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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  Previously  unpublished  manuscripts  in  English  pertaining  to  the  biological 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. 

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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 
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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, 


12 


Great  Basin  Naturalist  Memoirs 


No.  12 


Fig.  2  A-K.  Exitianus  mmor,  n.  sp.:  A,  head  and  thorax,  dorsal  view;  B,  head,  ventral  view;  C,  forewing;  D,  hind 
wing;  E,  genital  capsule,  lateral  view;  F",  male  plates,  ventral  view;  G,  style,  dorsal  view;  H,  connective,  dorsal  view; 
I,  aedeagus,  lateral  view;  J,  aedeagus,  dorsal  view;  K,  female  seventh  sternum. 

26. VI. 85,    Qadeer.    Seven   paratypes,    .same      head  projected  in  front,  broadly  rounded  at 


data  as  holotype,  all  in  Zoological  Museum, 
University  of  Karachi,  Karachi,  Pakistan. 

Exitianus  minor  Ahmed  &  Qadeer,  n.  sp. 

(Figs.  2  A-K) 

Length. — Male  4.00  mm,  female  4.5  mm; 


anterior  apex;  median  length  of  crown  less 
than  interocular  width;  transocular  width  of 
head  nearly  ecjual  to  transverse  width  of 
pronotum;  coronal  suture  present  up  to  more 
than  midlength;  ocelli  on  anterolateral  margin 


1988 


Ahmed  ETAL.:  New  Grassland  Cicadellidae  from  Pakistan 


13 


of  crown,  close  to  eyes;  crown  with  brownish 
markings  on  disc  and  transverse  brownish 
band  at  the  level  of  anterior  margin  of  eyes; 
ground  color  of  crown  more  or  less  pale 
brown;  frontoclypeus  nearly  parallel  sided, 
narrowed  at  apex;  anteclypeus  narrowed  ante- 
riorly, much  longer  than  broad;  lora  broad; 
pronotum  with  some  tiny  spots  in  middle  in 
anterior  region,  with  posterior  margin  nearly 
transverse;  scutellum  pale  brown,  with  oval- 
shaped,  brownish  spots  anterolaterally. 

Forewing  with  apex  broadly  rounded;  api- 
cal part  slightly  broader  than  basal  part;  ap- 
pendix broad,  extending  up  to  5th  apical  cell; 
apical  cells  5;  subapical  cells  3;  hind  wing  with 
4  closed  apical  cells. 

Male. — Plate  in  ventral  view,  broad 
basally,  narrowed  apically,  possessing  4-5 
macrosetae  near  apex  on  lateral  margin,  long 
row  of  9-10  macrosetae  from  near  base  to 
apex  on  ventral  surface;  pygofer  with  poste- 
rior margin  directed  into  posterodorsal  broad 
and  rounded  lobe,  lobe  possessing  4-5 
brownish  macrosetae;  style  in  dorsal  view 
massive,  membranous  in  middle,  sclerotized 
along  margins,  preapical  lobe  flattened,  al- 
most truncate  mesally,  possessing  3-4  macro- 
setae, apical  extension  prominent,  spinose, 
extreme  apex  directed  caudolaterad;  connec- 
tive Y-shaped,  with  arms  smaller  than  the 
stem,  converging  at  their  apex;  aedeagus  in 
lateral  view,  with  preatrium  reduced,  dorsal 
apodeme  present,  appearing  broad  in  dorsal 
view,  shaft  tubular,  curved  smoothly  dorsad, 
apical  part  narrowed. 

Female. — Seventh  sternum  with  posterior 
margin  possessing  three  lobes,  lateral  lobes 
more  projected  than  the  median. 

Exitianus  fninor  appears  close  to  E.  ka- 
rachiensis  Ahmed  and  E.  major  described 
hitherto  in  its  venation  and  general  pattern  of 
male  genitalia,  but  differs  in  its  shape  of  con- 
nective and  female  seventh  sternum. 

Holotype  (male). — Pakistan:  Karachi,  grass, 
12. V.  1985,  Qadeer.  Thirty  paratypes,  all  in 
Zoological  Museum,  University  of  Karachi, 
Karachi,  Pakistan. 

Chiasmus  Mulsant  and  Rey 

Distant  (1908)  recorded  Chiasmus  uzelli 
Melichar  from  Ceylon.  Pruthi  (1930)  de- 
scribed three  new  species  from  various  parts 
of  India  and  reassigned  Kartwa  mustelina 
Distant  to  the  genus  Chiasmus .  The  present 


description  of  two  new  species  in  Chiasmus  is 
the  first  record  of  the  genus  from  Pakistan. 

Chiasmus  karachiensis  Ahmed  & 
Qadeer,  n.  sp. 

(Figs.3A-J) 

Length. — Male  2.3  mm;  head  strongly 
projected  in  front,  apex  of  crown  narrowed 
and  rounded;  median  length  of  crown  much 
less  than  interocular  width;  transocular  width 
of  head  slightly  more  than  transverse  width  of 
pronotum;  coronal  suture  present  up  to  more 
than  midlength  of  crown;  ocelli  on  anterolat- 
eral margin  of  crown,  close  to  eyes;  color  of 
eyes  blackish;  crown  possessing  blackish 
brown  dots  throughout;  ground  color  of  crown 
pale  yellow;  pronotum  yellowish,  its  posterior 
margin  nearly  straight;  scutellum  dim  yellow; 
frontoclypeus  narrowing  anteriorly,  blackish 
in  color  throughout,  except  median,  longitu- 
dinal stripe  of  yellow  color;  anteclypeus  black 
in  middle;  lora  broad,  expanded. 

