonisia 325 MNHN Museum National d'Histoin- NaturelU-, Paris, France MRAC Musee Royal de I'Africiue Centrale, Tervuren, Belgium NCI National Insect Collection, Pretoria, South Africa RL R. Linnavuori Collection, Raisio, Finland Key to African Genera of Kermesiinae 1. Clypeus rounded, without lateral carinae. Forewing with only one branch to M apically . . Nisid — Clypeus with lateral carinae. Forewing with two branches of M at apex (Fig. 4) 2 2(1). Eyes large, twice as long as wide. Forewing with clavus more than half total length. Hind tarsal segments with segment 1 approximately ecjual- ing in length segments 2 + 3 ... Afronmii gen. n. — Eyes smaller, only slightly longer than wide. Forewing with clavus one-third total length. Hind tarsal segment 1 longer than 2 + 3 Kcrmesia Afronisia, gen. n. Type-species. — Eponisia albovittata Fen- nah. Description. — Small, slender meenoplids, length overall 3.5-5.9 mm. Head with vertex longer than wide, merging smoothly into frons with profile strongly rounded, postero- lateral areolets widely spaced. Frons over twice as long as broad; lateral carinae parallel for most of their length, diverging slightly on clypeus, interrupted at frontoclypeal suture. Clypeus with lateral carinae extending from lateral carinae of frons; median carinae weakly developed but present toward rostrum. Ros- trum long, extending beyond hind tro- chanters. Pronotum with well-defined ante- rior carina parallel to the anterior margin, carina also present on shoulder of pronotum. Mesonotum with median carina present but not sharply defined, lateral carinae obscure. Forewing: Relatively long and narrow, 2.5- 3.0 times longer than maximum width, clavus longer than half total length. Sc + R united for almost half wing length, R bifurcate at apex, M bifurcate at apex. Claval veins appear to join inner margin before apex of clavus but actually enter the apex. Hindwing: r-m crossvein at approximately half length of wing, R and M bifurcate at apex. Hindlegs; Hind tibiae with lateral spines. Posttibia with 8 spines grouped as 3 + 5. Apex of segment 1 with approximately 8 spines, median ones less sclerotized than other ones. Apex of segment 2 with similar arrangement of spines. Tarsal segment 1 approximately ecjual in length to segments 2 + 3. Male (;ENiTAi.iA. — Pygofer dorsoventrally elongated, in lateral view with dorsolateral angle slightly or strongly produced, either rounded or pointed. Anal segment with lat- eral margins diverging apically, posterolateral angles produced or rounded. Aedeagus (phal- lus) tubular, enlarged at apex, more or less trumpet-shaped. Periandrium (phallobase) with several pairs of appendages arising from base, inner pairs more slender than outer pairs. Parameres long and narrow, apical por- tion upcurved. Female genitalia. — Of the "reduced" meenoplid form. Distribution. — Africa south of Sahara. Diagnosis. — The genus may be distin- guished from Nisia and Kermesia by the fol- lowing characters. In ISlisia the lateral carinae of the clypeus are absent; they are present in Afronisia and Kermesia. In Kermesia the cari- nae extend smoothly from the frons; in Afro- nisia they are disjunct. The eyes oi Afronisia species are large, twice as long as wide, while in Kermesia and ISlisia they are more globular. The forewing venation in Afronisia differs, with the clavus more than half the length of the forewing and M bifurcate at its apex. In Kermesia M is also bifurcate, but the clavus is short, only one-third the length of the fore- wing. In Nisia M is single at its apex, with the clavus more or less half the length of the forewing. The hind tarsal segment 1 is longer than 2 + 3 in both Nisia and Kermesia and is more or less equal to the length of 2 + 3 in Afronisia. All of these features are at present only diagnostic. However, the appendages of the aedeagus are characteristic and are not present in other meenoplid genera but are here regarded as apomorphic for the genus. Oriental species oi Eponisia resemble Kerme- sia in some respects and differ markedly from Afronisia in the form of the male genitalia (Tsaur et al. 1987). The biology and host plants are little known. Linnavuori (1973) records A. albovit- tata (Fennah) and A. brunnescens (Synave) as being collected "in a swamp." Key to Species (Males) oi Afronisia 1. Forewing almost entirely dark brown, contrast- ing strongly with yellow, dorsal stripe from ver- tex to pronotum and mesonotum. Aedeagus 326 Great Basin Naturalist Memoirs No. 12 with 3 pairs of processes (Fig. 7) albovittata (Fennah) — Forewing variable