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Full text of "Great Basin naturalist memoirs"

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 Previously unpublished manuscripts in English pertaining to the biological 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 Cpra<aga//ia 

Kirkaldy (Homoptera; Cicadellidae). J. Washing- 
ton Acad. Sci. 29: 529-.543. 

25. Oman, P. W. 1940. Three new species of Delto- 

cephalus (Homoptera: Cicadellidae). Proc. Ento- 
mol. Soc. Washington 42: 201-203. 

26. Oman, P. W. 1941. Revision of the Nearctic Megoph- 

thalminae (Homoptera: Cicadellidae). J. Wash- 
ington Acad. Sci. 31: 203-210. 

27. Oman, P. W. 1943. A new leafliopper from Oceania 

(Homoptera: Cicadellidae). Pan-Pac. Entomol. 
19: 33-34. 

28. Oman, P. W. 1943. A new leafliopper of the genus 

Helochara (Homoptera: Cicadellidae). Proc. En- 
tomol. Soc. Washington 43: 74-75. 

29. Oman, P. W., and R. H. Reamer. 1944. Some new 

species of Cuerna (Homoptera: Cicadellidae). J. 
Kansas Entomol. Soc. 17: 119-128. 
.30. Oman, P. W., and Arthur D. Cushman. 1946. Collec- 
tion and preservation of insects. U.S. Dept. Agric. 
Misc. Publ. 601. 42 pp. 

31. Oman, P. W. 1947. The types of auchenorrhynchous 

Homoptera in the Iowa State College collection. 
Iowa State College J. Sci. 21: 161-228. 

32. Oman, P. W. 1947. Miscellaneous notes on Cicadelli- 

dae (Homoptera). J. New York Entomol. Soc. .55: 
.59-63. 

.33. Rlack, L. M., and P. W. Oman. 1947. Parthenogene- 
sis in a leafhopper, Agallia quadripunctata (Pro- 
vancher) (Homoptera: Cicadellidae). Proc. Ento- 
mol. Soc. Washington 49: 19-20. 

.34. Oman, P. W., and L. C. Christenson. 1947. Malaria 
and other insect-borne diseases in the South 
Pacific campaign, 1942-1945. III. Entomology, 
Amer. J. Trop. Med. 27: 91-117. 

.35. Oman, P. W. 1948. Notes on the beet leafhopper, 
Circiilifer tenellus (Raker), and its relatives (Ho- 
moptera: Cicadellidae). J. Kansas Entomol. Soc. 
21: 10-14. 

36. Oman, P. W. 1948. Distribution of Baldulus maidis 
(DeLong and Wolcott) (Homoptera: Cicadellidae). 
Proc. Washington Entomol. Soc. .50: 34. 

.37. Oman, P. W. 1949. A leafhopper injurious to culti- 
vated prune in the western United States. J. Econ. 
Entomol. 41:983. 

.38. Oman, P. W. 1949. The Nearctic leafhopper (Ho- 
moptera: Cicadellidae). A generic classification 
and check list. Memoir No. 3, Entomol. Soc. 
Washington. 2.53 pp. 

.39. Raker, W. L., P. W. Oman, and Thaddeus J. Parr. 
19.50. Sucking insects. Pages 105-152 in Craig- 
head, Insect enemies of eastern forests. USDA 
Misc. Publ. 6.57. 

40. Oman, P. W. 1951. Whv the flv? Pages 24-25 in Med. 

Rull. No. Ill, Vol. I, Hdcjrs. 3rd Army. 

41. Oman, P. W. 1952. How to collect and preserve 

insects for study. Pages 65-78 in Insects. U.S. 
Department of Agriculture Year Rook. 

42. Oman, P. W. 1952. Three new Errhontiis, with a key 

to the species (Homoptera: Cicadellidae). Pan- 



Pac. Entomol. 23: 13-15. 

43. Oman, P. W. 19.55. Review of. J. Linsley Gressit, 

Insects of Micronesia, introduction. Rull. Rrook- 
lyn Entomol. Soc. .50: 75-76. 

44. Oman, P. W. 19.57. Under separate cover — . Agric. 

Chem. 12: 33-34, 93, 95. 

45. Oman, P. W. 19.57. The relation of insect taxonomy to 

mosquito control. Mosq. News 17: 147-151. 

46. Oman, P. W. 19.59. Strength through union. Rull. 

Entomol. Soc. Amer. 5: 1-2. 

47. Oman, P. W. 1960. The relation of insect taxonomy to 

applied biology. Rull. Entomol. Soc. Amer. 6: 
3-5. 

48. Oman, P. W. 1961. What insects are positively photo- 

sensitive? (Summary). Pages 33-34 in Response of 
insects to induced light. ARS 20-10, July. 

49. Oman, P. W., Lev. F. Curl, Theodore R. Davich, 

Herbert L. Haller, Halbert M. Harris, Carl R. 
Huffaker, and John V. Osmun. 1961. Entomology 
in the Soviet Lhiion, report of a technical study 
group. USDA, ARS. 24 pp. 

50. Harper, P. A., W. G. Downs, P. W. Oman, and N. D. 

Levine. 1963. Chapter VIII, New Hebrides, 
Solomon Islands, Saint Matthias Group, and 
Ryukyu Island. Pages 399-495 in Preventive 
medicine in World War II. Vol. VI, Communica- 
ble diseases. Office of the Surgeon General, De- 
partment of the Army, Washington, D.C. 

51. Oman, P. 1964. World problems in entomology in 

"Farmer's World." Pages .546-5.54 in U.S. Depart- 
ment of Agriculture Year Rook. 

.52. Maramorosch, K., and P. Oman. 1966. U.S. -Japan 
joint conference on arthropod-borne plant virus- 
es. Riosci. 16: 608-610. 

.53. Oman, P. 1966. Current trends and broad objectives 
of USDA Entomology Research Division Pro- 
gram. Proc. North Central Rranch, Entomol. Soc. 
Amer. 21: 13-19. 

54. Oman P. 1966. Facts and concepts of insect control 

without pesticides. Pages 52-62 in C. W. Mc- 
Conib, Minutes of the Eastern Plant Roard 4Ist 
Annual Meeting. Virginia Reach, Virginia, 26-30 
April 1966. 

55. Oman, P. 1968. Prevention, surveillance and man- 

agement of invading pest insects. Rull. Entomol. 
Soc. Amer. 14: 98-102. 

56. Oman, P., and K. V. Krombein. 1968. Systematic 

entomology: distribution of insects in the Pacific. 
Science 161: 78-79. 

57. Oman, P. 1969 Criteria of specificit\' in virus-vector 

relationships. Pages 1-22 in K. Maramorosch, 
ed.. Viruses, vectors and vegetation. Interscience 
Publ., John Wiley, xii 4- 666 pp. 

58. Oman, P. 1969. Review of R. G. Fennah, Fulgoroidea 

(Homoptera) from New Caledonia and the Lo\alty 
Islands. Rull. Entomol. Soc. Amer. 15: 2.59-260.' 
.59. Oman, P. 1970. Taxonomy and nomenclature of the 
beet leiifliopper, Circulifer tenellus (Homoptera: 
Cicadellidae). Ann. Entomol. Soc. Amer. 63:507- 
512. 

60. Oman, P. 1970. Leaflioppers of the Agalliopsis no- 

vella complex (Homoptera: Cicadellidae). Proc. 
Entomol. Soc. Washington 72: 1-29. 

61. Oman, P. 1970. Leafhoppers of the A^a//(opsi.s' uari- 

(d>ilis 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, 



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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]. no fuscous spots on vertex. 

1 fuscous spots may be present. 
Character 2. 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. 



= ocellus approximately its diameter 
from eye. 

1 = ocellus more remote. 

--^ male plates unicolorous. 

1 male plates embrowned apically. 

^ male plates elongate. 

1 ^ male plates rounded apically. 

2 = male plates truncate apically. 

= male pygofer with caudal margin 
rounded. 

1 = male pygofer with caudal margin ex- 
tended. 

^ anal tube not reaching pygofer apex. 

1 = anal tube equal to or exceeding 
pygofer apex. 

= male pygofer not embrowned. 

1 = male pygofer embrowned caudoven- 

trally. 
Character 9. = male plates exceeding apex of 
pygofer. 

1 = male plates equal to apex of pygofer. 

2 = male plates not reaching apex of 

pygofer. 
Character 10. = styles not reaching apex of pygofer. 

1 = styles equal to apex of pygofer. 

2 = styles exceed apex of pygofer. 
Character 11. = styles not reaching apex of plates. 

1 = styles equal to apex of plates. 

2 = styles exceed apex of plates. 
Character 12. = styles without lateral flange. 

1 = styles with lateral flange. 

2 = styles with ventral arm thickened. 
Character 13. = connective shorter than style. 

1 = connective as long as or longer than 
style. 
Character 14. = aedeagus with caudal surface 
smooth. 
1 = aedeagus with caudal surface 
troughlike. 
Character 15. = shaft of aedeagus flared in caudal 
aspect. 
1 = shaft even-margined in caudal as- 
pect. 
Character 16. = aedeagus with no basal processes on 
shaft. 
1 = shaft with basal processes. 
Character 17. = aedeagus with caudal margin of shaft 
smooth. 
1 = caudal margin serrate. 
Character 18. = 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/«'/<u.s.s'pi»uwi<.s (squares), /. /iflp/i/s (circles), and/. /i«.sfi(/(/.v (triangles) 



1988 



CwiKLA: The Genus Ileopeltus 



51 





Figs. 23-26. Ileopeltus hapliis, n. sp.: 23, pygofer, left lateral aspect; 24, right style, dorsal aspect; 25, aedeagus, 
ventral aspect; 26, aedeagus and apex of connective, left lateral aspect. 



Color. — Ochraceous, without dark mark- 
ings. Eye grey. 

Male genitalia. — Pygofer with posterior 
margin rounded in lateral aspect, long, dor- 
sally directed process inserted on posteroven- 
tral margin, processes crossing each other in 
posterior aspect. Plate with lateral margin sin- 
uate. Style posteriorly directed, preapical an- 
gle rounded. Aedeagus with base expanded in 
lateral aspect, small flange present on right 
side in posterior aspect. Gonopore large on 
right side at base of shaft. 

Type. — Male holotype bearing the label 
"Surumu, Roraima [Brazil], IX-1966, M. Al- 
varenga and F. M. Oliveira Col." has been 
examined from OSUC. 

Distribution. — Known from the type lo- 
cality (Roraima, Brazil) and a second male 
specimen from Guarico, Venezuela, in 
USNM. This species may be restricted to the 
lowland tropics. 

Ileopeltus blockeri, n. sp. 
Figs. 33-37; Map 4 

Diagnosis. — Ileopeltus blockeri is near 7. 
hastulus and can be separated from it by the 



pygofer processes inserted near the middle in 
dorsal aspect. 

Length. — Male 4.2-4.7 mm, female un- 
known. 

Color. — Ochraceous, without dark mark- 
ings. Eye grey. 

Male genitalia. — Pygofer with posterior 
margin roundly produced, long acute process 
inserted on ventral margin, near middle in 
dorsal aspect, directed dorsally, crossing me- 
dian line in dorsal aspect. Plate with lateral 
margin insinuate, apex narrowly rounded. 
Style linear, apex slightly curved laterally. 
Aedeagus elongate compared to other mem- 
bers of the genus. 

Type. — Male holotype, Venezuela, Guar- 
ico, lOkmeastofCalabozo, 18-III-1982, G. F. 
Hevel and J. F. Hevel collectors. Type de- 
posited in USNM. 

Etymology. — This species is named in 
honor of the noted leafliopper worker, H. 
Derrick Blocker. 

Ileopeltus clavatus , n. sp. 

Figs. 38-42; Map 4 

Diagnosis. — Ileopeltus clavatus can be 



52 



Great Basin Naturalist Memoirs 



No. 12 




Figs. 27-.32. Ueopeltus hastulus (DeLong & Linnavuori): 27, pygofer, left lateral aspect; 28, pygofer, posterior 
aspect; 29, plates and valve, ventral aspect; 30, right style, dorsal aspect; 31, aedeagus, ventral aspect; 32, aedeagus and 
apex of connective. 



separated from other Ueopeltus species by the 
truncate stylar apex that is shghtly constricted 
subapically. 

Length. — Male 4.2-4.4 mm, female 4.5 
mm. 

Color. — Ochraceous, without dark mark- 
ings. Eye reddish brown. 

Male genitalia. — Pygofer with posterior 
margin truncate, stout process inserted on 
posteroventral margin in lateral aspect, apex 
of process crossing median line in dorsal as- 
pect. Plate triangular, lateral margin straight, 
fused to valve along anterior margin. Style 
directed posteriorly, apex truncate, with 
slight subapical constriction, preapical angle 
absent. Aedeagus elongate, only slightly 
asymmetrical. 

Female seventh sternum. — Posterior 
margin broadly convex, without excavations. 

Type. — Male holotype, Brazil, RR [Ro- 
raima], Boa Vista, 27-VII-1952, M. Alvarenga 
collector. One female paratype, same data as 
holotype. One male paratype, Venezuela, 
Guarico, 10 km east of Calabozo, 18-III-1982, 
G. F. HevelandJ. F. Hevel collectors. Holo- 
type and female paratype deposited in UPB 
and male paratype in USNM. 



Etymology. — Latin, clavat (clubbed) 
refers to the shape of the stylar apex. 

Ueopeltus aberrans (Oshorn), n. comb. 

Figs. 43-46; Map 4 

Chlorotettix aherrans Osborn, 1923:72-73. 
Doleranus cruzanus DeLong & Cwikla 1984:726. New 
synonymy 

Diagnosis. — Ueopeltus cruzanus is related 
to /. cuneus and can be separated from it by 
the narrow aedeagal shaft in ventral aspect. 

Length. — Male 5.5-6.0 mm, female un- 
known. 

Color. — Ochraceous. Crown with faint 
brown spot at apex of coronal sulcus, small 
triangular patch approximately halfway be- 
tween sulcus and eye. Pronotum ochraceous 
with irregular, light brown spots on anterior 
margin and six light orange bands running the 
length of pronottuii. Forewing brownish sub- 
h\'aline, dark brown coloration along commis- 
sure on clavus and lew brown spots on middle 
of wing. 

Male genitalia. — Pygofer with posterior 
margin truncate in lateral aspect, large, curved 
spine on posteroventral margin, spine curves 
medially then posteriorly, spines overlapping 



1988 



CwiKLA: The Genus Ileopeltus 



53 





34 





36 



Figs. 33-37. Ileopeltus blockeri, n. sp. ; 33, pygofer, left lateral aspect; 34, pygofer, dorsal aspect; 35, right style; 36, 
aedeagus, ventral aspect; 37, aedeagus and apex of connective, left lateral aspect. 




Map 4. Distribution of Ileopelttts blockeri (squdres), I. c/«ra/us (circles), and/. aZjerrans (triangles) 



54 



Great Basin Naturalist Memoirs 



No. 12 





Figs. 38-42. Ileopeltus clavatus , n. sp. : 38, pygofer, lateral aspect; 39, pygofer, dorsal aspect; 40, right style, dorsal 
aspect; 41, aedeagus, ventral aspect; 42, aedeagus and apex of connective, left lateral aspect. 



at middle. Plate triangular, slightly longer 
than pygofer, lateral margin straight, not ap- 
pearing fused to valve, two rows of setae 
present, one row of macrosetae plus row of 
small setaelike structures on lateral margin. 
Stylar apex elongate, preapical angle absent. 
Aedeagus U-shaped in lateral aspect, acute 
apically. Gonopore basal, on right side of 
shaft. 

Type. — Male holotype bearing the labels 
"Januaria, Minas Gerais, Brazil," "Dec 17, 
1907" and "Carn. Mus. Ace. 3702" was exam- 
ined from CMNH. The holotype oi criizanus 
DeLong & Cwikla from Santa Cruz, Bolivia, 
was examined from OSUC and found to match 
the type of/, aherrans. 

Distribution. — In addition to the type lo- 
cality and Santa Cruz, Bolivia, two males from 
Ceara and Sao Paulo, Brazil, were examined 
from BMNH. The specimen from Sao Paulo 
was collected from grass in a cloud forest. 
Collecting dates included late February and 
May. 

Ileopeltus cuneus (DeLong & 

Martinson), n. comb. 

Figs. 47-50; Map 5 

Chlorotettix cuneus DeLong & Martinson 1974:265. 

Diagnosis. — Ileopeltus cuneus is close to /. 



Cyclops and can be separated from it by the 
asymmetrical aedeagus and the unique 
pygofer process. 

Length. — Male 4.8 mm, female unknown. 

Color. — Ochraceous. Crown with median 
triangular patch at apex of coronal sulcus, faint 
brown spot halfway between sulcus and each 
eye. 

Male genitalia. — Pygofer produced into 
long process, curved medially then dorsally. 
Plate rounded apically, lateral margin slightly 
convex. Stylar apex linear, spatulate, U- 
shaped notch subapically on laterally surface 
in dorsal aspect. Aedeagus slightly expanded 
in posterior aspect, right side with two tooth- 
like projections near base. 

Type. — Male holotype bearing the labels 
"Piracicaba, Sao Paulo, Brazil, 1-4-1966," 
"collr. C. A. Triplehorn," and "Blacklight 
trap" was examined from OSUC. 

Distribution. — Known only from the holo- 
type. 

Ileopeltus Cyclops (Linnawiori), n. comb. 

' Figs. 51-,55; Map 5 

Doleranus cyclops Linnavuori 1959:275-276. 

Diagnosis. — This unique species can be 
separated from other Ileopeltus species by the 
median spot on the crown and the unique 
stylar apex. 



1988 



CwiKLA: The Genus Ileopeltus 



55 




Figs. 43-46. Ileopeltus aberrans (Osborn): 43, pygofer, left lateral aspect; 44, right style, dorsal aspect; 45, aedeagus, 
ventral aspect; 46, aedeagus and apex of connective, left lateral aspect. 



Length. — Male 4.9 mm, female unknown. 

Color. — Ochraceous. Crown with median 
triangular brown patch. Eye gray. 

Male genitalia. — Pygofer with posterior 
margin produced into acute process posteri- 
orly directed then curving medially, sclero- 
tized. Plate short, rounded apically. Stylar 
apex acute, distal third abruptly curved later- 
ally, preapical angle absent. Aedeagus only 
slightly asymmetrical, shaft narrow in poste- 
rior aspect. 

Type. — Male holotype bearing the labels 
"Loreto, Missiones [Lapsus calmi for Mi- 
siones? Possibly from Corrientes], ARC [Ar- 
gentina] XII-4-1931, A.A. Ogoblin" and 
"collected at light" has been examined from 
USNM. 

Distribution. — Known only from the holo- 
type. 

Cladistic Relationships among the Species 

Biological classifications should be based on 
hypothesized relationships estimated by cla- 
distic (or phylogenetic) analyses (Hennig 
1966). Such analyses are dependent upon the 
determination of plesiomorphic and apomor- 
phic character states. Hypotheses of character 



polarity are accomplished by comparing the 
taxa under question with their outgroup. 
Ileopeltus is thought to be closely allied to 
Chlorotettix by nature of both having the 
crown completely microsculptured. Because 
of this single synapomorphy, Chlorotettix is 
considered the sister-group to Ileopeltus . 

Twelve binary characters were chosen for 
the cladistic analysis and are summarized in 
Table 1. The resulting data matrix (Table 2) 
was analyzed by means of phylogenetic analy- 
sis using the parsimony (PAUP) program de- 
veloped by Swofford (1985). 

The paucity of characters for Ileopeltus spe- 
cies, I believe, is the result of a generalized 
reduction in complexity of characters. This 
becomes immediately obvious when the male 
genital structures are examined, e.g., the 
aedeagus without processes. These reduced 
characters most assuredly obscure the inter- 
relationships among the species and possibly 
at higher levels. Even with this reduction, 
most of the characters used in this analysis 
were genitalic and are discussed below. 

The hypothesized cladogram is presented 
in Figure 56. The most primitive lineage con- 
tains the species /. tethys and /. nanocanthus , 



56 



Great Basin Naturalist Memoirs 



No. 12 




Figs. 47-50. lleopettus citneus (DeLong & Martinson): 47, pygofer with anterior portion damaged, left lateral aspect; 
48, right style, dorsal aspect; 49, aedeagus, ventral aspect; 50, aedeagus and ape.x of connective, right lateral aspect. 




Map 5. Distribution oi Ileopeltus cuiwtis (sciuarcs) and /. ctjclops (circk 



1988 



CwiKLA: The Genus Ileopeltus 



57 




Figs. 51-55. Ileopeltus cyclops (Linnavuori): 51, pygofer, left lateral aspect; 52, right style, dorsal aspect; 53, 
aedeagus, ventral aspect; 54, aedeagus and apex of connective, left lateral aspect; 55, head, pronotum, and scutellum, 
dorsal aspect. 




Fig. 56. Hypothesized cladogram for the genus Ileopeltus. Numbers refer to characters listed in Table 1. 



58 



Great Basin Naturalist Memoirs 



No. 12 




Fig. 57. Ileopeltus tethys (Van Duzee), aedeagus, ventral aspect. Fig. 58. /. nanocanthus, n. sp. , aedeagus, ventral 
aspect. 



which share the synapomorphy of the concave 
plate and the style with the preapical angle 
absent. Both characters are homoplastic and 
are found throughout the cladogram. The re- 
maining species have a distinct process. 

Of those Ileopeltus species with prominent 
pygoferal processes, two major lineages are 
hypothesized. One contains those individuals 
with stout pygoferal processes; the other is 
defined by long, heavily sclerotized pro- 
cesses. 

The lineage with stout pygoferal processes 
contains four species. Ileopeltus dorsalus has 
a median black patch on the crown, while the 
remaining three species have the preapical 
angle of the style absent, a homoplastic char- 
acter found throughout the cladogram. 
Ileopeltus haplus and /. spinosus have their 
stylar apex truncate, another homoplastic 
character. 

The lineage that is defined by the long 
sclerotized process is also supported by the 
processes curved medially, a character which 
is later reversed. The lineage itself is repre- 



sented by an unresolved trichotomy contain- 
ing/, hastulus, I. blockeri, and the remaining 
Ileopeltus species. Ileopeltus hastulus is de- 
fined by the autapomorphy of a large gono- 
pore. The remaining four species have the 
preapical angle of the style absent. As men- 
tioned before, this is a homoplastic character. 
Ileopeltus aberrans, I. cy clops, and I. 
cuneus have a median black patch on their 
crown and the lateral margin of their plate 
concave. Both characters are homoplastic. 
Only autapomorphic characters define these 
species. 

Acknowledgments 

I thank the following individuals and their 
associated institutions for allowing me to ex- 
amine specimens under their care: M. D. 
Webb, Department of Entomology, British 
Museum (Natural History), London (BMNH); 
C. W. Young, Section of Entomology, Car- 
negie Museum of Natural History, Pittsburgh 
(CMNH); C. A. Triplehorn, Department of 



1988 



CwiKLA: The Genus Ileopeltus 



59 




Table 1. Characters, presumed plesiomorphic and 
apomorphic states for Ileopeltus species, and numbers 
that refer to character distribution presented in Table 2 
and Fig. 56. 



Fig. 59. Ileopeltus tethys (Van Duzee), aedeagus, right 
lateral aspect. Fig. 60. Ileopeltus nanocanthus, n. sp., 
aedeagus, right lateral aspect. 



Entomology, Ohio State University, Colum- 
bus (OSUC); the late Francisco Jose Fer- 
nandes-Yepez, Facultad de Agronomia, Uni- 
versidad Central de Venezuela, Maracay 
(UCV); Paul H. Freytag, Department of 
Entomology, University of Kentucky, Lex- 
ington (UKC); K. M. Rocha Zanol, Universi- 
dade Federal do Parana, Curitiba, Brazil 
(UPB); J. P. Kramer and R. Froeschner, De- 
partment of Entomology, Smithsonian In- 
stitution, Washington (USNM); H. Wolda, 
Smithsonian Tropical Research Institute, Bal- 
boa, Republic of Panama (HWC). H. Derrick 
Blocker (Kansas State University), P. H. 
Freytag (UKC), Sam Stribling (USNM), and 
G. M. Chippendale (University of Missouri, 
Columbia) kindly criticized an earlier draft of 
this manuscript. Niki K. J. Miller (Ohio State 
University) skillfully executed the habitus 
drawing. Special thanks are extended to Paul 
W. Oman, whose works on leafhopper sys- 
tematics have proved to be most inspiring. 







Plesio- 


Apo- 




Character 


morphic 


morphic 


1. 


Coloration 


unmarked 


crown marked 
with median 
triangular 
patch 


2. 


Pygoferal processes 


absent 


present 


3. 


Pygoferal processes 








elongate 


no 


yes 


4. 


Pygoferal processes 








stout 


no 


yes 


5. 


Pygoferal processes long 








and heavily sclerotized 


no 


yes 


6. 


Pygoferal processes 
strongly curved 








medially 


no 


yes 


7. 


Plates fused to valve 


no 


yes 


8. 


Plates with lateral 








margin 


straight 


concave 


9. 


Style with preapical 








angle 


present 


absent 


10. 


Stylar apex 


stout 


narrow 


11. 


Stylar apex 


rounded 


truncate 


12. 


Gonopore 


small 


large 



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Caldwell, J S , and L F Martorell 1950. Review of 
the Auchenorynchous Homoptera of Puerto Rico. 
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CwiKLA, P. S. 1988. A systematic study of the leafhopper 
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CwiKLA, P S , AND P. H Freytag 1983. External mor- 
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DeLong, DM, and P S. Cwikla 1984. New Neotropi- 
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DeLong, D M , andC Martinson 1974. New species of 
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DeLong, D M , and R E Linnavuorl 1978. Studies on 
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Hennig, W 1966. Phylogenetic systematics. University 
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60 



Great Basin Naturalist Memoirs 



No. 12 



Table 2. Data matrix used in constructing a cladogram for Ileopeltus. Characters number refers to information in 
Table 1: 0, plesiomorphic; 1, apomorphic. 



Character 


1 


2 


3 


4 


5 


6 


7 


8 


9 


10 


11 


12 


Chlorotettix 






































1. cuneus 


1 




1 







1 




1 


1 





1 


1 


I. Cyclops 


1 




1 












1 


1 


1 








I. aherrans 


1 




1 







I 








1 











1. blockeri 







1 







1 



















I. hastulus 







1 







1 
















1 


I. tethijs 






















1 


1 











I. nanocanthus 





















1 


1 











I. haplus 







1 


1 













1 





1 





I. dorsahis 


1 




1 


1 

























1. clavatus 







1 





1 


1 







1 











I. sphwsus 







1 


1 













1 





1 





I. ventriculus 







1 


1 













1 












20: 1-370. 

OSBORN. H 1923. Neotropical Homoptera of the 
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1935. Insects of Puerto Rico and the Virgin Is- 
lands. Homoptera (excepting the Sternorhynchi). 
Puerto Rico and Virgin Islands Sci. Surv. 14: 
111-260. 

Oman, P W 1949. The Nearctic leaflioppers (Homop- 
tera: Cicadellidae); a generic classification and 
check list. Mem. Entomol. Soc. Washington No. 
3. 2.53 pp. 



SwoFFORD, D L 1985. PAUP, phylogenetic analysis us- 
ing parsimony. Users manual. Illinois Natural 
History Survey, Champaign. 

Van Duzee. E P 1907. Notes on Jamaican Hemiptera: a 
report on a collection of Hemiptera made on the 
island of Jamaica in the spring of 1906. Buffalo Soc. 
Nat. Sci. Bull. 8: .3-79. 

WOLCOTT. G N 1923. Insectae Portoricensis. A prelimi- 
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Rico Univ. J. Agric. 7: 1-313. 

1936. Insectae Borinquenses. A revised annotated 

checklist of the insects of Puerto Rico. Puerto Rico 
Univ. J. Agric. 20: 1-600. 



SOME ASPECTS OF THE BIOLOGY, MORPHOLOGY, AND EVOLUTION OF 
LEAFHOPPERS (HOMOPTERA: CICADELLOIDEA AND MEMBRACOIDEA) 

J. W. Evans' 

Abstract — This article summarizes some observations of a varied nature on the biology, morphology, and evolution 
of the Cicadelloidea (Cicadellidae, Hylicidae, Eurymelidae) and Membracoidea(Membracidae, Aetalionidae, Biturri- 
tidae, Nicomiidae). These observations, made over a period of more than half a century, have previously been recorded 
at different times, but lie buried in the literature. It is hoped that their interest will justify repetition and draw attention 
to some promising lines of research. 



Biology 
Food Plant Associations 

As Southwood (1961) has pointed out, in- 
sects have a particularly close association with 
plants belonging to the predominant flora of 
the time. Thus, most Australian cicadelloids 
feed on eucalypts and numerous membracids 
on acacias. Of particular interest are relict 
insects that feed on representatives of ancient 
floras. Examples are Koebelia californica 
Baker on conifers in California and Cornutipo 
spp. (Eurymelidae) on representatives of the 
Proteaceae in Australia. The Cephalelini, a 
tribe of the Ulopinae which have an Antarctic 
distribution, provide another example, for 
their feeding seems to be restricted to an an- 
cient family of plants, the Restionaceae, in 
Australia, South Africa, and New Zealand. 

Changes of food plants have recently oc- 
curred in the Tartessinae (Cicadellidae). 
These are primarily eucalypt feeders, but 
some have become adapted to living in tropi- 
cal rain forests and have taken to feeding on 
other kinds of trees (F. Evans 1981). 

Sound Production and Courtship 

Ossiannilsson, in his notable study of the 
"songs" of Homoptera, was the first to draw 
attention to the presence of tymbals in insects 
of both sexes belonging to the Cicadelloidea, 
Cercopoidea, and Cicadoidea (Ossiannilsson 
1949). Previously, I had noted the presence of 
structures that I described as "resembling 
tymbals" on the first abdominal segment of 
Darthula hardwickii Gray and Aetalion retic- 



ulatum Linnaeus (Evans 1946b). In his discus- 
sion of the function of the songs of various 
Auchenorrhyncha, Ossiannilsson described 
some as being "calls of courtship." Subse- 
quently, I noted the presence of well-devel- 
oped tymbals in nymphs belonging to every 
instar but the first in both sexes of A. reticida- 
tiim (Evans 1957). As, presumably, nymphal 
song cannot have a sexual significance, it may 
serve an assembling purpose since aetalionids 
are gregarious insects. However, amongst the 
Eurymelidae, which are likewise gregarious, 
I have been unable to find any trace of tym- 
bals. Nevertheless, prior to mating, eu- 
rymelids indulge in a prolonged courtship 
(Evans 1931). 

Tymbal sound production occurs among 
representatives of all the superfamilies of the 
Auchenorrhyncha, and even in the 
Cicadoidea it is not confined to one sex since 
females of Tettigarcta can "sing." Thus, it 
must have been a feature acquired in early 
Mesozoic, if not Palaeozoic, times (Evans 
1941). Moreover, it would seem that during 
the early Mesozoic some Auchenorrhyncha 
were capable of stridulation. This is supposed 
because some Upper Triassic Homoptera 
(e.g. , Eoscartoides bryani Evans) have stridu- 
latory areas on the proximal costal area of their 
tegmina (Evans 1961). 

Oviposition 

When ovipositing, the majority of leafhop- 
pers insert their eggs into plant tissue, either 
singly or in batches. In the latter case, the 
eggs are covered with secretions that harden 



'Australian Museum, Sydney, N.S.W., Australia. Present address; 47 Bundarra Road, Bellevue Hill, Sydney 2023, Australia. 



61 



62 



Great Basin Naturalist Memoirs 



No. 12 



on exposure to the air. The eggs of aetahonids 
and of some bitturitids and membracids are 
contained in oothecae situated on the surface 
of the plants. Because the AetaHonidae are an 
ancient group of insects, it might be thought 
that such a method of oviposition predated 
egg insertion in plant tissue. However, this is 
improbable, as the last-named method is 
shared with insects belonging to the super- 
families of the Auchenorrhyncha; and eggs, 
whether laid inside plants or on their surface, 
are equally subject to heavy parasitism. 

Gregariousness and Ant Attendance 

While all eurymelids and aetalionids, many 
membracids, and possibly all bitturitids are 
gregarious, so far as I am aware, no cicadellids 
have this behavior pattern. Since female 
membracids sometimes remain with their 
eggs after oviposition, some authors, such as 
Haviland (1925), have credited them with ma- 
ternal care. Eurymelids of both sexes have 
been recorded as remaining with their eggs 
for a period. It is possible that the gregarious 
habit in this family may be associated with the 
fact that their nymphs, unlike those of other 
leafhoppers, lack the ability to jump. 

It would seem that all gregarious leafliop- 
pers are ant attended, but the Pogonoscopini, 
a tribe of the Eurymelidae, are the only ones 
that have developed the characteristics of true 
myrmecophiles; i.e., they have unusually 
long legs and small eyes and live in the nests of 
ants (Evans 1966). 

Morphology 
Color and Size 

Previously, 1 have suggested that early 
leafhoppers were brown, as, for instance, is 
Darthula hardwickii , and that green was the 
first alternative color to be acquired (Evans 
1966). Later evolutionary developments have 
given rise to insects that may be predomi- 
nantly black and have, or lack, yellow, red, or 
occasionally bluish markings. Some Cicadelli- 
dae from Madagascar have striking color pat- 
tern differences (Evans 1953). In others, spe- 
cies occur that have identical male genitalia 
and comprise populations with differing, but 
stable, color patterns (e.g., Eunjmela fenes- 
trata Le Peletier & Serville) (Evans 1933). 
Then, some species have individuals with a 
very variable color pattern (e.g., Euryme- 



loides punctata [Signoret]) (Evans 1966). 

The largest known leaffiopper, the aetal- 
ionid Darthula hardwickii Gray, has a length 
of 28 mm, of which the apical 12 mm in both 
sexes consists of the prolonged ninth abdomi- 
nal segment. The smallest leafhoppers are 
comprised in the Typhlocybinae (Cicadelli- 
dae), some species of which are no more than 
2 mm long. Evolutionary development seems 
to be frequently accompanied by increase in 
size. Thus, for example, in the Eurymelidae, 
the largest species are seemingly the most 
recently evolved (e.g., Eunjmelops generosa 
[Stal]). 

The two sexes of Cicadelloidea usually re- 
semble each other in coloration and size. An 
exception in respect to color is provided by 
the sole representative of the Tartessinae to 
occur in New Caledonia. This insect, Calo- 
tartessus stalii (Signoret), has males that are 
largely black and females that are predomi- 
nantly brown. In regard to size, the Stenoco- 
tini (Ledrinae) have males that are consider- 
ably smaller than females. 

The Head 

The most puzzling feature of the heads of 
Hemiptera is associated with the origin of 
their feeding apparatus. Insects in some other 
orders feed by suction, but with all these the 
nature of the transition from mandibulate to 
haustellate mouthparts can be readily under- 
stood. 

Previously, 1 have suggested that, though 
the ocelli are on the crown in representatives 
of the relict subfamily, the Ulopinae, this con- 
dition is a secondary one. If this is correct, 
even though insects with ventral ocelli do not 
seem to be disadvantaged, such a change of 
position would seem to be an adaptive one. 

Other primitive features of leaffiopper 
heads are the presence of a sensory pit on the 
maxillary plates, which may possibly be 
derived from the maxillary palp (Evans 1973), 
and the complete separation of the maxillary 
plates from the genae. 

Male Genitalia 

The male genitalia of insects are more sub- 
ject to change than any other parts of their 
bodies. For this reason, they are extensively 
used for species recognition. The nature and 
the extent of observable differences between 
the aedeagi of closely related insects vary 



1988 



EvANS; Leafhopper Biology, Morphology, and Evolution 



63 



widely and range from slight and constant to 
considerable and variable ones. The subgeni- 
tal plates and accessory processes are also sub- 
ject to considerable change of shape. 

While male genitalia differences are useful 
for species separation and genus recognition, 
they are sometimes also helpful for family 
recognition purposes. Thus, for example, 
while within the Cicadellidae the aedeagus 
invariably arises from a basal connective situ- 
ated between the paired parameres, in the 
Eurymelidae it lacks any association with the 
basal plate and the parameres. 

Inasmuch as simple male genitalia serve the 
same purpose as complex ones and pre- 
sumably in an equally satisfactory manner, 
and the structure of female genitalia is con- 
stant, it is difficult to understand the nature of 
the advantages conferred by increasingly 
complex male genitalia unless female insects 
avoid mating with males that have the wrong 
"key. " 

Adaptive Characters 

In addition to the ocelli noted above, other 
adaptive characters among leaflioppers are to 
be found in the Stenocotini, an endemic Aus- 
tralian subfamily of the Ledrinae, and in the 
Eurymelidae. Stenocotids, which live under 
bark of eucalypts, are flattened insects, and 
their nymphs are paper thin. While most eu- 
rymelids are wedge-shaped, one species, 
Platyeurymela semifascia (Walker), which, 
like stenocotids, lives under bark, is oval in 
shape and convex. 

Alary dimorphism and polymorphism are of 
widespread occurrence in the Heteroptera, 
they are rare in the Homoptera-Auchenor- 
rhyncha, and in the Cicadellidae they would 
seem to be particularly associated with insects 
living at high altitudes or in a marsh environ- 
ment. Thus, Monteithia spp. (Monteithiini, 
Ulopinae), which live at high altitudes in New 
Guinea, and of which fully winged insects 
have not been recorded, have males that dif- 
fer from females in the extent to which their 
wings are foreshortened (Evans 1968). Then, 
in Cephalelini, which inhabit both alpine and 
low-level marsh environments, both sexes oc- 
cur in a flightless and a fully winged condition. 
Some other leaflioppers with both brachyp- 
terous and fully winged individuals are 
Taslopa montana Evans (Ulopini, Ulopinae), 
Chiasmus varicolor (Kirkaldy) (Deltocephali- 



nae), and Euacanthella sp. (Euacanthellinae). 

The best-known examples of the acquisition 
of morphological features among leaflioppers, 
which are seemingly of nonadaptive signifi- 
cance, are to be found among the Mem- 
bracidae. This phenomenon has been dis- 
cussed by many authors (e.g., Haviland 1925) 
who observed these insects in the territory 
then known as British Guiana. She pointed 
out that the early stages of exaggerated prono- 
tal development in insects, which, in their 
extreme form, were either cryptic or mimetic, 
cannot have conferred any protective advan- 
tage, and yet the insects survived. Moreover, 
the pronota of many membracids seem to lack 
any degree of protective significance. 

Above I have suggested that the extreme 
differences found among leaflioppers belong- 
ing to the genus Colloborrhis Germar in 
Madagascar may have been initiated by 
"explosive speciation." I have also formerly 
suggested that the same phenomenon may 
have occurred among Membracidae isolated 
in the Neotropical Region during Tertiary 
times (Evans 1959). This is because, though 
doubtless enlarged pronota were a mem- 
bracid characteristic before this isolation took 
place, it is in South America only that the 
evolution of such a range of bizarre forms took 
place. 

The development in leaflioppers of bizarre 
structures, which apparently lack adaptive 
significance, is not confined to changes in the 
shape of the prothorax. These occur also in the 
heads of leaflioppers, as, for example, in those 
of Cornutipo tricornis Evans, Listrophora 
evansi Evans, and Wolfella caternaulti 
Spinola (all illustrated in Evans 1975a). 

Evolution 
Fossils and the Classification of Recent Forms 

An abundance of wings of Homoptera has 
been found in Permian and Triassic strata in 
both the northern and southern hemispheres. 
These seem to provide evidence that all the 
existing superfamilies of the Auchenorrhyn- 
clia were already differentiated by Triassic 
and possibly Permian times. If this suggestion 
is valid, it provides an example of unusual 
evolutionary stability (Evans 1964). 

In the forewings, or tegmina, of some 
Upper Permian Homoptera (e.g., Homa- 
loscytina plana Tillyard) six veins support 



64 



Great Basin Naturalist Memoirs 



No. 12 



their apices. These are as follows: Rib, Rs, 
Ml, M2, M3, and M4. In some recent Ci- 
cadelloidea, only four veins serve the same 
purpose, and it is of interest to note that the 
identity of these is different in each of the 
comprised families. Thus, in the Hylicidae, 
they are Rib, Rs, Ml + 2, and M3+4 + CuA; 
in the Eurymelidae, Rl, Ml + 2, M3+4, 
and CuA; and in the Cicadellidae, Rib, 
Rs + Ml+2, M3 + 4, and CuA. This circum- 
stance would seem to provide evidence of the 
separate, direct derivation of insects com- 
prised in the leafhopper families from a com- 
mon ancestral stock (Evans 1949, 1964). The 
recently found fossil insect placed in a new 
Homoptera family, the Jascopidae, belongs in 
my opinion to the Cicadellidae (Evans 1972). 

Parallel Evolution 

The most striking example of parallel evolu- 
tion in the family Cicadellidae is provided by 
the resemblance between insects comprised 
in the Cephalelini (Ulopinae) and the Para- 
dorydiini (Hecalinae). Leafhoppers in both 
tribes have species with short, triangular 
heads and others with long, narrow ones, the 
complete insect being seedlike in appearance. 
While the Paradorydiini, unlike the Cephale- 
lini, do not feed on the rushlike Restionaceae, 
some have been recorded from "rushes ' grow- 
ing in marshes. The Ulopinae and the Hecali- 
nae are not closely related and probably were 
differentiated during different geological pe- 
riods, the former being of Mesozoic and the 
latter possibly Tertiary origin. 

Explosive Speciation 

Above I have mentioned an instance of sup- 
posed "explosive speciation that has occurred 
in Madagascar with leafhoppers belonging to 
the genus Coloborrhis Germar (Ulopini, Ulo- 
pinae). The type species of the genus, C. cor- 
ticina Germar, has an extensive distribution 
in Africa from where no other species has 
been recorded. It is established also in Mada- 
gascar, from where no less than 16 other spe- 
cies, which could equally well be regarded as 
belonging to separate genera, have been de- 
scribed. These differ strikingly from one an- 
other in characters of the head and thorax. 
Thus, the head may be rounded anteriorly, or 
narrowly, or broadly spatulate, and it may 
have differently shaped prominences on the 
crown. Then, the pronotum may have a pair of 



lateral prominences of varying size, or be lat- 
erally humped, and it may be widest either 
anteriorly or posteriorly, while the scutellum 
may be of normal size or raised into a large, 
vertical crest. The tegmina may have basic 
cicadellid, or reticulate, venation. One spe- 
cies (C. rugosa Evans) has a characteristic 
shared only with the tegmina of Permian and 
Triassic Homoptera. This is the proximal 
arching of vein CuA so that it makes contact 
with vein M. Finally, the hind tibae may be 
narrowly rectangular in section, or broadly 
spatulate. It is of interest to note that some of 
the characteristics listed above are shared 
with the Ledrinae, while others are of a mem- 
bracidlike nature (Evans 1953, 1959). 

Supposed Sympatric Speciation 

During 1959 a symposium was held in Mel- 
bourne to celebrate the centenary of the Royal 
Society of Victoria. Its proceedings were later 
published under the title "Evolution of living 
organisms." At the end of the paper I con- 
tributed to this symposium (Evans 1962), I 
said: 

The concept of sympatric speciation is at the present 
time ahnost universally discredited and any mention of it, 
even as a possibility, might seem to have no place in a 
contribution purporting to be scientific. This is especially 
so when, as in the case of the present instance, a sugges- 
tion is made that is unsupported by experimental evi- 
dence. Nevertheless, in my opinion, it is possible that 
sympatric speciation may take place within a particular 
group of leaflioppers, the Typhlocybinae, and the reason 
for this opinion is because amongst these insects disconti- 
nuities between populations of an ecological and ethologi- 
cal nature would seem to be more readily capable of 
achievement than isolation of a geographical nature. 

Typhlocybids, which are of world-wide distribution, 
are particularly well represented in the Holarctic Region. 
They range in length from 2-4 mm. This group comprises 
some hundred genera, several of which contain many 
hundreds of distinct species. These leaflioppers differ 
greatly in the extent to which they are restricted in their 
feeding requirements. Some feed, mate and breed only 
on a single species of plant or on a limited range ol related 
plants. Others have wider feeding habits. The mere fact 
that related species have different ranges of food plants 
implies an ability, on occasion, to become accommodated 
to a new diet. 

Mention has already been made of the songs of Ho- 
moptera and it may be of significance that typhlocN bids 
have unusually large apodemes for the support of their 
tymbal muscles and hence, presumably, are particularK' 
vocal. It is accordingly suggested that an isolating factor 
which may have enabled the evolution of large numbers 
of sympatric species might be the s\nchronization of the 
ac(iuisition of a new call note with that of a new food plant. 