Forewing  with  apex  smoothly  rounded; 
maximum  width  of  wing  slightly  prior  to  mid- 
length;  appendix  broad,  extending  to  3rd  api- 
cal cell;  apical  cells  4;  subapical  cells  2;  hind 
wing  with  4  closed  apical  cells. 

Male. — Plate  in  ventral  view  broad  at  base, 
middle  and  apical  parts  much  narrowed, 
macrosetae  2-3  on  ventral  surface  in  apical 
half,  apex  of  plate  rounded  and  directed  cau- 
dad;  pygofer  in  lateral  view  with  posterior 
margin  narrowed  into  angular,  posterodorsal, 
much-projected  lobe,  processes  and  setae 
usually  absent;  style  in  dorsal  view  membra- 
nous in  middle,  sclerotized  along  margins, 
preapical  lobe  short,  narrow,  and  angular, 
apical  extension  dark  brown  in  color,  spinose 
and  curved  laterad;  connective  Y-shaped, 
with  arms  longer  than  the  stem,  arms  apposed 
most  of  their  anterior  part,  forming  nearly 
rounded  space  in  middle;  aedeagus  in  lateral 
view  with  preatrium  reduced,  dorsal  apo- 
deme absent,  shaft  tubular,  somewhat  curved 
dorsad  towards  apex,  processes  absent. 

C .  karachiensis  appears  close  to  C .  alata 
Pruthi  in  general  appearance  as  well  as  pat- 
tern of  venation  and  genitalia,  but  differs  in 
the  shape  of  pygofer. 

Holotype  (male). — Pakistan:  Karachi,  grass, 
1.XI.85,  Qadeer.  One  paratype,  same  data  as 
holotype,  both  in  the  Zoological  Museum, 
University  of  Karachi,  Karachi,  Pakistan. 


14 


Great  Basin  Naturalist  Memoirs 


No.  12 


Figs.  3  A-J.  Chiasmus  karachiensis ,  n.  sp.:  A,  head  and  thorax,  dorsal  view;  B,  head,  ventral  view;  C,  forewing;  D, 
hind  wing;  E,  genital  capsule,  lateral  view;  F,  male  plates,  ventral  view;  G,  style,  dorsal  view;  H,  connective,  dorsal 
view;  I,  aedeagus,  lateral  view;  J,  aedeagus,  dorsal  view. 


Chiasmus  lohata  Ahmed  &  Qadeer,  n.  sp. 

(Figs.  4  A-J) 

Length, — Male  2.5  mm,  female  3.0  mm; 
head  projected  and  narrowed  in  front,  apex 
rounded;  median  length  of  crown  nearly  equal 
to  or  slightly  less  than  interocular  width; 
transocular  width  of  head  equal  to  transverse 
width  of  pronotum;  coronal  suture  present  up 
to  more  than  midlength;  ocelli  present  on 


anterolateral  margin  of  crown,  close  to  eyes; 
crown  possessing  few,  dim,  brownish  spots  in 
middle  along  posterior  margin;  frontoclypeus 
broad,  narrowed  in  apical  1/3;  anteclypeus 
longer  than  broad;  lora  broad;  pronotum  pos- 
sessing rows  of  brownish  markings. 

Forewing  with  apical  margin  smoothly 
rounded;  apical  cells  4;  appendix  broad, 
reaching  3rd  apical  cell;   subapical  cells  2; 


1988 


Ahmed  et  al.:  New  Grassland  Cicadellidae  from  Pakistan 


15 


Figs.  4  A-J.  Chiasmus  lobata,  n.  sp.:  A,  head  and  thorax,  dorsal  view;  B,  head,  ventral  view;  C,  forewing;  D,  hind 
wing;  E,  genital  capsule,  lateral  view;  F,  male  plate,  ventral  view;  G,  style,  dorsal  view;  H,  connective,  dorsal  view; 
I,  aedeagus,  lateral  view;  J,  aedeagus,  dorsal  view. 


veins    with    brownish    shades    along    their 
length;  hind  wing  with  3  closed  apical  cells. 

Male. — Plate  in  lateral  view  broad  at  base, 
narrowed  in  apical  half,  apex  rounded,  pos- 
sessing row  of  2-3  macrosetae  on  ventral  sur- 
face in  apical  half,  another  row  of  7-8  micro- 
setae  on  lateral  margin  from  near  base  to  mid- 
length,  length  of  plate  more  than  the  pygofer, 
apposed  mesally  at  base;  pygofer  in  lateral 
view  with  posterior  margin  broad,  dented, 
more  projected  posterodorsally,  disc  with 
dorsoventral  row  of  4  macrosetae;  anal  tube 
moderately  developed;  style  in  dorsal  view 


with  preapical  lobe  narrowed,  angular,  apical 
extension  thin,  spinose,  directed  laterad; 
aedeagus  in  lateral  view  with  preatrium  re- 
duced, dorsal  apodeme  present,  shaft  tubu- 
lar, somewhat  curved  dorsad  in  middle,  apex 
rounded  and  thin,  directed  caudad;  connec- 
tive Y-shaped,  with  arms  much  longer  than 
stem,  arms  confluent  again  forming  space  in 
middle. 