in color but never entirely dark brown. Aedeagus with between 2 and 4 pairs of processes 2 2(1). Aedeagus with 2 pairs of large, sclerotized pro- cesses 3 — Aedeagus with 3 or 4 pairs of processes (Figs. 22, 35, 45) 4 3(2). Forewings overall pale whitish gray. Aedeagus with outer processes very large, curved, extend- ing beyond phallus (Fig. 56), short, central pro- cess present bredoi, n. sp. — Forewing with white veins, membranes pale brown (Fig. 15). Aedeagus with outer processes shorter than phallus, short, central process ab- sent muiri (Metcalf) 4(2). Aedeagus with 4 pairs of processes (Fig. 22) . . . flavescens (Synave) — Aedeagus with 3 pairs of processes 5 5(3). Aedeagus with outer pair of processes longer than inner pairs, curved inward and dorsad (Figs. 33-35). Pygofer with laterodorsal angles pointed (Fig. 31) hninnescens (Synave) — Aedeagus with outer pair of processes shorter than inner pairs, pygofer with laterodorsal an- gles rounded 6 6(5). Outer pair of aedeagal processes short and bifur- cate (Fig. 50) gemhuensis, n. sp. — Outer pair longer, about two-thirds length of inner pairs (Fig. 45) pallida (Linnavuori) Afronisia albovittata (Fennah), comb. n. Figs. 1-12 Eponisia albovittata Fennah 1955. Ann. Mus. Congo Tervuren in ser. 8. Zool. 40; 434. Length. — Male 4.2-4.4 mm, female 4.5-4.8 mm. Description. — Coloration: Overall dark brown species. Vertex and face with lateral edges of carinae dark brown, face between carinae brown or light yellow in some speci- mens, becoming yellow at junction of face and vertex. Antennae dark brown, eyes dark. Ver- tex pale yellow. Pronotum and mesonotum with central, yellow stripe (Fig. 1) with orange pigmentation following median carina. Fore- wing dark brown, inner margin of clavus and subapical line of transverse veins between R and Cu and between apex of clavus and Cuj white (Fig. 4). Legs brown. Abdomen and genital capsule dark brown. Male genitalia (Figs. 7-12). — Pygofer longer than broad, laterodorsal angles acute (Fig. 10). Anal segment with apical (posterior) margin concave between rounded posterolat- eral margins (Fig. 11), slightly deflexed in lateral view (Fig. 10). Aedeagus with perian- drium bearing three pairs of sclerotized pro- cesses from ventral portion (Fig. 7), the outer pair broader and slightly shorter than the in- ner pair and central pair very short. Long, trumpet-shaped central portion (phallus) ex- tending beyond processes. Parameres long, with upcurved, subspinose, apical process (Fig. 9), slightly widened before apical por- tion in dorsal view (Fig. 12). Distribution, — East Africa: Rwanda (type locality), Sudan (Equatoria Prov.) (Linnavuori 1973), Uganda, Kenya, Zaire. Material examined. — Males and females from Kenya, Uganda, Sudan (Equatoria Prov.)(BMMH). Remarks. — Fennah's original description (1955) states that the phallobase (perian- drium) possesses two pairs of sinuate spines, not three as actually present. However, Fen- nah's drawing was made from a lateral view, and the short, central (ventral) pair may not have been visible. The male genitalia were figured correctly by Synave (1957b) from ma- terial from the type locality, and for this rea- son the type-specimen was not reexamined. Diagnosis. — The dark coloration of the species and the pale yellow, dorsal stripe dis- tinguish it from all others in the genus. Afronisia muiri (Metcalf), comb. n. Figs. 13-21 Nisia albovenosa Muir 1927. Ann. Mag. Nat. Hist. (9)19: 200. [Preoccupied by Nisia albovenosa Distant, 1906] Nisia muiri Metcalf 1945: 228 replacement name for Nisia albovenosa Muir. Length. — Male 3.5-3.8 mm, female 4.5- 4.6 mm. Description. — Coloration: Lateral carinae of head and vertex dark brown. Face and clypeus brown, lateral carinae of clypeus pale. Vertex pale yellow-white. Pronotum and mes- onotum with lateral margins brown with cen- tral, pale stripe (Fig. 13), median carina nar- rowly orange. Forewing with veins white, membranes light brown, darker adjacent to veins (Fig. 15), granulations dark brown on white background. Abdomen dark brown, in- tersegmental membranes white. Legs light brown. Male genitalia (Figs. 16-21). — Pygofer longer than broad, laterodorsal angles almost rectangular (Fig. 18). Anal segment posteri- orly deflexed in lateral view (Fig. 18), slightly 1988 WiLSON: Genus Afronisia 327 Figs. 1-12. Afronisia allwvittata (Fennah): 1, head and thorax, dorsal view; 2, head, face; 3, head, lateral view; 4, forewing; 5, hindwing; 6, hindleg; 7, aedeagus, ventral view; S, aedeagus, dorsal view; 9, aedeagus and parameres, lateral