The reason it has been suggested that it is improbable 



1988 



EVANS: LEAFHOPPER BiOLOCJY , MOKPMOLOGY, AND EVOLUTION 



65 



that geographical isohition has been tlie principal factor 
enabhng prohfic speciation to take place among typhlocv - 
bids is because their small size and swarming habits make 
them particularly liable to transport in the upper air and 
hence to rapid and wide dispersal. 

My views on the above matter have not 
changed during the 25 years that have elapsed 
since I first expressed them. 

Some Characteristics of 
Island and Montane Faunas 

When the gene flow of insects is restricted 
following the isolation of small initial popula- 
tions, the process of evolution is often acceler- 
ated. Mention has already been made of de- 
velopments that have occurred on the island 
of Madagascar following the presumed isola- 
tion of a population of the ulopid Coloborrhis 
corticina . The occurrence of aberrant leaf- 
hoppers is not confined to large islands. Thus, 
on Juan Fernandez Island there occurs possi- 
bly the most grotesque of all leafhoppers, 
Evansiola kiischeli (China). The head of this 
species is twice as wide as long, and, when 
observed from above, projects laterally con- 
siderably beyond the sides of the rest of the 
body (Evans 1975a). 

Montane faunas in Australia and New 
Guinea comprise survivors from a time when 
prevailing climates were cooler than at pres- 
ent and from immigrant species belonging to 
later evolved groups. Examples belonging to 
the first category are provided by Taslopa 
montana Evans (Ulopinae, Ulopini), which 
has been recorded only from high altitudes in 
Tasmania and southeastern Australia, and 
Monteithia spp., which inhabit high-level en- 
vironments in New Guinea (Evans 1966, 
1968). A leafhopper in the second category is 
Atistroagalloides rosea Evans (Austroagal- 
loidinae) (Evans 1966). 

Relict Characteristics 

The relict characteristics of leafhoppers can 
be recognized by making a comparison of 
those features in which presumed recently 
evolved forms differ from those of presumed 
earlier origin. 

The head. — Below are given particulars of 
how recent Deltocephalinae differ in head 
structure from the ancient aetalionid, 
Darthula hardwickii. Not only are the Del- 
tocephalinae probably the most recently 
evolved of all cicadellids, but they are also the 



most widely distributed and contain the 
largest number of species. The characteristics 
of representatives of this subfamily, given be- 
low, precede those of D. hardwickii, which 
are given in parentheses. Wide maxillary 
plates with sensory pits, continuous with the 
genae (narrow maxillary plates with sensory 
pits, separated from the genae by a subgenal 
suture); hind margin of lora widely separated 
from the antennal bases (hind margins of lora 
adjacent to the antennal bases); frontoclypeus 
lacking any trace of epistomal suture and ex- 
tending posteriorly as far as the hind margin of 
the face (frons separated from the postclypeus 
by an epistomal suture anteriorly and the 
arms of the epicranial suture posteriorly); 
ocelli situated near the sides of the fronto- 
clypeus and close to, or on, the margin of the 
head separating the face from the crown 
(ocelli on the face of the head close to the sides 
of the epicranial suture and at a considerable 
distance from the hind margin of the face) 
(Evans 1975a). 

The thorax. — Some supposed relict char- 
acters are the small pronotal paranota, such as 
occur in the Myerslopiini (Ulopinae), and ex- 
tensions of the costal region of the tegminal 
pads of the nymphs of some ulopids, such as 
those of Coloborrhis corticina and also a few 
macropsids (Evans 1968). These paranota and 
the mesothorasic costal expansions are re- 
garded as relict features because the former 
were possessed by many Palaeozoic insects 
and the latter occur also in the Psylloidea, and 
in some Cicadelloidea and Membracoidea, 
which, for other reasons, are regarded as 
relict forms. 

The abdomen. — Among relict characters is 
the retention of tymbals in the nymphs as well 
as in the adults of some aetalionids, and the 
presence in the male genitalia of bisegmented 
subgenital plates. I have suggested that the 
latter, which occur in some Ulopinae, 
Macropsinae, and Agallinae, represent the 
gonocoxites and the gonostyli of the ninth ab- 
dominal segment (Evans 1975). 

Conclusions 

The observations recorded in this article 
serve to emphasize that the factors responsi- 
ble for evolutionary change are of varied na- 
ture. Some, involving major changes of struc- 
tural organization, present puzzling features. 



66 



Great Basin Naturalist Memoirs 



No. 12 



Others suggest that evolutionary develop- 
ments need not always be of an adaptive na- 
ture, that isolation of populations need not 
always be of a geographical nature, and that 
the development of bizarre forms is particu- 
larly associated with isolation of populations 
on islands. 

Literature Cited 

Evans, F. 1981. The Tartessinae of Australia and New 

Guinea (Homoptera, Cicadellidae). Pacificlnsects 

23: 112-188. 
Evans, J. W 1931. Notes on the biology and morphology 

of the Eurymelinae. Proc. Linn. Soc. N.S.W. 56: 

210-226. 
1933. A revision of the Eurymelini (Homoptera, 

Bythoscopidae). Trans. R. Soc. S. Australia 57: 

73-90. 
1941. The morphology of Tettigarcta toinentosa 

White. Pap. Roy. Soc. Tasmania 1940; 35-49. 
1946a. A natural classification of lealhoppers Qas- 

soidea, Homoptera) Pt. I. Trans. R. Soc. London 

96: 47-60. 
1946b. A natural classification of lealhoppers Pt. 

II. Trans. R. Soc. London 97: 39-54. 
1953. Les Cicadellidae de Madagascar. Mem. 

Inst. Sci. Madagascar E. 4: 87-137. 
1957. Some aspects of the morphology and inter- 
relationships of extinct and recent Homoptera. 

Trans. R. Entomol. Soc. London 169: 275-294. 
1959. Quelques nouveaux Cicadellides de Mada- 



gascar. Mem. Inst. Sci. Madagascar E. 11: 481- 
507. 

1961. Some Upper Triassic Heniiptera from 

Queensland. Mem. Queensland Mus. 14: 13-23. 

1962. Evolution in the Homoptera. Pages 250- 

259 in The evolution of living organisms. Mel- 
bourne University Press. 

1964. The periods of origin and diversification of 

the superfamilies of the Auchenorrhyncha as de- 
termined by a study of the wings of Palaeozoic and 
Mesozoic fossils. Proc. Linn. Soc. London 175: 
171-181. 

1966. The lealhoppers and froghoppers of Aus- 
tralia and New Zealand. Mem. Australian Mus. 
12: 1-347. 

1968. Some relict New Guinea lealhoppers and 

their significance in relation to the comparative 
morphology of the head and thorax of the Ho- 
moptera-Auchenorrhvncha. Pacific Insects 10: 
215-229. 

1972. Some remarks on the family Jascopidae (Ho- 
moptera, Auchenorrhyncha). Psyche 79: 120-121. 

1973. The maxillary plate of Homoptera- Auchen- 
orrhyncha. J. Entomol. 48: 43-47. 

197.5a. The external features of the heads of 

lealhoppers. Rec. Australian Mus. 29: 407-439. 

1975b. The structure, function and origins of the 

subgenital plates of leafhoppers. J. Australian En- 
tomol. Soc. 14: 77-80. 

Haviland, M D 1925. The Membracidae of Kartabo, 
British Guiana. Zoologica 6: 239-290. 

OssiANNlLSSON, F 1949. Insect drummers. Opusc. Ento- 
mol. Suppl. 10: 1-142. 

SouTHWOOD, T R E 1961. The numbers of species of 
insects associated with various trees. J. Anim. 
Ecol. 30: 1-8. 



REVISION OF THE GENUS CALLISCARTA (HOMOPTERA: 
CICADELLIDAE: NEOBALINAE)' 

Paul H. Freytag^ 

Abstract. — A taxonomic revision of the Neotropical genus Calliscarta Stal is presented. Most of the 19 species 
treated are described and illustrated, including 10 that are new to science. The new species include: ahrupta (Bolivia), 
acuta (Peru, Venezuela), delicata (Bolivia, Peru), elon^ata (Ecuador, Peru), expansa (Bolivia, Ecuador, Peru), lora 
(Peru), iimrginata (Honduras), inexicana (Mexico), ornata (French Guiana, Venezuela), riig,osa (Venezuela). Three 
cases of new synonymy are proposed; boliviana (Osborn) (- tiixfia Kramer), /a.sctafa (Oshorn) [= brunnea (Osborn)], 
decora (Fabricius) [= magnifica (Osborn)]. 



The genus Calliscarta as reviewed by Lin- 
navuori (1959) included nine species. Two 
species and the genus Idiotettix Osborn were 
placed in synonymy. Since then Linnavuori 
and Heller (1961) added one species, and 
Kramer (1963) added two species. In this 
study nine of these species are recognized as 
valid, three are placed in synonymy, and 10 
new species are added. The limited number 
of specimens in collections of this genus make 
it a very difficult genus with which to deal. No 
division of this genus is made at this time even 
though some species groups appear to repre- 
sent separate genera, such as the fasciata 
group. Much more material will be needed, 
especially from Central America, to know how 
to properly place many species. The problem 
of having only one sex of the majority of the 
species also makes it very difficult to under- 
stand relationships. For these reasons a phy- 
logenetic study must wait until the species are 
better known. 

Calliscarta Stal 

Calliscarta Stal 1869: 82 (type-species Cicada decora 
Fabricius); Dallas 1870: 497; Van Duzee 1890: 35; 
Van Duzee 1890: 79; Schulze, Kukenthal, and 
Heider 1927: 504; Neave 1939: 537; Evans 1947: 
230; Linnavuori 1959: 27; Linnavuori and Heller 
1961:3;Metcalfl967:946. 

Idiotettix Osborn 1929: 465 [type-species Thamnotettix 
magnificus Osborn { = Cicada decora Fabricius)] 
Oman 1938: 355; Neave 1939: 763; Metcalf 1944 
161; Evans 1947: 192; Evans 1951: 9; Nast 1952: 1 
Maldonado Capriles 1954: 247; Metcalf 1966: 225. 

Large, robust, wedge shaped. Head wider 
than pronotum; crown wide, same length near 



eyes as median, coronal suture short; ocelli on 
the margin about twice their width from eyes, 
face elongate; postclypeus convex and long, 
sutures along each side starting near ocelli, 
with antennal ledges from postclypeus to eye. 
Pronotum convex, lateral margins short, pos- 
terior margin slightly but evenly emarginate. 
Forewing truncate, appendix well developed, 
with three subapical cells. Femur of hind leg 
with spinulation 2-2-1. Male genitalia large 
and mostly sclerotized. Anal tube long, cylin- 
drical, heavily sclerotized. Pygofer always 
with a membranous ventral process that ex- 
tends into the genital capsule, or with a long 
ventral process. 

This genus has been previously placed in 
the subfamily Idiocerinae on the basis of the 
overall shape of the body and the large ap- 
pendix. It has also been placed in the subfam- 
ily Deltocephalinae, as the ocelli are on the 
margin of the head near the eyes. These and 
other characters, plus the fact that the major- 
ity of the species are brightly colored and 
larger than those found in the above-men- 
tioned subfamilies, caused Linnavuori (1959) 
to place this genus in the subfamily Neobali- 



Key to Known Males 

1. Aedeagus with extra dorsal apodemal process 
(Figs. 56, 57); pygofer with large ventral process 
(Fig. 55) fasciata (Osborn) 

— Aedeagus without extra apodemal process; 

pygofer with some type of apical process 2 

2(1). Aedeagus with shaft laterally compressed, knife- 
shaped, and with or without small lateral pro- 
cesses (Figs. 10, 11, 14, 15) 3 



'This paper is published with the approval of the Director of the Kentucky Agricultural Experiment Station as journal article No. 88-7-39. 
Department of Entomology, University of Kentucky, Lexington, Kentucky 40546-0091. 



67 



68 



Great Basin Naturalist Memoirs 



No. 12 



— Aedeagus with shaft tubular (Figs. 24, 2.5) . . 10 
3(2). Style with a lateral subapical tooth (Fig. 4) . . . 4 

— Style without subapical tooth 5 

4(3). Pygofer truncate with a small apical process 

(Fig. 1) decora (Fabricius) 

— Pygofer gradually narrowing to bluntly 
pointed apex, median of ventral margin with 

two small teeth 

tartessoides Linnavuori & Heller 

5(3). Pygofer with apical process bifurcate (Figs. 

13, 17, 18) boliviana (Osborn) 

— Pygofer with apical process not bifurcate (Fig. 
19) 6 

6. Pygofer with apical process dorsally produced 
and pointed (Fig. 5) 7 

— Pygofer with apical process bird-head-shaped 

(Fig. 9) 8 

7. Style thickened to near apex (Fig. 8); aedea- 
gus with small lateral processes (Figs. 6, 7) . 
elongata. n. sp. 

— Style narrowing to pointed apex (Fig. 46); 
aedeagus with conspicuous processes (Figs. 
44, 45) delicata, n. sp. 

8. Aedeagus with conspicuous processes (Figs. 
48, 49) ornata, n. sp. 

— Aedeagus without processes (Figs. 10, 11) . . . 9 

9. Style long and abruptly curved subapically 

(Fig. 12) invita (Nast) 

— Style short and evenly curved near apex (Fig. 
22) lora, n. sp. 

10(2). Style foot-shaped at apex (Fig. 26) 11 

— Style pointed at apex (Fig. 42) 13 

11(10). Aedeagus with shaft expanded at apex (Fig. 

32) ahrupta, n. sp. 

— Aedeagus with shaft not expanded at apex . . 12 

12(11). Aedeagus with small apical processes (Fig. 24) 

magna (Osborn) 

— Aedeagus with larger subapical processes 
(Fig. 28) expanda, n. sp. 

13(10). Aedeagus with processes expanded at base 

(Fig. 40) Columbiana (Nast) 

— Aedeagus with thin, spinelike processes (Fig. 
36) 14 

14(13). Aedeagus with gonopore basal to processes 

(Fig. 37) stigmata (Nast) 

— Aedeagus with gonopore apical to processes 
(Fig. 53) corvenda Kramer 

Key to Known Females 
1. Pronotum longitudinally banded 2 

— Pronotum not so marked 3 

2(1). Seventh sternum unusually long, over twice 

length of sixth (Fig. 61) fasciata (Osborn) 

— Seventh sternvmi shorter, about twice or less 
length of sixth (Fig. 65) ynexicana, n. sp. 

3(1). Seventh sternum truncate, with a nu'dian 

emargination rngosa, n. sp. 



— Seventh sternum not truncate, posterior mar- 
gin usually rounded 4 

4(3). Head with a frontal margin, bright yellow and 

black marginata, n. sp. 

— Head rounded to face 5 

5(4). Ovipositor short, less than 2 mm in length 

(Fig. 63) acuta, n. sp. 

— Ovipositor long, more than 2 mm in length 

(Fig. 62) ...6 

6(5). Seventh sterninn with median emargination 

of posterior margin V-shaped (Fig. 59) 

ornata, n. sp. 

— Seventh sternimi with median emargination 
of posterior margin U-shaped (as in Fig. 62) . 
decora (Fabricius) 

The decora species group is characterized 
by having orange transverse bands on the 
head, and the aedeagus laterally flattened and 
knife-shaped. The following seven species are 
placed in this group. 

Calliscarta decora (Fabricius) 
Figs. 1-4, 66 

Cicada decora Fabricius 1803: 69. (South America, possi- 
bly from Brazil; Stockholm Museum). 

Tettigonia decora: Signoret 1853: 327; Walker 1858: 200; 
Dohrn 1859; 88. 

Calliscarta decora: Stal 1869; 83; Dallas 1870: 495; Dis- 
tant 1908; 524; Evans 1947; 230; Linnavuori 19.59: 
29;Metcalfl967:946. 

Thamnotettix magnificus Osborn 1924: 424. (Cuatro 
Ojos, Santa Cruz, Bolivia; Carnegie Museum). 
Ncti' synontjmtj 

Idiotettix magnificus: Osborn 1929; 465; Oman 1938: 395; 
Evans 1947: 192; Nast 19,52: 1; Maldonado 
Capriles 19.54: 250; Metcalf 1966: 227. 

Calliscarta magnifica: Linnavuori 1959: 28. 

Idiotettix festivus Maldonado Capriles 19.54: 248. 
(Cunucunuma River, F. T. Amazonas, Venezuela; 
U.S. National Museum). 

Length of males 9.5-11.5 mm.; female 12 
mm. Head reddish yellow above antennal 
sockets, yellow below, with three orange 
transverse bands, one just above and one just 
below ocelli and the other above base of an- 
tennae. Pronotum reddish yellow with two 
darker red transverse bands, one along ante- 
rior margin and one across middle, and one 
yellowish green band along posterior margin 
continuing to anterior of lateral margins near 
eyes. Scutellum reddish brown. Forewing 
generally dark reddish brown, claval area 
lighter brown with foiu- large, yellowish green 
spots, costal margin red, and a row of si.\ to 
eight large, yellowish green spots through 
middle of wing from base to appendix, apex 



1988 



Freytag: Revision of Calliscarta 



69 






DECORA 





ELONGATA 




INVITA 




Figs. 1-4. Calliscarta decora (Fabricius), from Peru; 1, lateral view of male genital segments; 2, lateral view of 
aedeagus; 3, ventral view of aedeagus; 4, ventrolateral view of style. 

Figs. 5-8. Calliscarta elongata, n. sp. , holotype; 5, lateral view of male genital segments; 6, lateral view of aedeagus; 
7, ventral view of aedeagus; 8, ventrolateral view of style. 

Figs. 9-12. Calliscarta invito (Nast), from Ecuador: 9, lateral view of male genital segments; 10, lateral view of 
aedeagus; 11, ventral view of aedeagus; 12, ventrolateral view of style. All drawn to the same scale; the line equals 1 
mm. 



smoky brown. Ventral surface entirely yel- 
low. 

Male genitalia. — Pygofer truncate with a 
short, apical, spinelike process (Fig. 1). Geni- 
tal plates large, roundedly expanded at apex 
(Fig. 1). Style long, bent at right angle three- 
quarters of distance from base, apex pointed 
with a subapical tooth on inner margin, and a 
small but distinct spine at the bend (Fig. 4). 
Aedeagus short, laterally flattened, broad in 
lateral view, apex pointed, with a pair of small 
processes on caudal margin near gonopore 
(Figs. 2, 3). 

Female genitalia. — Seventh sternum 
large, twice length of sixth, with a shallow, 
median, U-shaped emargination similar to or- 
nata. 

Type. — Holotype male of decora not seen, 
but illustrated by Linnavuori (1959). Type is 
from South America, and Dohrn (1859) gave 
Brazil as the possible type locality. 

Distribution. — Known are specimens 
from: BOLIVIA: one female, Cuatro Ojos, 
November 1913, Steinbach Coll. , (holotype of 



magnificus) Carnegie Museum. Brazil: one 
male, ? (holotype oi decora) Stockholm Mu- 
seum. Ecuador: one male, Pompeya, 13-25 
May 1965, Luis E. PeiiaColl., Ramos Collec- 
tion; one male, Coca, May 1965, Luis E. Pena 
Coll., Ramos Collection; two males, Napo, 
Prov. Limoncocha, 10 June 1977, Dave L. 
Vincent Coll., U.S. National Museum. 
French Guyana: one male, Cabret Repub- 
hque, 29 August 1975, Guyane Frangaise 
Mission M. Boulard, et P. Pompanon Coll., 
Museum Paris. Guyana: one male, Bartica, 
20-30 March 1901, H. S. Parish Coll., (allo- 
type of magnificus) Ohio State University 
Collection. Peru: four males, Avispas, Octo- 
ber 1962, Luis E. Pena Coll., Ramos Collec- 
tion. Venezuela: one male, Cunucunuma 
River, F. T. Amazonas, 28 April 1950, J. Mal- 
donado Capriles Coll. (holotype o( festivus) 
U.S. National Museum; one male, Anacoco, 
Bolivar, 60 M, 6°5'N:6r8'W, 10-30 August 
1979, Expedicion La Salle, University of Cen- 
tral Venezuela Collection. 

Notes. — This species appears to be widely 



70 



Great Basin Naturalist Memoirs 



No. 12 



distributed in South America but is poorly 
represented in collections. Much confusion 
concerning the identification of this species 
has occurred because the color pattern is not 
fully developed in some specimens. How- 
ever, this species can be quickly recognized 
by the small spine on the male style, which is 
found on only one other known species, 
tartessoides. For now the female must be rec- 
ognized by the color pattern, which is the 
same as in the male. 

The types of magnificus and festivus were 
examined and found to represent the same 
species. Linnavuori (1959) synonymized fes- 
tivus with decora. 

Calliscarta tartessoides Linnavuori & Heller 

Calliscarta tartessoides Linnavuori & Heller 1961: 3. 
(Hacienda Mascoitania, Peru; Staatlichen Mu- 
seum fur Naturkunde, Stuttgart). 

This species was illustrated when de- 
scribed, and no additional information can be 
added at this time. It appears to be closely 
related to decora. 

Length of male 9.5 mm; female unknown. 
Brownish yellow. Head light yellow, with a 
reddish orange transverse band across vertex 
between eyes; face with an orange transverse 
band between ocelli and another across mid- 
dle of face, and a dark brown spot underneath 
each ocellus. Pronotum with anterior margin 
light yellow, posterior two-thirds darker mar- 
bled with darker orange. Scutellum faintly 
dark orange. Forewing mostly translucent, 
with veins brown; costal vein darker brown in 
middle. Ventral surface light yellow with 
some yellow-orange and brown markings on 
legs and abdomen. 

Male genitalia. — Pygofer narrowing to a 
posteriorly projecting, bluntly pointed apex, 
with two small spines on ventral margin near 
middle. Genital plates large and truncate at 
apex. Style long, bent laterally on apical 
fourth, apex pointed, a small subapical tooth 
on inner margin. Aedeagus laterally flattened, 
knife-shaped, without noticeable processes, 
gonopore on ventral margin near middle of 
shaft. 

Type. — Holotype (male), Peru, Hacienda 
Mascoitania, 10-12 November 1957, Urwald 
Coll., Staatlichen Museum fiir Naturkunde, 
Stuttgart. 

Notes. — This species is known only from 
the type specimen, which I have not seen. It is 



a species like decora, with a tooth on the style, 
but a color pattern that is more like magna. 
More material is needed to properly place this 
species. 

Calliscarta elongata, n. sp. 

Figs. 5-8 

This species resembles decora in general 
appearance, but with different male genitalia. 

Length of males 10-10.1 mm; female un- 
known. Head creamy yellow, crown having a 
red-violet tinge, with four orange transverse 
bands, one across middle of crown between 
eyes, one just below ocelli, two fainter bands 
across postclypeus; a black band on each side 
from base of antenna to anteclypeus along side 
of postclypeus. Pronotum orange, with a band 
along posterior margin and two central spots 
greenish yellow. Scutellum orange with a 
small yellow spot in middle of each lateral 
margin. Forewing orange anteriorly, fading to 
brown posteriorly, median of costal vein with 
a reddish tinge, four greenish yellow spots on 
clavus, three at base and one near middle, 
four yellow spots on corium, apical cells 
smoky brown with an L-shaped translucent 
spot pattern. Ventral surface generally 
creamy yellow, fore tibiae and tarsi brownish 
yellow. 

Male genitalia. — Pygofer narrowing to 
apex, apex a dorsally projecting, pointed pro- 
cess (Fig. 5). Genital plate enlarged and ex- 
panded on apical half, apex bluntly rounded 
(Fig. 5). Style stout, uniform width through- 
out, hooked apically, apex with a very small 
subapical tooth (Fig. 8). Aedeagus laterally 
flattened, knife-shaped, gonopore medially 
on ventral margin, a pair of small lateral pro- 
cesses basal to gonopore (Figs. 6, 7). 

Type. — Holotvpe (male), Ecuador, Napo, 
Lago Agrio (18 km E), 28 August 1975, Rio 
Aguarico, collected on gravel bar at black- 
light, Langley, Cohen, Cantor, Yingling 
Coll. , Ecuador-Peace Corps-Smithsonian In- 
stitution Aquatic Insect Survey, in the U.S. 
National Museum. Paratypes: Ecuador, one 
male, same data as holotype, except 23 Sep- 
tember 1975 and Andrea Langley Coll. , Uni- 
versity of Kentucky Collection; one male, 
same data as holotype, except 19 September 
1975 and Andrea Langley Coll., U.S. National 
Museum; Peru, Avispas, October 1962, Luis 
E. PenaColl., Ramos Collection. 



1988 



Freytac: Revision of Calliscarta 



71 



Calliscarta invita (Nast) 
Figs. 9-12 

Idiotettix invitus Nast 1952: 3. (Santa Inez, Ecnador, 
Polish Museimi of Zoology); Maklonado Capriles 
1954: 250; Metcalf 1966: 226. 

Calliscarta invita: Linnavuori 1959: 30; Linnavuori and 
Heller 1961: 3. 

Length of males 10-10.6 mm; female un- 
known. Head brownish yellow, with a tinge of 
violet on crown and postclypeus; an orange 
transverse band across middle of crown be- 
tween eyes, another orange transverse band 
below ocelli, two brownish orange transverse 
bands on postclypeus, a black band on each 
side from base of antennae to anteclypeus 
along sides of postclypeus. Pronotum mostly 
orange brown, lateral margins yellow. Scutel- 
lum light brown. Forewing light brown, with 
veins and obscure spots darker brown, me- 
dian of costa reddish orange, small, white, 
translucent spots on clavus and corium; apical 
area smoky brown with three translucent 
spots. Ventral surface mostly brownish yel- 
low, brown spot on propleura and sterna, fore 
legs with most of tibiae and entire tarsi brown, 
middle legs with base and apex of tibiae and 
entire tarsi brown. 

Male genitalia. — Pygofer truncately nar- 
rowed to apex, which is projected slightly dor- 
sad in the shape of a bird's head (Fig. 9). 
Genital plate large, expanded in lateral view, 
truncate (Fig. 9). Style elongate, even width 
throughout, hooked apically, with a pointed 
apex (Fig. 12). Aedeagus laterally flattened, 
knife-shaped, without processes, gonopore on 
ventral margin near base of shaft (Figs. 10, 
11). 

Type. — Holotype (male), Ecuador, Santa 
Inez, R. Haensch Coll., Polish Museum of 
Zoology. 

DiSTRlRUTlON. — Five males are known for 
this species. Linnavuori and Heller (1961) re- 
ported two males from Peru, Hacienda Mas- 
coitania, 10-12 September 1957, Urwald 
Coll., Staatlichen Museum fiir Naturkunde, 
Stuttgart. I have seen two males from Ecua- 
dor, Rio Margajita, Rio Pastaza, 1250 m, 20 
March 1939, F. M. and H. H. Brown Coll., 
University of Kansas Collection. 

Calliscarta boliviana (Osborn) 
Figs. 13-18 

Idiotettix bolivianus Osborn 1929: 466. (Coroico, La Paz, 
Bolivia, Ohio State University Collection); Oman 



1938: 395; Nast 1952: 1; Maldonado Capriles 1954: 

250; Metcalfl966: 225. 
Calliscarta boliviana. Linnavuori 1959: 29. 
Calliscarta ting,a Kramer 1963: 205. (Tingo Maria, 

Iluanuco, Peru, U.S. National Museum). NeuJ 

sifnoniimij 

Length of males 9.1-10 mm; female un- 
known. General color as in invita, pronotum 
with a band of yellow along posterior margin 
and lateral margins and white, translucent 
spots on forewings less conspicuous. 

Male genitalia. — Pygofer quite variable, 
narrowing to near apex, which projects dorsad 
in some type of bifurcate process (Figs. 13, 17, 
18). Genital plates greatly expanded on apical 
half (Fig. 13). Style elongate, narrowing 
slightly to apex, hooked apically, apex with a 
ventral subapical tooth (Fig. 16). Aedeagus 
laterally flattened, knife-shaped, gonopore on 
ventral margin near base of shaft, with a small 
pair of laterally projecting processes basal to 
gonopore (Figs. 14, 15). 

Type. — Holotype (male), Bolivia, Coroico, 
Ohio State University Collection. 

Distribution. — This species is known from 
10 male specimens. Besides the two types, I 
have seen: BOLIVIA, one male, Cristal Mayn, 
Chapare, 28 August 1949, L. E. Peiia Coll., 
University of Kansas Collection; Peru, four 
males, Avispas, September 1962, Luis E. 
Pefia Coll.; two males, Quincemil, August 
1962, Luis E. Peria Coll., Ramos Collection, 
and one from each locality in University of 
Kentucky Collection; Panama, El Volcan de 
Chiriqui, 26 February 1936, W. J. Gertsch 
Coll., American Museum of Natural History. 

Notes. — This species is quite variable in 
the shape of the apex of the pygofer, but it is 
the only species of the genus with the apex 
bifurcate. No two specimens seen have the 
same shape, and I consider this to be just a 
variation at this time; however, this may 
prove to be a complex of very closely related 
species. 

Calliscarta lora, n. sp. 

Figs. 19-22 

This species resembles invita in general ap- 
pearance, but with distinct male genitalia. 

Length of male 9.3-9.9 mm; female un- 
known. Head, pronotum, and scutellum as in 
elongata. Forewing with clavus mostly or- 
ange, with three yellow spots, two transverse 
spots anteriorly, one longitudinal spot along 
posterior part of claval suture; corium brown. 



72 



Great Basin Naturalist Memoirs 



No. 12 









14 15 



BOLIVIANA 



Figs. 13-16. Calliscarta boliviana (Osborn), liolotype: 13, lateral view of male genital segments; 14, lateral view of 
aedeagus; 15, ventral view of aedeagus; 16, ventrolateral view of style. 

Figs. 17—18. Lateral views of male genital segments of other specimens of C. boliviana from Bolivia. 

Figs. 19-22. Calliscarta lora, n. sp., holotype; 19, lateral view of male genital segments; 20, lateral view of aedeagus; 
21, ventral view of aedeagus; 22, ventrolateral view of style. All drawn to the same scale; the line equals 1 mm. 



median of costal vein with a reddish tinge, 
with four yellow spots, two at base, one (some- 
times two) above reddish tinge of costa, and 
one a continuation of longitudinal spot on 
clavus; apical cells smoky brown, with three 
small, translucent spots. Ventral surface 
creamy yellow, fore tibiae and tarsi brownish 
yellow. 

Male genitalia. — Pygofer narrowing to 
apex that extends as a large bird's-head- 
shaped process (Fig. 19). Genital plate gradu- 
ally expanding to a large, truncate apex (Fig. 
19). Style elongate, hooked apically, with a 
pointed apex (Fig. 22). Aedeagus laterally flat- 
tened, knife-shaped, gonopore on ventral 
margin near middle, without processes (Figs. 
20, 21). 

Type. — Holotype (male), Peru, Quincemil, 
August 1962, Luis E. Pena Coll., in the 
Ramos Collection but will be deposited in 
U.S. National Museum through courtesy of 
Dr. Ramos. Paratypes: Peru, two males, 
same data as holotype; one male, Quincemil, 
October 1962, Luis E. Peiia Coll.; two males, 
Avispas, September 1962, LuisE. Pena Coll., 
three in the Ramos Collection and two in the 
University of Kentucky Collection. 

Notes. — This species has a distinct color 
pattern and male genitalia that make it possi- 



ble to separate it from other known species 
that also occur in Peru. 



Calliscarta delicata. 
Figs. 43-46 



n. sp. 



This species resembles invita in general ap- 
pearance, but with distinct male genitalia. 

Length of male 8.5-10.1 mm; female 
unknown. Head yellow, with four orange 
transverse bands, one above ocelli, one be- 
low ocelli, one between antennae and one 
along lower margin of postclypeus. Pronotum 
orange, with green posterior margin extend- 
ing along lateral margins and a green trans- 
verse band in anterior half Scutellum mostly 
orange. Forewing generally smoky brown, 
with claval suture, anterior third of costa, and 
small spots at ends of claval veins green; costa 
red near middle; corium with two orange 
bands extending from near base to apical 
veins, one along claval suture, other along 
costa; apical cells smoky brown with three 
transverse, opaque spots. Abdominal seg- 
ments orange. 

Male genitalia. — Pygofer narrowing to 
somewhat expanded apex, with a dorsally pro- 
jecting, apical, pointed process (Fig. 43). 
Genital plate gradually expanded to a large, 
truncate apex (Fig. 43). Style narrowing and 



1988 



FreytaG: Revision of Calliscarta 



73 






MAGNA 



28 29 



EXPANDA 




Figs. 23-26. Calliscarta nuigna (0.shorn), from Brazil: 23, lateral view of male genital segments; 24, lateral view of 
aedeagus; 25, ventral view of aedeagus; 26, ventrolateral view of style. 

Figs. 27-30. Calliscarta expanda. n. sp., holotype: 27, lateral view of male genital segments; 28, lateral view of 
aedeagus; 29, ventral view of aedeagus; 30, ventrolateral view of style. 

Figs. 31-34. Calliscarta abrupta, n. sp., holotype; 31, lateral view of male genital segments; 32, lateral view of 
aedeagus; 33, ventral view of aedeagus; 34, ventrolateral view of style. All drawn to the same scale; the line equals 1 



curving laterally near pointed apex (Fig. 46). 
Aedeagus laterally compressed with a pair of 
long, curved processes from base of shaft, 
gonopore medially on ventral margin, apex 
bluntly pointed (Figs. 44, 45). 

Type. — Holotype (male), Peru, Tingo 
Maria, 20 June 1982, Fisk, Ohio State Univer- 
sity Collection. Paratypes: two males, same 
data as holotype, one in Ohio State University 
Collection and one in University of Kentucky 
Collection. Two males, Bolivia, El Palmar, 
Cochabamba, Chapare, 8-15 September 
1955, Luis E. PeiiaColl., Ramos Collection. 

Notes. — This species can be separated 
from invita by the longer processes on the 
aedeagus. 

The ornata species group is characterized 
by a single, transverse, black band on the face 
that replaces the orange transverse bands of 
the decora group, and by an aedeagus that is 
flattened and knife-shaped. Only one species, 
ornata, is placed here. 



Calliscarta ornata, n. sp. 
Figs. 47-50, 59, 67 

This species resembles invita in general ap- 
pearance, but with unique black band on face 
instead of usual orange bands. 

Length of male 9.4-10.1 mm; female 10- 
11 mm. Head creamy yellow with a reddish 
tinge; an irregular, black, transverse band 
below ocelli, a large, black spot in middle of 
postclypeus, a black, V-shaped band from an- 
tennal bases along sides of postclypeus over 
entire anteclypeus. Pronotum creamy yellow, 
with a large, irregular, transverse, brown 
band across middle, posterior margined with 
greenish yellow. Scutellum mostly creamy 
yellow. Forewing brown, with many large, 
greenish yellow spots, apical cells smoky 
brown, with two large, translucent spots. 

Male genitalia. — Pygofer narrowed to a 
truncate apex, with a large, nearly bird's- 
head-shaped apical process (Fig. 47). Genital 
plate large, greatly expanded at apex, which is 
bluntly rounded (Fig. 47). Style elongate, 



74 



Great Basin Naturalist Memoirs 



No. 12 







STIGMATA 





COLUMBIANA 





44 45 



DELICATA 




Figs. 35-38. Calliscarta stigmata (Nast), from Colombia: 3.5, lateral view of male genital segments; 36, lateral view of 
aedeagus; 37, ventral view of aedeagus; 38, ventrolateral view of style. 

Figs. 39-42. Calliscarta columbiana (Nast), from Colombia; 39, lateral view of male genital segments; 40, lateral 
view of aedeagus; 41, ventral view of aedeagus; 42. ventrolateral view of style. 

Figs. 43-46. Calliscarta delicata, n. sp., holotype; 43, lateral view of male genital segments; 44, lateral view of 
aedeagus; 45, ventral view of aedeagus; 46, ventrolateral view of style. All drawn to the same scale; the line equals 1 



hooked apically, with a pointed apex (Fig. 50). 
Aedeagus laterally flattened, knife-shaped, 
with a pair of recurved processes on ventral 
margin near base of shaft, gonopore on ventral 
margin just apical to bases of processes (Figs. 
48, 49). 

Female genitalia. — Seventh sternum en- 
larged, rounded posterior margin with a 
large, median, V-shaped emargination (Fig. 
59). Ovipositor extending twice its width be- 
yond pygofer. Pygofer narrowing to apex in 
ventral view. 

Type. — Holotype (male), French Guyana, 
Saint-Elie, 30 August-2 September 1975, 
Guyane Franyaise Mission, M. Boulard et P. 
Pompanon Coll., Paris Museum. Allotype 
(female), same data as holotype. Paratypes: 
French Guyana, one female, Garbet Lavaud 
(Rive Surinamienne), 24-26 September 1975, 
Itani (Guyanes) Mission, M. Boulard, P. Jauf- 
fret et P. Pompanon Coll.; one female, Em 
Gouchure de la Crique Oyaricoulet, 23 Sep- 
tember 1975, Itani (Guyanes) Mission, M. 
Boulard, P. Jauffret et P. Pompanon Coll.; 
one female, Alicoto-Oyapock-Guyane, 13 
November 1969, Piege lumineux, Guyane 
Mission, Balachowsky, Gruner Coll., Oct.- 



Nov. 1969, Paris Museum; Brazil, one fe- 
male, Chapada, September, C. F. Baker 
Coll.; one female, Chapada, October, C. F. 
Baker Coll., U.S. National Museum; Vene- 
zuela, one male, San Carlos de Rio Negro, 
T. F. Amazonas, 7-13 September 1982, A. 
Chacon, G. Yepez Gil Coll.; one female, Sta. 
Lucia, T. F. Amazonas, 15-21 September 
1982, A. Chacon, G. Yepez Gil Coll.; one 
male, one female, Rio Surukum, Carretera 
Sta., Elena Icabaru, Bolivar, 850 m, 19-31 
January 1985, F. Fernandez Y., Anibal 
Chacon, Jurg Demarmels Coll., University of 
Central Venezuela Collection. 

Notes. — This species can be easily sepa- 
rated from the known species of this genus on 
the basis of the dark band across the face. 

The magna species group is characterized 
by having the head mottled with brown and 
the aedeagal shaft tubular with apical or sub- 
apical processes. The following three species 
are placed in this group. 

Calliscarta magna, (Osborn) 

Figs. 23-26 

Idiocerus niap,nus Osborn 1923: 13. (San Antonio de Gua- 
pore, MattoGrosso, Brazil, Carnegie Museum). 



1988 



Freytag: Revision of Calliscarta 



75 







48 49 



OR N ATA 





52 53 



50 CORVENDA 





Figs. 47-50. Calliscarta oniata, n. sp., holotype: 47, lateral view of male genital segments; 48, lateral view of 
aedeagus; 49, ventral view of aedeagus; 50, ventrolateral view of style. 

Figs. 51-54. Calliscarta corvenda Kramer, from Venezuela: 51, lateral view of male genital segments; 52, lateral 
view of aedeagus; 53, ventral view of aedeagus; 54, ventrolateral view of style. 

Figs. 55-58. Calliscarta fasciata (Osborn), from Brazil: 55, lateral view of male genital segments; 56, lateral view of 
aedeagus with connector to anal tube; 57, ventral view of aedeagus and connector to anal tube; 58, ventrolateral view of 
style. All drawn to the same scale; the line equals 1 mm. 



Idiotettix magnus: Maldonado Capriles 1954: 247; Met- 

calf 1966: 227. 

Calliscarta rrmgna: Linnavuori 1959: 30. 

Length of male 9-10 mm; female unknown. 
Head, pronotum, and scutellum creamy yel- 
low with a reddish violet tinge, greatly mot- 
tled with brown. Forewing mostly translu- 
cent, veins dark brown; clavus with a large, 
fuscous spot at middle and apex; apical cells 
smoky brown, with two transverse, translu- 
cent spots. Ventral surface mostly creamy yel- 
low, with fore and middle legs with apex of 
tibiae and entire tarsi brownish yellow. 

Male genitalia. — Pygofer gradually nar- 
rowing to a pointed apex (Fig. 23). Genital 
plate large, greatly expanded on apical half 
(Fig. 23). Style elongate, hooked apically, 
apex bluntly pointed (Fig. 26). Aedeagus 
tubular, with pair of small, lateral, apical pro- 
cesses, gonopore subapical on ventral margin 
(Figs. 24, 25). 

Type. — Holotype (male), Brazil, Rondo- 
nia, Santo Antonio de Guapore (Rio Gua- 



pore), 26 July 1909, Haseman Coll., Carnegie 
Museum. 

Distribution. — Known only from Brazil on 
the basis of the type and the following speci- 
mens: Brazil, five males, Sao Felice, Camp 
IV, Moyen Xingu, (Bresil) Mission, M. Bou- 
lard, P. Jauffret et P. Pompanon Coll. , four in 
Museum Paris and one in University of Ken- 
tucky Collection. 

Calliscarta expanda, n. sp. 

Figs. 27-30 

This species resembles magna in general 
appearance, but with distinct male genitalia. 

Length of male 9.7-10.5 mm.; female un- 
known. Head, pronotum, and scutellum 
creamy violet, mottled with brown. Forewing 
brown, with most cells translucent, veins dark 
brown; apical cells smoky brown, with three 
or four large, translucent, transverse spots. 
Ventral surface creamy yellow heavily marked 
with brown. 

Male genitalia. — Pygofer narrowing to a 



76 



Great Basin Naturalist Memoirs 



No. 12 



pointed apex (Fig. 27). Genital plate large, 
expanded on apical half (Fig. 27). Style elon- 
gate, hooked apically, with apex bluntly 
pointed, notched subapically (Fig. 30). Aede- 
agus tubular, with subapical, large, lateral 
pair of processes, gonopore on ventral margin 
subapical to processes (Figs. 28, 29). 

Type. — Holotype (male), Ecuador, Napo, 
Lago Agrio, 19 September 1975, at black- 
light, Andrea Langley Coll., Ecuador-Peace 
Corps-Smithsonian Institution Aquatic In- 
sect Survey, U.S. National Museum. Para- 
types: Ecuador, one male, same data as holo- 
type, except 23 September 1975, (18 km E) 
Rio Aguarico; two males, same data, except 28 
August 1975, collected on gravel bar, Lang- 
ley, Cohen, Cantor, Yingling Coll., U.S. Na- 
tional Museum; Peru, one male, Loreto, 
Ucayali R. , Yarina Cocha, 24 September 
1953, Peter Hocking Coll., Field Museum; 
one male, Quincemil, August 1962, Luis E. 
Peria Coll. ; one male, Loromavo, September 
1962, Luis E. Peiia Coll., Ramos Collection; 
Bolivia, Rurrenbaque, 10-23 October 1956, 
Luis E. Peiia Coll., Ramos Collection. 

Notes. — This species is similar to magna, 
but the aedeagus is larger and with longer, 
more subapical processes. 

Calliscarta abnipta, n. sp. 

Figs. 31-34 

This species is related to magna, but with a 
distinctly different aedeagus. 

Length of male 9.8 mm; female unknown. 
Head, pronotum, and scutellum mostly yel- 
low tinted with red. Head with a small, brown 
spot behind each ocellus next to eye, another 
larger, brown spot below each ocellus next to 
eye. Postclypeus with brown dashes on each 
side and brown speckles over remainder of 
surface. Pronotum speckled with brown. 
Forewings mostly brown, claval area with 
opaque spots at ends of claval veins, costal 
vein yellow, two cell rows behind costal vein 
transparent, giving wing appearance of having 
a brown longitudinal band through middle, 
apical cells smoky brown with several large, 
opaque spots. 

Male genitalia. — Pygofer narrowing grad- 
ually to an upturned, pointed apex (Fig. 31). 
Genital plate enlarged on apical half, truncate 
(Fig. 31). Style nearly same width to apex, 
hooked apically, with apex foot-shaped (Fig. 
34). Aedeagus tubular, with shaft bent at right 



angle near middle, apex expanded with a pair 
of subapical large processes extending along 
sides of shaft halfway to base, gonopore on 
ventral margin just basal to processes (Figs. 
32, 33). 

Type, — Holotype (male), Bolivia, Rurren- 
abaque, 10-23 October 1956, Luis E. Peiia 
Coll., in Ramos Collection but is being de- 
posited in U.S. National Museum through 
courtesy of Dr. Ramos. 

Notes. — This species can be separated 
from the other species of the genus by the 
distinct aedeagus. 

The stigmata species group is characterized 
by having the head mottled with brown, and 
the aedeagal shaft tubular with processes near 
middle. The following four species are placed 
in this group. 

Calliscarta stigmata (Nast) 
Figs. ,3.5-38 

Idiotettix stigmatus Nast 1952: 2. (Hac. Pehlke [probably 
Magdalena], Colombia, Polish Museum of Zool- 
ogy); Maldonado Capriles 19.54: 250; Metcalfl966: 
227. 