Chiasmus  lobata  appears  quite  close  to  C . 
karachiensis  as  well  as  to  C .  niger  Pruthi  and 
C .  jagdishi  Pruthi  in  the  pattern  of  male  geni- 
talia, but  differs  from  them  in  the  peculiar 


16 


Great  Basin  Naturalist  Memoirs 


No.  12 


shape  of  pygofer,  particularly  the  posterodor- 
sal  margin. 

HoLOTYPE  (male). — Pakistan:  Karachi, 
15. XII. 85,  grass,  Qadeer.  One  paratype, 
same  data  as  holotype,  both  in  Zoological 
Museum,  University  of  Karachi,  Karachi, 
Pakistan. 

Goniagnathus  Fieber 

The  genus  Goniagnathus  Fieber  was  re- 
described  by  Dlabola  (1954).  Distant  (1918) 
described  three  species  of  the  genus  from  In- 
dia. Ahmed  and  Yunus  (1986)  reported  G. 
guttulinervis  Kirsch.  from  Pakistan.  Another 
species  of  the  genus  is  described  here  from 
grass. 

Goniagnathus  bifurcattis  Ahmed  & 

Qadeer,  n.  sp. 

(Figs.  .5  A-I) 

Length. — Male  5.0  mm;  head  with  ante- 
rior margin  broadly  convex,  smoothly 
rounded  in  front;  median  length  of  crown 
much  less  than  the  interocular  width;  transoc- 
ular  width  of  head  slightly  more  than  the 
transverse  width  of  pronotum;  coronal  suture 
up  to  more  than  midlength;  ocelli  on  antero- 
lateral margin  of  head,  close  to  apex;  ante- 
clypeus  longer  than  broad;  lora  broad;  prono- 
tum with  few  brownish  black  dots  along 
anterior  margin  and  on  posterior  margin; 
scutellum  grayish  black  throughout. 

Forewing  with  apex  broad  and  rounded; 
maximum  width  of  wing  in  basal  half;  ap- 
pendix broad,  extending  to  4th  apical  cell; 
apical  cells  5;  subapical  cells  3;  black  markings 
present  in  appendix,  2nd,  3rd,  and  4th  apical 
cells,  in  some  discal  subapical  cells  and  along 
all  the  veins;  hind  wing  with  4  closed  apical 
cells. 

Male. — Genital  segment  strongly  sclero- 
tized,  dark  brown  in  color;  plates  in  ventral 
view  fused  completely  to  form  a  single  ovid 
structure,  width  of  plate  decreasing  smoothly 
to  rounded  apex,  with  few  blackish  macrose- 
tae  and  marginal  microsetae  in  apical  part; 
pygofer  in  lateral  view  with  posterior  margin 
broad,  nearly  truncate,  slightly  sinuate  pos- 
terodorsally,  with  at  least  three  prominent 
groups  of  stout  macrosetae,  one  closely  placed 
group  posterodorsally,  one  mixed  group  of 
short  and  long  in  middle  of  posterior  margin  in 
middle,  and  one  group  in  posteroventral  re- 
gion of  disc,  microsetae  dispersed  throughout 


pygofer;  anal  tube  moderately  long,  possess- 
ing fine  microsetae;  posterior  margin  of 
pygofer  finely  dentate;  style  in  dorsal  view 
with  preapical  lobe  minutely  developed,  api- 
cal part  expanded,  with  extreme  apex 
obliquely  truncate;  connective  in  dorsal  view 
rod-shaped,  arms  indistinct,  connective 
curved  in  middle;  aedeagus  in  lateral  view 
with  preatrium  reduced,  dorsal  apodeme  well 
developed,  closely  associated  with  connec- 
tive; shaft  of  aedeagus  short,  tapered  apically, 
long  processes  arising  from  atrium,  subparal- 
lel  to  shaft,  exceeding  shaft  in  length;  gono- 
pore  subterminal  on  dorsal  surface. 

Goniagnathus  bifurcatiis  is  close  to  G.  gut- 
tulinervis Kirschbaum  in  the  general  pattern 
of  male  genitalia,  particularly  the  male  plate 
and  aedeagus,  but  differs  in  the  shape  of  style, 
pygofer,  and  connective. 

HoLOTi'PE  (male). — Pakistan:  Karachi,  grass, 
10. XI. 85,  Qadeer.  One  paratype,  same  data 
as  holotype,  both  in  Zoological  Museum,  Uni- 
versity of  Karachi,  Karachi,  Pakistan. 

Literature  Cited 

Ahmed.  M  1985.  Typhlocybinae  of  Pakistan.  Pakistan 
Agric.  Res.  Council,  Islamabad.  270  pp. 