view; 10, pygofer and anal segment, lateral view; 11, anal segment, dorsal view; 12, parameres, ventral view. 328 Great Basin Naturalist Memoirs 1988 Figs. 13-28. Figs. 13-21, Afronisia muiri (Metcalf); 13, head and thorax, dorsal view; 14, head, lateral view; 15, forewing; 16, aedeagus, ventral view; 17, aedeagus, lateral view; 18, pygofer and anal segment, lateral view; 19, anal segment, dorsal view; 20, paramere, ventral view; 21, paramere, lateral view. Figs. 22-28, Afronisia flavescens (Synave): 22, aedeagus, ventral view; 23, aedeagus, lateral view; 24, pygofer, lateral view; 25, anal segment, lateral view; 26, anal segment, dorsal view; 27, paramere, ventral view; 28, paramere, lateral view. 1988 WiLsoN: Genus Afronisia 329 produced posterior processes (Fig. 19). Ae- deagus with periandrium bearing two pairs of sclerotized processes (Fig. 16), outer pair large, curved, inner pair longer, thin (Fig. 17). Parameres with upturned apical portion (Fig. 21), subapical lobe in dorsal view (Fig. 20). Distribution. — South Africa. Material E.XAMiNED. — Holotype 9, South Africa, Port St. John, Pondoland, R. E. Turner, iv.l923 (BMNH); 1 9, same data as holotype; 1 c? 6 9 , Rustenburg, Transvaal, 10-13. xii. 1961 (Capener); 1 9, Trazaneen, Transvaal, 10. xii. 1918; 1 9, Umtentweni, Na- tal, 9-14.iii.1961 (Capener) (NCI, 2 9 BMNH); 1 d, Ladvsmith, 29. i. 1981 (Theron); 1 9, Dundee, 21. i. 1981 (Theron); 1 9, Dur- ban, 7. i. 1971 (Kluge) (BMNH). Remarks. — The species was originally de- scribed from one female. A second female from the same locality (Port St. John, Pon- doland) is also present in the BMNH collec- tion. Only two males were present among the specimens from South Africa. No further spe- cies have been found in South Africa. Com- parison of wing pigmentation supports the as- sociation of the males with the type female. Synave (1957b) repeated Muir s original de- scription but did not examine the type-speci- men. Diagnosis. — This is the only species known at present from South Africa. The forewing coloration is distinctive, and the male aedea- gus is characteristic with the two pairs of pro- cesses. Afronisia flavescens {Sy nave), comb. n. Figs. 22-28 Eponisia flavescens Svwdve 1971. Bull. Inst. r. Sci. nat. Belg. 47(39): 10. Length. — Male 4.2 mm, female 4.8 mm. Description. — Coloration: Male head and body orange, pale yellow stripe on pronotum and mesonotum with orange median pigmen- tation following median carina. Lateral cari- nae of head edged dark brown. Legs orange. Forewing with veins white, apices pale brown. Membranes suffused, very pale brown. Granulations with black points on white background. Female much paler than male. Overall stra- mineous. Male genitalia (Figs. 22-28). — Pygofer with laterodorsal angles acute (Fig. 24). Anal segment deflexed posteriorly in lateral view (Fig. 25), posterior margin scarcely produced (Fig. 26). Aedeagus with periandrium with four pairs of sclerotized processes (P'ig. 22), outer pair thickest, two inner pairs thinner but more or less equal in length, central pair about half the length. Central portion (phal- lus) longer than processes (Fig. 22). Par- ameres as in other species with upturned api- cal portion (Figs. 27, 28). Distribution. — Nigeria. Material examined. — Holotype d, Nige- ria, Badeggi, N.W. State (IRSNB). Paratype 9 (not male as stated in original description), Nigeria, Yankar, N.E. State (IRSNB). Diagnosis. — A pale species, similar in col- oration to A. hredoi n. sp. The only species to have four distinct pairs of aedeagal processes. Remarks. — Known only from the type ma- terial. Afronisia brnnnescens (Synave), comb. n. Figs. 29-37 Eponisia hninnescens Synave 1957a. Expl. Pare National de rUpemba. Fasc. 43(2): 7. Length. — Overall male 4.2-4.4 mm, fe- male 4.6-4.9 mm. Description. — Coloration: Male body and legs pale orange. Vertex and mesonotum with distinct, pale yellow stripe, darker orange lat- erally. Edges of lateral carinae of head dark brown extending to lateral ocellus, inner mar- gin (sensory pits) whitish. Antennae pale yel- low brown. Forewing: Membranes pale, stramineous, light brown, darker adjacent to veins. Veins white, with apices darker. Claval vein with sensory pits dark brown. Forewing narrow, parallel-sided, rounded at apex. Hindwing: Whitish, veins pale brown. Female paler generally than male. Male genitalia (Figs. 31-37). — Pygofer with laterodorsal angles extended, acute in profile (Fig. 31). Anal segment with postero- lateral apical lobes reflexed (Figs. 31, 32). Aedeagus with periandrium bearing