Calliscarta stigmata: Linnavuori 1959: 31. 

Length of male 8.5-9.4 mm; female un- 
known. Head, pronotum, and scutellum gray- 
ish yellow tinged with violet, mottled with 
brown; orange bands of crown and face ob- 
scure. Forewing brown, with numerous 
whitish, translucent spots. Ventral surface 
brownish yellow; legs banded with brown. 

Male genitalia. — Pygofer narrowing to a 
long, tubular, apical area, apex with small, 
dorsally projecting process (Fig 35). Genital 
plate long, expended on apical half, bluntly 
rounded at apex (Fig. 35). Style long, bent 
apically, with pointed apex (Fig. 38). Aedea- 
gus tubular, shaft elongate, apex bent dorsad 
and pointed, a pair of lateral processes at base 
of apical bend, gonopore basal to processes on 
ventral margin (Figs. 36, 37). 

Type. — Holotvpe (male), Colombia, Ha- 
cienda Pehlke, ' 1925, Pehlke Coll., Polish 
Museum of Zoology. 

Distribution. — This species is known from 
eight male specimens. The holotype and five 
paratypes are from Colombia and are in the 
Polish Museum of Zoology, and I have seen 
one male, Colombia, Alban, Cund., 10 Sep- 
tember 1965, J. A. Ramos Coll., Ramos Col- 
lection; and one male, Venezuela, Rio Frio, 



1988 



Freytag: Revision of Calliscarta 



77 






ORNATA 



MARGINATA 



FA SCI ATA 



Figs. 59-61. Calliscarta spp., ventral view offemale segments: 59, C. ornata, n. sp., allotype; 60, C. nuirg,inata, n. 
sp., holotype; 61, C.fasciata (Osborn), from Brazil. All drauni to the same scale; the line equals 1 mm. 



Tachira, 600 m, 11-14 December 1980, J. A. 
Clavijo, A. Chacn, J. AyalaColl., University 
of Central Venezuela Collection. 

Calliscarta columbiana (Nast) 
Figs. 39-42 

Idiotettix columbianns Nast 1952: 2. (Hac. Pehlke [proba- 
bly Magdalena], Colombia, Polish Museum of 
Zoology); Maldonado Capriles 1954: 250; Metcalf 
1966: 226. 

Calliscarta columbiana: Linnavuori 1959: 29. 

Length of male 8-9.1 mm; female un- 
known. General color pattern as in stigmata, 
except lighter in color and legs only faintly 
banded with brown. 

Male genitalia. — Pygofer narrowing to 
truncate apex with a small, dorsad projecting 
process (Fig. 39). Genital plates large, ex- 
panded, truncate at apex (Fig. 39). Style elon- 
gate, hooked apically, with pointed apex 
(Fig. 42). Aedeagus semitubular, apex bluntly 
pointed, with a pair of basally expanded, lat- 
eral processes halfway to base, gonopore on 
ventral margin at base of processes (Figs. 40, 
41). 

Type, — Holotype (male), Colombia, Ha- 
cienda Pehlke, 1921, Pehlke Coll., Pohsh 
Museum of Zoology. 

Distribution. — This species is known only 



from Colombia from eight male specimens. 
The holotype and six paratypes are in the 
Polish Museum of Zoology, and I have seen 
one male, Alto Rio Opon, Santander, January 
1950, L. RichterColl., Ramos Collection. 

Calliscarta corvenda Kramer 

Figs. 51-54 

Calliscarta corvenda Kramer 1963: 210. (Covendo, La 
Paz, Bolivia, U.S. National Museum). 

Length of male 9-9.8 mm.; female un- 
known. Color pattern similar to stagmata, ex- 
cept spots of forewings not forming rows or 
bands. 

Male genitalia. — Pygofer narrowing to 
inwardly hooked and pointed apex (Fig. 51). 
Genital plate large, expanded on apical half 
(Fig. 51). Style very elongate, hooked api- 
cally, with pointed, upturned apex (Fig. 54). 
Aedeagus semitubular, pair of lateral pro- 
cesses on ventral margin near middle of shaft, 
gonopore subapical on ventral margin (Figs. 
52, 53). 

Type. — Holotype (male), Bolivia, Coven- 
do, August 1921, Mulford Biol. Expedition, 
Wm. M. Mann Coll., U.S. National Museum. 

Distribution. — Four specimens are 
known, including the type and a paratype from 



78 



Great Basin Naturalist Memoirs 



No. 12 






RUGOSA 



ACUTA 



MEXICANA 



Figs. 62-65. Calliscarta spp. , ventral view of female segments: 62, C. rugosa, n. sp. , holotype; 6.3, C. acuta, n. sp. , 
holotype; 64, C acuta, n. sp., paratype; 65, C. mexicana, n. sp., holotype. All drawn to the same scale; the line equals 
1 mm. 



Bolivia. Other specimens: Venezuela, two 
males, Barinitas, 15 km SW Barinas, 25 Feb- 
ruary 1969, Duckworth and Dietz Coll., one 
in U.S. National Museum and one in Univer- 
sity of Kentucky Collection. 

Calliscarta rugosa, n. sp. 

Figs. 62, 68 

Resembling columbiana in general color 
pattern, but with lighter, more reddish brown 
markings. 

Length of female 10. 1 mm; male unknown. 
General color pattern as in columbiana, but 
with forewing darker and more reddish 
brown. 

Female genitalia. — Seventh sternum large, 
truncate, with a large, median, U-shaped 
emargination (Fig. 62). Ovipositor extending 
its length beyond pygofer. 

Type. — Holotype (female), Venezuela, 
Aragua, El Limon, 450 m, 23 August 1979, 
LuzdeMercurio, F. Fernandez Y. Coll., Uni- 
versity of Central Venezuela Collection. 

Notes. — This female may be the female of 
columbiana or corvenda. However, since it 
was collected along the northern mountains of 
Venezuela, it is just as likely to represent a 
separate species. The color pattern is suffi- 
ciently different from the known species of the 
stigmata species group to warrant its descrip- 
tion at present. 



The following two species are not placed in 
a species group at this time, as the males are 
unknown and the color patterns give little 
indication as to how these relate to other 
known species. However, they are closer to 
the preceding groups than to the group fol- 
lowing them. 

Calliscarta marginata, n. sp. 
Figs. 60, 69 

This species resembles ornata, but with a 
brighter color pattern and the anterior margin 
of the crown more distinct. 

Length of female 10.5 mm; male unknown. 
Head creamy yellow with a U-shaped band in 
middle of crown, a wider U-shaped band be- 
hind eyes over ocelli and along anterior mar- 
gin, two transverse bands between antennae 
and ocelli, two dashes in middle of lower part 
of postclypeus, and a V-shaped band from 
underneath eyes along lateral margins of post- 
clypeus down middle of anteclypeus, black. 
Pronotum black, with a yellow longitudinal 
band down median; also a large, yellow spot 
on each side with a smaller cream spot near 
anterior margin between yellow spot and 
band. Scutellum black, with cream spots. 
Forewing black, with six large, yellow spots or 
dashes on clavus, eight large, yellow spots or 
dashes on corium, and four smaller cream 
spots in apical cells. 



1988 




FreytaG: Revision of Calliscarta 

( 



79 



..Hill. TIUp*^ 



\ 



7 



68 






70 



Figs. 66-71. Calliscarta spp., lateral aspect: 66, C. decora (Fabricius), male from Peru; 67, C. ornata, n. sp., male 
paratype from French Guyana; 68, C. rugosa, n. sp. , female holotype; 69, C. marginata, n. sp. , female holotype; 70, C. 
acuta, n. sp., female holotype; 71, C mexicana, n. sp., female holotype. 



Female genitalia. — Seventh sternum 
large, with posterior margin with a median, 
V-shaped emargination (Fig. 60). Ovipositor 
extending beyond pygofer three times its 
width. 

Type. — Holotype (female), Honduras, 
Com., Lago Yojoa, 19 July 1974, C. W. and 
L. B. O'Brien and Marshall Coll., the Ohio 
State University Collection. 

Notes. — This species is quite distinct from 
other members of the genus both in color 
pattern and in having a slight anterior margin 
of the head. 

Calliscarta acuta, n. sp. 

Figs. 63, 64, 70 

Resembling marginata, this species is 



smaller and has a distinctly different color 
pattern. 

Length of female 8.9-9 mm; male un- 
known. Head creamy yellow, with a spot on 
either side of median near posterior margin of 
crown, a spot on either side of median on 
anterior margin of crown, an area around an- 
tennae down lateral margins of postclypeus, 
lora, and anteclypeus black. Pronotum 
creamy yellow with a large spot either side of 
median. Scutellum black with a median, or- 
ange spot. Forewing green, with three black 
bands, one at end of scutellum, one across 
middle, and one at end of clavus, apical cells 
black. 

Female genitalia. — Seventh sternum 



80 



Great Basin Naturalist Memoirs 



No. 12 



large, roundedly produced, with a small, me- 
dian, V-shaped emargination (Fig. 63). 
Ovipositor extending beyond pygofer nearly 
twice its width. 

Type. — Holotype (female), Venezuela, 
T. F. Amazonas, 20 December 1981, eacuri, 
G. Yepez Gil Coll., University of Central 
Venezuela Collection. Paratype (female), 
Peru, Quincemil, August 1962, Luis E. Peria 
Coll., Ramos Collection. 

Notes. — The striking color pattern makes 
this species easy to identify. The two speci- 
mens differ slightly in color pattern and in the 
shape of the female genitalia, but this is proba- 
bly individual variation as they come from 
quite different localities. 

The fasciata species group is characterized 
by the longitudinal bands on the pronotum 
and the unusual, extra apodemal process on 
the aedeagus. The following two species be- 
long to this group. 

Calliscarta fasciata (Osborn) 
Figs. 55-58, 61 

Idiotettix fasciatus Osborn 1929: 466. (Hohenau, Itapua, 
Paraguay, Ohio State University Collection); 
Oman 1938: 395; Evans 1947: 190;' Nast 1952: 1; 
Maldonado Capriles 19.54: 248; Metcalf 1966: 226. 

Calliscarta fasciata: Linnavuori 1959: 30. 

Idiotettix brunnetis Osborn 1929: 467. (Hohenau, Itapua, 
Paraguay, Ohio State University Collection); 
Oman 1938: .395; Nast 1952: 1; Maldonado 
Capriles 1954: 248; Metcalf 1966: 226. Neiv syn- 
onijmij 

Calliscarta hrunnca: Linnavuori 1959: 32. 

Idiotettix laiitus Nast 1952: 3. (Porto Epitacio, Sao Paulo, 
Brazil, Polish Museum of Zoology); Maldonado 
Capriles 19.54: 248; Metcalf 1966: 227. 

Length of male 8-9.5 mm; female 9.8 mm. 
Head yellow, with a transverse band above 
ocelli, one below ocelli, a spot on each anten- 
nal ledge, a pair of longitudinal bands on post- 
clypeus, and much of gena and lora orange; 
a row of brown or black transverse lines on 
each side of postclypeus. Pronotum yellow, 
with anterior margin, four wide, longitudinal 
bands, and lateral margins orange. Scutellum 
yellow, with orange-brown longitudinal band 
down median. Forewings orange fading to 
smoky brown at apex; claval and discal veins 
bordered with translucent puri:)le bordered 
with dark brown, apical half with large, irreg- 
ularly shaped white spots bordered with 
brown, apical cells with whitish, translucent 
spots. Ventral surface mostly yellow. 

Male genitalia. — Pygofer large, tnmcate. 



with a long process from ventral margin (Fig. 
55). Genital plate large, expanded apical half, 
bluntly rounded at apex (Fig. 55). Style short, 
slightly curved laterad, with a pointed apex 
(Fig. 58). Aedeagus stout, curved dorsad api- 
cally, with a pair of lateral flanges from base of 
shaft to near apex, extra apodeme at base con- 
necting aedeagus to anal tube (Figs. 56, 57). 

Female genitalia. — Seventh sternum 
greatly enlarged, rounded on posterior mar- 
gin with a V-shaped, median emargination 
(Fig. 61). Ovipositor extending beyond 
pygofer its own width. Pygofer narrowed on 
apical third. 

Type. — Holotype (male), Paraguay, Ho- 
henau, Ohio State University Collection. 

Distribution. — This species is known from 
several hundred specimens from Brazil, 
Paraguay, and Peru. The only known female is 
in the University of Kansas Collection. 

Notes. — Linnavuori (1959) placed lautus as 
a synonym of fasciatus. The type of brunnea 
represents the teneral form of this species; 
consequently, it is placed in synonymy. 

Calliscarta mexicana, n. sp. 

Figs. 65, 71 

This species resembles fasciata, but it is 
smaller and the apical area of the forewing is 
without large, transparent spots. 

Length of female 8 mm; male unknown. 
Head creamy yellow, with a large, orange 
transverse band across middle of crown, an 
orange transverse band just below ocelli with 
ends below ocelli dark brown, an orange- 
brown transverse band across postclypeus 
halfway between antennae and ocelli with ex- 
tensions down both sides of postclypeus as a 
brown longitudinal band to anteclypeus, and a 
dark brown area around bases of antennae 
extending along lateral margins of post- 
clypeus. Pronotum creamy yellow with four 
wide, orange-brown longitudinal bands, me- 
dian pair extending as brown longitudinal 
bands on scutelhuu. Forewing orange-brown 
with longitudinal bands of creamy white, one 
along costa, one along claval vein on clavus, 
one along commissure, as well as white areas 
along apical longitudinal veins. 

Female genitalia. — Seventh sternum 
slightly enlarged, with posterior margin 
roundly produced with a slightly rounded me- 
dian emargination (Fig. 65). Ovipositor ex- 
tending only slightly more than its width be- 
yond pygofer. 



1988 



Freytag: Revision of Calliscarta 



81 



Type. — Holotype (female), Mexico, Tama- 
zunchale, S. L. Potosi, 19 June 1941, Henry 
S. Dybus Coll., Field Museum. 

Notes. — This species is very similar to fas- 
ciata in overall color pattern. These two spe- 
cies form a species group that is quite different 
from the other species of this genus. 

Acknowledgments 

Material for this study has been gathered 
over the last 20 years from the institutions 
listed below. I certainly thank the curators for 
their patience over these long years, for the 
loan of these most valuable specimens, and 
also for the opportunity to study type mate- 
rial: American Museum of Natural History, 
New York City, Dr. Jerome G. Rozen, Jr., 
and Dr. Randall T. Schuh; Carnegie Mu- 
seum, Pittsburg, Dr. Chen W. Young; Field 
Museum of Natural History, Chicago, Dr. R. 
Wenzel; Ohio State University, Columbus, 
Dr. Charles A. Triplehorn; Museum National 
d'Histoire Naturalle, Paris, Dr. Michel Bou- 
lard; Ramos Collection, Mayaguez, Dr. J. A. 
Ramos; United States National Museum, 
Washington, D.C., Dr. James P. Kramer; 
University of Kansas, Lawrence, Dr. George 



W. Byersand Dr. Peter D. Ashlock; Universi- 
dad Central de Venezuela, Instituto de Zoolo- 
gia Agricola, Maracay, the late Dr. F. J. Fer- 
nandez- Yepes. 

Literature Cited 

epilations prior to 195(i arc cited in Metcalf (1964). 

Kramer, J. P. 1963. New Neotropical Neobalinae with 
keys to the genera and to the .species of Conala 
(Homoptera: Cicadellidae). Proceedings of the 
Entomological Society of Washington 65: 201- 
210. 

LiNNAVUORi, R. 1959. Revision of the Neotropical Delto- 
cephalinae and some related subfamilies (Ho- 
moptera). Annales Zoologici Societatis Zoologicae 
Botanicae Fennicae 'Vanamo' 20(1); 1-369. 

LiNNAVUORi, R , AND F Heller 1961. Beitrag zur Ci- 
cadelliden, Fauna von Peru. Stuttgarter Beitrage 
zur Naturkunde 67: 1-14. 

Metcalf, Z. P 1964. General catalogue of the Ho- 
moptera, Fascicle VI, Cicadelloidea, bibliography 
of the Cicadelloidea. Agricultural Research Ser- 
vice, United States Department of Agriculture. 
349 pp. 

1966. General catalogue of the Homoptera, Fasci- 
cle VI, Cicadelloidea, Part 16, Idioceridae. Agri- 
cultural Research Service, United States Depart- 
ment of Agriculture. 237 pp. 

1967. General catalogue of the Homoptera, Fasci- 
cle VI, Cicadelloidea, Part 10, Euscelidae. Agri- 
cultural Research Service, United States Depart- 
ment of Agriculture. 2,695 pp. 



A NEW SPECIES OF PAZU OMAN FROM EASTERN CALIFORNIA 
(HOMOPTERA: CICADELLIDAE: DELTOCEPHALINAE) 

Raymond J. Gill' 

Abstract — A new species of Pazu, P. monoinyo, from the Inyo-Mono County area of California is described. 
Additional occurrence records are provided for Pazu halli (Beamer), and some of its morphological characteristics are 
illustrated and compared with P. monoinyo. 



The monobasic genus Pazu was erected by 
Oman (1949) to include Hebecephalus halli 
Beamer (1946) from central Arizona. Pazu is 
distinguished from Hebecephalus primarily 
by the shape of the styles, which in Hebe- 
cephalus have the apical portion bent laterad 
to the shaft at right angles, forming an elon- 
gate, slender "toe." The type species oiPazu 
and the new species described here have 
more conventional, straighter styles (see Figs. 
IE and 2E). 

Specimens of this new taxon were collected 
from antifreeze (ethylene glycol) pitfall traps 
placed in sandy areas or on small sand dune 
habitats in a 78-km" area in Inyo and Mono 
counties near Bishop, California, and from a 
single location 42 km east of Bishop near Deep 
Springs. The area is semiarid Great Basin 
habitat irrigated in the lower areas by 
snowmelt runoff from nearby mountain peaks 
or from artesian springs. The traps were 
placed at these and other sand dune locations 
throughout the southwestern U.S. as part of a 
joint project between the California Depart- 
ment of Food and Agriculture and the 
U.S. Bureau of Land Management (BLM) 
(Andrews et al. 1979). The purpose of the 
project was to study the insect fauna associ- 
ated with sand dune habitats as part of an 
effort by the BLM to preserve sand dune habi- 
tats. 

The fact that all known specimens of P. 
monoinyo exhibit microptery and have been 
collected from just three pitfall trap locations 
indicates that the species may spend consider- 
able time on the soil surface and on the crowns 
of the host rather than on aerial plant parts. 
Whether a fully macropterous form capable of 



flight actually exists in certain situations is 
unknown. A macropter collected at Blythe, 
Riverside County, California, on 2-IV-63 is 
nearly identical in color pattern to P. mono- 
imjo; however, the specimen is missing the 
abdomen, thus making accurate identification 
impossible. If such a form does exist, it is 
probably not common and may explain why 
the species has previously gone undetected. 
In all collection locations the dry sand dune 
habitats were in close proximity to water and 
the lush vegetative growth associated with it. 
The dune site east of Bishop, California, is 
within 100 m of the Owens River; the Fish 
Slough and Deep Springs locations are adja- 
cent to a series of artesian springs. Since the 
species has not been collected from other sand 
dune localities, it would appear that the spe- 
cies lives and feeds in or on the crowns of 
plants, presumably grasses, near permanent 
sources of surface water. However, several 
attempts to re-collect the species at two of the 
known localities have failed. 

Pazu monoinyo, n. sp. 

Description. — Resembles P. balli (Beamer) 
but with darker, more distinct markings and 
different genitalia. 

External CHARACTERISTICS. — Groundcolor 
apparently cinereous, although the possible 
color-altering effects of the antifreeze and al- 
cohol preservatives leave the actual color in 
doubt. Markings fuscous. Crown with a 
broad, longitudinal pair of lines, each with a 
central cinereous line, so that the markings 
appear as two pairs of fuscous lines on either 
side of the midline; with a fuscous spot be- 
tween each ocellus and compound eye. Frons 



Analysis and IdentiPication, Division of Plant Industry, California Department of FcKid and Agriculture. 1220 N Street, Sacramento, California 95814. 



82 



1988 



GiLL: A New Pazu from California 



83 







B 



Fig. 1. Pazu rnonoimjo, n. sp.: A, adult female habitus; B, male aedeagus and connective, dorsal view; C, male 
aedeagus and connective, lateral view; D, apex of male abdomen, lateral view; E, apex of male abdomen, ventral view. 




with two pairs of arclike, subapical, fuscous 
markings and occasionally with a pair of 
rectangular, basal, fuscous spots next to 
clypeus; frons apparently never clouded with 
fuscous as in P. balli. Thorax with six fuscous, 
longitudinal lines, thickest lines medial, all 
lines lighter than those on crown. Abdominal 
dorsum with three pairs of longitudinal, fus- 
cous lines, the lateral pairs often coalescing to 
form bands twice the width of medial pair. 
Venter cinereous variously marked with fus- 
cous, especially on anterior median areas of 
sternites. Legs cinereous with two or more 
fuscous spots or bars on femora, other leg 
segments darker near spines and at spine 
bases. Elytra reduced (micropterous as delin- 
eated by Oman 1987), barely reaching ninth 
abdominal segment. Most elytral cells filled to 
a greater or lesser degree with fuscous; all 
veins or cells on either side of claval vein 
usually clear or white. Wings reduced, ca 1/2 x 



length of elytra. There appears to be no color 
or other dimorphisms between the sexes. 

Males — Length 3.4-3.7 mm (mean 3.6 
mm), specimens slightly shriveled due to 
preservation methods. Wing length 2.4-2.7 
mm (mean 2.6 mm); vertex length 0.4-0.6 
mm (mean 0.5 mm); transocular width 
1.0-1.3 mm (mean 1.2 mm); interocular 
width 0.4-0.6 mm (mean 0.6 mm). Pygofer 
shorter than in P. balli, more truncated, with 
just a vestige of a tooth on lateroventral cor- 
ners; weakly setose. Plates evenly tapered to 
acute apices and completely divided, not 
basally fused and truncate as in P. balli; spine- 
like setae uniseriate. Style apices J-shaped, 
medial apex long, slender, very slightly 
curved laterad; lateral tooth broadly triangu- 
lar. Connective linear, about as long as aedea- 
gal shaft. Aedeagus roughly V-shaped; ante- 
rior arm broader, apex curved posteriorly; 



84 



Great Basin Naturalist Memoirs 



No. 12 






Fig. 2. Pazu halli (Beamer): A, adult female habitus; B, male aedeagu.s and connective, dorsal view; C, male aedeagus 
and connective, lateral view; D, ape.x of male abdomen, ventral view. 



posterior arm ca 1/2 x width of anterior, gradu- 
ally and evenly tapered to apex, crescentic in 
lateral view. Aedeagal apex with three pairs of 
apicolateral flanges or blunt teeth. 

Females. — Length 4.6-5.0 mm (mean 4.6 
mm). Wing length 2.9-3.1 mm (mean 3.0 
mm); vertex length 0.7-0.9 mm (mean 0.7 
mm); transocular width 1.4-1.6 mm (mean 
1.4 mm); interoeular width 0.6-0.9 mm 
(mean 0.6 mm). Last ventral segment with 
broadly rounded, posteriorly directed median 
lobe. 

Etymology. — The name monoinyo is a 
noun in apposition referring to the two locali- 
ties where the species was collected. Mono 
and Inyo counties, California. 

Discussion. — Pazu monoinyo is similar to 
the generic type, P. halli, in color pattern and 
male genitalia. Color patterns differ in that 
the dorsal, longitudinal lines tend to be much 
darker and more distinct on the head and 
thorax in P. monoinyo, but much lighter and 
with a tendencv to coalesce in P. halli. The 



male genitalia differ in that the pygofer is 
much shorter in P. monoinyo; the plates are 
completely divided and have acute apices in 
P. rnonoinyo compared with basally fused, 
truncate plates in P. halli. The last ventral 
segment of the female is lobed posteriorly in 
P. monoinyo but deeply cleft in P. halli. All 
known specimens of P. monoinyo exhibit mi- 
croptery, whereas all those of P. halli are 
macropterous. 

Specimens of P. halli are known from 
Cochise and Pearce, Cochise County, Ari- 
zona. 

Material examined. — Male holotype and 
female allotype: "Calif, Mono Co., 9 mi N 
Bishop, Fish Slough, 4,200', sand dunes, V- 
12-82 to XII-20-82, Derham Giuliani, col- 
lected in ethylene glycol pit trap." Paratypes: 
"Calif, Mono Co., 9 mi N Bishop, Fish 
Slough, 4,200', .sand dunes, V-12-82 to XII- 
20-82, Derham Giuliani, collected in ethyl- 
ene glycol pit trap," 4 males and 2 females; 
"Calif, 3 mi E Bishop, 4,100', sand dunes, 



1988 



GiLL: A New Pazu from California 



85 



V- 12-82 to XII-10-82, antifreeze pit trap, D. 
Giuliani," 1 male and 4 females; "Inyo Go., 
Ga. , Deep Springs Valley, VI-9-78, antifreeze 
pit trap, D. Giuliani," 1 female. 

Type deposition. — The holotype and allo- 
type (both GAS Type # 16039) were deposited 
in the Galifornia Academy of Sciences, San 
Francisco (GAS). Twelve paratypes were de- 
posited as follows: 2 males, 2 females, San 
Francisco (GAS); 2 males, 4 females, Galifor- 
nia Department of Food and Agriculture, 
Sacramento (GDFA); 1 male, 1 female. 
United States National Museum, Washing- 
ton, D.G. (USNM). 

Acknowledgments 

Thanks must go first to Dr. Paul Oman, who 
has been such a great source of inspiration for 
all of us who work on the Gicadellidae. His 
publications have been extremely valuable to 
me in the course of my work as a taxonomist 
and identifier, and his 1949 book on the 
Nearctic lealhoppers was the primary build- 
ing block toward my understanding the Gi- 
cadellidae. 

I express gratitude to Dr. Leon W. Hep- 
ner, Mississippi State University, State Gol- 



lege, Mississippi, and to Dr. Steve Wilson, 
Gentral Missouri State University, Warrens- 
burg, Missouri, for reading the manuscript 
and offering useful suggestions. I also thank 
Dr. Robert Brooks, Snow Entomological Mu- 
seum, Lawrence, Kansas, for the loan of the 
type specimens oi Pazu balli. I am extremely 
grateful to Susan Sawyer, GDFA, for the 
habitus sketches of Pazu monoinyo and Pazu 
balli. 

Literature Gited 

Andrews, F G , A R Hardy, and D Giuliani 1979. The 
coleopterous fauna of selected California sand 
dunes. Report to the ELM on contract CA-960- 
128.5-1288 DEOO, California Department of 
Food and Agriculture, Sacramento. 142 pp. 

Reamer, R. H 1936. New leafhoppers from the western 
United States (Homoptera: Gicadellidae). Cana- 
dian Entomol. 68: 2.52-2.57. 

Oman, P W 1949. The Nearctic leafhoppers (Ho- 
moptera: Gicadellidae). A generic classification 
and checklist. Washington Entomol. Soc. Mem. 3: 
1-2.53. 

1987. Alary polymorphism in the Gicadellidae and 

its ecological implications. Pages 55-63 in M. R. 
Wilson and L. R. Nault, eds., Proc. 2nd Int. 
Workshop on Leafhoppers and Planthoppers of 
Economic Importance, Provo, Utah, USA, 28 
July-1 August 1986. GIE, London. 



EVIDENCE FOR AN INDO-PACIFIC ORIGIN OF HAWAIIAN ENDEMICS 
IN BALCLUTHA AND RELATED GENERA (CICADELLIDAE: MACROSTELINI) 

W. J. Knight' and M. D. Webb' 

Abstract — The 10 endemic species of Balclittha in Hawaii are reviewed and compared with the Pacific and 
American forms of the genus. Two new species, nigriventris and fiiscifrons , are described; kilaueae is transferred to a 
new genus Balolina n. comb. ; beardsleyi is synonymized with saltuella , n. syn. ; and hospes is relegated to a subspecies 
oiincisa, n. stat. Descriptions and notes, together with illustrations and a key, are given for each taxon. Evidence is 
presented to suggest that all the endemic species oi Balclittha have arisen within the islands from a single colonization 
and that the taxon showing the greatest similarity to them is the widespread Pacific species B. lucida. It is shown that 
the Hawaiian genus Nesolina is most closely related to the endemic species-group oi Balclutha and that Balolina is an 
independent colonization. 



The isolated position, volcanic origin, and 
diverse habitats of Hawaii, together with the 
division of the area into several large islands, 
have combined to produce a depauperate, yet 
large and highly endemic, biota (Simon 1987). 
This phenomenon is well illustrated by the 
plant-feeding insect family Cicadellidae. The 
family was studied in the islands by Kirkaldy 
(1910) and Osborn (1935) and subsequently 
reviewed by Zimmerman (1948). Namba 
(1956) later described several new species. 

In his review of the Hawaiian insect fauna, 
Zimmerman (1948) considered the generic 
affinities to be predominantly Pacific and only 
5% American. The Homoptera were consid- 
ered to be exclusively Pacific, although little 
evidence was provided. 

Compared with the Pacific and mainland 
America, the Hawaiian cicadellid fauna is very 
depauperate at the supraspecific level. Only 5 
of the 35 subfamilies are present, 4 of which 
are represented by 5 recently introduced spe- 
cies from North America. In contrast, the 
Deltocephalinae, a large, worldwide group 
with 17 currently recognized tribes, is repre- 
sented by 2 tribes. The endemic genera 
Nesophryne (2 species), Kirkaldiella (2 spe- 
cies), and Nesophrosyne (60 species) belong to 
the Opsiini and are related to the Pacific 
genus Orosius (Linnavuori 1960, Ghauri 
1966). Nesolina (1 species), Balolina n. gen. (1 
species), and the cosmopolitan genus Bal- 
clutha, comprising 105 species and repre- 
sented in Hawaii by 11 endemic and 2 cos- 



mopolitan species, belong to the Macros- 
telini, the major cicadellid group in the simi- 
larly isolated islands of Marquesas in the 
Pacific, and St. Helena in the Atlantic. This 
tribe contains approximately 30 genera world- 
wide, of which 10 are island endemics, 2 in 
Hawaii, 4 in the Marquesas, and 4 on St. 
Helena. Interestingly, although the main host 
plants of the group are Gramineae, in Hawaii, 
Marquesas, and St. Helena shrubs and tree 
ferns of southern temperate distribution are 
also utilized (Webb 1987). 

Materials 

The material studied is primarily from the 
Bishop Museum, together with some in the 
British Museum (Natural History), abbrevi- 
ated throughout the text to BPBM and 
BMNH, respectively. 

Methodology 

The present study investigates the origins 
of the Hawaiian Balclutha species. This has 
necessitated a reexamination of all the Hawai- 
ian species and a review of the variation in 
characters among Pacific and American spe- 
cies of the genus. Until a phylogenetic analy- 
sis of the genus and tribe is undertaken, any 
statements on the polarity of characters within 
these taxa would be premature. Our bio- 
geographical inquiry is therefore based on 
phenetic resemblances rather than on shared 



British Museum (Natural History), Cromwell Road, London SW7 5BD, Enfiland. 



86 



1988 



Knight, Webb: Hawaiian Endemics in Balclutha 



87 



apomorphies and should be seen as only ex- 
ploratory and preliminary in nature. 



Faunal Relationships of 
Hawaiian Balclutha 

The variation in morphological characters 
in Balclutha was reviewed by Knight (1987) 
for the south and west Pacific species and by 
Blocker (1967) for the American species. Both 
faunas are of comparable size, 30 and 32 spe- 
cies, respectively, but have only 3 species in 
common, saltuella, incisa, and lucida. 

Compared with the Pacific and American 
species, the 10 endemic Hawaiian species, 
nigriventris n. sp., fuscifrons n. sp., grandis 
Namba, phoxocephala Namba, timberlakei 
(Osborn), lohata Namba, usitata Namba, vol- 
canicola (Kirkaldy), plutonis (Kirkaldy), and 
peregrina (Kirkaldy), differ only slightly from 
each other in head shape and minor details X)f 
the male genitalia, suggesting that they prob- 
ably arose from one founder species. The 
character states common to the 10 endemic 
species are listed below, the condition 
present in the 10 endemic species shown in 
italics: 

1. Relative width of head and pronotum: head equal to 
or wider than the pronotum. 

2. Length of vertex: longer medially than next to eyes. 

3. Laterofrontal sutures: terminating at or near the 
ocelli. 

4. Distance between ocelli and eyes: ocelli separated 
from the eyes by a distance equal to 2-4 times their 
own diameter. 

5. Foretibia setal formula; 1.3. 

6. Hind femora setal formula: 2. i.i. 

7. Shape of male pygofer: simple, without ventral lobes 
or processes. 

8. Shape of subgenital plates: triangtdate, with apical, 
fingerlike process 1/4-1/2 total length of plate. 

9. Shape of styles: robust, with apical process and 
preapical lobe well developed. 

10. Relative length of stem and arms of connective: stem 
longer than arms. 

11. Shape of aedeagal shaft: simple, elongate, curving 
dorsally or ante r odor sally to point level with or 
slightly anterior to atrium. 

Comparison between the Hawaiian, 
Pacific, and American Species 

In the present section, the Pacific and 
American species of the genus are reviewed 
for each of the characters listed above. The 
alternative conditions of characters 3, 5, and 
7 are coded A and B for ease of reference in 



subsequent discussion. 

The relative width of the head (character 1) 
varies within the genus and may be wider, 
equal to, or narrower than that of the prono- 
tum. All three conditions are present in both 
the American and the Pacific faunas, half the 
species in each region showing the head ap- 
proximately equal to or wider than the prono- 
tum, as in Hawaii. 

The vertex (character 2) is usually short and 
uniform in length and is one of the main dis- 
tinguishing features of the genus. The medial 
prolongation of the vertex found in the Hawai- 
ian species occurs in the Pacific in only 
smaragdula , and in a few individuals offlagel- 
lata, bifasciata, rieki, and the cosmopolitan 
species punctata. It is far more prevalent 
among the American forms, being a character- 
istic feature of 7 species and occurring occa- 
sionally in 15 others. 

The laterofrontal sutures (character 3) usu- 
ally terminate at or near the ocelli (condition 
B) but are occasionally turned medially, ven- 
trad of the ocelli (condition A). With the ex- 
ception of simplex and the cosmopolitan 
saltuella , the Pacific species have the sutures 
terminating near the ocelli as in Hawaiian 
species. Both conditions also occur in the 
American fauna but with an approximately 
equal number of species of each. 

The ocelli (character 4) are usually small 
and situated on the anterior margin of the 
head, separated from the corresponding eye 
by approximately their own diameter. Most of 
the American and Pacific species have the 
ocelli in this position, or only slightly more 
remote from the eye. Only i7npicta and arc- 
tica in America, flagellata and cheesmanae in 
the Pacific, and the cosmopolitan species 
punctata, have the ocelli more than twice 
their own diameter from the eyes, and ap- 
proximating the condition found in Hawaiian 
species. The Pacific fauna shows greater flexi- 
bility in this character, however, with chlor- 
optera and hulhosa having the ocelli enlarged 
and more dorsad in position. 

Throughout the genus, the foretibia setal 
formula (character 5) may be 1. 1 (condition A), 
1.3 (condition B), or rarely 1.4. The condition 
in Hawaii (condition B) is similar to that in the 
majority of Pacific species. Only two Pacific 
species, simplex and the cosmopolitan saltu- 
ella, show condition A. In the American 
fauna, half of the species examined have con- 
dition A and half condition B. 



Great Basin Naturalist Memoirs 



No. 12 



The setal pattern at the apex of the hind 
femora (character 6) varies within the genus 
from 2. 1. 1 to 2.2. 1. The two conditions occur 
equally throughout both the Pacific and 
American faunas, unlike Hawaii where only 
the former is found. 

The male pygofer (character 7) varies from a 
simple, rounded shape (condition A), as found 
in Hawaii, to a more complex form with pos- 
teroventral lobes or processes (condition B). 
The simple form is rare outside Hawaii, occur- 
ring in only simplex in the Pacific and in the 
species lucida and saltuella, which occur in 
both regions. 

The subgenital plates (character 8) are char- 
acterized in all species by a membranous, fin- 
gerlike, apical segment. The shape of the 
plate found in the Hawaiian species occurs in 
the majority of Pacific and American species, 
being slightly more prevalent in the latter 
region. The remaining species in both regions 
have the basal part of the plate much reduced 
in size. 

The robust, well-developed styles (charac- 
ter 9) of the Hawaiian species occur in the 
majority of species in both the Pacific and 
America but are more widespread in the for- 
mer. Only two Pacific species, bulbosa and 
rieki, have a reduced style, compared with 
nine species in the Americas, plus the cosmo- 
politan saltuella in both regions. 

The Y-shaped connective (character 10) 
varies principally in the relative length of the 
stem and arms. The condition present in 
Hawaii, with the stem longer than the arms, is 
present in 50% of the species in the Pacific but 
in only 30% of the species in America. 

A simple, filamentous aedeagus (character 
11) is characteristic of the majority of species 
in the genus. In the Hawaiian species the shaft 
is directed dorsally or curved anterodorsally 
to a point approximately level with the well- 
developed, basal apodeme, a form that occurs 
in 50% of the fauna in both the Pacific and 
America. 

Faunal Similarity 

If we take the number of species in each 
region showing character states identical with 
those in Hawaii as a measure of similarity, 
then the Hawaiian species show greater simi- 
larity to the Pacific fauna with respect to the 
laterofrontal sutures, foretibial setal pattern, 
styles, connective, and to a lesser extent the 



ocelli and male pygofer. The American spe- 
cies show greater similarity only in the length 
of the vertex and to a lesser extent in the shape 
of the subgenital plates. The relative width of 
the head, the setal pattern on the hind 
femora, and the shape of the aedeagus provide 
little information in this respect. If we elimi- 
nate those characters that confer only minimal 
regional preference, such as the ocelli, male 
pygofer, and subgenital plates, and also those 
known to vary infraspecifically, such as the 
length of the vertex, we are left with only four, 
the laterofrontal sutures, the foretibial setal 
pattern, and the shape of the styles and con- 
nective, all of which support the theory of 
closer links with the Pacific. 

Further support for a Pacific link is pro- 
vided by the Sorensen Coefficient of Similar- 
ity (Southwood 1978) 

Cs = 2j/(a + b) 
where j is the number of character states com- 
mon to the two areas and a -I- b are respec- 
tively the number of character states in each 
area. Based on the endemic species in each 
area, the coefficient for Hawaii and the Pacific 
is 0.74 and for Hawaii and the Americas is 
0.58, showing a greater degree of similarity 
between the first two areas. 

From the above review it is seen that no 
one character is shared exclusively between 
Hawaii and either of the other two regions to 
provide evidence of affinity. However, if we 
consider a combination of characters, rather 
than single characters alone, then the picture 
becomes a little clearer. 

Table 1 shows the Pacific species and their 
similarity or dissimilarity to the Hawaiian en- 
demic species with regard to the 11 charac- 
ters. This reveals an obvious correlation be- 
tween characters 3, 5, and 7. All three 
characters show condition A" in the first two 
species and condition B" in the fointh species 
onward. An intermediate stage occurs in the 
third species, lucida , which has condition B in 
characters 3 and 5 but retains condition A in 
character 7. The American species show the 
same correlation between characters 3, 5, and 
7, but, except for lucida , the character states 
in the intermediate stage are reversed. This is 
shown in Table 2, where the intermediate 
stage is highlighted for each region. The simi- 
larity between the Hawaiian endemics and 
lucida in this respect is unique. 



; scttion, paragraphs 4, 6. ami H. 



1988 



Knight, Webb: Hawaiian Endemics in Balclutha 



89 



Table 1. A list of the Pacific species oi Balcltttha. showinjz; the similarities ( + ) and differences ( — ) between them and 
the Hawaiian endemics for the 1 1 listed characters. (For further explanation, see text.) 



Pacific 










Characters 












species 


1 


2 


3 


4 


5 


6 


7 


8 


9 


10 


11 


saltuella 


+ 


- 


- 


- 


— 


+ 


+ 


- 


- 


- 


+ 


simplex 


+ 


- 


- 


- 


- 


+ 


+ 


- 


+ 


- 


+ 


lucida 


+ 


— 


+ 


- 


+ 


+ 


+ 


+ 


+ 


— 


- 


bilobata 


- 


- 


+ 


- 


+ 


+ 


- 


- 


+ 


- 


- 


batuensis 


+ 


- 


+ 


- 


+ 


- 


- 


+ 


+ 


+ 


+ 


punctata 


- 


+ 


+ 


+ 


+ 


- 


- 


+ 


+ 


+ 


+ 


neglecta 


+ 


- 


+ 


- 


+ 


+ 


- 


+ 


+ 


- 


+ 


viridinervis 


+ 


- 


+ 


- 


+ 


- 


- 


+ 


+ 


+ 


- 


flageUata 


- 


± 


+ 


+ 


+ 


- 


- 


+ 


+ 


± 


- 


bifasciata 


+ 


■± 


+ 


- 


+ 


+ 


- 


- 


+ 


+ 


+ 


trilineata 


+ 


- 


+ 


- 


+ 


- 


- 


+ 


+ 


- 


- 


rieki 


+ 


■±_ 


+ 


- 


+ 


+ 


- 


- 


- 


- 


+ 


cheesinanae 


- 


- 


+ 


+ 


+ 


- 


- 


+ 


+ 


+ 


- 


yanchepensis 


-H 


- 


+ 


- 


+ 


- 


- 


+ 


+ 


+ 


+ 


smaragdula 


- 


+ 


+ 


+ 


+ 


+ 


- 


- 


+ 


+ 


+ 


alstoni 


+ 


- 


+ 


- 


+ 


- 


- 


+ 


+ 


+ 


+ 


kuroiwae 


+ 


- 


+ 


- 


+ 


? 


- 


- 


+ 


+ 


+ 


asyrmnetrica 


- 


- 


+ 


- 


+ 


? 


- 


+ 


+ 


- 


- 


chloroptera 


- 


- 


+ 


+ 


+ 


- 


- 


+ 


+ 


+ 


- 


bulbosa 


- 


- 


+ 


+ 


+ 


- 


- 


+ 


- 


+ 


- 


bacchusi 


+ 


- 


+ 


- 


+ 


+ 


- 


+ 


+ 


+ 


+ 


spiniloba 


+ 


- 


+ 


- 


+ 


? 


- 


+ 


+ 


+ 


+ 


incisa 


+ 


- 


+ 


- 


+ 


+ 


- 


+ 


+ 


+ 


- 


rosea 


+ 


- 


+ 


- 


+ 


+ 


- 


+ 


+ 


- 


+ 


pseudorosea 


+ 


- 


+ 


- 


+ 


+ 


- 


+ 


+ 


- 


+ 


rubrostriata 


+ 


- 


+ 


- 


+ 


+ 


- 


+ 


+ 


+ 


+ 


wilsoni 


- 


- 


+ 


- 


+ 


- 


- 


+ 


+ 


- 


- 


distenda 


- 


- 


+ 


- 


+ 


- 


- 


+ 


+ 


- 


- 


riibrinervis 


- 


- 


+ 


- 


+ 


- 


- 


+ 


+ 


- 


- 


noonadana 


+ 


- 


+ 


- 


+ 


- 


- 


+ 


+ 


- 


- 



Table 2. Character-state distribution for three characters in Pacific and American species of Balclutha. (For further 
explanation, see text.) 







Characters 




American 
species* 




Chi 


iracters 




species 


3 


5 


7 


3 




5 


7 


saltuella 


A 


A 


A 


saltuella 


A 




A 


A 


simplex 


A 


A 


A 


neglecta 


A 




A 


B 


lucida 


B 


B 


A 


chiasma 
sandersi 


A 
A 




A 
A 


B 
B 


remaining spp. 


B 


B 


B 


remaining spp. 


B 




B 


B 



*lucida. which occurs in Central and South America, has been omitted from this half of the table in the interests of clarity. 



Distribution of B. lucida 

Since lucida is the species most similar to 
the Hawaiian forms, it is of interest to examine 
its distribution relative to Hawaii. Figure 53 
shows its distribution, extending from the 
Aldabra Islands (Webb 1980 \filum]) in the 
west to Central and South America in the east 
(Blocker 1967 \jloridana ] and Knight 1987). It 
is a tropical and semitropical species clearly 



capable of movement over long distances and 
with the ability to colonize and establish itself 
on islands and mainland areas. Its presence in 
the Marshall and Marquesas islands places it 
within a comparatively short distance from 
Hawaii. The available data, however, provide 
no information on the center of origin for the 
species, although its absence from mainland 
Africa and Asia suggests that it is either the 
Pacific or Central America. 