1986.  Some  investigations  of  leafhoppers  of  grass- 
lands and  allied  crops  in  Pakistan  (in  press). 

1986.  Some  investigations  of  lealhoppers  of  grass- 
lands in  Lower  Sind,  Pakistan  (in  press). 

Ahmed.  M  ,  andS  Rao.  1986.  Some  commonly  found  leaf 
and  planthoppers  on  vegetable  plants  in  the  sub- 
urbs of  Peshawar.  NWFP,  Pakistan  (in  press). 

Andrezejewska.  L.  1984.  Ecological  structure  of 
Auchenorrhyncha  meadow  communities  under 
increasing  cultivation.  Mitt.  Sch.  Entomol.  Ges. 
57(4):  40,5-406. 

Blocker,  H  D  .  T  L  Harvey,  and  J  L  Launchbaugh. 
1972.  Grassland  leafhoppers.  1.  Leafhopper  popu- 
lations of  upland  seeded  pastures  in  Kansas.  Ann. 
Entomol.  Soc.  Amer.  6.5(1);  166-172. 

Distant.  \V  L  1908.  The  fauna  of  British  India  including 
Ceylon  and  Burma.  Rh\nchota4:  1.56-419. 

1918.  The  fauna  of  British  India  including  Ceylon 

and  Burma.  Rhynchota  7;  1-109. 

Dlabola,  J  1954.  Fauna  CSR  Svazek-Homoptera.  Prace 
Ceskoslocenske  Akad.  1954:  7-339. 

Metcalf,  Z.  P  1967.  General  catalogue  of  the  Ho- 
moptera.  Fasc.  VI(IO).  2695pp. 

C)nl\n,  P  W  1949.  The  Nearctic  le;iflioppers  (Homop- 
tera:  Cicadellidae).  A  generic  classification  and 
check  list.  Mem.  Entomol.  Soc.  Washington  3: 
1-253. 

Pruthl  H.  S  1930.  Studies  on  Indian  Jassidae  (Ho- 
moptera).  Mem.  Indian  Mns.  11(1):  1-68. 

Theron.  J  G  1982.  Grassland  le;iflioppers  (Hemiptera: 
Cicadellidae)  from  Natal,  South  Africa,  with  de- 
scriptions of  new  genera  and  species.  Phytophy- 
lactica  14;  17-.30. 


1988 


Ahmed  et  al.;  New  Grassland  Cicadellidae  from  Pakistan 


17 


Figs.  5  A-I.  Goniagnathus  bifurcatus ,  n.  sp. :  A,  head  and  thorax,  dorsal  view;  B,  head,  ventral  view;  C,  forewing;  D, 
hind  wing;  E,  genital  capsule,  lateral  view;  H,  aedeagus  and  connective,  lateral  view;  I,  aedeagus,  dorsal  view. 


CLASSIFICATION  OF  THE  SUBGENUS  ATHYSANELLA,  GENUS 
ATHYSANELLA  BAKER  (HOMOPTERA,  CICADELLIDAE,  DELTOCEPHALINAE)^ 

H.  Derrick  Blocker^  and  James  W.  Johnson^ 

Abstract. — The  subgenus  Athysanella  Baker  is  reviewed  and  a  possible  phylogeny  of  the  group  is  discussed.  A  total 
of  35  species  is  recognized.  Descriptions,  illustrations,  and  a  key  to  males  are  presented.  Twelve  new  species  are 
described:  aphoda,  cursa,  deserta,furnaca,  hemijona,  krameri,  pastora,  strobila,  stylosa,  vativala,  and  whitcombi  are 
from  the  western  United  States,  and  marthae  is  from  Mexico. 


The  subgenera  of  Athysanella  Baker  have 
been  reviewed  by  Blocker  and  Johnson 
(1988).  The  nominate  subgenus  revised 
herein  contains  35  species,  of  which  12  are 
new.  Distribution  of  this  subgenus  is  primar- 
ily in  the  western  half  of  the  United  States, 
with  4  species  recorded  from  Mexico  and  4 
from  western  Canada.  The  hosts  are  primarily 
shortgrasses. 

Blocker  (1984)  has  described  the  morpho- 
logical variation  that  occurs  in  both  sexes  be- 
cause of  incompletely  developed  genital 
structures.  Undeveloped  and  partially  devel- 
oped specimens  are  often  difficult  to  identify. 
We  have  attempted  to  illustrate  sufficient 
characters  to  allow  identification  of  males,  but 
the  user  must  attempt  to  make  identifications 
from  a  series  of  specimens  that  appear  to  be 
fully  developed.  As  an  example,  the  hind  tib- 
ial spur  of  partially  developed  males  may  be 
absent  or  reduced;  the  genitalia  also  may  be 
reduced  in  size. 

Females  are  particularly  difficult  to  deter- 
mine. It  is  best  to  try  to  associate  them  with  a 
male  and  then  check  the  shape  of  the  posterior 
margin  of  sternum  VII.  There  is  much  in- 
fraspecific  variation  in  specimens  of  both 
sexes.  Ball  and  Beamer  (1940)  should  be  con- 
sulted for  additional  and  original  descriptions; 
Blocker  and  Wesley  (1985)  and  Wesley  and 
Blocker  (1985)  should  be  consulted  for  distri- 
bution in  Canada- Alaska  and  Mexico,  respec- 
tively. 

Depositories 

Persons  and  institutions  (acronyms  are  used 
in  the  text)  furnishing  material  for  this  study 
include  (the  authors  regret  any  omissions): 


UAz       -  University  of  Arizona,  F.  Werner. 