three pairs of sclerotized processes, the outer pair longer and thicker than two inner pairs, curved inward and dorsad apically (Figs. 33-35), inner pair diverging apically (Figs. 33, 35), central pair short, about half the length of outer pairs. Phallus longer than pro- cesses, trumpet-shaped. Parameres slender, apical quarter directed dorsad (Fig. 36). Some 330 Great Basin Naturalist Memoirs No. 12 variation appears in length of processes (Fig. 33 illustrates the paratype. Fig. 35 is a speci- men from Cameroon). Distribution. — Central and west Africa: Cameroon, Sudan (Equatoria Prov.) (Lin- navuori 1973), Zaire (Katanga Prov., type lo- cality, Orientale Prov.), Central African Re- public (Bossangoa). Material EXAMINED. — Holotype 6 , Zaire, Kaziba (MRAC). Paratype d, Zaire, Kabwe (MRAC). Zaire, numerous specimens from Pare National de la Garamba (northeast Zaire) (IRSNB, MRAC); 1 6, Cameroon (Banyo) (BMNH); 1 6, Central African Republic (Bossangoa) (RL). Diagnosis — The male genitalia are very characteristic with the curved outer pair of processes. The species was found mixed in series with A. pallida and may be separated by the narrow, parallel-sided forewings as well as by the male genitalia. Afronisia pallida (Linnavuori), comb. n. Figs. 38-45 Eponisia pallida Linnavuori 1973. Notiil. ent. .53: 111. Length. — Overall male 4.5-5.0 mm, fe- male 5.2-5.8 mm. Description. — Coloration: Male/female, lateral carinae of head with edges dark brown, body and head pale yellow, mesono- tum orange laterally, pale, median stripe in- terrupted by orange, median carina. Legs pale yellow. Abdomen and genital capsule stramineous. Forewing with pale yellow membrane, white veins, apices of veins marked with brown (Fig. 38), apical cells darkened adjacent to veins, black tips to white sensory pits on claval veins. Forewing becom- ing broader toward apex. Male genitalia (Figs. 40-45). — Pygofer with laterodorsal angles broadly rounded in profile (Fig. 41), anal segment with postero- lateral lobes (Fig. 40) reflexed (Fig. 41). Aedeagus (Figs. 45, 46) with periandrium bearing three pairs of sclerotized processes and one central (ventral) process, outer pair shorter than inner pairs (Fig. 45), the more dorsal of the inner pairs converging apically (Fig. 45), the inner pair parallel for half their length before diverging (Fig. 45), single, cen- tral spine short. Two small spines on dorsal surface of phallus. Parameres with distinct ventral lobe one-third distance from apex with apical third curved upward and twisted (Figs. 42, 43). Distribution. — Central Africa: Sudan (Equatoria Prov., type locality) (Linnavuori 1973), Central African Republic, Zaire. Material e.xamined. — Holotype d, Su- dan, Equatoria, Yambio (RL). Paratype 9 , Sudan, Equatoria, Yambio (BMNH). Cen- tral African Republic (Bossangoa) (IRSNB). Zaire, numerous specimens from Pare Na- tional de la Caramba (northeast Zaire) (IRSNB, MRAC). Remarks. — Found in same localities as A. bninnescens since it is mixed in collections from Zaire. Diagnosis. — A pale species, externally similar to A. bninnescens but slightly larger and with forewings broadened rather than parallel-sided. Male genitalia very distinct with three pairs of long processes. Afronisia gembuensis, n. sp. Figs. 46-51 Length. — Male 4.7 mm, female 5.2 mm. Description. — Body color pale orange, re- sembling A. mtiiri, especially in darker pig- mentation of forewings. Male genitalia (Figs. 46-51). — Pygofer with laterodorsal angles rounded (Fig. 46). Anal segment with lateral margins diverging apically, posterolateral angles acute (Fig. 47), with apical margin concave. Aedeagus with periandrium with three pairs of sclerotized processes and one short, central process, outer pair short and bifurcate (Fig. 50), inner pairs long, with outer of these diverging, in- ner one almost parallel (Fig. 50), central pro- cess short. Parameres with pronounced ven- tral lobe two-thirds as long as its length, apical third thinner and directed dorsad, slightly twisted (Figs. 48, 49). Distribution. — Nigeria. Type material. — Holotype 6, Nigeria, N.E. State, Gembu-Yelwa,' 22.viii.1973, Lin- navuori (RL). Paratvpes, 3 (5 6 9, Nigeria, N.E. State, Gembu, 21-22. viii. 1973, Lin- navuori (RL, 16 19 deposited in each BMNH, MRAC). Diagnosis. — Somewhat similar to A. pall- ida in aspects of the male genitalia and partic- ularly in the shape of the laterodorsal angles of the pygofer and the shape of parameres. The aedeagal (periandrium) processes are, how- ever, diagnostic, particularly the bifurcated outer pair. 1988 