90 



Great Basin Naturalist Memoirs 



No. 12 



Relationship of Balclutha to Other 
Hawaiian Macrostelini 

In addition to Balclutha , two other macros- 
teline genera occur in Hawaii, both endemic. 

The monotypic genus Nesolina differs from 
Balclutha principally in the more elongate 
head and the possession of longitudinal brown 
stripes on the vertex and pronotum. In all 
other characters it closely resembles the en- 
demic species of Balclutha to which it is prob- 
ably most closely related. Despite this close 
similarity, the present status of Nesolina is 
preserved, pending a phylogenetic study of 
the Macrostelini. 

The relationship of the remaining Hawaiian 
macrosteline genus, Balolina n. gen., is un- 
certain. Although it is similar in general ap- 
pearance to Balclutha and the Pacific genus 
Nesoclutha and has the first valvulae fused as 
in Balclutha (Fig. 33), it differs in many signif- 
icant features, as listed below in the key and 
generic description. Except for the position of 
the ocelli, there is little to indicate that it is in 
any way related to the Hawaiian species of 
Balclutha . On the contrary, it appears to be 
an independent introduction to the islands. 

All three Hawaiian macrosteline genera be- 
long to a group having the hindwing veins sc 
and r united into a single vein, with a conse- 
quent reduction in the number of apical cells 
(Fig. 32). This is considered by Ossiannilsson 
(1983) to be justification for placing Balclutha 
in a separate tribe, the Balcluthini, which is 
also characterized (Oman 1949) by the posses- 
sion of a sulcate hind basal tarsus (Fig. 74). 
Both characters are found to some degree, 
however, in other genera, including Macro- 
steles . The phylogeny of the Macrostelini pro- 
posed by Triplehorn and Nault (1985) in- 
cludes Balclutha but omits several other gen- 
era, including the 10 island endemics. Some 
of these have been treated by Webb (1986, 
1987). 

Distribution of Hawaiian Macrostelini 

Table 3 summarizes previous (x) and new 
(N) distribution data and shows that the 
largest and youngest islands, Hawaii and 
Maui, respectively, support the most species, 
and that Kauai and Molokai, which differ con- 
siderably in size, have similar numbers of spe- 
cies. Further collecting on the islands may 



have profound effects on the distributional 
records and may even reveal further species. 

Keys and Descriptions 

The taxonomy of the Hawaiian Macrostelini 
has been treated by several authors (see in- 
troduction), but descriptions are usually 
incomplete and contain little information on 
intraspecific differences. Namba's (1956) revi- 
sion o( Balclutha was a considerable improve- 
ment on previous work, but a few of the char- 
acters referred to are unreliable. New keys to 
genera and species are given to take account of 
this and to accommodate the new taxa. All 
host records given below are taken from ei- 
ther Zimmerman (1948), Namba (1956), or 
specimens examined. 



Kev to the Genera of Hawaiian Macrostelini 



1. Head and thorax with longitudinal, brown 

bands; vertex elongate (Fig. 54) Nesolina 

— Head and thorax not marked as above; vertex 
short to moderately long (Figs. 7, 15) 2 

2(1). Vertex moderately long, ocelli situated almost 
to point midway between eye and midlength of 
foremargin of vertex (Fig. 67). Clypellus ex- 
tending well beyond margin of face, trans- 
clypeal suture obscure; laterofrontal sutures 
short (Fig. 68). Foretibia with apical setae simi- 
lar in length (Fig. 73). Hind femur with apical 
setal formula 2 + 2+1 (Fig. 69). Macrosetae of 
legs and genital segment nonhairy. Aedeagal 
shaft with pair of apical processes (Fig. 82); 
gonopore apical on ventral surface (Fig. 76). 
Connective with stem very short (Fig. 80). 
Subgenital plate with irregular macrosetae 
(Fig. 83). Second valvulae evenly tapered to 
apex; teeth extending over distal half of dorsal 
margin and onto apex of ventral margin (Fig. 
75) Balolina 

— Vertex short to moderately long; ocelli situated 
near eye or at point one-third distance from eye 
to midlength of foremargin of vertex (Figs. 2, 
15, 28). Foretibia with apical setae dissimilar in 
length (Fig. .35). Hind femur with apical setal 
formula 2+1 + 1 (Fig. 30). Male and female 
genitalia not as above Balclutha 

Key to Species of Hawaiian Balclutha 

1. Vertex of uniform length; ocelli 1.0-1.5 times 
own diameter from corresponding eye; coronal 
suture visible in facial view (Figs. 28, 29) 
Pygofer lobe with a ventral process (Figs. 37, 
46). Connective with arms ecjual in length to 
stem (Fig. 42) 11 

— Not as above 2 

2(1). Yellow to greenish yellow, marked with brown 
on first thoracic sternite (as in saltuella. Fig. 



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Knight, Webb: Hawaiian Endemics in Balclutha 



91 



Table 3. Distribution ofthe Macrostt'lini on the Hawaiian islands, from the oldest in the north (left) to the youngest 
in the south (right). 



Area (sq km) 



Kauai 

1437 



Oahu 
1564 



Molokai 

673 



Lanai 
365 



Maui 
1885 



Hawaii 
10437 



Nesolina 
line at a 

Balolina 
kilaueae 

Balclutha 
grandis 
saltuella 
phoxocephala 
incisa hospes 
timberlakei 
lobata 
usitata 
volcanicola 
plutonis 
peregrina 
nigriventris 
fuscifrons 

Total 



N 
N 
10 



11 



43), abdomen (at least dorsally), and sometimes 
legs 3 

— Not marked as above or if abdomen and first 
thoracic sternite marked with brown, then 
overall color yellow tinged with scarlet 5 

3(2). Large species, 4.0-4.4 mm in length. Aedea- 

gus and style as in Figures 3, 4, 18 

fuscifrons. n. sp. 

— Small to moderately large species, up to 3.5 
mm 4 

4(3). Vertex marked with stramineous or brown 
(Figs. 5, 7). Aedeagal shaft robust (Fig. 24) . . . 
volcanicola (Kirkaldy) 

— Vertex without markings. Aedeagal shaft slen- 
der (Fig. 19) nigriventris, n. sp. 

5(2). Large species 4.5-5.3 mm in length. Stem of 
connective broad. Aedeagus as in Figure 21 . . 
grandis Namba 

— Small to moderately large species, up to 4.0 
mm 6 

6(5). Preapical lobe of style large; apical process 
abruptly tapered distally (Figs. 16, 17). Female 
genital sternite marked with brown on disc 

(Fig. 8) lobata Namba 

— Not as above 7 

7(6). Moderately large species, 3.5-4.0 mm. 
Clypellus with sides concave. Connective 
elongate, longer than style (as in timberlakei. 
Figs. 13, 14). Aedeagal shaft slender, evenly 
curved dorsally (Figs. 25, 26) . plutonis (Kirkaldy) 

— Small species, 2.6-3.1 mm; without the above 
combination of characters 8 

8(7). Aedeagal shaft slender (Figs. 25, 27) 9 

— Aedeagal shaft robust (Figs. 22, 23) 10 

9(8). Clypellus with sides straight (Fig. 12). Aedea- 
gal shaft evenly curved throughout length (as in 



plutonis. Fig. 25) peregrina (Kirkaldy) 

— Clypellus with sides concave. Aedeagal shaft 

angularly curved basally (Fig. 27) 

usitata Namba 

10(8). Aedeagal shaft sinuate (Fig. 22). Connective 
elongate, longer than style (Figs. 13, 14) .... 
timberlakei (Osborn) 

— Aedeagal shaft not sinuate (Fig. 23). Connec- 
tive approximately equal in length to style . . . 
phoxocephala Namba 

11(1). Female pregenital sternite with disc marked 
with brown (Fig. 36). Male genitalia as in Fig- 
ures 37-40 incisa hospes (Kirkaldy) 

— Female pregenital sternite not marked with 
brown. Male genitalia as in Figures 41, 42, 45, 

46 saltuella (Kirschbaum) 

Balclutha Kirkaldy 

Balclutha Kirkaldy, 1900: 243. New name for Gnathodus 
Fieber, 1866; Blocker 1967; 4, Knight 1987: 1178. 
Type species: Cicada punctata Fabricius, 1775, 
by monotypy. 

Balclutha fuscifrons , n. sp. 
Figs. 1-4, 18 

Length. — S , 4.0-4.2 mm; 9 , 4.2-4.4 mm. 

Yellow tinged with green, abdomen heavily 
marked dorsally and ventrally with dark 
brown; clypeus pale to dark brown; legs and 
thoracic sternites variably marked with dark 
brown. 

Head approximately equal in width to 
pronotum. Vertex with medial length approx- 
imately 1.33 times length next to eye. Ocelli 
marginal, separated from corresponding eye 
by distance equal to 3-4 times own diameter. 



92 



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No. 12 




Figs. 1-17. Endemic Hawaiian Balclutlia species. 1-4, B. fuscifrons: 1, male pygofer, left lateral view; 2, head and 
thorax, dorsal view; 3, right style, dorsal view; 4, connective, dorsal view. 5-7, B. volcanicohi: 5. head and thorax, 
dorsal view (Kilauea); 6, right style, dorsal view; 7, head and th()ra.\, dorsal view (Wailua). 8, B. lohata. female 
pregenital sternite. 9-11, B. nigriventris: 9, right style, dorsal view; 10, connective, dorsal view; 11, head and thorax, 
dorsal view. 12, B. peregrina, face. 13-15, B. timhcrlakci: 13, right style, dorsal view; 14, connective, dorsal view; 15, 
head and thorax, dorsal view. 16-17, B. lohata, apex of right style, dorsal view. 



Face slightly wider than long; clypellu.s with 
sides concave, apex extending very slightly 
beyond margin of face; laterofrontal sutures 



convex, distally concave, reaching to corre- 
sponding ocellus. Setal formulae: foretibia, 
1.3 or 1.4; midtibia, 4.3, 4.4, 5.3, or 5.4; hind 



1988 



Knight, Webb: Hawaiian Endemics in Balclutha 



93 



femur, 2.1.1 or 2.1.1.1. 

Male. — Pygofer simple, posterior margin 
broadly and evenly rounded in lateral aspect. 
Subgenital plate tapered distally to moder- 
ately long, fingerlike process. Connective 
with arms shorter than stem. Style approxi- 
mately equal in length to connective, preapi- 
cal lobe elongate. Aedeagal shaft slender, 
more or less straight distally. 

Female. — Pregenital sternite with poste- 
rior margin more or less straight. Valvulae as 
in incisa (Figs. 33, 38). 

Host. — Unknown. 

Distribution. — Endemic (Maui). 

HOLOTYPE. — 6, Maui, Kula Pipe Line, 
4,200', vii.1956, R. Namba (BPBM). 
Paratypes: Maui: 13(5, 199, same data as 
holotype; 4c5, Holua, Haleakala Crater, 
vi.l983, D. E. Hardy; 4c?, 99, Waikamoi, 
4,000', D.E. Hardy (BPBM, BMNH). 

Remarks. — This species can be distin- 
guished by its large size, brown markings, and 
more or less straight aedeagal shaft. Its exter- 
nal appearance is similar to Balolina hilaueae, 
with which it may be confused. 

Balclutha volcanicola (Kirkaldy) 

Figs. 5-7, 24 

Nesosteles volcanicola Kirkaldy, 1910; 574: Osborn 1935; 

58, Fig. 24e. Lectotype S, Hawaii (BPBM), here 

designated [examined]. 
Balclutha volcanicola (Kirkaldv); Zimmerman 1948: 87, 

Fig. 27, Namba 1956; 107-108, Fig. 7. 

Host. — Cyperus (Cyperaceae), Eragrostis 
(Gramineae), Lythrum (Lythraceae). 

Distribution. — Endemic (Molokai, Maui, 
Hawaii). 

Material examined. — Lectotype <5, Ha- 
waii, Kilauea. Maui: \6 , Wailua (BPBM). 
Hawaii: Id, paralectotype, same data as lec- 
totype, M, 1 abdomen missing, Kilauea, on 
Lyf/inm (BPBM); 4c?, Kilauea (BMNH); 4(5, 
Kealakekua(BMNH). 

Remarks. — This species can be distin- 
guished by its small size (c?, 2.6 mm, 9 , 2.8 
mm), stramineous or brown markings on the 
head (Figs. 5, 7), brown markings on the first 
thoracic sternite and dorsum of the abdomen, 
the head being distinctly wider than the 
pronotum (Figs. 5, 7), 3.3 midtibial setal for- 
mula, robust aedeagal shaft (Fig. 24), and the 
moderately long preapical lobe on the style 
(Fig. 6). 

The medial length of the vertex varies from 



slightly longer than to 1.5 times length next to 
eyes (Figs. 5, 7). 

Balclutha nigriventris , n. sp. 
Figs. 9-11, 19 

Length.— (5, 3.0-3.3 mm; 9, 3.2-3.5 mm. 

Yellow to greenish yellow, variably tinged 
with orange to brown on head, venter, and 
legs. Dorsum of abdomen dark brown. 

External characters as in fuscifrons . 

Male. — Genitalia as in fuscifrons but with 
aedeagal shaft curved slightly more dorsally 
(Fig. 19). 

Female. — Genitalia with posterior margin 
of pregenital sternite more or less straight. 
Valvulae as in incisa (Figs. 33, 38). 

Host. — Deschampsia (Gramineae). 

Distribution. — Endemic (Maui, Hawaii). 

Holotype. — S, Hawaii: Hualalai, 6,000- 
7,000', 21. iv. 1944, N. L. H. Krauss (BPBM). 
Paratypes: Maui: 36, 39, Halemauu Trail, 
1.V.1945, 8,000' (1(5, on Deschampsia), 16, 
Holua, Haleakala Crater, 6,500', vi.l953. 
Hawaii: 16, 19, same data as holotype; 
19, Hualalai, Hinatapoula, 5,900', vii.1964 
(BPBM, BMNH). 

Remarks. — This species is similar to fus- 
cifrons but is significantly smaller, often lacks 
the brown marking on the clypellus, and has 
the shaft of the aedeagus slightly different in 
shape (compare Figs. 18, 19). 

Balclutha grandis Namba 

Fig. 21 

Balclutha grandis Namba, 1956: 105-106, Figs. 4a-b. 
Holotype cJ, Kauai (BPBM) [examined]. 

Host. — Unknown. 

Distribution. — Endemic (Kauai). 

Material examined. — Holotype 6 , Kauai: 
Alakai Swamp, viii.1953, 4,000' (BPBM). 

Remarks. — This species can be distin- 
guished by its large size (6 , 4.5 mm, 9, 5.3 
mm), the head being distinctly wider than the 
pronotum, 4.4 midtibial setal formula, broad 
stem of the connective, and the shape of the 
aedeagus (Fig. 21). Namba (1956) recorded 
1 c5 , 1 9 from the type locality. 

Balclutha lobata Namba 

Figs. 8, 16, 17, 20 

Balclutha lobata Namba, 1956; 106, Figs. 5a-b. Holotype 
6, Kauai (BPBM) [examined]. 

Host. — Cyathodes (Gramineae), Styphelia 
(Epacridaceae). 



94 



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No. 12 












Figs 18-27. Endemic Hawaiian Balclutha species, aedeagus, left lateral view, 18, B.fuscifrons; 19, B. ni^riventris; 
20, B. lobata; 21, B. grandis; 22, B. timbcrlakei; 23, B. phoxocephala; 24, B. volcanicola; 25-26, B. plutoms; 27, b. 



usitata. 



Distribution. — Endemic (Kauai, Oahu, 
Molokai, Maui, Hawaii). 

Material examined. — Holotype S, Kauai, 
Kokee, vii.1952, 3,600' (BPBM). Kauai: 8(5, 



49, Pihea, Kaunuohua Ridge, 4,260'; 6c?, 
19, Kokee, on Styphelia. Molokai: 16, Ko- 
moku; 9(5, 69, above Waikolu V. 1,400m; 
16, Kalaupapa Lookout; 26, Kewala Gulch, 



1988 



Knight, Webb: Hawaiian Endemics in Balclutha 



95 



3,500'. Maui: 19, Kula Pipe Line, 4, 22' 
(BMNH, BPBM). 

Remarks — This species can be distin- 
guished by its moderately large size (6, 
3.0-3.3 mm, 9, 3.3-3.4 mm), its dark ap- 
pearance resulting from the dark brown dor- 
sum of the abdomen and the greenish tinge of 
the forewings, the 3.3 midtibial setal formula, 
the disc of the female pregenital stemite 
marked with brown (Fig. 8j, and the large 
preapical lobe of the style (Fig. 16j. Some 
specimens (Molokai) have the lobe of the style 
narrower (Fig. 17). Contrar\- to \amba(1956). 
the head is approximately equal in width to 
the pronotum. 

Balclutha plutonis (Kirkaldy) 
Figs. 2.5, 26 

Nesosteles plutonis Kirkaldy, 1910: 574; Osbom 19.3.5: 
57-58, Fig. 24d. Lectotype 6 , Hawaii, here des- 
ignated [e.xamined]. 

Balclutha plutonis ( Kirkaldy i: Zimmerman 1948: 87, Fig. 
27, Namba 1956: 105. Fig. -3. 

Host. — Coprosma (Rubiaceael Vaccinium 
reticulatus (Ericaceae), Sophora iLegumi- 
nosae), Argyroxiphium virescens, Raillardia 
(Compositae), Deschampsia , Eragrostisi?) 
(Gramineae), Vincentia (Cx-peraceae). 

Distribution. — Endemic (Maui, Hawaii). 

Material EX.\MIN ED — Lectotype 6, Ha- 
waii, Kilauea, vii.1906. R. C. L. Perkins 
(BPBM). Numerous specimens from Maui 
(Haleakala, Puu Luau, Puu Xoauiau) and 
Hawaii (Pohakuloa, Kilauea, Mauna Loa, 
Hualalai)(BPBM, BMNH). 

Re.MARKS. — This species can be distin- 
guished by its moderately large size \ i . 3.5- 
3.6 mm, 9, 3.6-4.0 mm), its uniform yellow 
or greenish yellow color, usuall\- without 
browTi markings on dorsum of abdomen, and 
the elongate, evenly cur\ed shaft of the 
aedeagus. It is similar to peregrina but is 
larger, has the shaft of the aedeagus usually 
slightly longer, the connective longer than 
rather than equal in length to the styles, and 
the sides of the clypellus conca% e rather than 
straight. The lectot>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 



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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 



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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 



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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.] 



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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 



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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 



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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 



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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, 



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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 



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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 



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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 



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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. 



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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) 



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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. 



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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) 



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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 



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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 



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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 



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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 



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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. 



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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 



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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 



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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 



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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 



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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 



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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 



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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 



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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 



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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 



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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 



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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. 



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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. 

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BLACKrrH, R E., AND R. a Reyment 1971. Multivariate 
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BOULARD, M. 1982. Les cigales du Portugal, contribution 
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BoYCE, A J. 1964. The value of some methods of numeri- 



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Cheetam, A H 1968. Morphology and systematics of the 
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Claridge. M F . M R Wilson, and] S Singhrao 1979. 
The songs and calling sites of two European ci- 
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Gibson, A R , A J Baker, and A Moeed 1984. Morpho- 
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sis. Systematic Zoology 33(4): 408-421. 

Gomez-Menor, O J 1957. MonografiadeCicadidos (Ho- 
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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. 
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JOLICOEUR, P , AND J E MosiMANN 1960. Size and shape 
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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 
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OXNARD, C E 1978. One biologist's view of morphomet- 
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QuARTAU, J A 1983. A comparison of different numerical 
methods applied to the taxonomy of Batracomor- 
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lassinae): ordination techniques. Journal of the 
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9-35. 

1987. Ordination vs. cluster analysis in the numer- 
ical taxonomy of some leafhoppers (Homoptera, 
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pers of Economic Importance, CLE., London. 

QuARTAU, J A , AND R G Davies 1983. A comparison of 
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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 



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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 + (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 



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Great Basin Naturalist Memoirs 



No. 12 



2 + 2 + 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 + + 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 + 
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). 



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/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. 



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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. 



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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. 



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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 



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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 



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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 



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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 



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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. 



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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. 



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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 



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195 




tho'Sfe^l^trf '^pS.^r -^'^■' '': -r' ^™^''^^ ''- --' ^-- ^2' head and 

79, .a,e forewing'. .. ^.^^ (tuchT^^^^^^^^^^ -^-^-> lateral and caudal views; 



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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. 



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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 



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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) 



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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. 



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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. 



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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 



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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. 



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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. 



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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. 



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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. 



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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. 



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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. 



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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 



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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. 



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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 



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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. 



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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 



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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. 



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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, 



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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 



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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- 



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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. 



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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. 



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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. 



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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 



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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 



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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 



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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 



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Fig. 2. Habitus ofFlexamia species: A, alhida; B, shssonae; C, grammica; D, ritana; E, pijrops: F, areolata. 



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Whitcomb, Hicks: Flexamia Revision 



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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 



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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 



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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 



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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. 



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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. 



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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; 



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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 



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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). 



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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 



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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 



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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 <S: Kimble Co., Texas, 
Junction, 14 September 1987, R. F. Whit- 
comb and R. C. Chambers (1,300 ft, IPL 
003599, Bouteloua uniflora). Deposited 
USNM. Paratypes: 16 d, 19 9, same locality; 
Sutton Co., Texas, Roosevelt, 14 September 
1987, 14 d, 31 9 (1,400 ft, IPL 003593, 
003595, 003598). Deposited at KU, OSU, 
CNC, USNM, KSU, and BARC. 

Remarks. — Flexamia collorum is distinct 
from other members of the pectinata species 
group, from which it is readily separated by 
the short, avicephaliform style apices and by 
the extensively developed ridge on the inside 
of the male plates. It has been collected at 
several localities (Fig. 19) in the Texas hills 
(Edwards Plateau), where it appears to spe- 
cialize on Nealley grama, Bouteloua uniflora. 
Nealley grama differs from side-oats grama in 
having single spikelets at the base of the 
rachis; it also has a narrow geographic distri- 
bution in Texas and Mexico. Although this 
species appears to intergrade with curtipen- 
dula in the Mexican state of Coahuila (Johnson 
1943), it is readily distinguished from it in 
central Texas. 

The name collorum is an adjective meaning 
"of the hills," in recognition of the Texas hills 
where it was discovered. 

8. Flexamia jacala, n. sp. 

Length of 6 3.5 mm; head width of 6 1.08 
mm. Crown produced; length of crown 1.42 x 
interocular width, 0.75 x head width. 

Color gray with irregular, dark markings on 
dorsum and forewings; face without black in- 
terocular band; crown with oblique, rear and 
transverse, midlength markings. Venter and 
legs dark with irregular, fuscous markings. 



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Fig. 18. Flexamia collorum, 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, apex of right style, lateral aspect; H, female sternum VII; I, bases of first valvulae 
of female, dorsal aspect. 



Male. — Pygofer (Fig. 20D) as in ban- 
darita. Plates (Fig. 20G) extend to approxi- 
mately 2/3 length of pygofer, fused basally, 
contiguous medially for 2/3 of their length, 
converging to bluntly rounded apices; male 
genitalia similar to those of bandarita, but 
styles (Fig. 20E) straight in ventral aspect in- 
stead of sinuate, diverging apically; apices 
abruptly turning ventrad, appearing avi- 
cephaliform in lateral aspect (Fig. 20F), 
preapical lobe of style produced laterally and 
caudally. Connective (Fig. 20A) in lateral 
view with dorsal keels narrow; apodemal pro- 
cesses as in bandarita; aedeagal tip (Fig. 20C) 
slightly flared in ventral view. Gonopore sub- 
apical on caudoventral surface. 

Types. — Holotype S: Mexico, Jacala, Hi- 
dalgo, 13 August 1936, E. D. Ball. Deposited 
inUSNM. 

Remarks. — This species is known from a 
single male. Nothing is known of its biology. It 
is most closely related to bandarita, from 
which it can be distinguished by its styles. 



which are straight in ventral aspect instead of 
sinuate; also, in lateral aspect the apices are 
avicephaliform instead of chelate. The name 
jacala is a noun in apposition denoting the 
type locality. 

9. Flexamia niescalero, n. sp. 



Length of 6 3.6 (3.5-3.9) mm, 9 3.8 
(3.6-4.1) mm; head width of 6 1.07 mm, 9 
1.12 mm. Crown moderately produced; me- 
dian length of crown 1.42 x head width and 
0.66 X interocular width (d n = 6; 9 n = 16). 

Color pale with brown markings. Crown 
ivory with pair of oblique markings on rear 
and pair of transverse lines at midlength; 
forewings pale gray with irregular, dark mark- 
ings. Face variable, with or (more commonly) 
without black interocular band; specimens 
without band variably marked on face with 
brown or black. 

Male. — Pygofer (Fig. 21 E) with posterior 
lobe angulate on ventral and caudal margins, 



1988 



Whitcomb, Hicks: Flexamia Revision 



243 




Fig. 19. Geographic distribution of mescfl/ero (< 
paucijlora and Bouteloua uniflora, respectively. 



and collorum (■), with distribution of their hosts, Muhlenbergia 



slightly convex dorsally; plates (Fig. 21D) ex- 
tending to approximately 4/5 length of 
pygofer, fused basally, diverging gradually 
along entire length; lateral margins tapering 
mesad to acute apices, inner surface with lon- 
gitudinal groove and thickened mesal ridge 
along 3/4 of length, ending in small, em- 
browned tooth. Connective (Fig. 21A) in lat- 
eral view with dorsal keels narrow, approxi- 
mately 1/4 height of dorsal apodeme; 
apodemal processes as in bandarita; styles 
(Fig. 21 F) long in ventral aspect, slightly di- 
vergent distally, in lateral aspect with preapi- 
cal lobe moderately produced and acute. Api- 
cal 1/2 of style flattened and minutely serrate 
basally, narrowing to ventromesally directed, 
calipterate apices (Fig. 21G), much as in ban- 
darita. Aedeagus symmetrical, expanded api- 
cally in ventral aspect, appearing bluntly sag- 
gitate (Figs. 21C, 22), in lateral aspect slightly 
indented preapically; gonopore subapical on 
caudoventral surface. 

Female.— Sternum VII (Fig. 21 H) with 
posterior margin notched deeply, middle 
teeth very short and deeply inset, median 
incision deep, in some specimens divergent at 



margin. Ovipositor with bases of first valvulae 
(Fig. 211) with sinuate, recurved processes 
that converge medially. 

Types. — Holotype 6 : Eddy Co. , New Mex- 
ico, 23 August 1985. A. L. Hicks and R. F. 
Whitcomb, 4,600 ft, IPL 001977, Muhlenber- 
gia paticiflora. Deposited in USNM. 
Paratypes: 9 d, 16 9, same locality as holo- 
type. Deposited at KU, KSU, OSU, CNC, 
BARC, and USNM. 

Remarks. — Flexamia mescalero is the only 
member of the pectinata group that has not 
been collected on Bouteloua. The unique 
aedeagal apex (Figs. 21C, 22) and female ster- 
num VII (Fig. 21 H) are diagnostic for this 
species, give it unique status in the group, and 
could justify treatment as a distinct group (see 
cladogram of Fig. 58). The host of this species 
is New Mexican muhly, Muhlenbergia pauci- 
flora. Attempts to find other localities for this 
species have been unsuccessful. 

The name mescalero is a noun in apposi- 
tion honoring the Native American Mesca- 
lero Apache nation, which inhabited the 
mountains of southeastern New Mexico in 
the pre-Columbian era and which is today 



244 



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No. 12 




Fig. 20. Flexamia jacala, n. sp.: A, aedeagus and connective, lateral aspect; B, aedeagus and connective, dorsal 
aspect; C, apex of aedeagus, caudoventral aspect; D, male pygofer, lateral aspect; E, right style, dorsal aspect; F, apex 
of right style, lateral aspect; G, male plates and pygofers, ventral aspect. 



establishing high standards for judicious man- 
agement of tribal lands. 

II. The abbreviata Group 

The abbreviata group consists of a single, 
widespread species that specializes on 
Bouteloua spp. The morphology of the aedea- 
gal apex constitutes an autapomorphy that de- 
fines the monobasic group. 

10. Flexamia abbreviata (Osborn & Ball) 

Deltocephalus abbreviatus Osborn & Ball 1897: 206. 
Deltocephalus (Flexamia) abbreviatus, DeLong 1926: 33. 
Flexamius abbreviatus , DeLong and Sleesman 1929: 83. 

Important CHARACTERS. — Length of c? 3.1 
(2.8-3.4) mm, of 9 3.2 (2.9-3.6) mm. Head 
moderately produced (mean crown length 
1.26 X interocular width, 0.61 x head width) 
(d n = 20; 9 n = 20). Face without distinct 
interocular line (but very rarely with a brown 
band contrasting with a pale face). Face often 
with parallel, sinuate, brown lines. [6] pygo- 



fer (Fig. 62H) strongly constricted, posterior 
lobe strongly produced on upper portion of 
posterior margin. Plates (Fig. 8E) shield- 
shaped, about 0.65 x pygofer length. Aedea- 
gus (Fig. 7H) with elongate, anteapical 
gonopore on caudoventral surface, apex with 
pair of short, truncate processes curved 
cephalad, apodemal processes each with 
broad, membranous, apical expansion. [9] 
Sternum VII (Fig. lOE) with a medial, trun- 
cate projection, which has a median incision 
that is almost always decorated on either side 
by a pair of infuscated spots. 

Geographic distribution. — This species 
occurs (Fig. 23) from the Canadian prairie 
provinces to Montana, Utah, southwestern 
Nevada, Wisconsin, Illinois, Oklahoma, 
Texas, and Durango, Mexico (1 <5, 27 mi N 
LaZarca, 27 October 1981, M. W. Nielson). 

Biology. — Because of the dominance of its 
Bouteloua hosts, the densities it achieves on 
these hosts, and the accessibilitv of its hosts to 



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Fig. 21. Flexainia mescalero, 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, apex of right style, caudoventral aspect; H, female sternum VII; I, bases of first 
valvulae of female, dorsal aspect. 



collectors, this is the most frequently col- 
lected Flexamia species (Appendix I). In the 
northern and western parts of its range it is a 
specialist on blue grama {Bouteloua gracilis), 
and the range of this grass limits its distribu- 
tion (Fig. 23). In Chihuahuan grasslands and 
montane islands of southern Arizona and New 
Mexico, it is an inhabitant of mixed Bouteloua 
grasslands, where it may occur on blue, hairy 
(B. hirsute), or, more rarely, black (B. eri- 
opoda) grama. In the eastern prairie where 
the grasslands form a mosaic with patches of 
eastern deciduous forest, the only known host 
is hairy grama; the eastern boundary of the 
range of abbreviata is determined by this 
host. In the Edwards Plateau of Texas, abbre- 
viata occurs on B. hirsuta, B. pectinata, Muh- 
lenbergia reverchonii (seep muhly), and per- 
haps other chloridoid grasses. 

Oligophagy COEFFICIENTS. — Gramineae 
1.000; Chloridoideae 0.936; Bouteloua 0.836. 
Prairie: Bouteloua 0.780; B. gracilis 0.745 



(n = 428); mixed Bouteloua grasslands: B. 
gracilis 0.317, B. hirsuta 0.568 (n = 347). 

Remarks and diagnosis. — Males and fe- 
males of this species can be recognized by the 
combination of their small size, brown face 
without interocular band, and unique mor- 
phologies of the sternum VII or male plates 
and pygofer (Fig. 8E). The hind margin of the 
female sternum VII (Fig. lOE) contains four 
teeth, a condition we believe to be ho- 
mologous with similar structures in the pecti- 
nata group. Flexamia abbreviata occurs with 
curvata (host: Buchloe dactyloides) through- 
out much of the range oi Buchloe. Dark male 
specimens of curvata are occasionally con- 
fused with abbreviata but can be tentatively 
recognized without dissection by characters of 
the plates and pygofers. 

III. The zacate Group 

The zacate group consists of two species 
that specialize on bush muhly (Muhlenbergia 



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B 




Fig. 22. Scanning electron micrograph of aedeagus oi Flexarnia mescalero: A, lateral aspect (312X); B, ventral aspect 
of aedeagal apex (1680X). 



porteri). These sister species are distin- 
guished on the basis of their posteriorly pro- 
duced male pygofers, curved aedeagus lack- 
ing apical processes, and elongate, but 
rounded male plates. No other Flexarnia spe- 
cies has this combination of characters. 

Description of the zacate Group 

Medium-sized. Length of S 3.2-4.1 mm, 
of $ 3.6-4.1 mm. Dorsum brown. Crown 
moderately produced. [6] Plates elongate, 
but only about 2/3 length of pygofer, fused 
basally, narrowing continuously to apices. 
Pygofer (Figs. 25E, 621) with posterior lobe 
strongly produced posteriorly. Aedeagus and 
connective (Fig. 6C) distinctly articulated. 
Aedeagus curved in lateral aspect, without 
apical processes (Fig. 25A). [ 9 ] Sternum VII 



(Figs. 101, L) with deep, median incision. 

Key to Species oizacate Group 
1. Males 2 

— Females 3 

2(1). Aedeagus (Fig. 71) narrow in ventral and lateral 
aspects; gonopore (Fig. 24) long and narrow, 

extending along apical third of shaft 

zacate, n. sp. 

— Aedeagus (Figs. 6C, 7J) wide, scimitar-shaped 
in lateral aspect, diamond-shaped in ventral as- 
pect. Gonopore not elongate, near midpoint of 
shaft camjonensis Young & Beirne 

3(1). Hind margin of sternum Vll (Fig. lOL) with 
outer pair of teeth essentially absent, fused with 

medial teeth to form median projection 

zacate, n. sp. 

— Hind margin of sternum VH (Fig. 101) with 

outer teeth distinct, rounded 

camjonensis Young & Beirne 



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Fig. 23. Geographic distribution oiabbreviata. Western limit is defined by the range oi Boiitelotia gracilis. Eastern 
limit is defined by the range oi Botttelotia hirsuta. Host relationships in Mexico are unknown. 



1 1 . Flexamia canyonensis Young & Beirne 
Flexamia canyonensis Young & Beirne 1958: 18. 

Important CHARACTERS. — Length of 6 3.7 
(3.5-4.1) mm, of 9 3.9 (3.8-4.1) mm. Head 
moderately produced (1.29 x interocular 
width; 0.68 X head width) (d n = 4; 9 n = 5). 
Face with clypeus and dorsal portion of genae 
black. Clypeus, lora, and lower part of genae 
pale yellow. [6] Pygofer (Fig. 621) as in ab- 
breviata. Plates (Fig. 9C) gradually narrowing 
apically to rounded apices. Aedeagus sym- 
metrical, scimitar-shaped in lateral aspect, di- 
amond-shaped in ventral aspect. Gonopore 
on ventral surface of aedeagus, near midpoint 
of shaft. [ 9 ] Sternum VII with deep, median 
incision on hind margin. Ovipositor with 



bases of each first valvula (Fig. 63H) strongly 
recurved, processes sinuate in dorsal aspect. 

Geographic distribution. — Flexamia can- 
yonensis has been collected at Sabino Can- 
yon, Arizona (E. D. Ball), and in our own 
studies at Kingman, Portal, and Santa Rita 
Experimental Range, Arizona (Fig. 26). 

Biology. — Our collections of canyonensis 
are from bush muhly, Muhlenbergia porteri. 
This ^j ass is common in the type locality, and 
we assume that it is the sole host. Flexamia 
canyonensis appears to be restricted to the 
highlands of the Sonoran Desert. This species 
and its Chihuahuan sister, zacate, are among 
the few Flexamia species to colonize semi- 
arid habitats. Perhaps canyonensis is pro- 
tected from desiccation by the microclimate 



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Fig. 24. Scanning electron micrograph of aedeagal apex oiFlexamia zacate, ventral aspect (800X). 



produced by its host, which grows within pro- 
tective shrubs. 

Oligophagy coefficients. — Muhlenber- 
gia porteri 1.00 (n = 9, 3 series). 

Remarks and diagnosis. — Females can be 
easily recognized by their distinctive sternum 
VII (Fig. 101), which bears a deep, median 
incision. Males can be tentatively recognized 
by their characteristic face pattern and pro- 
duced, flattened pygofer. The aedeagus of 
canyonensis is unique. 

12. Flexamia zacate, n. sp. 

Length of 6 3.5 (3.2-3.9) mm, 9 3.8 
(3.6-4.0) mm; head width of 6 1.04 mm, 9 
1.08 mm. Crown produced; median length of 
crown 1.48 X head width and 0.71 .x interocu- 



lar width (d n = 37; 9 n = 12). 

Dorsum ivory. Crown with at least trace of 
oblique, basal and midlength, transverse 
lines. Face without black interocular line, 
black in interocular area with at most pale 
arcs, bottom of face brown, with irregular, 
pale markings. Venter stramineous with ir- 
regular markings. 

Male. — Pygofer (Fig. 25E) with weakly de- 
veloped constriction before posterior lobe, 
ventral lobe small, caudoventral margin pro- 
duced posteriorly, dorsal margin unevenly 
convex. Plates (Fig. 25D) extending to ap- 
proximately 2/3 length of pygofer, fused 
basally, contiguous medially for 1/2 length, 
acute apically; connective in lateral view (Fig. 
25A) with dorsal keels narrow, approximately 



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249 








bLK 



Fig. 25. Flexamia zacate, 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, female sternum VII; H, bases of first valvulae of female, dorsal aspect. 



1/4 height of dorsal apodeme, with distinc- 
tive, minute tooth on ventral surface, anterior 
to distinct joint with the aedeagus; apodemal 
appendages campanulate in caudal aspect, 
with very slender, anteapical processes di- 
rected mesad; styles (Fig. 25F) with apex digi- 
tate, curving sharply ventrad, each bearing 
small preapical tooth on mesal surface; preapi- 
cal lobe not produced. Aedeagus symmetri- 
cal, shaft slender and cylindrical without pro- 
cesses, curving gradually caudodorsad in 
lateral aspect, appearing straight in ventral 
aspect. Apex slightly flared in caudoventral 
aspect, flattened in lateral aspect, with apical 
margin concave. Gonopore (Figs. 24, 25C) 
long and narrow, extending along apical third 
of caudoventral surface of shaft. 

Female.— Sternum VII (Fig. 25G) with 
posterior margin produced medially, with 
deep, median incision; ovipositor with basal 



processes of first valvulae (Fig. 25H) with 
broad portions close to base. 

Types. — Holotype 6: Brewster County, 
Texas, Big Bend National Park (Panther Junc- 
tion), 26 August 1985 (3,600 ft, R. F. Whit- 
comb and A. L. Hicks; IPL 002093). De- 
posited USNM. Paratypes: 33 6 , 10 9 , same 
locality and date, IPL 002057, 25 August 
1985; 13 (?, 4 9, same locahty and date, de- 
posited USNM, BARC, CNC, KU, KSU, and 
OSU. Thirty-five immature specimens (IPL 
002057) have also been deposited with the 
paratypes. 

Remarks. — This species is a sister to can- 
yonensis but can be distinguished from it by 
the shape of the aedeagus, which, though 
curved, is much narrower in lateral aspect and 
is not diamond-shaped in ventral aspect. The 
length of the gonopore of zacate also separates 
it from canyonensis. Both canyonensis and 



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Fig. 26. Geographic distribution of canyonensis (•) and zacate (■) and their host, Muhlenbergia porteri. 



zacate are apparently specialists on bush 
muhly, Muhlenbergia porteri. This grass oc- 
curs chiefly in association with thorny shrubs 
such SIS Acacia, Prosopis, Larrea, or Opuntia, 
which defend it against large herbivores. The 
entire volume of the host shrub may be filled 
with the tangled culms of bush muhly; even- 
tually the host shrub dies, presumably from 
competition. Because of its unique habitat, 
zacate is difficult to collect without a vacuum- 
collecting device. The name zacate is a noun 
in apposition denoting various grasses of the 
Muhlenbergia or Sporobolus types. Bush 
muhly is known in Mexico as "zacate arana ' 
(spider grass). 

IV. The gramniica Group 

The grammica group consists of a single, 
distinctive species. The mor^Dhology of the 
aedeagal apex and the female sternum VII 
provides autapomorphies that define the 
group. 

13. Flexamia grammica (Ball) 

Deltocephalus gravimicus Ball 1900; 204. 
Deltocephalus (Flexamia) grammicus), DeLong 1926: 37. 
Flexamiiis graminicus , DeLong and Caldwell 19.37: 27. 

Important CHARACTERS. — Length of 6 4.5 



(4.1-5.0) mm, of 9 4.6 (4.3-5.0) mm. Head 
not strongly produced (mean crown length 
1.08 X interocular width, 0.54 x head width) 
(d n = 20; 9 n = 20). Dorsum (Fig. 2C) with 
three pairs of longitudinal, dark stripes, me- 
dian pair extending from hind portion of the 
crown posteriorly through the scutellum and 
wings. Face stramineous with single, black 
interocular line. [6] Pygofer (Fig. 620) with 
narrow, posterior lobe, dorsal more promi- 
nent than ventral constriction in lateral as- 
pect. Plates (Fig. 8F) broad, contiguous, par- 
allel-sided for much of their length, broadly 
rounded apically. Aedeagus asymmetrical 
with anteapical gonopore on anterodorsal sur- 
face, with pair of recurved processes (Fig. 
7E), one edentate on the side of shaft near 
gonopore, one at the shaft apex. [ 9 ] Sternum 
VII (Fig. lOF) broadly and shallowly concave 
on posterior margin. 

Geographic distribution. — Flexamia gram- 
mica occurs in southern Alberta and Sas- 
katchewan south through the Great Plains to 
Texas (Fig. 27). 

Biology. — In the northern part of its 
range, grammica is a specialist on prairie 
.sandreed, Calamovilfa longifolia. In the 
southern part of its range, it specializes on 
C. gigantea. 



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251 




Fig. 27. Geographic distribution oi Flexamia grammica, and two hosts, Calamovilfa longifolia (northern prairie) and 
C. gigantea (southern prairie and Southwest). 



Oligophagy coefficients. — Gramineae 
1.000; Chloridoideae 1.000; Calamovilfa spp. 
0.872 (n- 47). 

Remarks and diagnosis. — The habitus of 
grammica is distinctive, as is its size. 

V. The curvata Group 

The curvata group consists of two closely 
related sister species that specialize on buf- 
falograss, Buchloe dactyloides. One (curvata) 
is adapted to temperate regions, whereas the 
other (surcula) is adapted to the subtropical 
climate of south Texas and Mexico. In the 
following description synapomorphies are 
designated [s]. 

Description of the curvata Group 

Small, deltocephaline leafhoppers. Length 
of 6 2.4-3.5 mm, of 9 2.6-3.9 mm. Usually 
stramineous in color, but occasionally much 
darker. [S] Pygofer (Figs. 62K,L) somewhat 
produced caudodorsally. Plates (Figs. 9D,E) 
short, shield-shaped, about half length of 
pygofer. Aedeagus (Figs. 7F,G) [s] with two 
pairs of apical processes; style apex [s] foot- 
shaped in broadest aspect. [ 9 ] Female ster- 



num VII (Figs. 10J,K) with median incision or 
notches, with infuscation surrounding or bor- 
dering notch or incision. 

Key to Species oi curvata Group 
1. Males 2 

— Females 3 

2(1). Gonopore small, subcircular, opening near 
aedeagal apex; apical pair of processes short 
(Fig. 7G) surcula DeLong & Sleesman 

— Gonopore elongate; apical pair of processes 
much longer (Fig. 7F) curvata DeLong 

3(1). Female sternum VII (Fig. lOK) with completely 

infuscated, median incision .... curvata DeLong 

— Female sternum Vll (Fig. lOJ) notched, but 
without median incision; infuscated on each side 

of notches surcula DeLong & Sleesman 

14. Elexamia surcula DeLong & Sleesman 

Flexamius [sic] surculus DeLong & Sleesman 1929: 99. 

Important char.\cters. — Length of S 3.0 
(2.6-3.4) mm, of 9 3.4 (2.9-3.7) mm. Head 
moderately produced (mean crown length 
1.35 X interocular width; 0.68 x head width) 
((J n = 20; 9 n = 20). Face stramineous, 
except for broad, brown, basal, interocular 



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Fig. 28. Geographic distribution oi Flexamia curvata (•) and surcula (A) and their host, Biichloe dactyloides. 



line. [6] Pygofer (Fig. 62L) with posterior 
lobe strongly produced; style with apical por- 
tion foot-shaped, but longer than in curvata. 
Plates (Fig. 9E) as in curvata. Aedeagus (Fig. 
7G) with pair of recurved, apical processes, 
shorter than in curvata, and pair of anteapical 
processes. [9] Sternum VII (Fig. lOJ) infus- 
cated on either side of shallow notch. Oviposi- 
tor with each first valvula (Fig. 631) with basal 
portion curved dorsad through more than 90 
degrees, recurved portion oblique and cres- 
centiform in dorsal aspect. 