CAS       -  California  Academy  of  Science,  P.  Arnaud  and 

N.  Penny. 
CDA     -  California  Department  of  Agriculture,  R.  Gill 

and  J.  Sorensen. 
CNC     -  Canadian  National  Collection,  K.  G.  A.  Hamil- 
ton. 
IPL        -  Insect   Pathology   Laboratory,    US  DA,    Belts- 

ville,  A.  L.  Hicks  and  R.  F.  Whitcomb. 
KSU      -  Kansas  State  University. 
KU        -  Snow  Entomological  Museum,  University  of 

Kansas,  R.  W.  Brooks. 
MWN   -  Collection  of  M.  W.  Nielson. 
OhSU    -  Ohio  State  University,  P.  S.  Cwikla. 
OkSU    -  Oklahoma  State  University,  R.  A.  Drew. 
OrSU    -  Oregon  State  University,  P.  W.  Oman. 
UBC      -  University  of  British  Columbia,  G.  Scudder  and 

S.  Cannings. 
USNM  -  U.S.  National  Museum,  Natural  History,  J.  P. 

Kramer. 

Athysanella  (Athysanella)  Baker 

Athysanella  Baker,  1898;  185.  Type  species:  Athysanella 
magdalena  Baker,  1898,  by  original  designation. 

This  subgenus  is  characterized  by  the  pres- 
ence of  a  hind  tibial  spur  and  the  absence  of  a 
pygofer  process  (Blocker  and  Johnson  1988). 
It  is  a  sister  of  the  subgenus  Gladionura. 

Species  of  the  subgenus  Athysanella: 

aphoda  Blocker,  n.  sp.  Nevada. 

aspera  Ball  and  Beamer,  1940.  California. 

bifida  Ball  and  Beamer,  1940.  Colorado,  Montana,  New 

Mexico,    North    Dakota,    South    Dakota,    Texas, 

Wyoming,  Canada. 
cursa  Blocker,  n.  sp.  Arizona,  Colorado,  New  Mexico, 

Utah. 
deserta  Blocker,  n.  sp.  California. 
foeda  Ball  and  Beamer,  1940.  Wyoming. 
fredonia  Ball  and  Beamer,  1940.  Arizona,  Colorado,  New 

Mexico,  Nevada,  Utah. 
furnaca  Blocker,  n.  sp.  California. 
gardenia  Osborn,  1930.  Colorado,  Wyoming. 


'Contribution  88-.312-]  from  the  Kansas  Agriculture  Experiment  Station. 
^Department  of  EntomoloKy,  Kansa.s  State  University.  Manhattan,  Kansas  66506 
Department  of  Entomology,  University  of  Missouri,  Columbia,  Missouri  65221. 


18 


1988 


Blocker,  Johnson:  Subgenus  Athysanella 


19 


frigida  Oshorn,  1930. 

glohosa  Ball  and  Beanier,  1940.  Arizona. 

hemijona  Blocker,  n.  sp.  California. 

incerta  Ball  and  Beamer,  1940.  Colorado,  Kansa.s,  Mon- 
tana, Nebraska,  New  Mexico,  Wyoming. 

incongrua  Baker,  1898.  Colorado,  Iowa,  Kan.sas,  Ne- 
braska, New  Hampshire,  New  Mexico,  Okla- 
homa, South  Dakota,  Texas,  Wyoming. 

kadokana  Knull,  1951.  Colorado,  Montana,  Nebraska, 
South  Dakota,  Wyoming,  Canada. 

kanahana  Ball  and  Beamer,  1940.  Arizona,  Kansas,  New 
Mexico,  Utah. 

krameri  Blocker,  n.  sp.  California. 

laeta  Ball  and  Beamer,  1940.  Arizona,  New  Mexico. 

magdalena  Baker,  1898.  Arizona,  California,  Colorado, 
Montana,  New  Mexico,  Nevada,  Oklahoma, 
Texas,  Wyoming,  Mexico. 

inarthae  Blocker,  n.  sp.  Mexico. 

parca  Ball  and  Beamer,  1940.  Colorado,  New  Mexico. 

pastora  Blocker,  n.  sp.  New  Mexico. 

plana  Ball  and  Beamer,  1940.  California. 

planata  Ball  and  Beamer,  1940.  California. 

robusta  Baker,  1898.  Colorado,  Montana,  Nebraska, 
North  Dakota,  Wyoming,  Canada. 

montana  Osborn,  1930. 

rostrata  Ball  and  Beamer,  1940.  California. 

salsa  Ball  and  Beamer,  1940.  Kansas,  Texas,  Mexico. 

strobila  Blocker,  n.  sp.  California,  Idaho,  Nevada,  Ore- 
gon. 

stylosa  Blocker,  n.  sp.  New  Mexico. 

supina  Ball  and  Beamer,  1940.  Texas. 

tenera  Ball  and  Beamer,  1940.  Nevada. 