WILSON: Genus Afronisia 331 Figs. 29-45. Figs. 29-37, Afronisia bnmnescens (Synave); 29, head, lateral view; 30, forewing; 31, male pygofer and anal segment, lateral view; 32, anal segment, dorsal view; 33, aedeagus, ventral view, paratype; 34, aedeagus, lateral view; 35, aedeagus, ventral view; 36, paramere, lateral view; 37, paramere, ventral view. Figs. 38-45, Afronisia pallida (Linnavuori): 38, forewing; 39, head, lateral view; 40, male anal segment, dorsal view; 41, pygofer and anal segment lateral view; 42, paramere, ventral view; 43, paramere, lateral view; 44, aedeagus, lateral view; 45, aedeagus, ventral 332 Great Basin Naturalist Memoirs No. 12 Figs. 46-51. Afronisia gemhuensis , n. .sp. : 46, male pygofer and anal segment, lateral view; 47, anal segment, dorsal view; 48, paramere, ventral view; 49, paramere, lateral view; 50, aedeagus, ventral view; 51, aedeagus, lateral view. Afronisia bredoi, n. sp. Figs. 52-57 Length. — Male 5.2 mm, female 5.7-5.9 mm. Description. — Coloration male/female: body and legs light orange dusted with wax. Head with lateral carinae of vertex and face edged with dark brown, face suffused with light brown. Mesonotum/pronotum light or- ange laterally, central, pale yellow stripe with median carina light orange. Antennal scape very pale brown, pedicel darker. Forewing: Overall pale grey-white, mem- branes darker, particularly adjacent to veins. Apices of veins at margin darker, especially in female. Forewing distinctly wider toward apex. Male genitalia (Figs. 52-57). — Pygofer with laterodorsal angles rounded (Fig. 52). Anal segment with lateral margins diverg- ing apically, posterolateral angles produced (Fig. 53). Aedeagus with periandrium bearing two pairs of sclerotized processes, and a sin- gle, central one, outer processes very large, curved inward beyond the phallus, with short processes arising laterally at base. These are asymmetric in specimen examined (Fig. 56). Inner processes shorter than phallus, more or less parallel, central spine short. Parameres with apical third curved smoothly upward in lateral view (Fig. 57), twisted apicallv (Fig. 54). Distribution. — Zaire (Elisabethville). Type material. — Holotype 6, Zaire, la- beled Congo Beige, Elisabethville, iv.l939, H. J. Bredo (MRAC). Paratypes: 2 9, same data as holotype (MRAC). Dia(;n()SIS — A pale-colored species resem- bling A. pallida. The long, curved, outer ap- pendages of the male aedeagus are diagnostic and resemble no other species. Remarks on Eponisia alhiiicriosa Muir 1934 This species is considered the junior syn- onym of Kermesia alhinervosa Muir 1927 (type locality. Sierra Leone). The male holo- type and female allotype were very badly damaged in transit from Honolulu to London 1988 WiLSON: Genus Afronisia 333 Figs. 52-57. Afronisia brecJoi, n. sp. ; 52, male pygofer and segment, lateral view; 53, anal segment, dorsal view; 54, paramere, ventral view; 55, paramere, lateral view; 56, aedeagus, lateral view; 57, aedeagus, ventral view. in 1927 (together with other African meen- oplid types described in the same paper). For- tunately, the male genitalia of the holotype were intact and other specimens have been associated. It is with these specimens that the female holotype o( Eponisia albinervosa from Uganda has been compared. The synonymy is as follows: Kermesia albinervosa Muir 1927. Ann. Mag. Nat. Hist. (9)19: 203. Eponisia albinervosa Muir 1934. Ann. Mag. Nat. Hist. (10)14: 564. syn. n. Acknowledgments I am pleased to thank the following for the loan of material used in this study: Dr. Jan Van Stalle (IRSNB, Brussels, Belgium), Dr. H. Andre (MRAC, Tervuren, Belgium), Dr. R. Linnavuori (Raisio, Finland), the late Dr. J. Theron, deceased January 1988 (Stellenbosch University, South Africa), and Dr. 1. M. Mil- lar (NCI, Pretoria, South Africa). I am very grateful to Mr. W. R. Dolling, Dr. W. J. Knight, and Mr. M. D. Webb for making useful comments and criticism of the manu- script. Literature Cited Distant, W L 1906. Fauna of British India including Ceylon and Burma. Rhynchota. Heteroptera- Homoptera 3. Taylor and Francis, London, UK. 503 pp. Emeljanov. a. F 1984. [A contribution of knowledge of the families Kinnaridae and Meenoplidae (Ho- moptera, Fulgoroidea).] Ent. Obozreniye 63(3): 468-483. [Translated in Ent. Rev. 64: 49-65, 1985.] Fennah, R G 1955. Contributions a I'etude de la faune entomologique du Ruanda-Urundi (Mission P. Basilewsky 1953). LXXVIII. Homoptera, Ful- goroidea. Ann. Mus. Congo Tervuren in ser. 8 Zool. 40: 427-446. 