Geographic distribution. — Subtropical 
Texas, and probably coastal prairies of north- 
eastern Mexico (Fig. 28). 

Biology. — Flexatnia surcula is restricted to 
subtropical grasslands. We have collected it 
from Buchloe in the Rio Grande Valley in 
Cameron and Hidalgo counties of south 
Texas. Collection records from December, 
January, and February indicate that this spe- 
cies does not diapause and probably breeds 
more or less continuously throughout the sea- 
son as host condition permits. 

Remarks and diagnosis. — This species is 
closely related to curvata. We have referred a 
specimen labeled "paratype" from San Anto- 
nio, Texas, in the OSU collection to curvata. 



The sternum VII of surcula females lacks the 
deep, median incision oi curvata. 

15. Flexatnia curvata DeLong 

Deltocephalus (Flexamia) ciirvattis DeLong 1926: 34. 
Flexamhis ciirvatus, DeLong and Caldwell 1937: 27. 

Important characters. — Length of c? 3.1 
(2.4-3.5) mm, of 9 3.4 (2.6-3.9) mm. Head 
moderately produced (mean crown length 
1 . 33 x interocular width; 0. 63 x head width) ( 6 
n = 21; 9 n = 23). Face usually pale brown, 
often with traces of indistinct, brown interoc- 
ular band. [6] Pygofer (Fig. 62K) with poste- 
rior lobe simple, somewhat produced cau- 
dodorsally. Plates (Fig. 9D) short, shield- 
shaped. Style foot-shaped in broadest aspect. 
Aedeagus (Fig. 7F) subsymmetrical, recurved 
anterior processes extending basad almost to 
midlength of shaft, pair of shorter apical pro- 
cesses not in bilaterally symmetrical plane; 
gonopore an elongate channel on caudoven- 
tral surface. Recurved anteapical processes 
extending basad almost to middle of shaft; 
pair of shorter apical processes extending 
basad. [9] Sternum VII (Fig. lOK) with hind 
margin concave on each side of median inci- 
sion. Ovipositor with base of each first valvula 



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253 



(Fig. 63F) bearing posteriorly directed, slen- 
der process. 

Geographic distribution. — Flexamia cur- 
vata occurs throughout much of the temper- 
ate range of its host, Buchloc dactyloides (Fig. 
28). 

Biology. — This species often occurs in 
large populations on Biichloe, where it is a 
member of a guild that also includes Athy- 
sanella, Pohjamia, Laevicephalus, and Gillet- 
tiella species (Whitcomb et al. 1987). In the 
subtropical part of the range of bufFalograss 
(in south Texas and, presumably, Mexico), 
curvata is replaced by surcula. 

Oligophagy coefficients. — Gramineae 
1.000; Chloridoideae 0.986; Biichloe dacty- 
loides 0.925 {n = 560). 

Remarks and diagnosis. — This species is 
readily recognized by the combination of its 
small size and external genitalic characteris- 
tics. Most specimens are light stramineous, 
with conspicuous, black spots in the middle 
and apical cells of the forewing (Fig. 4B); these 
specimens are unmistakable. Very large male 
specimens of curvata might be confused with 
inflata, whose plates and pygofer are superfi- 
cially similar. However, inflata, which like 
curvata is usually light stramineous in color, 
never has a dark spot in the center of the 
forewing. Dark specimens of curvata are oc- 
casionally referred erroneously to abbreviata. 

VI. The picta Group 

This group consists of two species, each 
with a distinctive habitus, which can be unam- 
biguously designated as sister species on the 
basis of the morphology of the male aedeagus 
and pygofer, and the indistinct articulation 
between the aedeagus and connective. In ad- 
dition, both species feed on Aristida. One 
(pyrops) has, perhaps, the most distinctive 
habitus in the genus (Fig. 2E), but picta can 
also be recognized without dissection. In the 
following description synapomorphies defin- 
ing the group are designated [s]. 

Description of the picta Group 

Medium-sized or large, deltocephaline 
leafhoppers. Length of 6 2.9-4.0 mm, of 9 
2.9-4.9 mm. Grown variably produced (in 
pyrops, reaching more than 2 x interocular 
width). [ 6 ] Pygofer [s] notched on dorsal mar- 
gin, posterior lobe angulate, sharply pro- 
duced. Plates small in relation to large pygo- 



fers; [s] joint between aedeagus and connec- 
tive indistinct. Aedeagus symmetrical, shaft 
elongate, [s] apex capitate, pair of slender, 
recurved, anteapical processes with entire 
edges. Gonopore subapical on caudoventral 
surface. 

Key to Species of picta Group 
1. Head produced, with crown more than twice 

interocular width (Fig. 2E) pyrops (Crumb) 

— Head only moderately produced . . picta (Osborn) 

16. Flexamia picta (Osborn) 

Deltocephalus pictus O.sborn 1907: 165. 
Deltocephalns funahuhi.s Crumh 1915; 189. 
Deltocephalus (Flexamia) pictus, DeLong 1926: .32. 
Flexamius [sic] pictus, DeLong and Sleesman 1929: 83. 

Important CHARACTERS. — Length of 6 3.5 
(2.9-4.0) mm, of 9 3.6 (2.9-4.2) mm. Head 
moderately produced (1.38 x interocular 
width; 0.64 X head width) (c? n = 24; 9 n = 
20). Face pale yellow, with broad interocular 
band. Forewings usually dark gray, contrast- 
ing with lighter crown and pronotum, dark- 
pigmented spot in apex. [ 6 ] Pygofer (Fig. 5A) 
similar to that of pyrops. Plates (Fig. 9A) very 
short, only about 1/3 length of pygofer, nar- 
rowed apically. Aedeagus and connective 
with indistinct joint with the connective, 
gonopore subapical on caudoventral surface. 
Aedeagal apex capitate; expanded portion 
with a few small teeth; pair of slender, re- 
curved, anteapical processes with entire 
edges arising on each side of gonopore. [9] 
Sternum VII with median projection. Ovipos- 
itor with base of each first valvula (Fig. 63J) 
curved through 180 degrees, obliquely bisin- 
uate in dorsal aspect. 

Geographic distribution. — This species 
occurs throughout much of the Southeast, 
from Massachusetts to Kansas, and south 
through the mixed prairie to Texas (Fig. 29). 

Biology. — Flexamia picta appears to be as- 
sociated with Aristida spp. Some of the colo- 
nized species are annuals. This is a unique 
situation in Flexamia hosts; all other known 
hosts are perennials. 

Oligophagy coefficients. — Gramineae 
1.000; Ghloridoideae 0.919; Aristida spp. 
0.870 (n = 285). 

Remarks and diagnosis. — It is important 
for students of the genus to learn the habitus 
of picta, which is very common throughout 
much of its range. This species can be recog- 
nized by the combination of a dark-pigmented 



254 



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No. 12 




Fig. 29. Geographic distribution of three Aristida speciahsts; Flexamia arizonensis (O), pyrops (A), and picta (•). 



spot in the apex of the forewing (Fig. 4C) in 
combination with the face characteristics. The 
face is usually sordid yellow but occasionally is 
light gray or very pale brown. A few dark 
individuals of certain other species (e.g., 
clayi) may have a similar wing spot, but if so, 
the face is never pale. The geographical distri- 
bution of picta (southeastern) is also usually 
indicative. The range of picta overlaps the 
ranges of flexulosa and arizonensis, which 
have superficially similar faces, only in central 
Texas, Oklahoma, and Kansas. Finally, the 
small male plates, which contrast with very 
large py gofers, are diagnostic for picta. Thus, 
dissection is usually unnecessary for recogni- 
tion of this species. 

17. Flexamia pyrops (Crumb) 

Deltocephahis pyrops Crumb 1915: 191. 
Deltocephalus (Acurhinus) pyrops, DeLong 1926: 21. 
Acurhinus pyrops, DeLong 1948: 226. 
Flexamia pyrops, Oman 1949: 167. 

Important CHARACTERS. — Length of 6 3.8 
(3.6-4.0) mm, of $ 4.2 (3.8-4.9) mm. Head 
very strongly produced (crown length 2. 19 x 
interocular width, 1.00 x head width) {6 n == 
20; 9 n = 20). Face stramineous, interocular 
line with three or more short, dentate projec- 



tions from lower margin. [6] Py gofer (Fig. 
5B) with distinct notch on dorsal margin, pos- 
terior lobe produced ventrad in conspicuous, 
acute process; plates (Fig. 9B) very short, no 
more than 1/3 length of py gofer, rounded api- 
cally. Joint between connective and aedeagus 
not well developed. Aedeagus symmetrical, 
gonopore on caudoventral surface, pair of re- 
curved, anteapical processes arising basad of 
gonopore. Apex of shaft (Fig. 7L) capitate, 
similar to that of picta, but with fewer, more 
distinct teeth. [9] Sternum VII (Fig. lOH) 
with hind margin conspicuously concave at 
middle. 

Geographic distribution. — Although this 
is a southeastern species, it occurs as far north 
as Pennsvlvania, Ohio, Illinois, and Kansas 
(Fig. 29).^ 

Biology. — Flexamia pyrops, like F. picta, 
is closely associated with Aristida spp., but 
the two species are seldom found together. 
The type oi pyrops from Tennessee was taken 
on "Aristida longespica. " In Illinois the host 
was A. dichotoma var. curtisii (Whitcomb 
1957). 

Remarks and diagnosis. — The habitus of 
pyrops is unique and diagnostic. 



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VII. The albida Group 

The albida group comprises two species 
that are very distinct in general facies and 
geographic distribution. Common character- 
istics of the species are their patterns of dorsal 
stripes, face patterns, and their bifurcate, api- 
cal, aedeagal processes (an apomorphy shared 
with serrata). The morphology of the male 
pygofers of the species may constitute a syn- 
apomorphy. The biology of albida and slos- 
sonae appears to be more complex than that of 
many other Flexamia species. The dorsal 
stripes, face patterns, and large size of these 
species suggest that they may have arisen 
from an ancestor related to Spartopyge. 

Description of the albida Group 

Medium-sized to large. Length of 6 
3.4-4.2 mm, of $ 3.6-4.8 mm. Dorsum with 
distinctive pattern of stripes extending from 
head across pronotum, scutellum, and 
forewings. Face pale with black interocular 
band. Male pygofer with poorly differentiated 
posterior lobe. Aedeagus with pair of bifur- 
cate, apical appendages, unpaired ventral 
process. 

Key to Species of the albida Group 

1. Forewings rounded, male plates (Fig. 8A) not 
exceeding pygofer, female sternum VII not 
trilobed; prairie albida (Osborn & Ball) 

— Forewings obliquely truncate, male plates ex- 
ceeding pygofer (Fig. 8B); female sternum VII 
trilobed (Fig. lOB); Florida slossonae (Ball) 

18. Flexamia albida (Osborn & Ball) 

Deltocephalus albidus Osborn & Ball 1897: 201. 
Deltocephalus (Flexamia) albidus, DeLong 1926: 36. 
Flexamius albidus, DeLong and Sleesman 1929: 83. 

Important CHARACTERS. — Length of c? 3.7 
(3.4-3.9) mm, of 9 4.3 (3.6-4.8) mm. Crown 
not strongly produced (median crown length 
1.08 X interocular width, 0.57 x head width) 
(6 n = 12; 9 n ^ 21). Face entirely pale. 
Pronotum (Fig. 2A) traversed by six longitudi- 
nal stripes, central pair extending to rear of 
crown, across scutellum, and as spots on base 
of forewing; pair of transverse lines midway on 
crown. Forewings subhyaline with several 
highly pigmented cells, conspicuously pig- 
mented, costal cross-veins, and conspicuous 
pigmentation along the inner edge of the 
forewing. [ 6 ] Pygofer (Fig. 62Q) with poste- 
rior lobe short, with small, rounded, ventrally 



produced lobe. Plates (Fig. 8A) elongate, un- 
notched. Aedeagus symmetrical, gonopore 
subapical on caudoventral surface. Aedeagal 
apex (Fig. 7B) with pair of distinctive, bifur- 
cate processes, each with few teeth on the 
dorsal margin. Apodemal processes greatly 
expanded at apex. [ 9 ] Sternum VII (Fig. lOA) 
with posterior margin shallowly concave on 
each side of produced median portion. 
Ovipositor with base of each first valvula (Fig. 
630) recurved, basal extremity rounded and 
directed caudodorsad. 

Geographic distribution. — ^This species 
occurs in the prairie of Minnesota and North 
Dakota, east to Illinois and south to the Black- 
land tall-grass prairie of Texas (Fig. 30). 

Biology. — Flexamia albida is a resident of 
tall-grass and mixed prairies. Many records 
are from Schizachyrium scoparium. How- 
ever, albida is not found in pure stands of S. 
scoparium, suggesting a life history more 
complex than that of other Flexamia species. 

Oligophagy coefficients. — Gramineae 
1.000; Panicoideae 0.674 (Schizachyrium sco- 
parium 0.674); Chloridoideae 0.196 (Boute- 
loua curtipendula 0. 196) (n = 46). 

Remarks and diagnosis. — The habitus of 
F. albida is unique, and its geographic range 
does not overlap that of the other striped Flex- 
amia species. 

19. Flexamia slossonae (Ball) 

Deltocephalus slossoni [sic] Ball 1905: 119. 
Deltocephalus (Flexamia) slossoni, DeLong 1926: 36. 
Deltocephalus (Secopennis) slossoni, DeLong and Slees- 
man 1929: 85. 
Flexamia slossoni, DeLong and Caldwell 1937: 27. 
Secopennis slossonae, Oman 1949: 168. 

Important characters. — Length of d 3.8 
(3.5-4.2) mm, of 9 4.2 (3.8-4.5) mm. Crown 
produced (median crown length 1.38 x inter- 
ocular width, 0.64 X head width) (c? n = 20; 9 
n = 20). Face ivory with black interocular 
band at base. Dorsum with longitudinal lines 
extending from rear of the head across prono- 
tum and* scutellum to rear of forewings (Fig. 
2B). Male plates (Fig. 8B) exceeding pygofer 
in length. [6] Aedeagus (Fig. 7C) symmetri- 
cal, gonopore an elongate slit extending 
through most of length of unpaired ventral 
process; apical processes bifurcate, without 
teeth. [ 9 ] Sternum VII (Fig. lOB) conspicu- 
ously tri-lobed. 



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Fig. 30. Geographic distribution oiFlcxamia serrata (■), alhida (•), slossonae (D), and ritana (O). 



Geographic distribution. — Flexamia slos- 
sonae is restricted (Fig. 30) to subtropical 
grasslands of south Florida. 

Biology. — DeLong (1926) reported find- 
ing adults and nymphs of slossonae on salt- 
grass (Distichlis spicata). We have found the 
species on the Muhlenhergia prairies of the 
Florida Everglades, in the absence of Distich- 
lis. It seems likely that this species is a hal:)itat 
rather than a host specialist. 

Remarks and diagnosis. — Even if slos- 
sonae were not confined to the subtropical 
region of .south Florida, its habitus would per- 
mit its unambiguous identification. The dorsal 
stripes of alhida (Fig. 2A) are somewhat simi- 
lar to those of slossonae, but in the latter 
species the stripes extend conspicuously to 
the rear of the forewings (Fig. 2B). Also, the 



forewings (Fig. 2B) are obliquely truncate, 
exposing the terminal abdominal segment, 
which is also striped. The forewings of ritana 
(Fig. 2D) are broadly truncate. 

VIII. The ritana Group 

The ritana group consists of a single spe- 
cies. The monobasic group is defined by the 
bizarre aedeagal apex, which we designate as 
an autapomorphy. 

20. Flexamia ritana Beamer 

Flcxainiii ritana Beamer 1936: 257. 

Important CHARACTERS. — Length of 6 4.2 
(3.9-4.4) mm, of 9 4.4 mm. Crown produced 
(mean crown length 1.67 x interocular width; 
0.73 X head width) (d n = 4; $ n = 1). Dorsum 



1988 



Whitcomb, Hicks; Flexamia Revision 



257 



with stripes extending from head to forcwings 
(Fig. 2D). Face pale white with broad, hiscous 
interocular band. [6] Plates (Fig. 8D) about 
1/2 length of pygofer, fused basally, tapering 
to bluntly acute apices. Aedeagus symmetri- 
cal, gonopore on caudoventral surface at base 
of unpaired processes (Fig. 7D); processes 
with entire edges, together lyriform; apode- 
mal processes with anteapical protuberances. 
[ 9 ] Sternum VII with conspicuous, median 
projection on hind margin. 

Geographic distribution. — Flexamia ri- 
tana has been collected only in the Santa Rita 
Mountains of Arizona (Fig. 30). 

Biology. — Despite many field hours ex- 
pended in search, ritana has not been found 
since its collection in 1935. The original field 
notes of R. H. Beamer are not explicit in 
pinpointing the locality (see notes, Appendix 
III). Our field studies have emphasized col- 
lections on relatively pure patches of grass. 
Considering the accumulating evidence that 
primitive Flexamia species may be habitat 
specialists, future efforts should perhaps be 
directed at diverse habitats rather than single 
host species. 

Remarks and diagnosis. — Superficially, 
ritana (Fig. 2D) resembles albida (Fig. 2A) 
but lacks transverse lines on the crown and 
has no darkly pigmented forewing cells. The 
forewings (Fig. 2D) are broadly truncate. The 
male plates and female sternum VII are en- 
tirely unlike those of either slossonae or al- 
bida. 

IX. The serrata Group 

This group consists of a single species. The 
defining autapomorphy of the group is the 
pair of dorsal, aedeagal processes. 

21. Flexamia serrata Beamer & Tuthill 
Flexamia serrata Beamer & Tuthill 1934: 4. 

Important characters. — Length of 6 3.9 
(3.6-4.3) mm, of 9 3.8 (3.2-4.4) mm. Crown 
moderately produced (mean crown length 
1.26 X interocular width, 0.66 x head width) 
{6 n = 20; 9 n = 20). Face pale with interocu- 
lar band consisting of parallel lines inter- 
rupted at middle. [6] Pygofer (Fig. 62P) with 
posterior margin of lobe subangulate above 
midlength. Plates (Fig. 8C) short, broadly tri- 
angulate, about half length of pygofer, acute at 
apices. Aedeagus (Fig. 7A) subsymmetrical. 



gonopore anteapical on caudoventral surface. 
Prominent, unpaired, ventral process basad 
of gonopore, extending basad along shaft, pair 
of shorter, recurved processes arising on each 
side of gonopore. Apodemal processes with 
mesal, anteapical protuberances. The un- 
paired aedeagal processes of some males from 
Linton, North Dakota (11 August 1977, 3 6, 
3 9, R. F. Whitcomb, IPL 000351, and 24 
July 1985, 34 c? , 16 9 , R. F. Whitcomb and 
E. A. Clark, IPL 001612) and Balta, North 
Dakota (1 6, 3 August 1985, K. G. A. Hamil- 
ton) are broken; such genitalia can be initially 
confusing. [9] Sternum VII (Fig. IOC) with 
broad, median, posterior projection. Oviposi- 
tor with base of each first valvula very strongly 
recurved, recurved portion approximately 
parallel to axis of valvula (Fig. 63 P). 

Geographic distribution. — This species 
(Fig. 30) occurs from the southern tier of west- 
ern Canadian provinces to Utah and North 
Dakota. 

Biology. — Flexamia serrata is apparently 
associated with Muhlenbergia richardsonis , 
where it often occurs (especially in Canada) 
with decora. 

Oligophagy coefficients. — Gramineae 
1.000; Chloridoideae 1.000; Muhlenbergia 
richardsonis 1.000 (n = 57). 

Remarks and diagnosis. — The face pattern 
of serrata (Fig. 3A) is unique. The range of 
this species overlaps considerably with that of 
stylata, which has superficially similar facial 
lines. However, in stylata the lines are not 
interrupted at the middle and contrast with a 
yellow rather than white face. The male plates 
oi stylata (Fig. 34E) and the unique morphol- 
ogy of the male pygofer (Fig. 5C) also differen- 
tiate it from serrata. Whereas all individuals 
of stylata have at least a trace of a transverse 
line midlength on the crown, serrata has a 
completely unmarked crown. 

X. The flexulosa Group 

The flexulosa species group consists of 
11 species. Evolution within this group has 
occurred on Aristida, Bouteloua, and, espe- 
cially, Muhlenbergia. We have found no syn- 
apomorphy to define the group, but we have 
been able to define four subgroups. The nomi- 
nate subgroup consists of two species. One of 
these, arizonensis, specializes on Aristida 
purpurea and/or other Aristida species. The 
other member of this subgroup is flexulosa, a 



258 



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No. 12 



blue grama (Bouteloua gracilis) specialist. 
This species is designated as a sister to ari- 
zonensis on the basis of the extremely short, 
apically divergent plates, which are synapo- 
morphic. The decora subgroup consists of 
three species. F. decora and youngi are iden- 
tified as sisters on the basis of their ventral, 
aedeagal processes, which are much longer 
than the paired processes. Both are specialists 
on Muhlenbergia richardsonis . The current 
geographic distribution of these species sug- 
gests that they represent vicariant segregates 
of an ancestral Muhlenbergia specialist. The 
third species, modica, is a specialist on Muh- 
lenbergia repens (a close relative of M. 
richardsonis ) in more mesic grasslands of the 
eastern Desert Plains. This species is linked to 
the decora subgroup by the morphology of the 
male pygofer and female sternum VII. The 
third subgroup consists of three species: 
arenicola, a specialist on Muhlenbergia pun- 
gens; stylata, a specialist on Muhlenbergia 
species in uplands of the northern prairie and 
the eastern Rocky Mountains; and celata, a 
specialist of chloridoid grasses in sandhills, 
including sand blowout grass, Redfieldia flex- 
uosa. The latter two species are designated 
sisters on the basis of their unique male 
pygofers, which are synapomorphic. A fourth 
subgroup consists of three species. One of 
these, inflata, a species of moist northern 
prairies, commonly occurs on Muhlenbergia 
asperifolia, which enables it to colonize ripar- 
ian areas of the Southwest. Flexamia texana, a 
rare species from south central Texas whose 
biology is unknown, is designated sister to 
beameri, a species from New York whose biol- 
ogy is also unknown. The triangulate male 
plates and boat-shaped pygofers are synapo- 
morphic for this subgroup. 

Description oi the flexulosa Group 

Small to large. Length of 6 2.9-4.6 mm, 
of 9 3.0-4.7 mm. Gray to stramineous. 
Crown not strongly produced in some species 
{decora, inflata), moderately produced in 
others. Face in many species iflexulosa and 
decora subgroups) white, ivory, or at the 
darkest, very pale yellow, contrasting sharply 
with dark, usually black interocular band. 
Face patterns in stylata subgroup similar, but 
with interocular band of parallel, discrete 
lines. In other species {inflata subgroup), face 
stramineous with no interocular band or at 



most indistinct band not contrasting with 
lower face. Male plates triangulate, short or 
elongate, often apically divergent, in some 
species fused basally. Aedeagus and connec- 
tive articulated. Aedeagus with paired apical 
processes and ventral unpaired process in 
sagittal plane, or displaced from sagittal plane 
{inflata subgroup). Gonopore apical or sub- 
apical on caudoventral surface. Female ster- 
num VII with median projection. 

Key to Males of the/7erH/osa Group 

1. Pvgofer produced caudallv in spinelike process 

(Fig. 5C) ■ 2 

— Pygofer not so 3 

2(1). Shaft of aedeagus thick, conspicuously curved, 
expanded apicallv with long, apical appendages 
(Fig. 31H) stijlata (Ball) 

— Shaft of aedeagus slender, not conspicuously 
curved or expanded apically, with shorter apical 
appendages (Fig. 311) . . . celata Lowry & Blocker 

3(1). Pygofer with posterior lobe produced, heavily 
sclerotized, and angled on ventral margin (Figs. 
32E; 62MM) arizonensis Young & Beime 

— Pygofer with posterior lobe rounded ventrally . . 4 

4(3). Aedeagus with greatly elongate, unpaired ven- 
tral process and with shaft straight or arched 
caudodorsally 5 

— Aedeagus with paired and unpaired processes 
more or less same length or if one greatly elon- 
gate, then shaft sigmoidal 6 

5(4). Aedeagal shaft (Fig. 31G) curved, unpaired pro- 
cess straight decora Beamer & Tuthill 

— Aedeagal shaft (Figs. 31F, 38A,B) straight, un- 
paired process recurved youngi, n. sp. 

6(4). Aedeagal shaft at least weakly sigmoidal in lat- 
eral aspect 7 

— Aedeagal shaft arched caudodorsally, not sig- 
moidal 9 

7(6). Aedeagus with paired processes less than 1/4 
length of unpaired process, and not diverging 
from plane of shaft in lateral aspect (Fig. 31E) 
inflata (Osborn & Ball) 

— Paired processes longer, and diverging from 
plane of shaft in lateral aspect 8 

8(7). Aedeagal processes irregular, apex strongly 
asymmetrical (Fig. 31A); south central Texas 
texana Young & Beirne 

— Aedeagal apex subsymmetrical (Fig. 31 D), pro- 
cesses regular; New York beaineri, n. sp. 

9(6). Pygofer in lateral aspect with caudoventral mar- 
gin from ventral lobe to caudal apex straight, not 
curved; paired processes of the aedeagus in lat- 
eral aspect diverging dorsally from the plane of 
the shaft; dorsal keels of the connective narrow; 
plates (Fig. 33A) small, short flextdosa (Ball) 



1988 



WhITCOMB, HlCKS: FLEXAMIA REVISION 



259 




Fig 31. Aedeagal apices of the Flexamia flexulosa group: A, texana- B, imputans; C, areolata; D,beamen; E, inflata; 
F youngU G, decora; H, stylata; I, celata: J, arenicola; K, i?exu/o5a, L, arizonensis; M, modtca. Aspects: CL, 
caudolateral CV, caudoventral; DS, dorsal; LT, lateral; VN, ventral. Redrawn from Young and Beime (1958). 



260 



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No. 12 









DlX 



Fig. 32. Male plates and pygofers oi some flcxulosa group species, lateral aspect: A, ijoungi, B, mocUca, C, arenicola, 
D, decora, E, arizonensis, F,fJexuIosa. 



— Pygofer in lateral aspect with caudoventral margin 
curved from ventral lobe to caudal apex; paired 
processes of aedeagus in lateral aspect diverging 
weakly or not at all from the plane of the shaft; 
dorsal keels wider in lateral aspect; plates (Figs. 
33C,F) comparatively long 10 

10(9). Apices of paired processes curving ventrad in 
lateral aspect; processes and shaft of aedeagus 
comparatively robust . . . inodica Beamer & Tuthill 

— Apices of paired processes curving dorsad in 
lateral aspect; processes and shaft comparatively 
slender arenicola Lowry & Blocker 

Key to Females oi the flexulosu Group 
1. Ovipositor with base of each first valvula curved 



2(1). 



dorsad and laterad, forming angle of more than 
45 degrees with long axis of valvula and extend- 
ing laterad beyond lateral margin of valvula 
when viewed from above (Figs. 63Q,R,AA) .... 2 

Ovipositor with recurved portion of each first 
valvula not extending laterad beyond lateral 
margin of valvula (or only very slightly so); if 
longer than broad, then with long axis forming 
angle of less than 45 degrees with axis of valvula 
in dorsal aspect 4 

Recurved base of each first valvula very slender 
in dorsal aspect, posterior margin concave (Fig. 
63AA) stylata (Ball) 

Recurved base of each first valvula broader in 



1988 



Whitcomb, Hicks: Flexamia Revision 



261 




Fig. 33. Male plates and pygofers of the Flexamia flexulosa group: A, flexulosa, B, arizonensis; C, arenicola; D, 
decora; E, youngi; F, modica. 



dorsal aspect, anterior margin straight or convex 3 

3(2). Recurved base of each first valvula more distant 
from broadest part of valvula and with dorsal 
portion often lightly sclerotized and with small, 

anterior point (Fig. 63Q) 

decora Beamer & Tuthill 

— Recurved base of each first valvula without 
small, anterior point and closer to broadest part 

of valvula (Fig. 63R) . . . modica Beamer & Tuthill 

4(1). Recurved portion of each first valvula short, 
with acute, dorsal projection directed anterolat- 
erad (Fig. 63T) inflata (Osborn & Ball) 

— Recurved portion of first valvula without such 
projection 5 

5(4). Recurved portion of first valvula concave poste- 
riorly in dorsal aspect (Fig. 63Y) 

arizonensis Young & Beirne 

— Recurved portion of each first valvula not so ... 6 

6(5). Recurved portion of each first valvula directed 

almost caudally (Fig. 63BB) flexulosa (Ball) 

— First valvula not so 7 

7(6). Upper New York State beameri, n. sp. 

— South central Texas texana Young & Beirne 



22. Flexamia arizonensis Young & Beirne 

Flexamia arizonensis Young & Beirne 1958: 30. 

Important CHARACTERS. — Length of 6 3.3 
(3.0-3.8) mm, of 9 3.6 (3.2-4.0) mm. Head 
variably produced (1.40 x interocular width; 
0. 70 X head width) {6 n = 20; 9 n = 20). Face 
pale white with black interocular band. [6] 
Pygofer with posterior lobe (Fig. 62MM) api- 
cally truncate, ventral portion extending ven- 
trad as short, heavily sclerotized process. 
Plates short, less than 1/2 length of pygofer, 
apically divergent. Aedeagus symmetrical, 
paired apical processes arising on each side of 
subapical gonopore, unpaired ventral process 
about one-third length of shaft. [ $ ] Sternum 
VII with broad, median projection. Oviposi- 
tor with each first valvula (Fig. 63Y) curved at 
base more than 90 degrees, recurved portion 
concave caudally in dorsal aspect. 

Geographic distribution. — ^This species 
is more widely distributed than had been pre- 
viously supposed, occurring (Fig. 29) from 



262 



Great Basin Naturalist Memoirs 



No. 12 






Fig. 34. Male plates and pygofers of the Flexamiaflexulosa, imputans, and areolata groups, ventral aspect: A, inflata; 
B, texana; C, heameri, D, celata; E, stylata; F, imputans; C, areolata. 



western Nebraska, Colorado, Utah, central 
Texas, and Arizona to northern Mexico 
(Sonora and Chihuahua). Two females from 
southern California (Pine Flats Camp, Indio, 
L. H. Banker, 12 July 1941 [KU]) may repre- 
sent this species. 



Biology. — This species is a specialist on 
perennial Aristida spp., especially A. pur- 
purea. 

Olicophagy coefficients. — Gramineae 
1.000; Chloridoideae 1.000; Aristida spp. 
0.932. 



1988 



Whitcomb, Hicks: Flexamia Revision 



263 




^A 




{ 


\ -"1""' 




iL'^'--' 


—?f.- 


'^'":"""r"' 


■ 7 . 


J— -'-''' 



Fig. 35. Geographic distribution o( Flexamia flexulosa and its host, Bouteloua gracilis. 



Remarks and diagnosis. — Prior to the 
study of Young and Beirne, arizonensis had 
been confused with flexulosa. Males of these 
species can be distinguished by the lateral 
aspect of their undissected pygofers (Fig. 
32E,F; see discussion under flexulosa). Al- 
though the dorsum of arizonensis is usually 
pale, several specimens from southern Ari- 
zona show dorsal stripes that are reminiscent 
of those ofritana. Since the external genitalic 
characteristics of arizonensis are very distinct 
from those oi ritana, such specimens pose no 
problem for identification. However, they are 
important in suggesting an evolutionary link 
between the ritana and flexulosa groups. 

23. Flexamia flexulosa (Ball) 

Deltocephalus flexulosus Ball 1899: 189. 
Deltocephalus (Flexamia) flexiilosus , DeLong 1926; 31. 
Flexamius flexulosus, DeLong and Sleesman 1929: 84. 

Important CHARACTERS. — Length of S 3.2 
(2.9-3.7) mm, of 9 3.5 (3.0-4.1). Head vari- 
ably produced (1.26 x interocular width; 0.60 
xhead width) (d n = 40; 9 n = 44). Face pale 
with black interocular stripe. [6] Pygofer 
(Fig. 62NN) with posterior lobe rounded api- 
cally. Plates very short, less than 1/2 length of 



pygofer, apically divergent. Aedeagus sym- 
metrical, gonopore anteapical on caudoven- 
tral surface; paired apical processes short, 
minutely serrulate along one edge; unpaired 
ventral process short, grooved, less than one- 
fourth length of the shaft, acute at apex. [ 9 ] 
Sternum VII with hind margin produced to 
form convex lobe; ovipositor with each first 
valvula curved through more than 90 degrees 
at base, recurved portion slender, almost par- 
allel to long axis of valvula, with narrow pro- 
jection extending caudad at basal extremity. 

Geographic distribution. — This species 
occurs (Fig. 35) from eastern Nevada, Utah, 
Wyoming, Montana, and the southern tier of 
Canadian provinces from Alberta to Mani- 
toba, south through the Great Plains to west- 
ern Texas, Arizona, and northern Mexico 
(Chihuahua). 

Biology. — Although flexulosa and ahhre- 
viata are both specialists of blue grama 
(Bouteloua gracilis), their ranges do not com- 
pletely overlap. Whereas y/(?x«/osa appears to 
be adapted solely to blue grama in cool, dry 
climates, abbreviata is adapted to warmer 
grasslands, especially the mixed Bouteloua 



264 



Great Basin Naturalist Memoirs 



No. 12 



grasslands of Texas, New Mexico, and Ari- 
zona. Such differences may be important 
clues to the evolutionary origins of these Flex- 
amia species. 

Oligophagy coefficients. — Gramineae 
1.000, Chloridoideae 0.973; Bouteloua gra- 
cilis 0.889 (n - 380). 

Remarks and diagnosis. — Young and 
Beirne (1958) consider flexulosa to be "greatly 
variable." Actually, we have found substantial 
variability in all Flexamia species for which 
large sample sizes were available; flexulosa 
seems to us to be no more or less variable than 
other species. The face pattern of flexulosa 
(black interocular band contrasting with a pale 
face) is, in fact, remarkably stable. Unfortu- 
nately, this contrasting pattern is shared by 
other members of the flexulosa group (ari- 
zonensis, modica, arenicola, decora, and 
youngi). Although the definitive identity of 
these species should be ascertained by exami- 
nation of genitalic characteristics, flexulosa is 
an abundant species, so it is important to de- 
fine external characters that permit it to be 
sorted from less common species. The lateral 
aspect of the male pygofer (Figs. 32A-F) is a 
useful character in sorting white-faced mem- 
bers of the flexulo.sa group. In flexulosa the 
posterior margin of uncleared male pygofers 
appears rounded. The pygofer (Fig. 32E) of 
the widely distributed arizonensis is truncate 
apically, but it is necessary to clear the speci- 
men to see the distinctive, heavily sclerotized 
ventral process. In regions where both decora 
SLud flexidosa occur, decora can often be rec- 
ognized by its characteristic habitus (see dis- 
cussion under that species), and by its pygofer 
(Fig. 32D), which appears narrower than that 
of other white-faced species. The pygofer of 
rnodica (Fig. 32B), an uncommon but locally 
abundant (New Mexico mountains) specialist 
of Muhlenbergia repens, is also diagnostic. 
The pygofer of arenicola (Fig. 32C) is also 
more or less diagnostic; this insect is an inhab- 
itant of Muhlenbergia pungens in the Ne- 
braska Sand Hills and the Four Corners re- 
gion of the Colorado Plateau (Fig. 36). Thus, 
the combination of male pygofer, geographic 
range, and host data (if available) provides an 
excellent character set that permits tentative 
recognition of pale-faced males of the flexu- 
losa group. Nevertheless, the identity of 
these species, induding flexulosa , should be 



confirmed by examination of characters of the 
genitalia. 

24. Flexamia arenicola Lowry & Blocker 
Flexamia arenicola Lowry & Blocker 1987; .59. 

Important characters. — Length of d 3.7 
(3.5-3.9) mm (Sand Hills form), 3.4 (3.2-3.8) 
mm (Anasazi form); of 9 4.0 (3.8-4.3) mm 
(Sand Hills form), 3.7 (3.3-4.1) mm (Anasazi 
form). Crown moderately produced (median 
crown length 1. 19 x [Sand Hills form] or 1.35 x 
[Anasazi form] interocular width; 0.63 x head 
width [both forms]) ((5 n - 20, 9 n - 7 [Sand 
Hills form]; d n = 20, 9 n = 18 [Anasazi 
form]). Crown moderately produced (median 
crown length 1.25 x interocular width; 0.67 x 
head width). Face pale, with conspicuous, 
black interocular band. [6] Pygofer (Fig. 
62LL) broadly ovate; plates (Fig. 33C) broadly 
triangulate, diverging apically. Aedeagus 
(Fig. 31J) symmetrical with gonopore anteapi- 
cal on the caudoventral surface, lateral, apical 
processes approximately one-fourth shaft 
length and curved laterodorsad; unpaired 
process approximately the same length and 
directed anteriorly. [ 9 ] Sternum VII with 
posterior margin produced medially with 
small, median notch. Ovipositor with re- 
curved process at base of first valvula (Fig. 
63S) not exceeding lateral margin, process ex- 
tending caudad, sinuate and digitate apically. 

Geographic distribution. — There are two 
disjunct populations of arenicola (Fig. 36). 
One population occurs in the sandhills of 
western Nebraska and northeastern Colo- 
rado. The other population occurs in the Colo- 
rado Plateau and adjacent regions of New 
Mexico. The disjunct distribution of arenicola 
can be readily explained by the essentially 
disjunct distribution of its host (Fig. 36). 

Biology. — Flexamia arenicola has been 
found almost exclusively on sandhill muhly, 
Muhlenbergia pungens. 

Oligophagy coefficients. — Gramineae 
1.000; Chloridoideae 1.000; Muhlenbergia 
pungens 0.989 {n 91). 

Remarks and diagnosis. — The unpaired 
processes of the aedeagus in males of the Four 
Corners population (which we term the 
"Anasazi form" in honor of the pre-Columbian 
residents of the area) is uniformly absent, but 
on close inspection it can be seen to be bro- 
ken; this presumably occurs during copula- 
tion. The occurrence of the Anasazi form is 



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Fig. 36. Geographic distribution of Flexamia arenicola (Sand Hills 
Muhlenbergia pungens. 



[•] and "Anasazi" [■] forms) and the host. 



patchy, and populations usually are low. Be- 
cause the region occupied is now undergoing 
desertification and salinization as a result of 
climatic changes during the interglacial era, 
this form may be a relict population that, like 
the host population, is currently stressed by 
climatic change. Presumably, the outcome of 
these selective pressures could lead to extir- 
pation or to evolution of a new species that 
would be better adapted to semiarid environ- 
ments. The characteristics of the male pygofer 
(Fig. 32C) in lateral aspect, in combination 
with the face characteristics, are diagnostic for 
arenicola. Lowry and Blocker (1986) distin- 
guished arenicola from flexiilosa by the ex- 
panded dorsal keels of the arenicola connec- 
tive, the more robust aedeagal shaft, and the 
longer aedeagal processes. They distin- 
guished arenicola from modica by its larger 
size and nearly symmetrical aedeagus with 
lateral, dorsally curved, apical processes. In 
addition, the pygofer of modica (Fig. 32B) is 
very different from that of arenicola. Some 
characters in arenicola are similar to those of 
celata and stijlata, but the general facies of the 
latter two species separate them from areni- 
cola. 



25. Flexamia celata Lowry & Blocker 

Flexamia celata Lowry & Blocker 1987; 57. 

Important CHARACTERS. — Length of d 4.1 
(3.9_4.4) mm, of 9 4.5 (4.4-4.7) mm. Crown 
not produced (1. 13 x interocular width; 0.58 x 
head width) (d n = 19; 9 n = 6). Face, as in 
stijlata, pale with broad interocular band con- 
sisting of parallel, discrete lines. [6] Pygofer 
(Fig. 62PP) with central margin strongly pro- 
duced to form spinelike process, as in stijlata. 
Plates (Fig. 34D) extending to approximately 
2/3 length of pygofer, fused mesally for 1/2 
length, narrowed apically to rounded, lateral 
lobe, apices meeting in V-shaped notch. 
Aedeagus (Fig. 311) symmetrical, shaft slen- 
der, neither conspicuously curved nor ex- 
panded apically, ventral and paired apical 
processes approximately equal in length and 
less than 1/4 length of shaft; gonopore apical 
on caudoventral surface. [ 9 ] Sternum VII 
with posterior margin medially produced, 
with slight median notch; ovipositor with 
basal processes of first valvulae recurved and 
extending laterad beyond lateral margin. 

Geographical distribution. — Nebraska 
Sand Hills and southwestern Kansas (Fig. 37). 



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Fig. 37. Geographic distribution ofFlexamia stylata (•) and celata (■) and a host oicelata, Redfleldia flexuosa. 



Biology. — This species occurs on sand 
blowout grass, Redfieldia flexuosa, the pri- 
mary colonist of bare sand in the Sand Hills. It 
may also occur on other sandhill grasses. It has 
been recorded from Calmyiovilfa longifolia in 
southwestern Kansas (I (?, 3 9 , Garden City, 
Finney County, 12 June 1948, R. H. Beamer). 

Oligophagy coefficients. — Gramineae 
1.000; Redfieldia flexuosa 0.612; Muhlenber- 
gia pungens 0.265 (n = 49). 

Diagnosis and remarks. — This is a close 
sister species with stylata, under which diag- 
nostic characters have been discussed. The 
present-day area of endemism was apparently 
created by the southern interface of the ice 
sheet during the last glacial maximum (Wright 
1970), suggesting that celata could represent a 
recent speciation. On the other hand, its mor- 
phological characters are much closer to the 
ground plan for the flexulosa group than are 
those oi stylata. The aedeagal symmetry and 
wide pygofer of celata suggest a relationship 
with arenicola, which occurs with it in the 
Nebraska Sand Hills. 

26. Flexamia stylata (Ball) 
Deltocephalus stylatus Ball 1899: 190. 



Deltocephaltis (Flexamia) stylatus, DeLong 1926: 34. 
Flexamius stylatus, DeLong and Sleesman 1929: 84. 

Important CHARACTERS. — Length of c? 4:1 
(3.7-4.6) mm, of 9 4.2 (3.8-4.6) mm. Crown 
not strongly produced (1.17 x interocular 
width; 0.54 X head width) (c? n = 33; 9 n = 
33). Face pale, except for broad interocular 
band consisting of discrete, parallel lines. [6] 
Plates (Fig. 34E) with conspicuous, sub- 
rectangular apices; pygofer (Fig. 6200) with 
posterior margin strongly produced, gradu- 
ally narrowed, forming curved, spinelike pro- 
cess. Aedeagus asymmetrical; shaft subcylin- 
drical; gonopore subapical, at base of three 
processes on caudoventral surface; pair of api- 
cal processes directed laterobasad, unpaired 
ventral process extending basad to midlength 
of shaft, but not parallel to axis of shaft, the 
three processes appearing subequal in length 
in caudoventral aspect; paired processes ap- 
pearing shorter in lateral aspect, each slightly 
serrate on dorsal edge. [ 6 ] Sternum VII as for 
group; ovipositor with base of each first 
valvula (Fig. 63AA) curved dorsad through 
slightly more than 90 degrees, recurved por- 
tion extending laterad beyond lateral margin 



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267 



of valvula in dorsal aspect. 

Geographic distribution. — Southern tier 
of Canadian provinces south through the 
western Great Plains, eastern Rocky Moun- 
tains, northern New Mexico, and Arizona 
(Fig. 37). 

Biology. — This species has been collected 
from Muhlenhergia on upland slopes in Mon- 
tana and Wyoming. In northern New Mexico 
the host is Muhlenhergia wrightii. This host 
may also account for records from northern 
Arizona. 

Diagnosis and remarks. — The male plates 
of this species are unique. The face pattern 
(interocular bands with discrete lines) that 
stylata shares with celata is very distinctive; 
superficially similar species almost always 
have bands with coalesced lines. Finally, sty- 
lata specimens are larger than those of most 
other species. Females of stylata and celata 
are similar but can be distinguished by charac- 
teristics of the bases of the first valvula of the 
ovipositor. Also, stylata is a widely dis- 
tributed Muhlenhergia specialist, whereas 
celata is restricted largely to the Nebraska 
Sand Hills, where it is a rare inhabitant of 
Redfieldia flexiiosa and other chloridoid 
grasses. 

27. Flexainia decora Reamer & Tuthill 

Flexamia decora Beamer & Tuthill 1934: 2. 