(ereforans  (Gillette  and  Baker),  1895.  Colorado,  Montana, 
North  Dakota,  Nebraska,  Utah,  Wyoming,  Can- 
ada. 

utahna  Osborn,  1930.  Arizona,  California,  Colorado, 
Idaho,  Nevada,  Utah,  Oregon,  Washington, 
Canada,  Mexico. 

vativala  Blocker,  n.  sp.  Nebraska,  North  Dakota. 

whitcombi  Blocker,  n.  sp.  Nevada. 

yumana  Osborn,  1930.  Arizona,  California. 

Key  to  Athysanella  Males,  sensu  stricto 

1.  Style    rounded    apically;    pygofer    variable; 

plates  embrowned  apically 2 

—  Style  widened,  usually  bifid  apically;  pygofer 
usually  broadly  rounded  apically;  plates  vari- 
able . ' ' 5 

2(1).        Pygofer  extended  apically;  connective,  plates, 

and  vertex  variable   3 

—  Pygofer  truncate  apically;  connective  shorter 
than  style;  plates  truncate  apically;  vertex 
usually  with  fuscous  spots  ....  gardenia  Osborn 

3(2).  Aedeagus  with  basal  process,  unique;  plates 
truncate  apically;  connective  as  long  as  style; 
vertex  without  fuscous  spots  ....    robusta  Baker 

—  Aedeagus  simple  or  with  ventral  serrations; 
connective  shorter  than  style;  vertex  usually 
with  fuscous  spots  4 

4(3).       Aedeagus  with  shaft  simple,  elongate;  plates 

rounded  apically,  embrowned  .  .   pastora,  n.  sp. 

—  Aedeagus  with  shaft  shortened,  expanded  and 
serrate  ventrally;  plates  truncate  apically  .  .  . 
globosa  Ball  &  Beamer 


5(1).  Aedeagus  with  conspicuous  apical  hook;  style 
exceeding  apex  of  plate;  vertex  with  or  with- 
out fuscous  spots  6 

—  Aedeagus  with  poorly  developed  or  no  apical 
hook  (apical  serrated  flanges  may  be  present); 
style  variable;  vertex  with  or  without  fuscous 
spots 22 

6(5).       Aedeagus  with  single  apical  hook 7 

—  Aedeagus  with  semicircular  apical  hook  ....    20 

7(6).  Aedeagus  not  widest  basally  (may  have  crests 
at  midlength),  not  tapered  apically;  plates 
rounded  apically;  vertex  with  or  without  fus- 
cous spots 14 

—  Aedeagus  widest  in  basal  1/2,  shaft  tapered  in 
apical  1/2,  apical  hook  conspicuous;  plates 
usually  truncate  apically;  vertex  with  or  with- 
out fuscous  spots  8 

8(7).  Aedeagus  with  distinct  lateral  tubercles  at 
base  of  shaft;  style  with  ventral  apical  arm 
widened,  dorsal  arm  slender;  vertex  without 
fuscous  spots    planata  Ball  &  Beamer 

—  Aedeagus  without  distinct  tubercles;  style 
with  ventral  and  dorsal  arms  variable;  vertex 
with  or  without  spots 9 

9(8).        Style  without  apical  lateral  flange,  dorsal  arm 

short,  wide;  plates  rounded  apically   

krameri,  n.  sp. 

—  Style  with  distinct  apical  lateral  flange  or  ven- 
tral arm  expanded  laterally,  dorsal  arm  vari- 
able, elongate;  plates  truncate  apically 10 

10(9).  Aedeagus  with  apex  of  dorsal  apodeme  ex- 
panded; style  with  dorsal  arm  elongate 11 

—  Aedeagus  with  dorsal  apodeme  rounded; 
style  variable  12 

11(10).   Style  with  dorsal  arm  slender;  vertex  with  no 

fuscous  spots;  smaller  in  overall  size 

whitcombi,  n.  sp. 

—  Style  with  dorsal  arm  more  robust;  vertex 
usually  with  fuscous  spots;  larger  in  size  .... 
plana  Ball  &  Beamer 

12(10).  Aedeagus  with  shaft  robust,  short;  style  with 

dorsal  apical  arm  longer,  variable  apically    .  .    13 

—  Aedeagus  with  shaft  less  robust;  style  with 

dorsal  arm  rounded  apically 

aspera  Ball  &  Beamer 

13(12).  Style  with  dorsal  arm  bulbous  apically;  aedea- 
gus with  apical  hook  conspicuous,  with  slight 
processes  at  base aphoda,  n.  sp. 

—  Style  with  dorsal  arm  acute  apically;  aedeagus 
with  apical  hook  less  developed,  no  process  at 
base strobila,  n.  sp. 

14(7).     Style  bifid  apically;  aedeagus  with  apical  hook 

short;  vertex  with  or  without  fuscous  spots  .  .    15 

—  Styles  not  bifid  apically  or  dorsal  arm  elon- 
gate; aedeagus  with  apical  hook  variable;  ver- 
tex with  or  without  fuscous  spots 17 

15(14).  Aedeagus  short,  shaft  serrate  ventrally, 
widest  at  midlength;  style  without  distinct 
lateral  apical  flange;  vertex  without  fuscous 
spots supina  Ball  &  Beamer 


20 


Great  Basin  Naturalist  Memoirs 


No.  12 


—  Aedeagus  longer,  shaft  not  serrate  or 
widened;  style  variable;  vertex  with  or  with- 
out fuscous  spots   16 