1957. Results from the Danish expedition to the French Cameroons 1949-50. XXIV Fulgoroidea. Bull, de IT. F.A.N, ser. A. 19: 1274-1311. 1958. Fulgoroidea (Homoptera) from West Africa. Bull, de ri. F.A.N, ser. A. 20: 460-538. KiRKALDY, G W 1906. Leaf-hoppers and their natural enemies, Pt. IX. Leafhoppers-Hemiptera. Bull. Hawaiian Sugar Planters Assoc, Div. Ent. 1: 271-479. 334 Great Basin Naturalist M emoirs No. 12 1907. Leaf-hopper.s — supplement (Hemiptera). Bull. Hawaiian Sugar Planters A.SSOC., Div. Ent. 3: 1-186. LiNNAVUORl, R 1973. Hemiptera of the Sudan, with re- marks on some species of the adjacent countries 2. Homoptera Auchenorrhyncha: Cicadidae, Cer- copidae, Machaerotidae, and Fulgoroidea. Notul. Ent. 53:65-137. Metcalf. Z. P 1945. Meenoplidae. Pages 219-2.38 in General catalogue of the Hemiptera. Ease. IV. Pt. 6. Smith College, Northampton, Massachusetts, USA. Mum, F 1927. New species of African Meenoplidae (Eul- goroidea, Homoptera). Ann. Mag. Nat. Hist. (9)19; 197-208. 1934. New and little-known Fulgoroidea (Ho- moptera). Ann. Mag. Nat. Hist. (10)14; .561-586. Synave, H 1957a. E.xploration de Pare National de I'Upemba. Mission G. F. de Witte. 2 Meenopli- dae. Ease. 43; 79-81. 1957b. Pare National Albert 1. Mission G. E. de Witte 1933-35, Meenoplidae (Homoptera, Eul- goroidea). Ease. 90(2); 7-.30. 1971. Contribution a la connaissance des ful- gorides du Nigeria (Homoptera) (recoltes J. T. Medler). Bull. Inst. r. Sci. nat. Belg. 47: 1-34. TsAUR. S C , C T Yang, and M R Wilson 1987. Meenoplidae of Taiwan (Homoptera: Eul- goroidea). Monograph of Taiwan Museum No. 6; 81-117. December 1986 (1987), Wilson, M R 1987. African Derbidae (Homoptera, Ful- goroidea); taxonomic notes with descriptions of new species collected mainly from coconut. J. Nat. Hist. 21:567-595. DELPHACIDAE OF ALASKA (HOMOPTERA: FULGOROIDEA) Stephen W. Wilson' Abstract. — Fifteen species, in 10 genera, of Delphacidae are recorded from Alaska. One, Kusnezoviella niacleani, is described as new to science. Acanthodelphax analis (Crawford), Chilodelphax magnifrons (Crawford), Javesella arcanastyla (Beanier), and J. atrata (Osborn) are new combinations. Delphacudes saileri Beamer is a synonym of Javesella sinullima (Linnavuori). Of the 15 species, 1 is recorded only from Alaska, 3 are Holarctic, 4 are Nearctic, and 7 are Palearctic and restricted to tundra habitats, with 5 of these apparently found only north of the Brooks Range. Little information is available on species diversity and distribution of much of the Alaskan insect fauna (MacLean and Hodkin- son 1980). For example, only seven species of Delphacidae have been described from Alaska. Delphacodes stejnegeri (Ashmead) was described by Ashmead (1904) and re- described and illustrated by Dozier (1926). Vilbaste (1980) synonymized this species with Javesella pelhicida (Fabricius). Delphacodes atrata Osborn was recorded from Alaska by Beamer (1951). Delphacodes saileri Beamer (1952b) was described from Alaska but, as in- dicated below, is a synonym oi Javesella simil- lima (Linnavuori). DuBose (1960) recorded the Holarctic D. pelhicida from Alaska. Del- phacodes uniformis (Walker) was listed from Alaska by Metcalf (1943); however, Muir (1919) stated that this insect is not a delphacid. Megamelus flavus Crawford was recorded from Alaska by Beamer (1955). The present paper provides a key and infor- mation on distribution for the 15 delphacid species found in the Canadian National Col- lection, Ottawa, and material collected by MacLean in the northern part of the state. The name used by the first describer and the first placement in the current genus are given for each species. Because the species have been described in detail elsewhere, only the new species o( Kusnezoviella is described. Ac- curate identification of delphacids relies on use of characters of the male genitalia; for this reason, all species treated here are illustrated. 2(1) 3(1). 4(3). 5(3). 6(5). Key to the Alaskan Delphacidae Frons with 2 longitudinal, median carinae (Fig. 1) 2 Frons with 1 longitudinal, median carina .... 