Important characters. — Length of c? 3.6 
(3.0-4.2) mm, of 9 3.9 (3.4-4.2) mm. Head 
not strongly produced (median crown length 
1.07 X interocular width, 0.62 x head width) 
(c? n = 20; $ n = 20). Face pale with broad 
black interocular stripe. [6] Pygofer (Fig. 
62FF) with slightly truncate rear margin. 
Plates (Fig. 33D) broadly divergent, with nar- 
row apices. Aedeagus (Fig. 31G) symmetrical. 
Gonopore anteapical on caudoventral surface; 
pair of retrorse processes arising on each side 
of gonopore, each serrate on dorsal edges. 
Unpaired ventral processes, with entire 
edges, extending basad more than half length 
of shaft. [ 9 ] Sternum VII with hind margin 
concave at each side of median convex projec- 
tion, notched apically, with teeth on each side 
of notch. Ovipositor with each first valvula 
(Fig. 63Q) with base expanded dorsally and 
laterally. 

Geographic distribution. — Flexamia de- 
cora, as defined herein, is a specialist oi Muh- 



lenhergia richardsonis in the New Mexico 
and Colorado Rockies north to the Peace 
River region of northwestern Alberta, and 
east to Manitoba and North Dakota (Fig. 39). 
Its host occurs in eastern North America but 
does not occur south of Maine. We have ex- 
amined specimens reported by Young and 
Beirne (1958) from Kentucky and Virginia. 
The Kentucky specimen is a female referable 
to inflata. The Virginia specimen labeled only 
"Oct. 12, Va., ED. Ball" appears to have been 
recently relabeled (and therefore possibly 
mislabeled), perhaps at the time of acquisition 
of the Ball collection. There are therefore no 
reliable records for decora east of the mixed- 
grass prairie. 

Biology. — Flexamia decora appears to be 
largely a specialist of Muhlenhergia richard- 
sonis. Large populations are not uncommon, 
and decora is the most abundant Flexamia 
species in the CNC (Appendix I). 

Oligophagy coefficients. — Gramineae 
1.000; Chloridoideae 0.874; Muhlenhergia 
richardsonis 0.874. [Note: M. richardsonis 
has been present in all communities where 
decora occurs.] 

Remarks and diagnosis. — Many individu- 
als of decora have a rather distinctive habitus, 
with very dark fore wing markings, especially 
on the apical portions, and along the veins of 
the forewing that contrast sharply with the 
lighter pronotum and crown. Unfortunately, 
paler specimens are not rare; the identity of 
these specimens is often indicated by the ex- 
tremely short crown length. Flexamia decora 
and its close sister species, youngi, are appar- 
ently restricted to Muhlenhergia richard- 
sonis. 

28. Flexamia youngi, n. sp. 

Length of c? 3.7 (3.4-3.9) mm, 9 3.8 
(3.7-4.2) mm; head width of 6 1.12 mm, 9 
1.18 mm. Crown not produced; median 
length of crown approximately 1.11 x head 
width and 0.58 x interocular width (c? n = 16; 
9 n = 20). 

Color Stramineous. Crown with at most a 
trace of basal or midlength markings. Prono- 
tum with or without traces of wide, brown 
stripes. Face with black interocular line con- 
trasting with white lower face. Venter strami- 
neous. 

Male. — Pygofer (Fig. 38D) as in decora 
but with posterior lobe less angulate, more 



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Fig. 38. Flexamia youngi, n. sp.: A, aedeagus and connective, lateral aspect; B, aedeagus and connective, dorsal 
aspect; C, male plates and pygofers, ventral aspect; D, male pygofer, lateral aspect; E, right style, dorsal aspect; F, 
female sternum VII; G, bases of first valvulae of female, dorsal aspect. 



broadly rounded ventrally; plates (Fig. 38C) 
extending approximately 2/3 length of pygo- 
fer, fused basally for 2/3 length, connective in 
lateral aspect (Fig. 38A) with dorsal keels 
broad, approximately 1/3 height of dorsal 
apodeme; apodemal processes sinuate in ven- 
tral view, bell-shaped in caudal aspect; styles 
as in other members of decora subgroup; 
aedeagus (Fig. 38B) asymmetrical, shaft elon- 
gate, apparently twisted 1/8 turn counter- 
clockwise in caudal aspect, tapering gradually 
in lateral aspect, slightly enlarged apically in 
ventral aspect; three apical processes extend- 
ing basad; unpaired process recurved, entire, 
with median ventral groove almost 1/2 length 
of shaft. Paired processes recurved, serrate on 
dorsal edges, approximately 1/3 length of un- 
paired process. Gonopore small, circular, and 
apical. 

Female. — Sternum VII (Fig. 38F) with 
posterior margin produced medially, pro- 



nounced median notch with infuscated spots 
on either side. Ovipositor with base of first 
valvula (Fig. 38G) recurved, appearing tear- 
shaped and only slightly exceeding lateral 
margins when viewed dorsally. 

Types. — Holotype 6 : White Pine County, 
Nevada, Baker, 10 August 1986, R. F. Whit- 
comb (IPL 002590, Muhlenhergia richord- 
sonis). Deposited USNM. Paratypes: 10 6, 
119, same collection data; 1 9 , Mt. Wheeler, 
White Pine County, Nevada, 10 August 1986, 
8,500 ft, IPL 002569; 4 d , 4 9 , Spring Valley, 
White Pine Countv, Nevada, 11 August 1986 
(5,800 ft, R. F. Whitcomb, IPL 002596). De- 
posited in BARC, CNC, KSU, and USNM. 
Paratypes (KU) were also collected by R. H. 
Beamer (1 9, Deeth, Elko Co., Nevada, 21 
July 1947); and D. H. Lindsay (1 d, 3 9 
Soldier Summit, Utah, 13 August 1936). 

Remarks. — This species, apparently a 
Great Basin endemic (Fig. 39), is closely 



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269 




Fig. 39. Geographic distribution o( Flexamia decora (•), youngi (O), and modica (I 
richardsonis (decora and youngi) and M. repens (modica). 



and their hosts, Mtihlenbergia 



related to decora but can be distinguished 
from it by the shape of the aedeagus, which is 
straight and twisted rather than curved and 
symmetrical, and by the recurved, apical pro- 
cesses. The host so far as known is (like de- 
cora) Mtihlenbergia richardsonis. This spe- 
cies is named in honor of D. A. Young, whose 
classic monograph, with B. P. Beirne, estab- 
lished the modern generic concept of Flex- 
ainia. 

29. Flexamia modica Beamer & Tuthill 
Flexamia modica Beamer & Tuthill 1934; 3. 

Important CHARACTERS. — Length of 6 3.4 
(3.1-3.7) mm, of 9 3.8 (3.4-4.0) mm. Crown 
moderately produced (1.26 x interocular 
width, 0.62 X head width) (c? n = 20; 9 n = 
20). Face ivory, with broad interocular band. 
[6] Pygofer (Fig. 62JJ) as in decora. Plates 
(Fig. 33F) short, about 2/3 length of pygofer, 
broadly triangulate, widely divergent at 
apices. Aedeagus (Fig. 31 M) asymmetrical, 
gonopore triangular; unpaired ventral process 
less than half length of shaft, base appearing 
fiised to shaft in lateral aspect. [9] Sternum 



VII with broad median projection; ovipositor 
with base of each first valvula turned dorsad 
through 90 degrees and bilobed, recurved 
portion extending strongly laterad and ex- 
ceeding lateral margin of valvula in dorsal as- 
pect. 

Geographic distribution. — High desert 
plains of southern New Mexico and adjacent 
counties of Trans-Pecos Texas and Arizona 
(Fig. 39). It is abundant in the Sacramento 
Mountains of southeastern New Mexico on its 
host Muhlenbergia repens. It is much less 
common in the Gila Mountains, where its host 
is more patchy, and occurs rarely in the cen- 
tral Guadalupe Mountains (Guadalupe Moun- 
tains National Park, Culberson Co., Texas) 
and northeastern Arizona (St. Johns, Apache 
Co.). 

Biology — Apparently a specialist oi Muh- 
lenbergia repens. Our records show seven se- 
ries from this host 16-18 June 1985 (R. F. 
Whitcomb and K. A. Allred, approximately 
440 S, 9, and immatures). M. repens is 
closely related to M. richardsonis (Morden 
1985, Morden and Hatch 1987). It is likely 
that modica arose from the decora lineage via 



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No. 12 



a host transfer from M. richardsonis. 

Oligophagy coefficients. — Gramineae 
1.000; Chloridoideae 1.000; Muhlenbergia 
repens 1. 000 {n = 377). 

30. Flexamia inflata (Osborn & Ball) 

Deltocephalus inflatus Osborn & Ball 1897: 202. 
Deltocephalus (Flexamia) inflatus, DeLong 1926: 3.5. 
Flexamius inflatus, DeLong and Sleesman 1929: 84. 

Important characters. — Length of 6 3.8 
(3.1-4.4) mm, of 9 4.0 (3.6-4.6) mm. Crown 
short (1.11 X interocular width; 0.58 x head 
width) ((5 n = 26; 9 n ^ 26). Face strami- 
neous, with or (more frequently) without a 
basal, pale brown interocular band. [6] 
Pygofer (Fig. 62GG) with upper portion of 
posterior lobe strongly produced posteriorly. 
Plates (Fig. 34A) short, about 1/2 length of 
pygofer, apically divergent. Aedeagus (Fig. 
31 E) asymmetrical; shaft elongate, slender, 
gradually broadened apically; gonopore apical 
on caudoventral surface; pair of short, re- 
curved processes arising on each side of 
gonopore, slender, unpaired ventral process 
almost half length of shaft, slightly expanded 
anteapically, acute at apex, which extends to 
right of shaft, all processes with edges entire. 
[9] Female sternum VII broadly, medially 
produced, medially notched with traces of 
teeth between notch and lateral margin; 
ovipositor with base of each first valvula (Fig. 
63T) curved dorsad, small projection on dorsal 
edge extending dorsolaterad, entire curved 
portion more heavily sclerotized and appear- 
ing almost as separate sclerite in ventrolateral 
aspect. 

Geographic distribution. — This species 
is distributed (Fig. 41) from Utah, Texas, and 
New Mexico north to British Columbia and 
Manitoba east to Pennsylvania. 

Biology. — This species may be adapted to 
northern mixed grasslands where pooid 
grasses, Juncus spp. , and Muhlenbergia grow 
in intermixed stands. In the Southwest, at 
least, inflata is a specialist on Muhlenbergia 
asperifolia ; occasional collections have also 
been made on Distichlis spicata. We do not 
have host records from the Northwest, but the 
northern limit of the distribution of inflata 
coincides roughly with that of Muhlenbergia 
asperifolia. In Illinois it occurs on Juncus 
tenuis. In Maryland it occurs on J uncus tenuis 
and on the introduced grasses Cynodon dacty- 



lon, Zoysiajaponica, and Eragrostis curvula. 
This species has been reared on /. tenuis 
(Whitcomb 1957). Populations occasionally 
turn up for a time on Festuca or Poa but never 
seem to persist. We have observed no mor- 
phological indications of host races, although 
it is possible that the Juncus populations may 
be reproductively isolated from other popula- 
tions. Rearing studies may be required to re- 
solve the status of the biology oi inflata. 

Oligophagy coefficients. — Prairie and 
savanna: Gramineae 0.725, Juncaceae 0.275; 
Chloridoideae 0.725, Muhlenbergia asperifo- 
lia 0.562; Eragrostis curvida 0.066 (n = 379). 
Other records: Manitoba (Woodside, 24 Au- 
gust 1981, 16 d , 16 9 , Poa pratensis, K. G. A. 
Hamilton, CNC). 

Remarks and diagnosis. — Although defin- 
itive identification of inflata requires exami- 
nation of the genitalia, it is important to learn 
to sort the majority of specimens of this often 
abundant species on the basis of habitus 
alone. The vast majority of specimens of in- 
flata are very lightly pigmented; these can be 
recognized by their light stramineous color in 
combination with a black spot on the apical 
forewing cell (Fig. 4A), by the lack of a spot in 
the corium, and by their crown, which is one 
of the shortest in the genus. 

31. Flexamia beameri, n. sp. 

Length of 6 3.2 (3.1-3.3) mm, 9 3.4 
(3.3-3.4) mm; head width of 6 0.93 mm, 9 
1.06 mm. Crown moderately produced; me- 
dian length of crown 0.67 x head width and 
1.39 X interocular width (d n = 6; 9 n = 2). 

Color stramineous with dark markings on 
corium and apical cell of forewings; face as in 
inflata and texana; head with light markings, 
pronotum with wide, faint stripes; venter and 
legs with irregular, fuscous markings. 

Male. — Pygofer (Fig. 40D) as in texana; 
plates extending to approximately 2/3 length 
of pygofer, fused basally for 3/4 length; con- 
nective in lateral aspect (Fig. 40A) with dorsal 
keels broad, approximately 2/3 height of dor- 
sal apodeme; apodemal processes narrowly 
campanulate in caudal aspect; style (Fig. 40E) 
as in inflata and texana; aedeagus (Figs. 
40A,B) slightly asymmetrical, shaft sigmoidal 
in lateral view, ventral unpaired process not 
quite in median plane of shaft and appearing 
slightly longer than strongly recurved, 
paired, lateral processes; all processes with 
edges entire; gonopore circular and apical. 



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271 










Fig. 40. Flexamia heameri, n. sp.: A, aedeagus and connective, lateral aspect; B, aedeagus and connective, dorsal 
aspect; C, male plates and pygofers, ventral aspect; D, male pygofer, lateral aspect; E, right style, dorsal aspect; F, 
female sternum VII; G, bases of first valvulae of female, dorsal aspect. 



Female. — Sternum VII (Fig. 40F) similar 
or identical to texana, with median notched 
projection adjoined on each side by serrulate, 
infuscated projections; ovipositor with re- 
curved portion of the bases of each first 
valvula (Fig. 40G) narrowing to diverging 
apices in dorsal aspect. 

Types. — Holotype 6: Otter Lake, Oneida 
County, New York, 25 June 1946, R. H. 
Reamer. Deposited KU. Paratypes: 5 S and 2 
9 , same locality, deposited KU and USNM. 

Remarks. — This species, known from a sin- 
gle collection at its type locality (Fig. 41), is a 
sister to texana, a species also known from a 
single collection made long ago. Flexamia 
beameri can be readily separated from inflata 
and texana by the shape and length of the 
aedeagal processes. The aedeagal characters 
oi beameri resemble those of texana, but the 
processes of beam£ri are not twisted. Pre- 
sumably, the ranges oibeameri and texana are 



very different. In contrast to the hot climate of 
south central Texas, the type locality of beam- 
eri is located in one of the coolest regions of 
New York (see Notes, Appendix III). Since we 
have not yet encountered these species in the 
field, their biology is a mystery. This species 
has been named in honor of R. H. Reamer, 
who described several interesting species of 
the genus and who collected the holotype and 
paratypes at a geographic site far from his 
home base. 

32. Flexamia texana Young & Reirne 

Flexamia texana Young & Beirne 1958: 29. 

Important characters. — Length of 6 
3.1 (2.9-3.2) mm, of 9 3.2 (3.0-3.5) 
mm. Crown not strongly produced 
(1.30 X interocular width, 0.63 x head width) 
(c? n = 8; 9 n = 2). [(?] Pygofer (Fig. 62HH) 
with upper margin flat in lateral aspect, pos- 
terior margin gradually rounded. Aedeagus 



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No. 12 




Fig. 41. Geographic distribution oi Flexamia inflata (•), beameri (A), and tcxana (O). 



(Fig. 31A) asymmetrical; unpaired ventral and 
pair of lateral processes, all about equal in 
length, arising near gonopore; ventral process 
not in sagittal plane, lateral processes not in 
symmetrical planes. [ 9 ] Sternum VII as for 
group; ovipositor with base of each first 
valvula (Fig. 63CC) curved dorsad through 
about 90 degrees in lateroventral aspect, base 
acute in lateral aspect, regularly convex in 
dorsal aspect. 

Geographic distribution. — Known only 
from the type locality (Uvalde, Texas). 

Biology. — The host oitexana is unknown. 
Uvalde is situated just south of the southern 
edge of the Edwards Plateau. We have sur- 
veyed the main, dominant, warm-season 
grasses of the Edwards Plateau searching for 
this species. Although these searches turned 
up a new species, collorum, they failed to 
locate texana. The grasslands of the mesquite- 
acacia savannah south of Uvalde have been 
largely plowed for agricultural crops. Perhaps 
E. D. Ball was afforded an opportunity to 
collect in these grasslands before they were 
sacrificed for agriculture. If this is the explana- 
tion for the rarity of this species, we may 
experience continuing difficulty in our search 



for this elusive species. On the other hand, 
discovery of the sister species heameri, whose 
type locality suggests a marshy wetland, raises 
the possibility that texana may turn up in 
wetlands in south Texas, which were over- 
looked in our searches. 

XI. The imputans Group 

The imputans group consists of a single spe- 
cies, which is apparently a Muhlenbergia spe- 
cialist in the northern prairie. This species 
retains the plesiomorphic articulation be- 
tween connective and aedeagus; it could oth- 
erwise be regarded a close sister species to 
areolata, with which it shares a distinctive 
aedeagal morphology. These two species are 
both greenish dorsally and have black faces. 

33. Flcxamia imputans (Osborn & Ball) 

Deltoccphditis i)nput(in.s O.slwrn 6c Ball I89S: 75. 
Deltoceplialti.s (Ficxinnia) iiti})utiius, DeLong 1926: 29. 
Flexainius [.sic] imputans, DeLong and Caldwell 1937; 27. 

Important CHARACTERS. — Length of 6 3.4 
(3.1-3.7) mm, of 9 3.7 (3.3-3.9) mm. Crown 
moderatelv produced (1.33 x interocular 
width; 0.62 X head width) (d n = 10; 9 n = 



1988 



Whitcomb, Hicks; Flexamia Revision 



273 




Fig. 42. Geographic distribution oi Flexamia imputans and a probable host, Muhlenbergia cuspidata. 



14). Face usually entirely black but occasion- 
ally with oblique, pale streak on each gena. 
[6] Genitalia (Fig. 31B) similar to those of 
areolata but with hind margin of pygofer (Fig. 
62V) convex and with distinct joint between 
connective and aedeagus. [9] Sternum VII 
(Fig. lOP) with hind margin broadly, slightly 
convexly produced with slight median notch; 
ovipositor with bases of first valvulae (Fig. 
63II) not recurved. 

Geographic distribution. — Northern 
Great Plains from the Dakotas south to Kan- 
sas, east to Iowa and Wisconsin (Fig. 42). 

Biology. — This species is apparently a 
Muhlenbergia specialist and may occur 
largely on Muhlenbergia cuspidata. It is one 
of the rarest of the Flexamia species and, 
given the extensive turnover of its original 
upland prairie habitat to croplands, may be- 
come increasingly rare. 

Remarks and diagnosis. — This species is 
unique in its lack of markings on head and 
pronotum, in combination with its black face. 

XII. The areolata Group 
This group consists of a single species. 
34. Flexarnia areolata (Ball) 



Deltocephalus areolatus Ball 1899: 188. 
Deltocephahis (Flexamia) areolatus, DeLong 1926: 25. 
Flexamia areolatus, DeLong and Sleesman 1929: 84. 

Important gharacters. — Length of (5 2.8 
(2.6-3.2) mm, of 9 3.6 (3.2-4.0) mm. Crown 
strongly produced (1.77 x interocular width; 
0.75 X head width) (c? n - 20; 9 n = 22). Face 
entirely black. [6] Pygofer (Fig. 62Z) with 
dorsal portion of posterior lobe produced cau- 
dodorsad, posterior margin oblique and 
slightly concave. Plates (Fig. 34G) elongate, 
gradually narrowed from base to apex, fused 
basally for about 1/3 length. Aedeagus and 
connective fused. Aedeagus (Fig. 31C) asym- 
metrical; three tapering apical processes, two 
short, with serrate edges, extending basad 
along right side of shaft; third process extend- 
ing basad for about half the length of right side 
of shaft, minute projections in middle, 
broader than other two processes, bearing 
gonopore as elongate slit through almost en- 
tire length. [6] Sternum VII (Fig. lOQ) with 
posterior margin concave on each side of api- 
cally notched, median, convex projection; 
ovipositor with first valvulae not curved dor- 
sad at bases. 



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Fig. 43. Geographic distribution ofFlexamia areolata and its host, Eragrostis spectabilis. 



Geographic distribution — This species 
occurs from Connecticut to Wisconsin, south 
to Florida and Texas, covering much of the 
geographic range of its host (Fig. 43). One of 
the three specimens from the original cotype 
series is labeled "Phoenix, Arizona, May, 
1897." One specimen in the CNC is labeled 
"San Diego, Calif., 19-6-71, GL1251, H. H. 
Ross." Nevertheless, occurrence of areolata 
in the Southwest requires confirmation. 

Biology. — We consider areolata to be a 
specialist of Eragostis spectabilis. DeLong 
(1949) believed that the host was "Panicum or 
a closely related genus." Panicum capillare, 
an annual that commonly occurs with E. 
spectabilis in sandy habitats, has a superfi- 
cially similar inflorescence characterized by 
wide panicles that redden when mature. As if 
this were not confusing enough, Leptoloma 
cognatum, a representative of a genus that is 
indeed closely related to Panicum, not only 
occurs with the above two plant species in the 
same sandy habitats, but also has low, wide 
panicles that redden when mature. Unsure 
whether we may have overlooked compo- 
nents of this confusing plant association, we 
have returned to several communities where 



areolata occurs and have found E. spectabilis 
growing in the absence of the other two 
grasses. Furthermore, we have collected this 
species in the Nebraska Sand Hills on E. 
spectabilis, northwest of the range of Lep- 
toloma cognatum. Osborn (1928) recorded 
areolata from Eragrostis pectinacea in North 
Carolina. Because E. pectinacea is an annual, 
this report requires confirmation. 

Remarks and diagnosis. — The habitus of 
areolata is unique. 

XIV. The prairiana Group 

The prairiana group is named for its 
most plesiomorphic member, F. prairiana, 
a widespread resident of tall-grass prairie 
and southwestern semiarid grasslands. The 
group consists of 10 species that share a simi- 
lar general facies. All members of the group 
are medium-sized and brown but lack distinc- 
tive markings. Definitive identification re- 
quires examination of genitalia; the aedeagus 
and connective are fused, a condition we de- 
fine as synapomorphic. In prairiana a trace of 
the plesiomorphic articulation remains and 
constitutes an autapomorphy defining that 
species. Species of this complex are largely 



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inhabitants of prairie, savanna, or forest 
glades. Members of the group speciahze 
largely on andropogonoid grasses {Andro- 
pogon, Schizachyriiwi, and Botliriochloa 
spp.). Exceptions are (1) atlantica, which is 
often associated with switchgrass {Panicwn 
virgatU7n) or, in the Southwest, chloridoid 
grasses such as Sporoboliis spp.; and (2) 
producta, a southern coastal species that ap- 
pears to be associated with chloridoid grasses. 
Both atlantica and producta regularly colo- 
nize an introduced chloridoid grass, Cynodon 
dactylon. 

There are two sets of closely related sister 
species. One is sandersi-delongi. Ross and 
Cooley (1969) separated delongi from sandersi 
on the basis of the greater divergence of its 
apical, paired processes and lesser appression 
of the unpaired process to the aedeagal shaft 
(Figs. 44B,C). Hamilton and Ross separated 
satilla from clayi on the basis of a narrower 
aedeagal shaft (Figs. 44H,J). The distinctions 
between the species of these two sets are fine; 
some workers would consider the differences 
to represent infraspecific geographic varia- 
tion. However, we have retained satilla and 
delongi because, in each case, they can be 
unambiguously identified, and a geographic 
and historical basis can be presented to ex- 
plain both their divergence and maintenance 
of reproductive isolation. In the case of de- 
longi, which probably specializes on Schiz- 
achyriiim scoparium, the geographic range 
(Fig. 50) was almost entirely glaciated during 
the most recent glacial maximum. In contrast, 
sandersi occupies a region that was never 
glaciated, where it apparently specializes 
largely on Andropogon virginicus. These cir- 
cumstances provide a hypothetical scenario 
for speciation and an explanation of ecological 
factors that may now enforce isolation be- 
tween the two species. Similarly, satilla, de- 
scribed by Hamilton and Ross (1975) from 
south Georgia, is probably a Gulf Coast spe- 
cies (Fig. 51) that is at least partially sympatric 
with producta. Discovery of its sister species, 
clayi, as far north as Maine suggests that dif- 
ferentiation of these species, hke that of 
sandersi and delongi, may be a postglaciation 
event that is phenologically enforced by con- 
temporary climate. As closely related as clayi 
and satilla are, the sister set that they com- 
prise is also very closely related to graminea. 
A more distant pair of sister species is that of 



prairiana and reflexa. The most significant 
apomorphy shared by these sister species is 
the dorsal position of the gonopore (Figs. 
44D,G). These species also share a common 
general facies and cannot be distinguished on 
the basis of external characters. Each of these 
sister species has a variable aedeagal morphol- 
ogy, and we found that careful study was re- 
(juired to separate them. Both species are 
residents of tall-grass prairie, where they oc- 
cur on Andropogon and Schizachyrium spe- 
cies. Other species of the prairiana group are 
more distantly related. 

Description of prairiana Group 

Medium-sized. Length of 6 3.0-4.2 mm, 
of 9 3.0-4.6 mm. All species with similar 
general facies. Brown, or in darkest speci- 
mens, black on face or venter. Vertex moder- 
ately produced. Crown often stramineous 
with conspicuous transverse line at mid- 
length. Male plates elongate, in some species 
notched apically. Aedeagus and connective 
fused, or with at most (in prairiana) a faint 
trace of the plesiomorphic articulation. Ae- 
deagus with paired, apical, serrate processes 
and an unpaired, lateral or dorsal process par- 
tially fused to shaft in some species, or in 
others entirely fused or absent. Aedeagus 
asymmetrical in all but one species (dakota). 
Gonopore in primitive species oval and an- 
teapical, in more advanced species located on 
the unpaired process or, in the most highly 
specialized species, replacing the unpaired 
process as a spiral groove originating dorsally 
on the shaft. Female sternum VII produced 
posteriorly at middle in a convex, apically 
notched protuberance. 

Key to Males of the prairiana Group 
1. Aedeagus with 3 processes on shaft 2 

— Aedeagus with 2 processes on shaft 6 

2(1). Pygofer with posterior lobe strongly expanded 

on ventral margin (Fig. 5D) . . . producta (Walker) 

— Pygofer with ventral margin of posterior lobe 
not strongly expanded 3 

3(2). Gonopore dorsal, anteapical, located on a pro- 
cess or at the base of one (Figs. 44D,G) 4 

— Gonopore on dorsolateral surface of shaft at base 

of retrorse process 5 

4(3). Gonopore near apex of aedeagal shaft, at base of 

a process (Fig. 44G) prairiana DeLong 

— Gonopore located on a process (Fig. 44D) .... 
reflexa (Osborn & Ball) 



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Fig. 44. Aedeagal apices of the Flexamia prairiana group: A, atlantica- B, sandersi- C, delongi- D, reflexa; E, 
producta; F, graminea; G, prairiana; H, c/flyi, I, dakota; J, satilla. Aspects: AD, anterodorsal; CD, caudodorsal; CE, 
cephalic; CV, caudoventral; DS, dorsal; LT, lateral; VN, ventral. Redrawn from Young and Beirne (1958) and Hamilton 
and Ross (1975). 



5(3). Apical processes of aedeagus more divergent, 
each with 6-9 teeth; when viewed in a position 
with apical processes horizontal, a space is visi- 
ble between the base of the gonopore extension 
and the shaft (Fig. 44C) . . . delongi Ross & Cooley 

Apical process of aedeagus less divergent, each 
with 3-5 teeth; when viewed as above, no space 
is visible between the base of the gonopore ex- 
tension and the shaft (Fig. 44B) 

saiidersi (Osborn) 

6(1). Gonopore spiral 7 

— Gonopore not spiral 8 



7(6). Aedeagal appendages of aedeagus without 
prominent, anteapical processes; aedeagal shaft 
usually not strongly expanded at base of 
gonopore (Fig. 44F) graminea (DeLong) 

— Apodemal appendages of aedeagus each with 
prominent, anteapical protuberance directed 
mesad; aedeagal shiift strongly expanded at base 

of gonopore (Fig. 44H) 9 

8(6). Aedeagal shaft with apical process on each side; 
gonopore anteapical on anterodorsal surface 
(Fig. 441) dakota Young & Beirne 

— Aedeagal shaft with apical processes arising on 



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one side; gonopore anteapical on lateral surface 
of'shaft (Fig. 44A) atlantica DeLong 

9(7). Shaft of aedeagus comparatively robust (Fig. 

44H); north Georgia to Maine 

claiji Young & Beirne 

— Shaft of aedeagus slender (Fig. 44J); Gulf Coast 

(Florida, south Georgia, Mississippi) 

satilla Hamilton & Ross 

Key to Females of the prairiana Group 

1. Ovipositor with base of each first valviila re- 
curved dorsally in dorsal aspect 2 

— Ovipositor with base of each first valvula not 
recurved dorsally 5 

2(1). Recurved portion of each first valvula parallel to 
anteriormost border of each valvula in dorsal 
aspect, appearing as rim above the valvula (Fig. 
63LL) 3 

— Recurved portion appearing wider in dorsal as- 
pect prairiana DeLong 

3(2). Anteriormost border of each first valvula ap- 
pearing somewhat angular in dorsal aspect, re- 
curved portion very narrow and not produced 
dorsad (Fig. 63LL) graminea DeLong 

— Anteriormost border of each first valvula 
rounded, recurved portion much broader and 
produced dorsad (Fig. 63GG) 4 

4(3). Range north Georgia to Maine 

clayi Young & Beirne 

— Range Gulf Coast satilla Hamilton & Ross 

5(1). Bases of first valvulae obliquely truncate, valvu- 
lae apparently membranous basally (Fig. 63HH) 
reflexa (Osborn & Ball) 

— Bases of first valvulae convex basally 6 

6(5). First valvulae broadest near bases in dorsal as- 
pect (Fig. 63EE,KK) 7 

— First valvulae broadest more posteriorly in dor- 
sal aspect 8 

7(6). Face often with brown interocular line contrast- 
ing with pale color of remainder; if without line, 
then irregularly darkened; Southeast (Fig. 49) 
producta (Walker) 

— Face tan to dark brown with alternating sinuate, 

dark and light lines; dry prairie (Fig. 49) 

dakota Young & Beirne 

8(6). Face with black interocular line, or at least with 
this area black in addition to other black marks 
on face atlantica DeLong 

— Face brown or tan; interocular area never black 9 

9(8). Range southern Illinois southeastward 

sandersi (Osborn) 

— Range northern Illinois northward to Ontario, 

northwest to North Dakota 

delongi Ross & Cooley 

35. Flexamia prairiana DeLong 
Flexamia prairiana DeLong 1937: 32. 

Important CHARACTERS. — Length of 6 3.6 



(3.2-3.9) mm, of 9 3.9 (3.4-4.6). Crown 
moderately produced (0.60 x head width, L40 
X interocular width) (d n 73; 9 n 32). 
Face (Fig. 3D) usually pale in lower half, with 
broad, black interocular band; in darkly 
marked specimens almost completely black 
(Fig. 3C). Habitus characteristic of group. 
Male plates (Fig. 45A) elongate, unnotched. 
[6] Pygofer (Fig. 62W) with upper part of 
hind margin produced posteriorly. Aedeagus 
(Figs. 6F, 44G) asymmetrical, shaft slender, 
elongate, gradually broadened apically; gono- 
pore oval, anteapical on anterodorsal surface. 
Three retrorse apical processes diverging 
slightly from shaft, about one-third length of 
shaft. Unpaired process arising basad of 
gonopore, margins entire with dorsal, longi- 
tudinal groove on apical half; pair of distad, 
ventral, lateral processes, serrate on ventral 
margin. [ 9 ] Sternum VII as for group; ovipos- 
itor with each first valvula (Fig. 63 JJ) recurved 
slightly dorsad at base. 

Geographic distribution. — Widely dis- 
tributed (Fig. 46) from Manitoba to Colorado, 
Arizona, and Mexico, east to Missouri and 
Illinois. Two females from Michoacan, Mex- 
ico (21 October 1981, M. W. Nielson), have 
been tentatively referred to prairiana. A sin- 
gle male (Baker 2361, USNM) is recorded 
from Auburn, Alabama, 10 September 1897. 

Biology. — Southwestern populations usu- 
ally occur on Bothriochloa species. In the 
tall-grass prairie the usual hosts are Schiz- 
achyrium scoparium and Andropogon ger- 
ardii. Most reported hosts are andropogonoid 
grasses, but in tall-grass prairie prairiana may 
move short distances to more suitable hosts in 
drought periods or in late summer. Thus, 
oligophagy coefficients calculated for prairi- 
ana tend to be among the lowest for all Flex- 
amia species (Whitcomb et al. 1988). The 
green coloration of live specimens of prairi- 
ana fades to orange upon drying. In midsum- 
mer the intensity of coloration varies greatly. 
In the fall, coloration may be very dark; un- 
cleared, late-season specimens have often 
been misidentified. 

Oligophagy coefficients. — Tall-grass and 
mixed-grass prairies north of Texas: Grami- 
neae 0.922; Panicoideae 0.730; Schiza- 
chyrium scoparium 0.438; Andropogon ger- 
ardii0.1A\{n -527). 

Remarks and Diagnosis. — Because this 
species is one of the most abundant of Flex- 
amia species, it is important to recognize it 



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Fig. 45. Male plates and pygofer.s of the Flexamia imiiriana group, ventral aspect: A, prairiana; B, reflexa; C, 
atlantica; D, producta; E, dakota- F, sandersi; G, delun^i; H, graminea; I, clayi. 



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Fig. 46. Geographic distribution of Flexamia prairiana. 



(tentatively) without dissection. The prairi- 
ana face type is almost always distinguishable 
from the faces ofgraminea, sandersi, delongi, 
or dakota, which may occur with it in Andro- 
pogon prairies. Dissected male specimens re- 
veal an aedeagus that might be confused only 
with that of reflexa. Female specimens may 
prove difficult to identify because the bases of 
the first valvulae vary geographically. In 
southwestern populations this structure is 
rarely sclerotized, as illustrated by Young and 
Beirne (1958). 

36. Flexamia reflexa (Osborn & Ball) 

Deltocephalus reflexus Osborn & Ball 1897: 203. 
Deltocephalus (Flexamia) reflexus, DeLong 1926: 28. 
Flexamius reflexus, DeLong and Caldwell 1937: 27. 

Important CHARACTERS. — Length of S 3.5 



(3.1-4.0) mm, of 9 3.7 (3.2-4.2) mm. Crown 
moderately produced (1.49 x interocular 
width; 0.68 X head width) (c? n = 20; 9 n = 
15). Face variable, often pale on lower half 
with broad interocular band; band may be 
obscured in dark specimens. [6] Pygofer 
(Fig. 62X) with posterior lobe strongly pro- 
duced, truncate apically. Plates (Fig. 45B) 
elongate, without notches or lateral projec- 
tions. Connective in lateral aspect with dorsal 
keels extending almost as far dorsad as aedea- 
gal apodeme. Aedeagus (Fig. 44D) asymmet- 
rical, shaft elongate, gradually tapered; gono- 
pore anteapical on anterodorsal surface, asso- 
ciated with short, apically grooved, unpaired 
process. Paired apical processes serrate along 
ventral edge, extending basad less than half 
length of shaft in lateral aspect. [ 9 ] Ovipositor 



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Fig. 47. Geographic distribution of Flexamia reflexa and a host, Andropogon gerardii. 



with first valvulae membranous basally, not 
curved dorsad; sclerotized portion obliquely 
truncate in dorsal aspect. 

Geographic distribution. — Tall-grass 
prairie (Cwikla and Blocker 1981), savanna, 
and glades and prairie mosaic of deciduous 
forest, Kansas and Iowa to Texas, Ontario, 
Maryland, and Florida (Fig. 47). 

Biology. — This species is associated with 
Andropogon gerardii in the East (Maryland 
and West Virginia) and may be associated with 
that host in the tall-grass prairie. However, in 
Texas it occurs in mixed-prairie associations in 
which big bluestem is a minor component. 

Diagnosis and remarks. — Given the pau- 
city of specimens of reflexa in collections, it 
is remarkable that it was the second member 
of the prairiana group to be described and 
that it is the type species of the genus. Almost 
all uncleared specimens "identified" as this 
species in museum drawers turn out to be 
other prairiana group species. The abundant 
prairiana, which occurs through much of its 
range on httle bluestem {Schizachyrium sco- 
parium), is much more apt to be encountered 
in the prairie. Since we have found no reliable 
external characters for sorting reflexa from 



prairiana, a search for this species involves 
sifting through numerous prairiana speci- 
mens. The position of the gonopore (some 
distance from the aedeagal apex, situated 
basally on a short, appressed, unpaired pro- 
cess) contrasts sharply with the simpler, oval, 
anteapical gonopore oi prairiana . Some spec- 
imens of atlantica and producta may also re- 
semble reflexa in general facies, but their gen- 
italia bear little resemblance to those of 
reflexa. 

37. Flexamia atlantica (DeLong) 

Deltocephahis (Flexamia) atlanticus DeLong 1926: 29. 
Deltocephalus atlanticus, DeLong and Caldwell 1937; 27. 

Important characters. — Length of (5 3.7 
(3.2-4.2) mm, of 9 4.1 (3.6-4.5) mm. Crown 
moderately produced (1.31 x interocular 
width; 0.59 X head width) (d n 20; 9 n = 
20). Face black in upper half, often forming 
interocular band contrasting with pale lower 
half. [6] Pygofer (Fig. 62Y) with posterior 
lobe narrow, without ventral lobe. Plates 
(Fig. 45C) elongate, notched apically. Aedea- 
gus (Fig. 44A) asymmetrical, shaft nearly uni- 
form in width, gonopore simple, anteapical on 



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Fig. 48. Geographic distribution of Flexamia atlantica and a host, Panicum virgatum. 



the right side. Pair of elongate, retrorse, ser- 
rate, apical processes extending basad approx- 
imately half length of shaft. [9] Sternum VII 
as for group; ovipositor with base of each first 
valvula (Fig. 63DD) not curved dorsad. 

Geographic distribution. — Prairie from 
Manitoba, North Dakota, Wyoming, and New 
Mexico east to Illinois (Fig. 48). Also along the 
Atlantic coastal prairie of the Northeast, and 
very patchily in the Southeast. 

Biology. — In tall-grass prairie atlantica is 
usually associated with switchgrass {Panicum 
virgatum). This grass, a dominant in the 
northeastern coastal prairie, probably ac- 
counts for the occurrence of atlantica there. 
In mixed-prairie and southwestern grass- 
lands, atlantica occurs on native Sporobolus 
spp. and is the major Flexamia colonist in 
the extensive stands of two exotic chloridoid 
grasses (weeping love grass [Eragrostis cur- 
vula ] and bermudagrass [Cynodon dactijlon ]) 
in Texas and Oklahoma. 

Oligophagy coefficients. — Texas and 
Oklahoma: Gramineae 0.994; Chloridoideae 
1.000; Eragrostis curvula 0.243; Cynodon 
dactylon 0.633 (n = 161). Kansas, Missouri, 
and Nebraska: Gramineae 1.000; Panicoideae 



0.876; Panicum virgatum 0.783; Sporobolus 
spp. 0.124 (n = 177). 

Diagnosis and remarks. — Flexamia at- 
lantica is the only pale-faced Flexamia prairie 
species with apically notched plates. Also, 
individuals of atlantica tend to be larger than 
those of reflexa or prairiana. In the South- 
east, atlantica may occur with producta, 
which tends to be smaller, and which usually 
has a much more produced crown. Although 
these characters permit tentative recognition, 
definitive recognition is readily accomplished 
by examining the aedeagal morphology, 
which is unique. 

38. Flexamia producta (Walker) 

lassus productus Walker 1851: 891. 
Deltocephalus visendus Crumb 1915: 189. 
Deltocephahis (Flexamia) productus, DeLong 1926: 43. 
Flexamia producta, DeLong and Knull 1945: 35. 

Important CHARACTERS. — Length of c? 3.2 
(3.0-3.5) mm, of 9 3.5 (3.2-3.9) mm. Crown 
usually conspicuously produced (1.62 x inter- 
ocular width; 0.75 x head width) (cJ n = 20; 9 
n = 20). Face variable but usually with clearly 
delimited interocular band; band occasionally 
obscured in darker specimens. [6] Plates 



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Fig. 49. Geographic distribution of Flexamia dakota {•) and producta (■) and the host of dakota, Scizachyrium 
scoparium, which does not occur on the Atlantic coastal plain. 



(Fig. 45D) elongate, notched at apex. Pygofer 
(Fig. 62AA) with posterior lobe produced and 
rounded apically, ventral portion extending 
mesad as pronounced lobe, serrate along mar- 
gin, the two lobes often overlapping in ventral 
aspect (but not necessarily in prepared speci- 
mens). Aedeagus (Fig. 44E) asymmetrical api- 
cally; shaft elongate, slender, slightly swollen 
anteapically; gonopore irregularly oval, on left 
side of dorsal surface of shaft; pair of processes 
arising asymmetrically at apex of shaft, ex- 
tending basad almost half of shaft length, each 
process laterally compressed and conspicu- 
ously toothed along ventral edge; with short, 
small, curved process arising near proximal 
portion of gonopore, extending sinistrad. [ 9 ] 
Female sternum VII as for group, ovipositor 
with bases of first valvulae (Fig. 63EE) not 
curved dorsad, broad basally in dorsal aspect. 

Geographic distribution.— Gulf Coast 
and Atlantic seaboard from Mississippi to 
Maryland (Fig. 49). 

Biology. — This species has been collected 
commonly in coastal prairies, but there are 
few host records. We have recorded it from 
several exotic chloridoid grasses (e.g., Zoysia 



japonica and Cynodon dactylon) in Maryland 
but have little field experience in the coastal 
prairies where it is most abundant. We note 
that Sporobolus virginicus is a common domi- 
nant in southeastern coastal prairies; this chlo- 
ridoid grass should be examined as a possible 
natural host. On the other hand, there are 
indications that cicadellid host specialization 
decreases in mesic and wet prairies (Whit- 
comb et al. 1987, 1988). Given its occurrence 
in wet southeastern grasslands, producta may 
turn out to be a habitat specialist. 

Diagnosis and remarks,— The produced 
crown of this species affords a tentative indica- 
tion of its identity. Unfortunately, sotilla and 
sandersi, which occur with producta, also 
have produced crowns that approach the 
length/width ratios oi producta. Although the 
latter species are similar to producta in most 
aspects, their faces are without conspicuous 
interocular bands. It is therefore possible to 
sort, from an assemblage of southeastern Flex- 
amia, a large subset of producta specimens 
that have either an extremely produced crown 
or a distinct interocular band (or both). Unfor- 
tunately, this will leave a mixture of atypical 



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producta specimens and specimens of 
sandersi and satilla, or perhaps even clmji. 
The aedeagal characters of producta readily 
diagnose the species. 

39. Flexamia dakota Young & Beirne 
Flexamia dakota Young & Beirne 19.58: 44. 

Important CHARACTERS. — Length of 6 3.0 
(2.8-3.6) mm, of 9 3.3 (3.0-3.6) mm. Crown 
moderately produced (1.40 x interocular 
width; 0.64 X head width) {6 n = 21; 9 n - 
18). Face brown to tan, with parallel lines. [ 6 ] 
Pygofer (Fig. 62BB) with upper portion of 
posterior lobe produced posteriorly. Plates 
(Fig. 45E) with apical third narrowed. Aedea- 
gus (Fig. 441) symmetrical; shaft elongate, 
slender, narrower near midlength than at 
base or apex. Gonopore anteapical on 
anterodorsal surface; pair of recurved pro- 
cesses extending basad amost half length of 
shaft; each process serrate on ventral margin. 
[ 9 ] Sternum VII as for group. Ovipositor with 
bases of first valvulae broadest near base (Fig. 
63KK). 

Geographic distribution. — Semiarid 
Great Plains of western North and South Da- 
kota, eastern Wyoming, Nebraska Sand Hills 
to Oklahoma, and the Edwards Plateau of 
Texas (Fig. 49). A presumably disjunct popu- 
lation occurs in the Loess Hills of western 
Iowa. This species has recently been recorded 
from Mexico (2 6 , 42 km S Piedras Negras, 
Coahuila, 13 October 1987, A. L. Hicks and 
J. E. Lowry). 

Biology. — This species is apparently a spe- 
cialist on little bluestem (Schizachyrium sco- 
parium) in dry prairie. 

Oligophagy coefficient, — Gramineae 
1.000; Panicoideae 0.990; Schizachyrium sco - 
pariwm 0.990 (n - 209). 