16(15).   Style  with  conspicuous  apical  lateral  flange; 

vertex  without  fuscous  spots 

foeda  Ball  &  Beamer 

—  Style  without  apical  lateral  flange;  vertex  with 
fuscous  spots    parca  Ball  &  Beamer 

17(14).   Style  with  dorsal  arm  elongate;  vertex  without 

fuscous  spots    rostrata  Ball  &  Beamer 

—  Style  without  elongate  dorsal  arm;  vertex  usu- 
ally with  fuscous  spots 18 

18(17).  Aedeagus  with  shaft  shortened,  widest  in  api- 
cal 1/2,  apical  hook  short;  style  with  conspicu- 
ous lateral  flange  19 

—  Aedeagus  with  shaft  longer,  not  widest  in  api- 
cal 1/2,  apical  hook  longer;  style  without  lat- 
eral flange fredonia  Ball  &  Beamer 

19(18).  Aedeagus  with  pronounced  keels  on  ventral 
margin  of  shaft,  shorter;  hind  tibial  spur  1/2 
1st  tarsomere;  from  SW    cursa,  n.  sp. 

—  Aedeagus  without  pronounced  keels  on  shaft, 
longer;  hind  tibial  spur  longer;  from  Califor- 
nia (east  face  of  Sierras)  hemijona,  n.  sp. 

20(6).  Style  with  apical  dorsal  arm  truncate;  vertex 
commonly  with  fuscous  spots;  plates  rounded 
apically incerta  Ball  &  Beamer 

—  Style  with  apical  dorsal  arm  rounded  or  acute; 
vertex  usually  without  fuscous  spots;  plates 
variable 21 

21(20).   Plates  rounded  apically;  style  with  ventral 

apical  arm  thickened,  well  developed 

kadokana  KnuU 

—  Plates  truncate  apically;  style  with  apical  lat- 
eral flange;  ventral  apical  arm  not  as  well  de- 
veloped; vertex  occasionally  with  fuscous 
spots salsa  Ball  &  Beamer 

22(5).  Vertex  usually  without  fuscous  spots;  aedea- 
gus with  serrated  ventral  margin;  plates  not 
elongate  23 

—  Vertex  usually  with  fuscous  spots,  or  if  absent, 
then  aedeagus  without  ventral  serrations; 
plates  may  be  elongate  29 

23(22).  Style  distinctly  bifid  apically;  connective 
shorter  than  style;  aedeagus  without  apical 
crests  on  shaft    25 

—  Style  with  only  dorsal  arm  developed;  con- 
nective variable  in  length;  aedeagus  with 
crests  on  shaft    24 

24(23).  Aedeagus  with  apical  crests  on  shaft;  connec- 
tive equal  to  style  in  length;  vertex  occasion- 
ally with  fuscous  spots 

kanabana  Ball  &  Beamer 

—  Aedeagus  with  medial  crests  on  shaft;  connec- 
tive shorter  than  style;  vertex  without  spots 
furnaca,  n.  sp. 

25(23).  Style  with  apical  lateral  flange,  apical  arms 
equal  in  length;  aedeagus  with  shaft  elongate 
'. 26 


—  Style  without  flange,  dorsal  arm  longer; 
aedeagus  with  shaft  shortened,  acute  apically 

!    27 

26(25).  Style  with  conspicuous  lateral  flange;  plates 
short,  truncate  apically;  vertex  length  less 
than  interocular  width    nmrthae,  n.  sp. 

—  Style  with  lateral  flange  smaller;  plates 
rounded  apically,  vertex  length  may  equal 
interocular  width bifida  Ball  &  Beamer 

27(25).   Style  conspicuously  bifid,  dorsal  arm  slender, 

tapered iitahna  Osborn 

—  Style  less  bifid,  dorsal  arm  robust   28 

28(27).   Style  with  dorsal  arm  acute  apically;  vertex 

occasionally  with  spots   yumana  Osborn 

—  Style  with  dorsal  arm  bulbous  apically;  vertex 
without  spots deserta,  n.  sp. 

29(22).  Tibial  spur  2/3  length  of  1st  tarsomere;  plates 
elongate,  acute  apically,  exceeding  apex  of 

styles;  outer  arm  of  style  acute  apically 

inagdalena  Baker 

—  Tibial  spur  variable;  plates  shorter;  style 
equal  to  or  exceeding  apex  of  plate,  outer  arm 
variable 30 

30(29).   Style  with  dorsal  arm  acute  apically 31 

—  Style  with  dorsal  arm  rounded  or  truncate  .  .    32 

31(30).  Aedeagus  with  shaft  approximately  1  1/2X 
length  of  dorsal  apodeme,  with  minute  serra- 
tions on  ventral  margin  .  .  .    laeta  Ball  &  Beamer 

—  Aedeagus  with  shaft  2X  length  of  dorsal 
apodeme,  with  conspicuous  serrations  on 
ventral  margin sttjlosa,  n.  sp. 