3 Large pits present on frons, pronota, mesonota, and abdomen (Figs. 1, 2); aedeagus strongly decurved with poorly developed teeth (Figs. 2, 4) Achorotile sitbarctica Scudder Pits absent; aedeagus straight with well- developed teeth (Figs. 6, 7) .... C riomorphiis ivilhelmi Anufriev & Averkin Pronotal lateral carinae straight, extending to, or almost to, posterior margin of pronotum (Fig. 12); metatibial spur with or without black-tipped teeth 4 Pronotal lateral carinae curving laterad, not extending to posterior margin of pronotum; metatibial spur with black-tipped teeth, these sometimes weak 5 Metatibial spur with well-developed, black- tipped teeth; pygofer with lateral, inflated lobes (Figs. 9, 11) . Megamelus flavus CravAord Metatibial spur lacking teeth; pygofer not in- flated laterally (Fig. 13) ' .... N othodelphax ebiirneocarinatiis (Anuiriev) Broad, pale stripe usually bordering longitu- dinal, median carinae on vertex, pronotum, and/or niesonotum; pygofer with broad lobe laterally on caudal margin (Figs. 16, 21) 6 Broad, pale stripe not present; pygofer with- out broad lobe 7 Styles each with irregular teeth on broad apex (Fig. 20) Unkanodes excisa (Melichar) Styles each with acute apex (Fig. 23) Chilodelphax magnifrons (Crawford) 'Department of Biology, Central Missouri State University, Warrensburg, Missouri 64093 335 336 Great Basin Naturalist Memoirs No. 12 Figs. 1-20. Figs. 1-5, Achorotile subarctica Scudder; 1, head, frontal view; 2, male genitalia, lateral view; 3, anal tube, dorsal view; 4, aedeagus, lateral view; 5, styles, caudal view. Figs. 6-8, Criomorphiis wilhelmi Anufriev & Averkin; 6, male genitalia, lateral view, 7, aedeagus, lateral view; 8, styles, caudal view. Figs. 9-11, Megamelus flavus Crawford: 9, male genitalia, lateral view; 10, aedeagus, lateral view; 11, male genitalia, caudal view. Figs. 12-15, Nothodelphax ehurneocarinatus (Anufriev): 12, head and thorax, dorsal view; 13, male genitalia, lateral view; 14, aedeagus, lateral view; 15, styles, caudal view. Figs. 16-20, t/n/ca norfe.s excisa (Melichar): 16, pyfoger and style, lateral view; 17, anal tube, lateral view; 18, 19, aedeagus, lateral view; 20, styles, caudal view. 7(5). Anal tube with spines crossing (Fig. 27) .... Ribautodelphax albostriatus (Fieber) — Anal tube with spines parallel or diverging . . 8 8(7). Pygofer with caudally directed, median pro- jection ventral to base of styles (Fig. 29) .... Acanthodelphax analis (Crawford) — Pygofer without median projection 9 9(8). Apices of styles converging (Fig. 34) Kusnezoviella rnacleani, n. sp. — Apices of styles diverging 10 10(9). Aedeagus strongly decurved (Fig. 36) Javesella pellucida (Fabricius) — Aedeagus recurved or straight (Figs. 40, 43, 47,50,53) 11 11(10). Aedeagus evenly forked (Fig. 40) javesella obscurella (Boheman) — Aedeagus not forked (Figs. 43, 47, 50, 53) . . 12 12(11). Aedeagus with projection in basal half on ven- tral aspect (Figs. 50, 53) or with broad, weakly toothed region in basal half on ventral aspect (Fig. 47) 13 — Aedeagus not as above (Fig. 43) Javesella atrata (Osborn) 13(12). Aedeagus with large, ventral, basal projection (Figs. 50, 53) 14 — Aedeagus without large, basal projection (P'ig. 47) Javesella simillima (Linnavuori) 14(13). Aedeagal, ventral, basal projection acute, aedeagal dorsal margin slightly cur\'ed (Fig. 50) Javesella discolor (Boheman) — Aedeagal, ventral, basal projection rounded, aedeagal dorsal margin constricted (Fig. 53) Javesella areanastijla (Beamer) Achorotile subarctica Scudder Figs. 1-5 Achrotile [sic] subarctica Scudder, 1963:169 Distribution records for specimens used in this study are: Alaska: Brooks Range, Spruce 1988 WiLsoN; Delphacidae OF Alaska 337 Tree Mine, 68°03'N, 149°W, 800 m, 4 July 1982, Seppo Keponen, ex. B. tidris Oman 1933. Colorado. Aceratagallia robusta Oman 1933. USA. Aceratagallia sordida Oman 1933. Hondiuas, Mexico, Texas. Aceratagallia texana Oman 1933. Texas. Aceratagallia truncata Oman 1935. California. Aceratagallia unica Oman 1938. Argentina. Aceratagallia vastitatis Oman 1933. USA. Aceratagallia vidgaris Oman 1933. Kansas. Agallia abnormis Oman 1938. Argentina. Agallia acuticatida Oman 1933. Haiti. Agallia atromuculata Oman 1938. Bolivia. Agallia basalis Oman 1938. Bolivia. Agallia basifusca Oman 1938. Bolivia. Agallia biftircata Oman 1938. Bolivia. Agallia bicolor. Oman 1933. West Indies. Agallia bidigitata Oman 1933. Guatemala. Agallia blanda Oman 1938. Argentina. Agallia brevicaudata Oman 1938. Brazil. Agallia caudata Oman 1938. Bolivia. Agallia clara Oman 1938. Brazil. Agallia configa rat a Oman 1933. Puerto Rico. Agallia constricta cubana Oman 1933. Cuba Agallia corumba Oman 1938. Brazil. Agallia depleta Oman 1938. South America. Agallia didactylata Oman 1933. Arizona. Agallia dorsata Oman 1938. Argentina. Agallia extensa Oman 1938. Bolivia. Agallia fumida Oman 1938. Colombia. Agallia hyalina Oman. 1938. Brazil. Agallia hystricula Oman 1933. Costa Rica. Agallia incerta Oman 1938. Argentina. Agallia incisa Oman 1938. Brazil. Agallia incongrua Oman 1938. Argentina. Agallia ingens Oman 1933. Central America. Agallia inornata Oman 1938. Brazil. Agallia manaosa Oman 1938. Brazil. Agallia minuenda Oman 1938. Brazil. Agallia munda Oman 1933. West Indies. Agallia neoalbidula Oman 1938. Brazil. Agallia nigerrima Oman 1938. Brazil. Agallia obesa Oman 1933. USA. Agallia pumila Oman 1971. USA. Agallia pene constricta Oman 1933. Honduras. Agallia qnadrata Oman 1938. Brazil, Bolivia. Agallia quadripunctata excavata Oman 1933. USA. Agalliana ancora Oman 1934. Bolivia. Agalliana ensigera Oman 1934. South America. 