Diagnosis and remarks. — This species of- 
ten occurs in the absence of other specialists of 
andropogonoid grasses. However, in the Ed- 
wards Plateau of Texas, dakota occurs with 
graminea (which it closely resembles in gen- 
eral facies), prairiana, and, rarely, reflexa. 
Although it can be tentatively sorted from 
the latter two species on the basis of face 
pattern, we have found no reliable external 
character to sort it from graminea. The sym- 
metrical aedeagus of dakota, which has only 
paired apical processes, is unique. Young and 
Beirne (1958) described only males; Lowry 
and Blocker (1987) described the female. 



40. Flexamia sandersi (Osborn) 

Dritoci'pftalus sandersi Oshorn 1907: 164. 
Deltocephaliis (Flexamia) sandersi, DeLong 1926: 27. 
Flexamia sandersi. DeLong and Sleesman 1929: 83. 
Flexamius bidentata DeLong 1935: 155. 

Important characters. — Length of d 3.2 
(2.9-3.5) mm, of 9 3.4 (3.2-3.9) mm. Crown 
produced (1.48 x interocular width; 0.69 x 
head width) (d n - 11; 9 n - 10). Face (Fig. 
3H) varying from dark above and shading to 
paler apically to brown or black throughout, in 
former case with dark area shading gradually 
along lower margin; no interocular band, [c?] 
Pygofer (Fig. 62CC) truncate on dorsal por- 
tion of posterior margin. Plates (Fig. 45F) 
elongate, apically notched, in some speci- 
mens bidentate. Aedeagus (Fig. 44B) asym- 
metrical; shaft elongate, gradually tapered; 
gonopore on dorsolateral surface of shaft at 
base of retrorse, unpaired process, which is 
closely appressed to the shaft, has one 
minutely serrate margin, and extends basad to 
midlength of shaft; pair of apical, retrorse pro- 
cesses, dorsal slightly longer than ventral, 
each acute apically and with distinctly serrate 
margin. [ 9 ] Sternum VII as for group; ovipos- 
itor with base of each first valvula (Fig. 63FF) 
not curved dorsad. 

Geographic distribution. — Eastern Kan- 
sas to Louisiana, east to southern Illinois, 
Massachusetts, and South Carolina (Fig. 50). 

Biology. — The principal host appears to be 
broomsedge (Andropogon virginicus), but 
this conclusion requires further study. 

Diagnosis and remarks. — Throughout 
much of its range, sandersi occurs with clayi. 
These species usually have been collected 
in the summer before their host grasses 
have flowered. Many host records are listed 
simply as "Andropogon sp. ' In the Maryland 
Piedmont sandersi occurs abundantly on 
broomsedge in the absence of clayi; in Mary- 
land we have found the latter species only in 
the Appalachians. We have found no reliable 
external character that separates sandersi 
from clayi, satilla, graminea, or delongi. 
However, the characters of the aedeagus de- 
scribed above and in the key readily separate 
sandersi from all other prairiana group spe- 
cies, with the exception of delongi. We have 
discussed the relationship of these very 
closely related sister species in the corre- 
sponding section under delongi. 



284 



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No. 12 




Fig. 50. Geographic distribution of Flexamia sandersi (I 
Andropogon virginicus. 



and delongi (•) and the presumed major host oi sandersi. 



41. Flexamia delongi Ross & Cooley 

Flexamia delongi Ross & Cooley 1969; 246. 

Flexamia sandersi. Young and Beirne 1958: 164. (In part.) 

Important CHARACTERS. — Length of d 3.7 
(3.4-4.1) mm, of 9 3.8 (3.4-4.4) mm. Crown 
variably produced (1.34 x interocular width; 
0.65 X head width) (c? n = 17; 9 n = 21). Face 
varying from dark above, shading to paler api- 
cally to black or brown throughout, dark area 
often fading gradually along lower margin, 
interocular band absent. [6] Pygofer and 
plates as in sandersi. Aedeagus (Fig. 44C) 
asymmetrical; shaft elongate, not cylindrical, 
gradually tapered; gonopore on dorsolateral 
surface at base of retrorse process, which ex- 
tends basad to midlength of shaft; pair of lat- 
eral, apical, retrorse processes extending 
more than one-third but less than one-half 
length of shaft, each with a distinct, coarsely 
serrate margin and with very acute apex. [ 9 ] 
Sternum VII as for group; ovipositor with base 
of each first valvula as for sandersi. 

Geographic distribution. — Northern In- 
diana and Illinois, Wisconsin, and Ontario, 
west to South Dakota, Saskatchewan, and 
Manitoba (Fig. 50). 



Biology. — This species is presumably a 
specialist of little bluestem, Schizachyrium 
scoparium. 

Diagnosis and remarks. — The pair of sis- 
ter species consisting of delongi and sandersi 
is (with the exception of satilla and clayi) the 
most closely related of all Flexamia sister 
pairs. Ross and Cooley (1969) acknowledged 
this closeness, pointing out that delongi pre- 
viously had been considered a variant of 
sandersi. Ross and Cooley examined material 
from the East but did not examine specimens 
from Canada or from the Dakotas. Thus, their 
interpretation of delongi as a geographic re- 
placement for sandersi made the identity of 
the northern prairie material a crucial test. 
We examined specimens from these regions 
from the USNM and CNC and found them to 
be referable to delongi. We retain herein the 
concept of delongi, because it can be de- 
fended on morphological, biological, and his- 
torical grounds. The species occurs in a region 
that was largely or completely glaciated dur- 
ing the Wisconsinan. The principal (or sole) 
host of sandersi {Andropogon virginicus) 
does not occur in this region. Thus, san- 
dersi and delongi appear to utilize different 



1988 



Whitcomb, Hicks: Flexamia Revision 



285 



hosts, whose phenological asynchrony may 
reinforce reproductive isolation. F'inally, al- 
though genitalic differences between delongi 
and sandersi are slight, they are consistent. 
The apical, aedeagal processes of delongi are 
more divergent than those of sandersi; each 
process has 6-9 teeth. Further, the gonopore 
extension is more divergent from the shaft 
than in sandersi. 

In the northern prairies delongi may occur 
on little bluestem with graminea, prairiana, 
or dakota, but its aedeagal structure readily 
separates it from these species. 

42. Flexamia graminea (DeLong) 

Deltocephalus (Flexamia) graniineus DeLong 1926; 30. 
Flexamius graminetts, DeLong and Caldwell 1937: 27. 

Important CHARACTERS. — Length of 6 3.6 
(3.3-4.0)mm.,of 9 3.7(3.4-4.1) mm. Crown 
moderately produced (1.40 x interocular 
width; 0.68 X head width) (c? n - 25; 9 n - 
20). Face variable, from entirely stramineous 
to stramineous with clypeus darkened, from 
tan to almost black with pale arcs; interocular 
band absent. [S] Pygofer (Fig. 62EE) with 
posterior lobe not well differentiated ven- 
trally, posterodorsal portion produced cau- 
dodorsad and truncate apically. Plates (Fig. 
45II) divergent at apex, often appearing to be 
notched. Aedeagus (Fig. 44F) asymmetrical; 
shaft elongate, slender, twisted in apical half; 
gonopore consisting of a spiral groove origi- 
nating at shaft midlength; pair of asymmetri- 
cal, retrorse, apical processes, with one 
minutely serrate edge. [ 9 ] Sternum VII as for 
group. Ovipositor with base of each first 
valvula (Fig. 63LL) angular in dorsal aspect, 
narrowly curved dorsad anterolaterally, re- 
curved portion extending only slightly dor- 
sad. 

Geographical distribution. — Tall-grass 
prairie of North Dakota, Kansas to Blackland 
Prairie, and south Texas east to Missouri and 
Illinois (Fig. 51). A specimen from the Ball 
collection is labeled "DC." If this is an abbre- 
viation for "District of Columbia," and if the 
hastily scrawled, penciled label is correct, 
then it would be necessary to explain a widely 
disjunct distribution for graminea. Recently, 
little bluestem and several unusual prairie 
forb species have been found in a band of 
savannalike habitat along the north bank of 
the Potomac River between Great Falls Park 
(Maryland) and the District of Columbia 



(D. Boone, personal communication). The 
possibility that graminea (and perhaps other 
prairie insects) may have colonized such a 
habitat cannot be completely discounted. 

Biology. — This species appears to be a spe- 
cialist of little bluestem (Schizachyrium sco- 
pariiwi), but in late summer it may take 
refuge on other hosts. 

Oligophagy coefficient. — Schizachyri- 
um scoparium 0.91. 

Remarks and diagnosis. — Throughout 
most of the tall-grass prairie, graminea occurs 
regularly with prairiana on little bluestem. It 
can be distinguished from prairiana by its 
consistent lack of an interocular band and by 
its notched plates. However, through por- 
tions of its range it may also occur with 
sandersi, dakota, or delongi, or in mixed- 
prairie habitats with atlantica or reflexa. The 
structure of its aedeagus separates it readily 
from these species, but it is most closely re- 
lated to clayi and satilla. For differential diag- 
nosis of these three species, see the corre- 
sponding section under clayi. 

43. Flexamia clayi Young & Beirne 

Flexamia clatji. Young and Beirne 1958: 64. 

Important characters. — Length of 6 3.4 
(3.3-4.0) mm, of 9 3.7 (3.1-3.8) mm. Crown 
variably produced (1.44 x interocular width; 
0.71 X head width) (c? n = 20; 9 n = 20). Face 
brown to black above, usually shading gradu- 
ally to paler below. [6] Pygofer (Fig. 62DD) 
with posterior lobe more distinct than in 
graminea. Plates (Fig. 451) divergent apically, 
outer margins widened at midlength. Aedea- 
gus (Fig. 44H) much as in graminea but with 
shaft usually much more expanded at base of 
gonopore. [9] Sternum VII as for group; 
ovipositor with base of each first valvula 
(Fig. 63GG) rounded, with recurved portion 
broader than in graminea. 

Geographic distribution. — Appalachian 
region from Maine to north Georgia, west to 
Illinois (Fig. 51). 

Biology. — Some records for clayi are for 
broomsedge {Andropogon virginicus); others 
are for little bluestem {Schizachyrium scopar- 
ium). Many are for "Andropogon." Since 
broomsedge does not occur in Maine, at least 
some of the records for little bluestem must be 
correct. Many records, including our own, are 



286 



Great Basin Naturalist Memoirs 



No. 12 




Fig. 51. Geographic distribution of Flexamia ^raminea (•), clayi (■), and satilla (D). 



inadequate to discriminate between these two 
andropogonoid grasses. 

Remarks and diagnosis. — We are unaware 
of any region in which graininea, clayi, or 
satilla occur together. As in the case of 
sandersi and delongi, they may represent vi- 
cariant populations that have become repro- 
ductively isolated and diverged morphologi- 
cally. The initial description of clayi from 
Kentucky, and subsequent collections of the 
species from Georgia and Tennessee, ob- 
scured the extent of the range o( clayi. Our 
collections of the species from Ohio and the 
Maryland and West Virginia Alleghenies, and 
identification of material from Maine and New 
Hampshire as clayi present a different pic- 
ture, portraying the species as a cool-temper- 
ate species of the eastern mountains. This 
contrasts with that o{ graminea, whose range 
is essentially the tall-grass prairie, and satilla, 
whose known range is the Gulf Coast (Florida 
and south Georgia to Mississippi). 

Members of this complex can be readily 
separated from other prairiana group species 
by aedeagal characters. The aedeagal shaft of 
clayi is much more expanded basad of the 
gonopore than the usual condition in grami- 



nea. However, some specimens oi graminea 
from south Texas have a basally expanded 
gonopore that could be confused with that of 
clayi. The most consistent character that sepa- 
rates clayi and graniinea is the condition of the 
anteapical lobes of the apodemal processes. In 
clayi the processes each have a prominent 
anteapical lobe, which is directed mesad. The 
shaft in satilla is much narrower than that of 
clayi. 

44. Flexamia satilla Hamilton & Ross 

Flexamia satilla Hamilton & Ross 1975: 606. 

Important CHARACTERS. — Length of c? 3.4 
(3.3-3.4) mm, of 9 3.6 mm. Crown produced 
(1.49 X interocular width; 0.74 x head width). 
Face variable, but without distinct interocular 
band; usually dark brown or nearly entirely 
black. [6] Genitalia much as in clayi, but with 
aedeagal shaft ( Fig. 44J) not as expanded at 
base of gonopore. [9] Sternum VII as for 
group; ovipositor with base of first valvula 
rounded, with recurved portion broad and 
extending dorsad. 

Geographical distribution. — The type 
locality in south Georgia is just north of 



1988 



Whitcomb, Hicks: Flexamia Revision 



287 



Okefenokee Swamp. We have found other 
specimens, collected by Beamer, from the 
Gulf Coast in Florida and Mississippi. 

Biology, — This species was collected on an 
andropogonoid grass, probably Andropogon 
virginicus (K. G. A. Hamilton, personal com- 
munication). 

Remarks and diagnosis. — This species is 
sympatric with producta in the Gulf Coast 
region, and two specimens had been filed in 
the KU collection in the producta drawer. 
These specimens were tentatively separated 
from producta by the face, which was black, 
without a conspicuous interocular band. 
Unfortunately, some specimens with equally 
dark faces turned out, when they were 
cleared, to be producta. It is therefore easy to 
understand how this species escaped recogni- 
tion for many years. 

The degree of genitalic difference between 
clayi and satilla falls well within that consid- 
ered by some workers to represent geo- 
graphic variation. However, we feel that 
satilla is subject to the same phenological con- 
straints as producta. The hypothesis that the 
graminea complex in the East is divided into 
Appalachian and Gulf Coast species seems 
reasonable, and, given our ability to distin- 
guish the species on (admittedly minor) mor- 
phological grounds, justifies retention of 
satilla. 

This species is safely separated from its 
sympatric cohort, producta, only by genitalic 
examination, although the latter species (un- 
like satilla) often has a distinct interocular 
band. For distinctions within the clayi com- 
plex, see the comparable section under clayi. 

Biology of Flexamia 

Flexainia species are residents of North 
American grasslands, where they specialize 
predominantly on dominant, warm-season 
grasses (Panicoideae and, especially, Chlori- 
doideae). Several species, however, appear to 
breed on sedges (Cyperaceae) or rushes (Jun- 
caceae). The genus is divided into 13 species 
groups; host choice is related to membership 
in these groups. 

No Flexamia species are known to be uni- 
voltine; as a result, they are inhabitants of 
mesic grasslands and are poorly adapted for 
semiarid grasslands. The habitats of the genus 



include true prairie, short-grass plains, 
semitropical grasslands, high desert plains or 
montane grasslands, riparian grasslands, and 
saline or nonsaline lakeshores. 

In semiarid grasslands, grass hosts that sup- 
port Flexainia species tend to be capable of 
growth with minimal precipitation, but are 
unlikely to undergo repeated episodes of dor- 
mancy through the summer. Perhaps this is 
why, despite their proclivity for chloridoid 
grasses, not a single Flexamia species special- 
izes on Hilaria or Sporoholus species. This is 
true despite the fact that some members of 
these grass genera are among the major domi- 
nants in southwestern grasslands and com- 
prise a significant fraction of the regional 
grassland biomass. [Note: As a control to this 
"natural experiment," Hilaria and Sporoholus 
species have acquired specific non-Flexamia 
cicadellid guilds of their own (Whitcomb et al. 
1987).] 

The relation of Flexamia species to their 
hosts ranges from habitat specialization (in 
primitive species), to various degrees of inter- 
and intrageneric oligophagy, to absolute mon- 
ophagy. Narrow oligophagy is the most com- 
mon strategy. Body size of Flexamia species 
appears to be correlated with size of the host 
plant. Intraspecific variation in body size may 
reflect host plant quality; individuals from 
populations at the range periphery are often 
small. 

Flexamia species respond to several aspects 
of grassland structure. Disturbances such as 
mowing or burning tend to reduce popula- 
tions or to extirpate them altogether; for this 
reason Flexamia is an excellent genus for as- 
sessing habitat quality of grassland reserves 
(Whitcomb 1987). On a local level, popula- 
tions of Flexamia species are almost always 
much higher in more mesic segregates of 
grasslands defined by minor topographic vari- 
ation than in adjacent xeric sites (Whitcomb et 
al. 1987). On a regional level the degree of 
fidelity to single hosts appears to be least in 
mesic prairie or, perhaps, in southeastern 
grasslands (Whitcomb et al. 1987). The struc- 
ture of grassland dominance hierarchies may 
determine host choice. 

Habitat patchiness has had a profound ef- 
fect on Flexamia species on both historical and 
contemporary time scales. For monophagous 
species, their hosts are evolutionary islands. 
Contemporary host patchiness may affect 



288 



Great Basin Naturalist Memoirs 



No. 12 



presence or absence of Flexamia species 
within their ranges. However, no Flexamia 
species is able to track its host throughout the 
entire range of the host. This not only is the 
result of patchy host occurrence at the range 
periphery, but is often reinforced by climate. 
In many instances host colonization appears to 
be blocked by climatic factors. For example, 
the Flexamia complex that colonizes side-oats 
grama {Bouteloua curiipendula) is divided 
into six species, none of which is known to 
overlap in range. The isolating mechanisms 
are those associated with the different climatic 
regimes of prairie, Chihuahuan and Sonoran 
desert, interdesert grasslands of the Gila 
Mountains of southeastern New Mexico, and 
the eastern and western Mexican grasslands. 
Buchloe dactyloides , Schizachijrium scopar- 
ium, and Miihlenbergia porteri are similarly 
partioned, as is Bouteloua gracilis (although 
less spectacularly so). The possible role of host 
biotype (in its inherent genetic composition), 
apart from the purely phenologically deter- 
mined availability of growing host, has not 
been determined. 

We have seen no evidence that Flexamia 
species are highly fecund; they are rarely cap- 
tured at lights or in aerial traps. There is no 
evidence of seasonal movement under condi- 
tions that permit observation of immigration 
of Athysanella macropters or Laevicephalus 
adults into suitable grassland. We therefore 
predict that the life history strategy of 
Flexamia species will be found to resemble 
K- rather than r-selection (MacArthur and 
Wilson 1967). 

Eggs of most species are presumably 
oviposited in host tissue. In temperate re- 
gions it is reasonable to assume that eggs laid 
in the field in autumn are in diapause. In 
central latitudes first-generation adults ap- 
pear in early June. In subtropical latitudes, 
such as south Florida and south Texas, collec- 
tion records from December, January, and 
February indicate that reproduction may oc- 
cur throughout the year. 

Species Concept in Flexamia 

The species concept that emerges from our 
treatment of Flexamia is a composite of indi- 
vidual decisions concerning species. We faced 
diflPicult decisions in several Flexamia species 
groups. 



In the prairiana group there are three 
sets of close sisters. In one of these sets 
{prairiana -reflexa), the types differ so sub- 
stantially that no immediate problem is evi- 
dent. When specimens from different geo- 
graphic regions were examined, however, 
problems emerged. The holotype of prairi- 
ana, from Illinois, represents the northeast- 
ern periphery of the range of this species (Fig. 
46). Females from the southwestern part of 
the range, especially those from the montane 
grasslands of the Chihuahuan Desert region 
or the Trans-Pecos shrub savanna, differ from 
northern individuals in certain features of the 
first valvulae; these populations colonize 
Bothriochloa spp. rather than little bluestem 
{Schizachyrium scoparium). However, we 
found no characters of the male genitalia that 
distinguish the Bothriochloa populations. 
Further, as discussed by Young and Beirne 
(1958), the male aedeagal characteristics of 
reflcxa and prairiana vary intraspecifically. It 
appears that both species, although wide- 
spread, are distributed patchily in some parts 
of their ranges. Under these circumstances it 
is reasonable to suspect the existence of repro- 
ductively isolated populations. For example, 
we would not be surprised if the Chihuahuan 
populations of prairiana proved to be repro- 
ductively isolated from the tall-grass prairie 
populations. 

Our decision to retain species concepts pro- 
posed by Ross and Cooley (1969) and Hamil- 
ton and Ross (1975) defining the sandersi-de- 
longi and satilla-clayi sister pairs has been 
discussed under the species descriptions of 
delongi and satilla. In each case we elected to 
retain the newly described species on the 
criterion of probable phenological isolation 
and consistency of (admittedly) minor mor- 
phological differences. 

The Anasazi form of arenicola presents a 
more difficult problem. This population, cen- 
tered in the Four Corners area of the South- 
west, is reproductively isolated from popula- 
tions oi arenicola in the Nebraska Sand Hills. 
Further, it has a distinctive morphological 
characteristic — the unpaired, aedeagal pro- 
cess is invariably broken, presumably in copu- 
lation. Although this character tends to define 
the Anasazi form, we ha\e found one speci- 
men oi arenicola from Nebraska with a broken 
aedeagus. Thus, despite our strong suspicion 



1988 



Whitcomb, HickS; Flexamia Revision 



289 



that a certain amount of morphological diver- 
gence has occurred between the two popula- 
tions, we have found no consistent character 
to separate the two populations. Because we 
feel that a taxonomic designation is inappro- 
priate, we refer to the Four Corners popula- 
tion simply as the Anasazi form o{ arenicola . 

The celata-stylata and beaincri-texana sis- 
ter pairs also merit brief mention. The distinc- 
tive process of the male pygofer that occurs in 
both members of the celata-stylata set sug- 
gests that the divergence is relatively recent. 
Because the area of present-day occurrence of 
celata was largely created by the most recent 
glacial maximum (Wright 1970), we at first 
thought that this speciation event was Holo- 
cene in origin, as the minor degree of morpho- 
logic divergence would suggest. The geo- 
graphic proximity of the ranges of the two 
species seemed to suggest that speciation oc- 
curred by dispersal of stylata or a similar an- 
cestor into sandhill grasslands that emerged 
postglaciation. However, the morphological 
evidence contradicts this tidy explanation. 
The aedeagal symmetry o( celata and the pres- 
ence of presumably plesiomorphic, tapered, 
notched plates similar to those of arenicola or 
decora demand that celata be considered the 
most closely related to the ancestral lineage. 
Perhaps during various glacial episodes sandy 
grasslands were much more common than 
they are today. The existence of three other 
uncommon sand-inhabiting Flexamia species 
{grammica, arenicola, and areolata), in com- 
bination with other studies (Whitcomb et al. 
1986, 1987, 1988) indicating that only domi- 
nant grasses accumulate specialists, supports 
this hypothesis. 

The divergence of texana and beameri is 
also unclear. This murkiness is perhaps an 
artifact of the rarity of the species; each is 
known from single collections of E. D. Ball 
and R. H. Beamer, respectively. We there- 
fore know little of the ecology of these species. 
Each of these species occurs just outside the 
periphery of the known range of inflata, in 
one case in a cold region in New York, in the 
other a hot region of south central Texas. The 
propensity of inflata to form ephemeral, iso- 
lated colonies on a variety of hosts (Whitcomb 
et al. 1986) ought, theoretically, to spawn pe- 
ripheral new species. Although beameri and 
texana appear to be closely related, the bio- 
geographic data suggest that an independent 



origin for the two species from the inflata 
lineage should not be completely discounted. 

Species problems in the prairiana and flex- 
iilosa groups, though interesting and chal- 
lenging, pale when compared to the pectinata 
group. This group, prior to this study, con- 
sisted of only two species. We now report 
expansion of the group to nine species. Three 
of the new species (mescalero, jacala, and 
collorum) differ profoundly from previously 
recognized species. The remaining four spe- 
cies {gila, bandarita, minima, and zamora) 
are closely related members of a complex that 
includes the previously recognized pectinata 
and doeringae. It is likely that all of the spe- 
cies are specialists of side-oats grama {Bou- 
teloiia curtipendula). This grass host is impor- 
tant (in many regions dominant) throughout 
most of the prairie, from North Dakota to 
Ohio to Kansas to north Texas to eastern New 
Mexico; throughout this region of importance 
the characters of the styles, aedeagal tips, and 
female sternum VII of pectinata are relatively 
constant. Similarly, populations of doeringae 
in southeastern Arizona (where the climate 
has a strong Sonoran influence) show an in- 
traspecifically constant profile in the morphol- 
ogy of the aedeagus, style, and female ster- 
num VII. We now report discovery of a 
species (bandarita) from the Chisos Moun- 
tains and Marathon Basin of Texas which, in 
some respects, is intermediate between pecti- 
nata and doeringae. Flexamia bandarita oc- 
curs in an area that is rich in endemism, as 
attested by our description herein ofzacate, a 
new Chihuahuan sister of the Sonoran can- 
yonensis. 

We were delighted to find this endemic 
until we examined Flexamia specimens taken 
from side-oats grama in the intervening re- 
gion between the Chisos Mountains and 
southeastern Arizona that could be referred 
to neither bandarita nor doeringae. Rather, 
they possessed a mosaic of characteristics rep- 
resentative not only of bandarita and doe- 
ringae, but of pectinata as well. With collec- 
tions from three widely separated localities in 
New Mexico and Chihuahua, Mexico, totaling 
eleven individuals, we were compelled to rec- 
ognize another new species (gila). 

Although we are uncomfortable separating 
a large complex occurring on a single host 
into a number of species, we would be even 
more uncomfortable illustrating the variation 



290 



Great Basin Naturalist Memoirs 



No. 12 



within a single hypothetical polymorphic 
"species" and attempting to explain why we 
included such divergent characters under a 
single name. In our view such a treatment 
would require redefinition of the species con- 
cept in deltocephaline leafhoppers. 

Once we recognized the existence of a spe- 
cies complex of side-oats grama specialists, it 
was easier for us to accept the results of our 
reexamination of mini7na DeLong & Hersh- 
berger and zatnora DeLong & Hershberger 
from Mexico. In these examinations we were 
assisted by the availability of eight previously 
unidentified specimens referable to minima 
from Monterrey, Mexico, a location consider- 
ably north of the type locality. These speci- 
mens and the type of minima reveal a mor- 
phologic profile similar in some respects to 
zamora and pectinata, but with substantial 
differences in the male plates and styles and 
the female sternum VII. Similarly, on the ba- 
sis of the female sternum VII and, to a lesser 
extent, the male plates and styles, zatnora can 
be separated from all other members of the 
complex. 

Construction of a map (Fig. 15) of the geo- 
graphical distribution of members of the com- 
plex clarifies the taxonomic status of the spe- 
cies. The distribution of the complex, like that 
of the host (side-oats grama), encompasses a 
large percentage of North American grassland 
area. However, the six species of the complex 
occur in very different vegetational regions, 
some of which are recognizable at the biome 
level. Furthermore, side-oats grama is rare in 
the Texas mesquite-acacia savanna, so there is 
an essential discontinuity in host distribution 
that probably inhibits gene flow between the 
United States and Mexican populations of 
side-oats grama specialists. Part of the barrier 
between pectinata and minima and zamora is, 
in fact, the area occupied by doeringae, han- 
darita, and gila. Therefore, the existence of 
reproductively isolated populations of Flex- 
amia specialists on side-oats grama in grass- 
lands of the United States and Mexico is prob- 
able, and the reinstatement of minima and 
zamora is justified. 

The concept of climatically limited special- 
ists that geographically partition a single grass 
host is supported to some extent by studies in 
Mexico (Triplehorn and Nault 1985) of Dal- 
bulus species, most of which appear to be 
Tripsacum specialists. Further support is 



given by the recent demonstration (Hicks et 
al. 1988) that in New Mexico blue grama 
(Bouteloua gracilis) is colonized by seven 
Athijsanella species, but that the distribution 
of each species is confined to regions that 
are readily interpretable in terms of recog- 
nized biogeographic regions (e.g., Colorado 
Plateau, high plains, short-grass prairie, 
Chihuahuan grasslands). We believe that 
Bouteloua curtipendula is also partitioned by 
its Flexamia specialists. 

In an ideal world we would have postponed 
all decisions concerning this complex and in- 
stead organized an expedition to the South- 
west and Mexico to collect Flexamia from 
side-oats grama. However, publication of the 
conclusions herein need not discourage such 
exploration; perhaps it will encourage it. 

In summary, we have defined Flexamia 
species as populations with consistent and 
readily demonstrable morphological differ- 
ences whose reproductive isolation from pos- 
sibly conspecific populations can be inferred 
on biogeographic grounds. 

Phylogeny of Flexamia 

Young and Beirne (1958) established the 
groundwork for a Flexamia phylogeny by their 
careful descriptions of genitalic and other 
characters of the genus. Also, although they 
did not propose a formal phylogeny, they dis- 
cussed species relationships at some length. 
With a minimum of assumptions, these pro- 
posals and relationships can be presented as a 
cladogram (Fig. 52). 

H. H. Ross was intrigued by Flexamia and 
in 1970 published a preliminary phylogeny of 
the genus in which the species were identified 
only by number and no apomorphies were 
described. Although Ross did not present 
an explicit proposal, his unpublished notes, 
which specify apomorphies and synapomor- 
phies, make it possible to diagram his phy- 
logeny also (Fig. 53). 

OuTGROUPS — Ross considered Spartopyge 
Young & Beirne and Aflexia Oman to repre- 
sent Flexamia outgroups. Young and Beirne 
(1958) felt that Aflexia was not especially 
closely related to Flexamia. Although the 
habitus of Alapus DeLong and Sleesman is 
very different (Beamer and Tuthill 1934) from 
that o{ Spartopyge (Fig. 54), its genitalic char- 
acters (Figs. 55F,G) suggest relationship with 
this genus. 



1988 



Whitcomb, Hicks. Flexamia Revision 



291 




•I s 







•S ■§ 



■§ i 



\ y 



ANCESTOR 



Fig. 52. Species relationships in Flexamia : proposal of Young and Beirne (1958). These authors proposed an ancestral 
condition in which the Flexamia aedeagus possessed two pairs of recurved, bilaterally symmetrical appendages at or 
near the apex of the aedeagal shaft. From this ancestral condition, the authors proposed three or four independent lines 
of development, without specifying a phylogenetic sequence in which the lines diverged. In one line of development 
(I), the apical pair of processes tended to fuse and the anteapical to remain separate {picta and pyrops). In a second line 
of development (II), the pairs of processes are (1) either preserved, as in surcula and curvata , or (2) the apical processes 
are reduced and the basal processes absent (abbreviata), or both processes are absent (3) but the location of the 
gonopore (in camjonensis) suggests a derivation from the condition in abbreviata. Young and Beirne suggested that 
pectinata and doeringae, lacking any trace of aedeagal processes, represent the culmination of one of the developmen- 
tal lines. We take the liberty of placing these two species (4) at the culmination of line II. In the final lineage the 
important event was (5) the development of an unpaired ventral aedeagal process. In IIIA this occurred either (6) with 
additional processes, as in serrata , or (7) with branching of more apical processes, as in slossonae and albida . In albida 
(8) the unpaired ventral process is adherent to the shaft. The final sublineage (IIIB) encompasses most species of the 
genus. In the more generalized species, the gonopore is small, not elongated, and located on the caudoventral surface 
of the aedeagus near the apex (9). Young and Beirne placed seven species {decora, shjlata, modica, ritana, arizonensis, 
flexulosa, and inflata) in this cluster. In prairiana a phyletic torsion of the shaft has occurred (10) so that the gonopore 
is anteapical on the anterodorsal surface of the shaft. This species has a pronounced groove on the unpaired process, 
which in imputans constitutes the gonopore (11). In reflexa the unpaired process is fused basally to the aedeagal shaft 
(12) but is free at its extremity; the gonopore is circular, but at some distance from the base of the process. In sandersi 
the gonopore has become elongate (13). In producta the movement of the gonopore, to the left side of the dorsal surface 
of the shaft, far removed from the apex, approaches a maximum (14). In graminea and clayi the unpaired process has 
almost completely disappeared and the gonopore is spiral (15). In grammica, dakota, and atlantica the anteapical 
processes have been lost; these species were tentatively placed in line IIIB. 



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No. 12 



o g 5 

2 S = 



E E C -S o. 



:2 I a E 



u 



X 



Fig. .53. Flexatnia phylogeny proposed by Ross (1970, and unpublished notes). The tree diagram was transposed to a 
cladogram. Identification of species and apomorphies used by Ross to construct the cladogram are taken from 
unpubhshed notes. The apomorphies and conserved plesiomorphies designated were (1) mesal brush on pygofer; (2) 
thin aedeagal shaft with basal processes; (3) lack of aedeagal processes; (4) width of aedeagus, thick (4) or thin (4'); (5) 
length of male style; (6) angulate pygofer; (7) four aedeagal processes; (8) aedeagus with mesal process; (9) acute aedeagal 
apex; (10) long, mesal aedeagal process; (11) unpaired aedeagal process bifurcate; (12) aedeagal processes twisted; (13) 
dorsal location of gonopore; (14) fusion of connective and aedeagus; basal location of median process and gonopore 
(somewhat = 15, more so = 16); widening (17) of median process to bean-shaped structure appressed to shaft; (18) loss 
of medial process. 



Future phylogenetic hypotheses concern- 
ing Flexamia should be broadened to include 
a much wider range of outgroups than those 
discussed above. Candidates for study should 
perhaps include Enantiocephalus Haupt, a 
Eurasian genus (Figs. 55A,B). K. G. A. 
Hamilton (personal communication) has sug- 
gested that Mocuellus Ribaut and related gen- 
era should not be overlooked as outgroups. 
Finally, we will soon add a new genus (in 
preparation), presumably also related to Flex- 
amia. This genus consists, so far as known, of a 
single species that specializes on Muhlenber- 
gia arenacea in saline flats in semiarid regions 
of New Mexico and Arizona. The former 
genus Acurhinus {-^ Donjdium) may be more 
distant from Flexamia than once thought (Lin- 
navuori 1959). Acurhinus macukitus (Osborn), 



known only from three females in the Ohio 
State University collection, is currently re- 
garded as a species of Hododoecus Jacoli, an 
essentially African genus of the Stirellini (Lin- 
navuori and DeLong 1978). It is evident that 
a phylogeny of deltocephaline genera, espe- 
cially those with linear connectives articu- 
lated with the aedeagus, would greatly en- 
hance all phylogenies of deltocephaline 
species. 

In this study we have chosen two Sparto- 
pijge species [miranda (Knull) and mexicana 
(DeLong & Hershberger)] and Alapus elon- 
gattis Beamer & Tuthill for outgroup compari- 
son. 

Phylogeny of Flexamia croups. — We 
have developed an intuitive phylogeny utiliz- 
ing many of the characters and concepts 



1988 



Whitcomb, Hicks; Flexamia Revision 



293 







Fig. 54. Spartopyge Young & Beirne, the principal Flexainia outgroup. Spartopyge mexicana (DeLong & Hersh- 
berger): A, habitus; B, face; C, male plates, ventral aspect; D, female sternum VII; E, S. miranda (KnuU), female 
sternum VII. 



developed by Young and Beirne (1958) and 
Ross (1970, unpublished notes). Details of our 
reconstruction follow: 

A. Designation of sister species. We first 
identified synapomorphies that define close 



sister species sets. We also assigned polarities 
within these sets by defining significant aut- 
apomorphies of one of the set members. 

The sets and defining synapomorphies [s] 
and autapomorphies [au] were: 



294 



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No. 12 










Fig. 55. Male genitalia of genera related to Flexamia. Enantiocephalus corntitus (Herrich-Schaffer): A, aedeagus and 
connective, lateral aspect; B, male pygofer, lateral aspect. Spartopyge mexicana (DeLong & Hershberger): C, 
aedeagus, lateral aspect; D, male pygofer, lateral aspect. Spartopyge miranda (Knull); E, male pygofer, lateral aspect. 
Alapus elongatus Beamer ix Tuthill: F, aedeagus and connective, lateral aspect; G, male pygofer, lateral aspect. 



surctila —> 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 



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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 



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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 



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Fig. .58. See facing page. 



discovery of many new species from the 
Southwest, that further acquisitions of Flex- 
amia or related genera from Mexico may 
change our perceptions dramatically. Also, a 
wider selection of outgroups would clarify the 
status of the early Flexamia lineages. 

Young and Beirne (1958) proposed that 
grooves oil the unpaired aedeagal process 
were vestiges of a plesiomorphic condition in 
which the halves of the processes were sepa- 
rate. This condition occurs in several species 
{youngi, imputans, prairiana), but we have 
been unable to link these species in any trans- 
formation that is consistent with other trans- 
formations. 

Finally, we propose, with some confidence, 
that the articulation between aedeagus and 
connective was subject to fusion or partial 



fusion in four nodes of our model. These were 
in the lineages leading to picta-pyrops, to are- 
olata, to slossonae, and to the prairiana 
group. Given the different products of each of 
these fusions, and the position on the tree in 
which they occurred, we have no problem in 
treating each as a synapomorphy. 

The genus Flexamia is rich in characters 
and should provide an excellent model for the 
PHYSYS program. It is likely that careful 
study and numerous iterations using such a 
program will further improve phylogenetic 
hypotheses concerning Flexamia. 

blogeographic implications of the 
Phylogenetic Model 

Ross (1965, 1970) felt that many speciation 



1988 Whitcomb.HicksFlexamia Revision 301 

Fig. 58. Phylogeny of species of groups MX. The following events, inelnding synaponiorphies [s] and aiitaponior- 
phies [au] are postulated; (1) In the plesiomorphic state of the pectinata-abhrcviata subtree, the male plates were broad 
(Fig. 11), the male pygofers constricted (Fig. 62F), the gonopore oval on the caudoventral surface, and the aedeagus 
(Figs. 12A-H) without apical processes; (2) in one subtree of the pectinata group (comprising eight of its nine species), 
the aedeagal tip remained unmodified but flared (Figs. 12A-H). In all three species of the gi/a division (3) of this group, 
the female sternum VII developed [s] long, median, paired teeth (Figs, 14D-F); in doeririf^ae (4) these teeth [au] 
became associated into a medial process (Fig. 14F). In gila (5) [au] the male plates are elongate (Fig. 1 1 F), in handtirita 
(6) [au] shortened (Fig. II D). Evolution of this cluster, oi' pectinata, and, perhapsalsoof muiima and zamora. occurred 
on Bouteloua curtipendnla. The three species of the minima cluster are defined [auj by (8) subrectangular plates 
{pectinata: Fig. IIC), (9) short plates {minima: Fig. IIA), and (10) outer teeth of hind margin of female sternum VII 
longer than median teeth {zamora: Fig. 14B). Divergence from this cluster (1 1) involved [s] development of avicephali- 
form style tips. In coUorum (12), which appears to be a specialist of Bouteloua uniflora, a close relative of B. 
curtipendula, [au] the plates (Fig. IIG) are extremely short. In jacala the plates (Fig. IIH) remained long and broad 
(13); this is the only Flexamia species with both long, broad plates and avicephaliform style tips. In another species 
cluster a tendency toward development of aedeagal processes developed. In mescalero (15) [au] the aedeagus (Fig. 22) 
is slightly capitate rather than flared. This specialist oi Muhlenhergia pauciflora retained relatively broad plates (Fig. 
Ill) and constricted pygofers (Fig. 62B) typical of the pectinata group; we therefore regard it as a monobasic subgroup 
of group I. A monophyletic cluster of three species is presumed to have arisen (16) from this lineage. Each of these 
species has [s] a pygofer with a conspicuously produced posterior lobe (Figs. 62H,I). In abbreviata (group II) the 
aedeagal apex [au] bears (Fig. 7H) two minute, paired processes (17). In this species and in zacate (18) the gonopore 
(Fig. 24) is elongate. F. zacate is the only species of the genus that lacks aedeagal processes but has an elongate 
gonopore. In canyonensis the gonopore [au] is located (19) approximately at the midpoint of the aedeagal shaft, 
presumably as a consolidation of the elongate gonopore type of zacate. The shape [au] of the aedeagal shaft in 
camjonensis (Fig. 6C) is also unique. The sister species zacate and canyonensis are both specialists of Muhlenber^ia 
porteri. In a very important divergence (20), development of two pairs of aedeagal processes is hypothesized. In 
grammica (22), a Calamovilfa specialist, the plesiomorphic habitus of the Spartopyge dorsum and face were retained. 
Two of the four hypothesized aedeagal processes may have been lost. The pygofers and male plates also suggest 
relationships with Spartopyge , but in contrast to this outgroup, aedeagal processes are present. The relationship of 
grammica to other Flexamia species with aedeagal processes is unclear, given the [au] specific asymmetry of these 
processes (Fig. 7E) and the location of the gonopore, which has moved to a dorsal position. It is possibly significant [s?] 
that the style apices of the grammica and curvata groups are foot-shaped (21). In the curvata set (23) of sister species 
{Buchloe specialists) [s] two pairs of processes are present. In curvata (25) [au] the gonopore is elongate. In the picta 
lineage {Aristida specialists) two pairs of processes are present but [s] the apical pair has been modified into flangelike, 
toothed structures (26). Several autapomorphies [au] characterize (28) pyrops (see text). From this subtree the 
remaining species of Flexamia were derived; this derivation was featured (29) by an apomorphic fusion of the anteapical 
pair of processes into a single, unpaired process located on the ventral surface of the aedeagus in the sagittal plane. 



events involving phytophagous insects could The progression of evolutionary divergence 
be explained by host transfer; the genus Flex- in Flexamia can best be described in terms of 
amia was one of the genera that he chose for the phylogeny we propose. In Figures 64-67 
the study of host choice and speciation in the we illustrate the postulated biogeographic 
Cicadellidae. Ironically, Ross happened to and host transfer events, 
work largely in central, northern, and south- We believe that the most closely related 
eastern grasslands. In these regions the study outgroup is Spartopijge. We have no informa- 
of cicadellid host relationships is more diffi- tion on the host relationships of 7ne.ricana, but 
cult than in the Southwest, and, as it turned miranda occurs on Bouteloua curtipendula in 
out, almost all of the undescribed species in Arizona. The general facies o{ Spartopyge ap- 
the genus occurred outside the geographic pears to be homologous with that of some 
regions in which he conducted his extensive Flexamia groups (albida, grammica, and ri- 
fieldwork. tana groups). We propose that diversification 
Today, in retrospect, we report that the of the Spartopyge lineage in Mexican grass- 
choice of Flexamia was a wise one; however, lands led to Alapus and to Flexamia. 
fewer speciations appear to be due to host We propose that the initial divergence (Fig. 
transfer than Ross might have predicted. Sis- 65) of the Flexamia lineage resulted from spe- 
ter species have in many cases proved to be cializationofageneralist ancestor on side-oats 
specialists of the same host or closely related grama, Bouteloua curtipendula. This lineage, 
hosts. Indeed, entire groups of species have influenced by host patchiness and/or regional 
proved to have similar (often congeneric) pat- climatic differences, diverged by dispersal or 
terns of host selection. vicariance into at least six species. These are 



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No. 12 






5 



VII 



VIII 



13 



14 
12 



18 



16 



19 



17 



15 



10 11 



IX 



21 



9 20 



22 



25 



27 



28 



XI 



26 



XII 



24 



23 



XIII 



Fig. 59. Phylogeny of species of groups VII-XI. The following events, including synapomorphies [s] and autapomor- 
phies [au] are hypothesized. Plesiomorphically, this cluster retained (1) certain features of the Sparlopyge habitus (i.e., 
dorsal stripes and a pale face with black interocular band). However, prior to divergence, two pairs of aedeagal 
processes had developed, and, subsequently, the anteapical pair had fused to form a single, unpaired ventral process 
(Figs. 7A-C, 31). The lineage diverged into the (2) albida (VU), (5) serrata (VIII), and (6) ritana (IX) groups. The 
relationships of these groups is discussed in the legend to Figure 56. A large clade that then diverged (8) was associated 
mainly with Muhlenbergia. In all species of this subgroup the plesiomorphic character of a pale face with black 
interocular band was retained, but the dorsal stripes were lost. The subgroup is defined [s] by the triangulate male 
plates (Figs. 33A-F, 34D,E), which are strongly divergent apically (but see serrata [Fig. 8C]; the plates of this species 
suggest a link to the decora subgroup). In the ijoung,i sister species set (host: Muhlenbergia richardsonis ), the unpaired 
ventral process (Figs. 3IF,G) retained its plesiomorphic position and length (20), but in the remaining species [s] the 
process was shortened (9). In modica the aedeagus is subsymmetrical (11) with [au] the short, unpaired process (Fig. 
31M) slightly displaced from the sagittal plane and associated with the shaft basally; the pygofer is relatively 
undifferentiated. In the lineage that diverged subsequently, the pygofer (Fig. 62) became substantially modified (10). 
We hypothesize that it first became ovate, a condition that still persists [au] in arenicola (16). In stylata and celata [s] it 
developed [s] a distinctive posterior process (17: see te.xt for further synapomorphies), and in arizonensis (au) it 
developed a conspicuous ventral process (14). The male plates oiflexidosa and arizotwnsis (Figs. 33A, B) are in each case 
(13, 14) di.stinctive [au]. In the inflata subgroup of group X (24), the beginning of phyletic torsion of the aedeagus is 
evident (Fig. 61 B). In this subgroup the face is modified; traces of the interocular band are present in some individuals, 
but, when present, it is brown and contrasts much less sharply with the remainder of the face. The plates are short, and 
[s] the dorsal margin of the pygofer is essentially flat, giving the pygofers a boatlike outline in lateral aspect (Figs. 
62GC,HH,II). Two apparent (but see text) sisters, texana and beameri, are identified by [au] unique aedeagal 
configurations (Figs. 31A,D). From the moderately displaced aedeagus of the inflata group, further torsion (29) led to 
groups XI-XIII. 