346 Great Basin Naturalist Memoirs No. 12 Agalliana fusca Oman 1934. South America. Agalliana grossa Oman 1934. Argentina. Agalliana mediana Oman 1938. Argentina. Agalliana puella Oman 1938. Colombia. A galliopsis abietaria Oman 1970. USA. Agalliopsis acuminata Oman 1934. Mexico. Agalliopsis agrestis Oman 1934. Mexico. Agalliopsis ancistra Omin 1970. USA. Agalliopsis ancoralis Oman 1933. Mexico. Agalliopsis atra Oman 1934. Arizona, Mexico. Agalliopsis brunnea Oman 1933. Mexico, Honduras. Agalliopsis cervina Oman 19.3.3. Kansas. Agalliopsis cincta Oman 1933. Florida. Agalliopsis confonnis Oman 1933. Dominican Republic. Agalliopsis disparillis Oman 1938. Colombia. Agalliopsis distincta Oman 1934. Mexico. Agalliopsis dubiosa Oman 1933. Arizona. Agalliopsis elegans Oman 1938. Brazil. Agalliopsis exilis Oman 1933. Colorado. Agalliopsis florida Oman 193.5. Florida. Agalliopsis fuscosignata Oman 1933. Arizona. Agalliopsis fuscosignata minor Oman 1933. Arizona. Agalliopsis gracilis Oman 1934. Mexico. Agalliopsis hamatilis Oman 1970. Mexico. Agalliopsis huachiicae Oman 1933. Arizona. Agalliopsis inscripta Oman 19.34. Guatemala. Agalliopsis longipennis Oman 1934. Mexico. Agalliopsis magnifica Oman 1933. Mexico. Agalliopsis majesta Oman 1934. Mexico. Agalliopsis neopepino Oman 193.5. Cuba. Agalliopsis nervata Oman 1934. Mexico. Agalliopsis novella emulata Oman 1938. Argentina. Agalliopsis novella vicosa Oman 1938. Brazil. Agalliopsis novellina Oman 1935. USA. Agalliopsis oculata Oman 19.33. California. Agalliopsis ornata Oman 1938. Brazil. Agalliopsis ornaticollis Oman 19.38. Argentina. Agalliopsis peneoculata Oman 1933. USA. Agalliopsis peruviana Oman 1938. Peru. Agalliopsis pulchella Oman 1938. Argentina. Agalliopsis punctata Oman 1934. Mexico. Agalliopsis reflexa Oman 1970. Venezuela. Agalliopsis sijnilis Onvdn 1934. Mexico. Agalliopsis sonorensis Oman 1970. Mexico. Agalliopsis Stella Oman 1970. USA. Agalliopsis teapae Oman 1934. Mexico. Agalliopsis tincta Oman 1938. Brazil. Agalliopsis variabilis Oman 19.33. USA. Agalliopsis vonella Oman 19.38. Colombia. Agalliota clavata Oman 1938. Brazil. Agalliota dentata Oman 1938. Brazil. Agalliota parallela Oman 1938. Brazil. Agalliota parma Oman 1938. Brazil. Agalliota scutaria Oman 1938. Argentina. Agelina punctata Oman 1938. South America. Agudus tijpicus Oman 1938. South America Alaca longicauda Oman 1938. Argentina. Attenuiptjga balli Oman 1985. USA. Baldulus montanus Oman 1934. Arizona. Baroma reticulata Oman 19.38. Argentina. Bonamus lineatus Oman 1938. Argentina. Brincadorus laticeps Oman 19.38. South America. Caphodus maculatus Oman 1938. Brazil. Cariancha cariboba Oman 1938. Brazil. Ceratagallia arroija Oman 1939. California. Ceratagallia artemisia Oman 1939. Idaho, Utah. Ceratagallia califa Oman 1939. California. Ceratagallia canona Oman 1939. California. Ceratagallia delta Oman 19.39. California. Ceratagallia loca Oman 1939. California. Ceratagallia loma Oman 19.39. California. Ceratagallia longipes Oman 1939. California. Ceratagallia Ittdora Oman 1939. California. Ceratagallia lupina Oman 1939. California. Ceratagallia neodona Oman 1939. Nevada. Ceratagallia neovata Oman 1939. California. Ceratagallia pera Oman 19.39. California. Ceratagallia pudica Oman 1939. Arizona. Ceratagallia socala Oman 1939. California. Ceratagallia tristis Oman 1939. California. Cortona minuta Oman 1938. Argentina. Cuerna alba Oman & Beamer 1944. New Mexico. Cuerna alta Oman & Beamer 1944. New Mexico. Cuerna arida Oman & Beamer 1944. Arizona. Cuerna balli Oman & Beamer 1944. Arizona. Cuerna curvata Oman & Beamer 1944. California. Cuerna dixiana Oman & Beamer 1944. Utah, Nevada. Cuerna gladiola Oman & Beamer 1944. California. Cuerna mexicana Oman & Beamer 1944. Mexico. Cuerna obesa Oman & Beamer 1944. New Mexico, Texas. 1988 OMAN: Taxa in the Homopteha 347 Cuerna ohtusa Oman & Beanu'r 1944. Arizona. Cuerna occidentalis Oman & Beamer 1944. California. Cuerna yuccae Oman & Beamer 1944. California, Utah, Arizona. Cumora annulata Oman 1938. Argentina. DeltoccpJiaUts