1988 



Whitcomb, Hicks: Flexamia Revision 



303 



Si 
I 



5 






2 



I 

Si 



S 



5 



5 

o 
■4i 




1 


>4J 

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. 



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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). 



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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 



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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. 



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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 



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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. 

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315 



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1173: 1-53. 



316 



Great Basin Naturalist Memoirs 



No. 12 



Appendices 



Appendix 1. Specimens examined\ 



Species 


Holotype 


Paratype(s) 


barc 


CNC 


KU 


OSU 


USNM 


minima 


OSU(h) 


none 











1 


8 


zamora 


OSU(h) 


OSU(p) 











9 





pectinata 


USNM(c) 


USNM(c) 


57 


5 


150 


45 


75 


gila 


USNM(h) 


P^ 


9 














handarita 


USNM(h) 


P^ 


5 








6 





doeringae 


KU(h) 


USNM(p) 


35 





68 


24 


91 


collorum 


USNM(h) 


P^ 


81 














jacala 


USNM(h) 


none 














1 


mescalero 


USNM(h) 


P^ 


26 














abbreviata 


USNM(c) 


USNM(c) 


765 


141 


661 


74 


409 


zacate 


USNM(h) 


P^ 


96 














canyonensis 


USNM(h) 


USNM(p) 


9 











2 


surcula 


osu 


OSU 


72 





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 





50 





18 


serrata 


KU(h) 


KU(p) 


56 


98 


20 





14 


ritana 


KU(h) 


USNM(p) 








17 





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 











5 


celata 


USNM(h) 


USNM(p) 


49 





4 





5 


stylata 


USNM(l) 


USNM(c) 


83 


67 


66 


9 


70 


decora 


KU 


USNM(p) 


95 


246 


177 


3 


28 


youngi 


USNM 


P^ 


30 





5 








modica 


KU 


USNM(p) 


377 





110 


3 


8 


inflata 


USNM(c) 


USNM(c) 


665 


103 


257 


146 


155 


beameri 


KU 


USNM(p) 








7 








texana 


USNM 


USNM(p) 














10 


imputans 


OSU(c) 


OSU(c) 


4 





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 





2 


producta 


(BNM) 


USNM(c') 


4 


5 


136 


117 


267 


sandersi 


OSU(h) 


OSU(h^) 


45 


3 


60 


31 


126 


delongi 


(INHS) 


none 





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) 





2 


2 









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. <i}andiilosa (^ 
Betula glandulosa Michx.) (1 male); White 
Mts., Eagle Summit, 900 m, 65°30'N, 
145°25'W, 13 July 1982, Seppo Keponen, ex. 
B. glandulosa (1 male); Mt. Fairplay, Taylor 
Highway, 3,600 ft, 10 July 1962, P. J. Skitsko 
(1 male, 4 females); Happy Val, 15 June 1982, 
S. F. MacLean, tussock tundra (1 male, 2 
females, 2 nymphs). Other records are from 
Anufriev and Averkin (1982a, 1982b), Anu- 
friev and Emeljanov (1981), and Scudder 
(1963). 

Distribution. — NearctiC; Canada: Al- 
berta, British Columbia, Northwest Territo- 
ries; USA: Alaska. PalearctiC: Mongolia, 
USSR. 

Criomorphiis wilhelmi Anufriev & Averkin 

Figs. 6-8 

Crioinorphus horealis (Sahlberg), Anufriev 1972:613, 

1977:864 nee Sahlberg 
Criomorphiis wilhelmi Anufriev and Averkin, 1982a: 131; 

new name for C horealis Anufriev nee Sahlberg 

Distribution record for the specimen used 
in this study is: ALASKA; Unalakleet, 21 June 
1961, R. Madge (1 male). Other records are 
from Anufriev (1972, 1977) and Anufriev and 
Averkin (1982a). 

Distribution. — NearctiC: Alaska. Pale- 
ARCTIC: Mongolia, USSR. 

Megamehis flavus Crawford 

Figs. 9-11 

Megamehis nottihis flavus Crawford, 1914:609 
Megamehis flavus Crawford; Beamer 1955:31 

Specimens from Alaska were not available 
for this study; illustrations were made from a 
specimen with the following collecting data: 
WYOMING: Bighorn Nat'l. Park, Meadowlark 
Lake, 23 August 1954, M. W. Sanderson (1 
male). Beamer (1955) recorded this species 
from Alaska but gave no further data. Other 
records are from Anufriev and Averkin 
(1982a), Beamer (1955), and Scudder (1964). 

Distribution. — NearctiC: Canada: Al- 
berta, British Columbia, Manitoba, North- 
west Territories, Quebec, Saskatchewan; 
USA: Alaska, Colorado, Wyoming. PalearC- 
TIC; Mongolia. 

Nothodelphax ehurneocarinatus (Anufriev) 

Figs. 12-15 

Tyrphodelphax ehurneocarinatus Anufriev, 1979:295 



Nothodelphax ehurneocarinatus (Anufriev): Emeljanov 

1 982:90 
Nothodelphax ehurneocarinatus (Anufriev): Anufriev and 

Averkin 1982a: 1.37 

Distribution records for specimens used in 
this study are: ALASKA; Alder Woodland, Arc- 
tic Circle, 66°33'N, 150°45'W, 17 June 1982, 
S. F. MacLean (7 males, 18 females); Brooks 
Range, Happy Valley, 69°05'N, 149°W, 320 
m, 6 July 1982, Seppo Keponen, ex. B. glan- 
dulosa (1 male); Mt. Fairplav, Taylor Hwy., 
3,600 ft, 10 July 1962, R. J. Skitsko (12 males, 
54 females); Old Man Camp, 17 June 1982, S. 
F. MacLean (5 males, 21 females). Anufriev 
(1979) records it from sedge tussock bogs. 

Distribution. — NearctiC; USA: Alaska. 
PalearctiC: USSR. 

Unkanodes excisa (Melichar) 
Figs. 16-20 

Liburnia excisa Melichar, 1898:67 
Elymodelphax excisa (Melichar): Wagner 1963:167 
Unkanodes {= Ehjmodelphax), Dlabola 1965:86 

Distribution records for specimens used in 
this study are: ALASKA: Cape Thompson, 6 
August 1979, S. F. MacLean (7 males, 8 fe- 
males, 2 nymphs). Other records are from 
Nast (1972). This species was reported from 
Elymus arenarius L. in coastal habitats by 
Ossiannilsson (1978). 

Distribution. — NearctiC; USA: Alaska. 
PalearctiC: Denmark, Finland, East Ger- 
manv. West Germany, Poland, Sweden, 
USSR. 

Chilodelphax magnifrons (Crawford), 
n. comb. 

Figs. 21-23 

Megamehis magnifrons Crawford, 1914:614 

Eurijsa magnifrons (Crawford): Muir and Giffard 1924:8 

Beamer (1952a) included four North Ameri- 
can species in Eurysa but suggested they 
might belong in Eunjhregma. Examination of 
PalearctiC Eurysa and Eurybregma indicates 
clearly that, based on features of the male 
genitalia, this species belongs in neither 
genus. It appears similar to Chilodelphax sil- 
vaticus Vilbaste as described and illustrated 
by Kwon (1982); I tentatively place it in 
Chilodelphax Vilbaste (1968). 

Distribution records for specimens used in 
this study are: Alaska: Ferry, 4 June 1981, 
S. F. MacLean (9 males, 7 females). Other 
records are from Beamer (1952a). 



338 



Great Basin Naturalist Memoirs 



No. 12 




Figs. 21-37. Figs. 21-23, Chilodelphax magnifrons (CrdvAord). 21, male genitalia, lateral view, 22, aedeagus, lateral 
view; 23, styles, caudal view. Figs. 24-28, Ribautodelphax albostriatus (Fieber): 24, pygofer, lateral view; 25, 26, 
aedeagus, lateral view; 27, anal tube, caudal view; 28, styles, caudal view. Figs. 29-31, Acanthodelphax analis 
(Crawford): 29, male genitalia, lateral view; 30, anal tube and aedeagus, lateral view; 31, styles, caudal view. Figs. 
32-34, Kusnezoviella macleani, n. sp.; 32, male genitalia, lateral view; 33, anal tube and aedeagus, lateral view; 34, 
styles, caudal view. Figs. 35-37, Javesella pellucida (Fabricius); 35, male genitalia, lateral view; 36, aedeagus, lateral 
view; 37, styles, caudal view. 



Distribution. — NearctiC: Canada: British 
Columbia; USA: Alaska, Colorado, Montana, 
Wyoming. 

Ribautodelphax albostriatus (Fieber) 

Figs. 24-28 

Delphax albostriata Fieber, 1866:525 
Ribautodelphax albostriatus (Fieher): Wagner 1963: 170, 
176 

Distribution records for specimens used in 
this study are: Alaska: Inspiration Base Camp 
(64°N, 148°40'W), 7 June 1981, S. F. 
MacLean (1 male); Umiat, 8 July 1959, R. 
Madge (2 males), 20 July 1959 (1 male), J. E. 
H. Martin, 13 July 1959(1 male), 16 July 1959 
(2 males), 23 July 1959 (2 males). Other 
records are from Nast (1972). This species was 
reported from dry grass fields bv Ossian- 
nilsson (1978). 



Distribution. — NearctiC: USA: Alaska. 
PalearctiC: Austria, Belgium, Cyprus, 
Czechoslovakia, Denmark, Finland, France, 
East Germany, West Germany, Hungary, 
Italy, Mongolia, Netherlands, Norway, 
Poland, Portugal, Romania, Spain, Sweden, 
Switzerland, Tunisia, USSR, Yugoslavia. 

Acanthodelphax analis {Crdwford), n. comb. 

Figs. 29-31 

Megaini'lus analis CrdvAord, 1914: 620 

Delpluicodes analis (Crawford): Muir and Ciffard 1924:24 

Distribution records for specimens used in 
this study are: ALASKA: Inspiration Base Camp 
(64°N, 148°40'W), 7 June 1981, S. F. 
MacLean (7 males); Hope, Kenai Peninsula, 
24 May 1951, W. J. Brown, coll. (3 males, 1 
female, 2 nymphs). Other records are from 



1988 



WiLSON: Delphacidae of Alaska 



339 



DuBose (1960) and Wilson and McPherson 
(1980). 

Distribution. — NearctiC: Canada: Al- 
berta; USA: Alaska, Illinois, Michigan, Min- 
nesota, New York, North Carolina, Wiscon- 
sin. 

Kusnezoviella Vilbaste 

Kusnezoviella Vilbaste 1965: 16. Type specie.s Lihurnia 
dimidiatifrons Kusnezov, by original designation 

The following description is based on a 
translation from the original Russian supplied 
by Vilbaste. 

Vertex slightly longer than wide, widening 
anteriorly. Frons with lateral margins almost 
parallel, converging posteriorly; median 
carina distinct, forking at juncture with ver- 
tex. Antennal scape length subequal to width 
at apex; pedicel ca 2X length of scape. Beak 
extending to mesocoxae. Pronotal carinae dis- 
tinct; lateral carinae curving posterolaterad 
and disappearing between eye and posterior 
margin of pronotum. Mesonotal carinae di- 
verging at a 50-60-degree angle. Brachypter- 
ous forewings ca 2X longer than wide. Meta- 
tibia with 7 teeth (5 + 2) in transverse row at 
apex on plantar surface; spur foliaceous, with 
18 teeth becoming larger apically, apical tooth 
small. 

Pygofer, in lateral view, with caudal margin 
oblique, slightly concave in middle. Di- 
aphragm with dorsoposteriorly directed ex- 
tension bearing teeth laterad. Anal tube with 
pair of elongate, acute spines; bases of spines 
well separated, distance between them ca 1/2 
length of spine. Styles directed posterodor- 
sally, apex acute, base broad. Aedeagus di- 
rected caudally, with row of teeth around 
gonopore, gonopore on dorsal aspect. 

Kusnezoviella macleani, n. sp. 

Figs. .32-.34 

Vertex pale beige, one dark brown spot 
laterally at base on each side, two dark brown 
spots medially anterior to V-shaped, trans- 
verse carina. Frons with carinae pale beige, 
area between carinae dark brown to pale in 
middle. Clypeus dark brown with pale, me- 
dian carina. 

Pronota and mesonota pale beige medially, 
dark brown lateral to lateral carinae. Brach- 
ypterous forewing hyaline, light brown. Legs 
straw-colored with brownish, longitudinal 
stripes. 



Abdomen blackish, marked with pale 
beige. 

Male genitalia. — Pygofer subcyHndrical; 
in lateral view, height ca 1.5X width; in caudal 
view, with weak dorsolaterally flaring lobes on 
lateral margin in dorsal 1/2; ventral margin of 
diaphragm opening with subtriangular exten- 
sion, in lateral view, with one small spine on 
either side, in caudal view. Anal tube sub- 
cylindrical, two short, acute spines extending 
from dorsocaudal margin; each spine sepa- 
rated at base by distance subequal to length of 
spine. Styles converging at apices, narrowing 
toward subacute tips; apical 1/2 avicephali- 
form. Aedeagus subcylindrical, slightly later- 
ally compressed, recurved; gonopore apical; 
apex appears to be broken but is identical in 
every specimen; with ca 6-7 teeth extending 
from near apex on ventral margin anterodor- 
sally and 3 teeth on dorsal margin at point of 
downward curvature of aedeagus. 

Types. — Holotype (male) brachypter with 
label: "Atkasuk, AKA. , mouth of Meade R. , 18 
June 1977, S. F. MacLean, coll., 
770617-03T." Allotype (female) brachypter, 
same data; in the Canadian National Collec- 
tion, Ottawa. ParatypeS: Alaska: Atkasuq, 
mouth of Meade R., 17 June 1977, S. F. 
MacLean, coll. (1 male), 18 June 1977 (1 
male); Umiat, 23 July 1959, J. E. H. Martin (3 
males); in the University of Kansas collection 
and collection of S. W. Wilson. These speci- 
mens were swept from two species of shrub 
willow (Salix alaxensis [Anderss.] Coville and 
S. glauca L.) (MacLean, personal correspon- 
dence); it is possible that their actual host 
plant may be a grass or sedge found under or 
near the willows. 

Javesella pellucida (Fabricius) 

Figs. 35-37 

Fulgora pellucida Fabricius, 1794:7 

Delphacodes pellucida (Fabricius): Muir and Giffard 

1924:20 
Javesella pellucida (Fabricius): Fennah 1963:15 

Distribution records for specimens used in 
this study are: Alaska: Big Delta, 3 July 1951, 
Mason and McGillis (2 males); Cooper Land- 
ing, Kenai Peninsula, 7 June 1951, W. J. 
Brown (1 male); King Salmon, Naknek R., 3 
July 1952, W. R. Mason (1 male), 23 July 1952 
(2 males), 1 August 1952 (1 male), 8 August 
1952 (1 male); Shaw Creek, M. 289 Rich. 
Hwy., 11 July 1951, Mason and McGillis 



340 



Great Basin Naturalist Memoirs 



No. 12 




Figs. 38-.51. Figs. 38-41, Javesella obscurella (Boheman); .38, male genitalia, lateral view; 39, anal tube, lateral view; 
40, aedeagus, lateral view; 41, styles, caudal view. Figs. 42-44, Javesella atrata (Osborn): 42, male genitalia, lateral 
view; 43, aedeagus, lateral view; 44, styles, caudal view. Figs. 45-48, Javesella simillima (Linnavuori): 45, pygofer, 
lateral view; 46, anal tube, lateral view; 47, aedeagus, lateral view; 48, styles, caudal view. Figs. 49-51, Javesella 
disco/or (Boheman): 49, male genitalia, lateral view; 50, aedeagus, lateral view; 51, styles, caudal view. 



(1 male); Umiat, 4 July 1959, R. Madge (2 
males), 8 July 1959 (1 male); Unqlakleet, 8 
July 1981, R. Madge (1 male). Other records 
are from Dubose (1960), Muir and Giffard 
(1924), Nast (1972), and from my collection. 
Javesella pellucida is a vector of viral patho- 
gens of several cereals (Wilson and O Brien 
1987); studies of the biology of this planthop- 
per were summarized by Mochida and Kisi- 
moto (1971). 

Distribution.— Nearctic: Canada: Al- 
berta; USA: Alaska, Connecticut, Illinois, 
Maine, Massachusetts, New Hampshire, 
North Carolina, North Dakota, Oregon. 
PalearctiC: Algeria, Austria, Belgium, 
Czechoslovakia, Denmark, Finland, France, 
East Germany, West Germany, Great 
Britain, Hungary, Iceland, Ireland, Italy, 
Japan, Libya, Mongolia, Morocco, Nether- 
lands, Norway, Poland, Romania, Spain, 
Sweden, Switzerland, Turkey, USSR, Yugo- 
slavia. 



Javesella obscurella (Boheman) 

Figs. 38-41 

Dclphax obscurella Boheman, 1847:.53 

Javesella obscurella (Boheman): LeQuesne 1964:57 

Distribution records for specimens used in 
this study are: ALASKA: Atkasuq, 16 June 1977, 
mouth of Meade R., S. F. MacLean (1 male, 1 
female), 17 June 1977 (1 male); 24 July 1977 (2 
males); 31 July 1977 (3 males, 2 females); 
Brooks Range, Happy Valley, 69°05'N, 
149°W, 320 m, 6 July 1982, Seppo Keponen, 
ex. B. glandulosa (3 females); Cape Thomp- 
son, 23 Julv 1961, R. Madge (7 males); Umiat, 
4 July 1959, R. Madge (1 male), 8 July 1959 (1 
male); USSR: Chaun Bay, 14 August' 1977, S. 
F. MacLean, Jr., ex. Poa and Festuca (1 male, 
3 parasitized males, 3 females, 6 nymphs). 
Other records are from Nast (1972). Records 
from New York given in Metcalf (1943) come 
from before the widespread use of genitalic 
features in delphacid identification and, thus, 



1988 



WILSON; Delphacidae of Alaska 



341 



are probably in error. This species is a vector 
of viral pathogens of cereals (Wilson and 
O'Brien 1987); the biology of it on cereal crops 
was summarized by Ossiannilsson (1978). 

Distribution.— NearctiC: USA: Alaska. 
PalearCTIC: Austria, Belgium, Bulgaria, 
Czechoslovakia, Denmark, Finland, France, 
East Germany, West Germany, Great 
Britain, Hungary, Ireland, Italy, Mongolia, 
Netherlands, Norway, Poland, Portugal, Ro- 
mania, Sweden, Switzerland, Turkey, USSR, 
Yugoslavia. 

Javesella atrata (Osborn), n. comb. 
Figs. 42-44 

Delphacodes atrata Osborn, 1938:344 

Specimens from Alaska were not available 
for this study; illustrations were made from a 
"paramorphotype" with the following collect- 
ing data: OHIO: Rome, 19 July 1946, R. H. 
Beamer. Beamer (1951) recorded this species 
from Ketchikan, Alaska, as well as the other 
localities given below. 

Distribution.— NearctiC; USA: Alaska, 
Connecticut, Illinois, New Hampshire, New 
York, Ohio. 

Javesella simillima (Linnavuori) 

Figs. 45-48 

Calligypona simillima Linnavuori, 1948:45 
Delphacodes saileri Beamer 1952b: 114. New synonymy 
Javesella simillima (Linnavuori): Nast 1972:68 

I was unable to determine who first placed 
this species in Javesella. Based on information 
given by Hulden (1974), it appears to be Lin- 
navuori (1969), a paper which was unavailable 
to me. In the foregoing synonymy I hst Nast 
(1972) as my reference for first placement in 
Javesella. 

Distribution record for the specimen used 
in this study is: ALASKA: Umiat, 4 July 1959, R. 
Madge (1 male). Other records are from 
Beamer (1952b) and Nast (1972). This species 
is reported from Eriophorum and Carex by 
Ossiannilsson (1978). 

Distribution— NEARCTIC: USA: Alaska. 
PALEARCTIC: Finland, USSR. 

Javesella discolor (Boheman) 
Figs. 49-51 

Delphax discolor Boheman, 1847:61 

Javesella discolor (Bohemain): LeQuesne 1964:57 

Distribution records for specimens used in 
this study are: Alaska: Umiat, 3 July 1959, 




Figs. 52-54. Javesella arcanastyla (Beamer); 52, male 
genitalia, lateral view; 53, aedeagus, lateral view; 54, 
styles, caudal view. 



J. E. H. Martin (1 male); 4 July 1959, R. 
Madge (4 males), 8 July 1959 (3 males). Other 
records are from Nast (1972). This species has 
been reported to be a vector of viral pathogens 
of cereals (Wilson and O'Brien 1987); host 
plants were summarized bv Ossiannilsson 
(1978). 

Distribution —NEARCTIC; USA: Alaska. 
PalearctiC: Algeria, Austria, Belgium, 
Czechoslovakia, Denmark, Finland, France, 
East Germany, West Germany, Great Brit- 
ain, Ireland, Italy, Mongolia, Netherlands, 
Norway, Poland, Romania, Sweden, Switzer- 
land, USSR. 

Javesella arcanastyla (Beamer), n. comb. 
Figs. 52-54 

Delphacodes arcanastyla Beamer, 1948:101 

Distribution record for the specimen used 
in this study is: ALASKA; Matanuska, 9 June 
1944, J. C. Chamberlain, Rotary Traps, 
46-7695. Other records are from Beamer 
(1948) and the Michigan State University col- 
lection. 

Distribution.— NEARCTIC: USA: Alaska, 
Michigan, Washington, Wisconsin, Wyoming. 

Comments on Distribution 

Localities represented in the material used 
for this study range from the Kenai and Alaska 
peninsulas in the south through east central 
Alaska extending to ca 100 km from the north- 
ern coast. Three other localities represent 
coastal sites, one on the north coast in the 
western one-third of the state, one on the west 
coast above the Arctic circle, and one on the 



342 



Great Basin Naturalist Memoirs 



No. 12 



central west coast. Only one record is from the 
southeastern portion of the state. 

One of the 15 delphacid species is recorded 
only from Alaska (K. macleani); 3 species are 
Holarctic in distribution (A. subarctica,] . pel- 
lucida, and M. flatus), 4 are Nearctic (A. 
analis, C. magnifrons, J. arcanastyla, and/. 
atrata), and 7 are Palearctic (C. wilhehni, J. 
discolor, J. ohscurella, J. simiUiyyia, N. ebur- 
neocarinatus, R. albostriatus, and V. excisa). 
One of the seven Palearctic planthoppers (N. 
eburneocarinatus) appears to have an amphi- 
Beringean distribution; five of the seven have 
only been collected north of the Brooks 
Range; all seven are apparently restricted to 
tundra habitats in Alaska. The diversity of the 
Alaskan delphacid fauna is comparable to that 
reported by Vilbaste (1980), who listed 10 
species from the Kamchatka Peninsula. 

Acknowledgments 

Thanks are tendered to Dr. G. G. E. Scud- 
der, Department of Entomology, University 
of British Columbia, Vancouver, Canada, who 
recommended me as someone who might be 
able to identify Alaskan delphacids; Dr. S. F. 
MacLean, Jr., Department of Zoology, Uni- 
versity of Alaska, Fairbanks, for providing de- 
tailed information on the collecting localities; 
Dr. K. G. A. Hamilton, Biosystematics Re- 
search Centre, Agriculture Canada, Ottawa, 
for the loan of material; the late Dr. J. Vil- 
baste, Institute of Zoology and Botany, 
Academy of Sciences, Tartu, Estonian SSR, 
who suggested the generic placement of the 
new species and provided an English transla- 
tion of his description; Drs. M. Asche, H. 
Hoch, and R. Remane, Fachbereich Biologie- 
Zoologie, Philipps Universitiit, Marburg, 
West Germany, for their invaluable aid in 
identifying Palearctic material; and Dr. L. B. 
O'Brien for commenting on the manuscript. I 
appreciate support for work at Philipps Uni- 
versitat, where some of this research was 
done, from the German Academic Exchange 
and the Graduate School, Central Missouri 
State University. 

Literature Cited 

Anufriev, G, a. 1972. New and little known leaf-hoppers 
from the family Delphaeidae (Homoptera) from 
the Maritime District. Zool. Zh. 5L 612-615. 



1977. Delphacids (Homoptera, Auchenorrhyn- 

cha) of the Kurile Islands Fauna. Zool. Zh. 61: 
8.5.5-869. 

1979. Notes on the genus Tijrphodclphax Vil- 
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species (Homoptera, Delphaeidae). Reichen- 
bachia 17: 29,5-301. 

Anufriev, G A , and A M. Averkin 1982a. Cicadinea of 
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1982b. On delphacids of the Komsomolsky Re- 
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Anufriev, G. A , and A. F. Emeljanov. 1981. The genus 
Achorotile Fieb. (Homoptera, Delphaeidae) and 
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ASHMEAD, W H 1904. The Homoptera of Alaska. Harri- 
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Beamer, R H 1948. Some new species of Delphacodes 
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19.51. Some species of Delphacodes, new and old 

(Homoptera-Fulgoridae-Delphacinae). J. Kansas 
Entomol. Soc. 24: 11-15. 

19.52a. The genus Eurtisa in America north of 

Mexico (Homoptera: Fulgoridae, Delphacinae). 
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1952b. One old and five new species of delphacine 

fulgorids (Homoptera: Fulgoridae). J. Kansas En- 
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19.5.5. A revision of the genus Mc^amclus in Amer- 
ica north of Mexico (Homoptera, Fulgoridae, Del- 
phacinae). J. Kansas Entomol. Soc. 28: 29-46. 

Boheman, C H 1847. Nya Svenska Homoptera. Handl. 
Svenska Vet. Akad. 1847: 23-67. 

Crawford, D L 1914. A contribution toward a mono- 
graph of the homopterous insects of the family 
Delphaeidae of North and South America. Proc. 
U.S. Nat. Mus. 46: ,557-640. 

Dlabola. J 1965. Ergebnisse der zoologisehen 
Forschungen von Dr. Z. Kaszab in der Mongolei. 
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Entomol. Mus. Nat. Pragae, Praha 11(100): 
79-136. 

DoziER, H L 1926. Notes on new and interesting del- 
phacids. J. New York Entomol. Soc. 34: 2,57-263. 

DuBoSE, W P. 1960. The genus Delphacodes Fieber in 
North Carolina (Homoptera: Delphaeidae). J. El- 
isha Mitchell Sci. Soc. 76: .36-63. 

Emeljanov, A F. 1982. Fulgoroidea (Homoptera) col- 
lected in the Mongolian People s Republic by the 
entomofaunistical group of the Soviet-Mongolian 
complex biological expedition in 1970-1975. Ins. 
Mongolia 8: 69-122. 

Fabricius, J C 1794. Ryngota. Entomologia systematica 
emendata et aucta. Secundiun classes, ordines, 
genera, species adjectis synoimis, locis, observa- 
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Fennah, R. G. 1963. New genera of Delphaeidae (Ho- 
moptera: Fulgoroidea). Proc. Ro\al Entomol. 
Soc, London (B) .32: 1,5-16. 

Fieber, F X 1866. Grundzuge zur generischen Theilung 
der Delphacini. Verb. Zool. Bot. Ges. Wien 16: 
517-5,34. 



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WiLsoN: Delfiiacidae OF Alaska 



343 



HULDEN. L. 1974. The Javesclhi discolor group (llo- 
moptera, Delphacidue) of North Europe, witli de- 
scription of a new species. Not. Entomol. 54; 
114-116. 

KWON, Y. J 1982. New and little known planthoppers of 
the family Delphacidae (Homoptera; Auchenor- 
rhyncha). Korean). Entomol. 12; 1-11. 

LeQuesne, W. J. 1964. Some ta.\onomic observations on 
the British Delphacidae (Hemiptera). Proc. Roval 
Entomol. Soc. , London (B) 33; 56-58. 

LiNNAVUORl, R. 1948. Neue oder bemerkensverte 
Zikadenfunde aus Finnland, nebst einer Beschrei- 
bung einer neuen art. Ann. Entomol. Fenn. 14: 
45-47. 

1968. HemipterlV. Anim. Fenn. 13; 1-312. 

MacLean, S F , JR , AND 1 D HoDKiNSON 1980. The 
distribution of psyllids (Homoptera: Psylloidea) in 
arctic and subarctic Alaska. Arct. Alp. Res. 12: 
369-376. 

Melichar, L 1898. Eine neue Homopteren-Art aus 
Schleswig-Holstein. Wiener Entomol. Zeit. 17; 
67-69. 

Metcalf, Z. p. 1943. General catalogue of the Hemip- 
tera. Fasc. IV, Fulgoroidea. Part 3, Araeopidae 
(Delphacidae). Smith Coll., Northampton, Massa- 
chusetts. 552 pp. 

MocHiDA, O., AND R KisiMOTO 1971. A review of the 
studies on Javesella pellucida (F.) (Hom., Del- 
phacidae) and associated subjects. Rev. Plant 
Prot. Res. 4: 1-57. 

MuiR, F 1919. Notes on the Delphacidae in the British 
Museum collection. Canadian Entomol. 51; 6-8. 

MuiR, F., AND W. M GiFFARD 1924. Studies in North 
American Delphacidae (Homoptera). Bull. Expt. 
Sta. Hawaiian Sugar Planters" Assoc, Entomol. 
Ser. Bull. 15; 1-53. 



Nasi. J 1972. Palaearctic Auchenorrhyncha (Homop- 
tera), an annotated check list. Polish Acad. Sci., 
Polish Sci. Publ., Warsaw. 550 pp. 

OsBORN, H 1938. The Fulgoridae of Ohio. Bull. Ohio 
Biol. Surv. 6; 283-349. 

OssiANNlLS.SON. F. 1978. The Auchenorrhyncha (Ho- 
moptera) of Fennoscandia and Denmark. Part 1: 
Introduction, infraorder Fulgoromorpha. Fauna 
Entomol. Scand. 7; 5-222. 

SCUDDER. G G E 1963. Studies on the Canadian and 
Alaskan Fulgoromorpha (Hemiptera). \. The gen- 
era Achrotile Fieber and Laccocera Van Duzee 
(Delphacidae). Canadian Entomol. 95: 167-177. 

1964. Studies on the Canadian and Alaskan Ful- 
goromorpha (Hemiptera). II. The genus 
Meg,amelus Fieber (Delphacidae). Canadian En- 
tomol. 96; 813-820. 

ViLBASTE, J. 1965. On the Altaic cicaden fauna. Tartu. 144 
pp. 

1968. On the cicaden fauna of the Primorsk region. 

Tallinn. 179 pp. 

1980. On the Homoptera-Cicadinea of Kam- 
chatka. Ann. Zool. 35; 367-417. 

Wagner, W 1963. Dynamische Taxionomie, angewandt 
auf die Delphaciden Mitteleuropas. Mitt. Ham- 
burg Zool. Mus. Inst. 60; 111-180. 

Wilson, S. W, and J E McPherson. 1980. The distribu- 
tion of the Fulgoroidea of the eastern United 
States (Homoptera). Trans. Illinois Acad. Sci. 
73(4): 7-20. 

Wilson, S W , and L B OBrien 1987. A survey of 
planthopper pests of economically important 
plants (Homoptera; Fulgoroidea). Pages 343-360 
m Proc. 2nd Int. Workshop on Lealhoppers and 
Planthoppers of Economic Importance. 



TAXA IN THE HOMOPTERA, 
AUCHENORRHYNCHA DESCRIBED BY PAUL W. OMAN 



HOMOPTERA 



Fossil Cicadelloidea 

Mesojassoides Oman 1937. Colorado. 
Mesojassoides gi^antea Oman 1937. 
Colorado. 

Fulgoridae 

Pohlicia texana Oman 1936. Texas. 

Cicadellidae 

Subfamily 

Dorycephalinae 
Neocoelidiinae 
Nioniinae 
Xerophloeinae 

Tribe 

Acinopterini 

Cochlorinini 

Dorycephalini 

Mesaniiinae 

Neocoelidiini 

Nionini 

Scaphoideini 

Scaphytopiini 

Xerophloeini 

Genera and Subgenera 

Aceratagallia (Bergallia) Oman 1938. 

Argentina. 
Aflexia Oman 1949. Illinois. 
AgaUiana Oman 1933. Brazil. 
Agalliopsis (Agallaria) f)man 1949. 

Indiana. 
Agalliota Oman 1938. Mexico. 
Agelina Oman 1938. Argentina. 
Agudus Oman 1938. Argentina. 
Alaca Oman 1938. Argentina. 
Alhjgiella Oman 1949. California. 
Ankosiis Oman & Musgrave 1975. 

Oregon. 
Aplanus Oman 1949. Colorado. 
Arrugada Oman 1938. Bolivia. 
Ataniis Oman 1938. Bolivia. 
AuridUts Oman 1949. Colorado. 
Bahita Oman 1938. Brazil. 
Balduhi.s Oman 1934. Arizona. 
Baroma Oman 1938. Argentina. 
Bathijsmatophorns (Htjlaius) Oman & 

Musgrave 1975. Oregon. 
Bemda Oman 1938. Bolivia. 
Bolarga Oman 1938. Bolivia. 
B o na mu s OnvAw 1938. Argentina. 
Bonncijana Oman 1949. Utah. 
Brasa Oman 1938. Hispaniola. 



Brazosa Oman 1939. Brazil. 
BrendaOmAw 1941. California. 
B rincadorus OmAn 1938. Brazil. 
Bijthonia Oman 1938. Bolivia. 
C(d}rulus Oman 1949. Colorado. 
Caladonus Oman 1949. California. 
Cantura Oman 1949. Eastern USA and 

Canada. 
Caphodus Oman 1938. Argentina. 
Cariancha Oman 1938. Brazil. 
Cazenus Oman 1949. New Mexico. 
Cerrillus Oman 1938. Brazil. 
Cetexa Oman 1949. Colorado. 
CocUdiana Oman 1938. Brazil. 
Colimona Oman 1949. Arizona. 
Conala Oman 1938. Brazil. 
Cortona Oman 1938. Argentina. 
Cn7jr«,sOman 1949. Minnesota. 
Crumhana Oman 1949. Tennessee. 
Citmora Oman 1938. Argentina. 
Daltonia Oman 1949. Texas. 
Danhara Oman 1949. Washington, D. C. 
Decua Oman 1949. Arizona. 
Deltanus Oman 1949. Texas. 
Dcltella Oman 1949. Florida. 
Deltoccphahis (Haldorus) Oman 1938. 

Bolivia. 
Dcstria Oman 1949. Florida. 
DikrcUa Oman 1949. New Mexico. 
Dorycephalus (Attenuipyga) Oman 1949. 

Florida. 
Egentts Oman 1939. Argentina. 
Endria Oman 1949. Virginia. 
Errhoinus Oman 1938. Washington. 
Errhomtis (Carsonus) Oman 1938. 

Colorado. 
Eidonus Oman 1949. California. 
Etiragcdlia Oman 1938. Brazil. 
Etisama Oman 1949. Arizona. 
Eiisora Oman 1949. Colorado. 
Ext rusanus Oman 1949. New York. 
F(dtala Oman 1938. Argentina. 
Fitcluina Oman 1949. New York. 
Floridonus Oman 1949. Florida. 
Fridonu.s Oman 1949. Texas. 
Frigartus Oman 1949. Colorado. 
Fn.scflH(/.s' Oman 1939. California. 
Garapita Oman 1938. Argentina. 
Giprits Oman 1949. California. 
Hchcxa Oman 1949. Utah. 
Hecullus Oman 1949. Colorado. 
Hcgira Oman 1938. Argentina. 



344 



1988 



OMAN; Taxa in the Homoptf.ra 



345 



Hordnia Oman 1949. C'alifornia. 
Kanorba Oman 1938. Brazil. 
KeonoUa Oman 1949. Washington. 
Ktinzeana Oman 1949. Arizona. 
Lemellus Oman 1949. Colorado. 
Limhanus Oman 1949. California. 
Lijcioides Oman 1949. Arizona. 
Manzutus Oman 1949. Arizona. 
Marathonia Oman 1949. Te.xa.s. 
Mexara Oman 1949. Arizona. 
Nggosiana Oman 1949. Southea.stern U.S. 
Neobala Oman 1938. Bolivia. 
Neonirvana Oman 1936. Coasta Rica. 
Neopsis Oman 1938. Jamaica. 
Nesaloha Oman 1943. Canton Island. 
Nigridonus Oman 1949. Arizona. 
Niirentts Oman 1949. Arizona. 
Onura Oman 1938. Argentina. 
Orocastus Oman 1949. Montana. 
Pasarenus Oman 1949. Colorado. 
Pazu Oman 1949. Arizona. 
Pecomts Oman 1949. Arizona. 
Phlepsanus Oman 1949. Arizona. 
Quontiis Oman 1949. Colorado. 
Rosejius Oman 1949. Iowa. 
Sahara Oman 1949. Te.xas. 
Tehisus Oman 1949. Colorado. 
Texananus (Excidtanus) Oman 1949. 

Florida. 
Thatuna Oman 1938. Idaho. 
Tiaja Oman 1941. California. 
Verdanus Oman 1949. Alaska. 
Zabrosa Oman 1949. Brazil. 

Species and Subspecies 

Aceratagallia ahrupta Oman 1933. 

Arizona. 
Aceratagallia accola Oman 1933. USA. 
Aceratagallia (Bergallia) attenuata Oman 

1938. Argentina. 
Aceratagallia aplopappi Oman 1933. 

USA. 
Aceratagallia arida Oman 1933. USA. 
Aceratagallia calcaris OmAit 1933. USA. 
Aceratagallia catalina Oman 1938. Brazil. 
Aceratagallia chileana Oman 1938. Chile. 
Aceratagallia cinerea Oman 1933. USA. 
Aceratagallia compacta Oman 1933. 

California. 
Aceratagallia confusa Oman 1938. 

Argetitina. 
Aceratagallia curta Oman 1933. Arizona. 
Aceratagallia cnrvata Oman 1933. USA. 
Aceratagallia dondia Oman 1933. USA. 
Aceratagallia fuscoscripta Oman 1933. 

USA. 
Aceratagallia grisea Oman 1935. Arizona. 
Aceratagallia helveola Oman 1933. USA. 
Aceratagallia humilis Oman 1933. USA. 
Aceratagallia lateralis Oman 1938. 

Argentina. 
Aceratagallia lobata Oman 1933. USA. 
Aceratagallia nana Oman 1933. Arizona. 
Aceratagallia nanella Oman 1933. 

Arizona. 
Aceratagallia neosignata Oman 1938. 

Argentina. 



Aceratagallia nitidula Oman 1933. 

Arizona. 
Aceratagallia obsctira Oman 1933. USA. 
Aceratagallia ovata Oman 1935. Arizona. 
Aceratagallia pallida Oman 1933. Utah, 

Arizona. 
Aceratagallia ))(>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 <i.rex Oman 1940. Arizona. 
Deltoccphalus (lorstiihmm 1940. Utali. 
DeItocr))halus laredanus Oman 1934. 

Texas. 
Deltoccphalus lincatifron.s Oman 1931. 

Colorado. 
Deltoccphalus plaijensis Oman 1940. 

Arizona. 
Dikraneura cedrelae Oman 1937. Puerto 

Rico. 
Dikraneura lentrosemac Oman 1937. 

Puerto Rico. 
Dorijcara platt/rhi/ncha setosa Oman 

1985. USA. " 
Egenus acuminatus Oman 1938. 

Argentina. 
Empoasca curvcola Oman 1936. 

Argentina. 
Empoasca dolonis Oman 1936. Argentina. 
Empoasca ensiformis Oman 1938. 

Oregon. 
Empoasca fuscoviridis Oman 1938. 

Colorado. 
Empoasca indenta Oman 1938. Utah. 
Empoasca insularis Oman 1936. Puerto 

Rico. 
Empoasca longispina Oman 1936. 

Argentina. 
Empoasca missiona Oman 1936. 

Argentina. 
Empoasca ncaspersa Oman 1938. Western 

USA. 
Empoasca ornata Oman 1936. Bolivia. 
Empoasca papayae Oman 1937. Puerto 

Rico. 
Empoasca peregrina Oman 1936. Peru. 
Empoasca perclegans Oman 1936. 

Neotropical. 
Empoasca phaseola Oman 1936. Costa 

Rica. 
Empoasca rid)raza Oman 1936. South 

America. 
Empoasca setigera Oman 1936. 

Argentina. 
Empoasca trifurcata Oman 1936. 

Argentina. 
Empoasca vastitatis Oman 1938. New 

Mexico. 
Empoasca vermispina Oman 1938. 

California. 
Empoasca xerophila Oman 1938. 

Colorado. 
Euragallia alhopunctata Oman 1938. 

Brazil. 
Euragallia attenuata Oman 19.38. Brazil. 
Euragallia lata Oman 1938. Brazil. 
Euragallia magnicauda Oman 1938. 

Brazil. 
Euragallia major hreviata Oman 1938. 

Brazil. 



Errhimius (Carsonus) aridtis furcatus 

Oman 1938. Washington. 
ErHumius (Carsonus) aridus incertus 

Oman 19.38. California. 
Errtumius hrevis hrevis Oman 1987. 

Washington. 
Errhomus hrevis klickitat Oman 1987. 

Washington. 
Errhomus filamentus Oman 1952. 

Oregon. 
Errhomus idahoensis Oman 1987. Idaho. 
Errlumius (Carsonus) irroratus spicatus 

Oman 19.38. Oregon. 
Errhomus josephi Oman 1987. Oregon, 

Washington. 
Errhonnis lineatus umatilla Oman 1987. 

Oregon. 
Errhomus medialis medialis Oman 1987. 

Oregon. 
Errhomus medialis minutus Oman 1987. 

Oregon. 
Errhomus medialis truncus Oman 1987. 

Oregon. 
Errhomus ochoco. Oman 1987. Oregon. 
Errhomus paradoxus Oman 1987. 

Washington. 
Errhomus reflexus Oman 1987. 

Washington. 
Errhomus satus Oinan 1987. Washington. 
Errhomus serratus serratus Oman 1987. 

Oregon. 
Errhomus serratus instabilis Oman 1987. 

Washington. 
Errhomus serratus affinis Oman 1987. 

Idaho. 
Errhomus simcoe simcoe Oman 1987. 

Washington. 
Errhomus simcoe minor Oman 1987. 

Washington. 
Errhomus similis Oman 1952. 

Washington. 
Errhomus similis socius Oman 1987. 

Washington. 
Errhomus similis relativus Oman 1987. 

Washington. 
Errhomus sobrinus sobrinus Oman 1987. 

Washington. 
Errhomus sobrinus confinis Oman 1987. 

Washington. 
Errhomus sobrinus dubiosus Oman 1987. 

Washington. 
Errhomus sobrinus nanus Oman 1987. 

Oregon . 
Errhomus sobrinus kahlotus Oman 1987. 

Oregon. 
Errhomus solus Oman 1987. Montana. 
Errhomus variabilis variabilis Oman 

1987. Washington, Idaho. 
Errhonnis variabilis pallidus Oman 1987. 

Oregon. 
Errhomus variabilis calvus Oman 1987. 

Washington, British Columbia. 
Errhomus winquait Oman 1987. Oregon. 
Errhomus wolfei Oman 1952. 

Washington. 
Errhomus zonarius Oman 1987. 

Washington. 



348 



Great Basin Naturalist Memoirs 



No. 12 



Faltata brachyptera Oman 1938. 

Argentina. 
Fridonus concanus Oman. 1949. Texas. 
Garapita ^arbosa Oman 19.38. Argentina. 
Hebecephalus scriptanus Oman 1934. 

Arizona. 
Hebexa incognita Oman 1949. Utah, 

Washington. 
Hegira brunnea Oman 1938. Argentina. 
Helochara delta Oman 1943. California. 
Joruma neascripta Oman 1937. Puerto 

Rico. 
Kanorba reflexa Oman 1938. Brazil. 
Koebelia inyoensis Oman 1972. CaHfornia. 
Krisna in.^ularis Oman 1936. Puerto Rico. 
Laevicephahts angelus Oman 1937. 

CaHfornia. 
Laevicephahis aridus Oman 1934. 

Arizona. 
Laevicephahis b