32(30).  Style  with  outer  arm  rounded;  aedeagus  with 
shaft  only  slightly  widened  or  tapered  in  api- 
cal 1/2,  ventral  margin  with  or  without  serra- 
tions       33 

—  Style  with  outer  arm  truncate;  aedeagus  with 
shaft  widest  in  apical  1/2,  ventral  margin  not 
serrated    incongrua  Baker 

33(32).  Aedeagus  with  shaft  short,  widest  at  mid- 
length;  style  with  large  lateral  apical  flange 
tenera  Ball  &  Beamer 

—  Aedeagus  with  shaft  longer,  even-margined 
or  slightly  enlarged  apically;  style  without 
conspicuous  flange    34 

34(33).  Style  deeply  bifid  apically,  ventral  arm 
widened,  dorsal  arm  even-margined;  aedea- 
gus with  shaft  slightly  widened  apically  .... 
vativala,  n.  sp. 

—  Style  less  bifid,  with  inconspicuous  apical  lat- 
eral flange,  dorsal  arm  tapered;  aedeagus  with 

shaft  even-margined 

terebrans  (Gillette  &  Baker) 

Character.s  Used  in  Character  Code 

Character].       0      no  fuscous  spots  on  vertex. 

1       fuscous  spots  may  be  present. 
Character  2.       0      hind  tibial  spur  1/2  1st  tarsomere  or 
less. 
1  =  spur  approximate]}  length  of  1st  tar- 
somere. 


1988 


Blocker,  Johnson:  Subgenus  Athysanella 


21 


character  3. 

Character  4. 
Characters. 

Character  6. 

Character  7. 
Character  8. 


0  =  ocellus  approximately  its  diameter 
from  eye. 

1  =  ocellus  more  remote. 

0  --^  male  plates  unicolorous. 

1  male  plates  embrowned  apically. 

0  ^  male  plates  elongate. 

1  ^  male  plates  rounded  apically. 

2  =  male  plates  truncate  apically. 

0  =  male  pygofer  with  caudal  margin 
rounded. 

1  =  male  pygofer  with  caudal  margin  ex- 
tended. 

0  ^  anal  tube  not  reaching  pygofer  apex. 

1  =  anal  tube  equal  to  or  exceeding 
pygofer  apex. 

0  =  male  pygofer  not  embrowned. 

1  =  male  pygofer  embrowned  caudoven- 

trally. 
Character  9.       0  =  male  plates  exceeding  apex  of 
pygofer. 

1  =  male  plates  equal  to  apex  of  pygofer. 

2  =  male  plates  not  reaching  apex  of 

pygofer. 
Character  10.     0  =  styles  not  reaching  apex  of  pygofer. 

1  =  styles  equal  to  apex  of  pygofer. 

2  =  styles  exceed  apex  of  pygofer. 
Character  11.     0  =  styles  not  reaching  apex  of  plates. 

1  =  styles  equal  to  apex  of  plates. 

2  =  styles  exceed  apex  of  plates. 
Character  12.     0  =  styles  without  lateral  flange. 

1  =  styles  with  lateral  flange. 

2  =  styles  with  ventral  arm  thickened. 
Character  13.     0  =  connective  shorter  than  style. 

1  =  connective  as  long  as  or  longer  than 
style. 
Character  14.     0  =  aedeagus  with  caudal  surface 
smooth. 
1  =  aedeagus  with  caudal  surface 
troughlike. 
Character  15.     0  =  shaft  of  aedeagus  flared  in  caudal 
aspect. 
1  =  shaft  even-margined  in  caudal  as- 
pect. 
Character  16.     0  =  aedeagus  with  no  basal  processes  on 
shaft. 
1  =  shaft  with  basal  processes. 
Character  17.     0  =  aedeagus  with  caudal  margin  of  shaft 
smooth. 
1  =  caudal  margin  serrate. 
Character  18.     0  =  aedeagus  with  dorsal  apodeme  sim- 
ple. 
1  =  dorsal  apodeme  enclosing  shaft 
basally. 

Athysanella  gardenia  Osborn 
Figs.  1-5 

Athysanella  gardenia  Osborn  1930:701. 
Gladionura  frigida  Osborn  1930:709. 

Length  of  male  2.5  to  2.9  mm,  female  3.2  to 
3.4  mm;  head  width  of  male  1.0  to  1.05  mm, 
female  1.05  to  1.15  mm;  pronotal  width  of 
male  0.95  to  1.05  mm,  female  1.0  to  1.1  mm; 
interocular  width  of  male  0.45  to  0.55  mm. 


female  0.5  to  0.55  mm;  vertex  length  of  male 
0.35  to  0.4  mm,  female  0.35  to  0.45  mm; 
pronotal  length  of  male  0.3  to  0.4  mm,  female 
0.3  to  0.35  mm.  Vertex  length  0.76  to  0.8 
interocular  width;  pronotal  length  0. 85  to  0. 88 
vertex  length. 

Character  code:  1-1-1-1-2-0-0-0-2-1-2-0-0- 
1-0-0-0-0. 

Lectotype,  male  (No.  43183),  and  lectoallo- 
type,  female  (No.  43183),  from  Garden  of  the 
Gods,  Colorado,  (Webster)  No.  7104,  in 
USNM. 

This  species  is  related  to  pastora  but  can  be 
distinguished  by  the  apex  of  the  style,  which  is 
narrowed  (Fig.  2),  and  the  posterior  margin  of 
the  plate,  which  is  embrowned  and  has  a  me- 
dian projection.  The  style  occasionally  has 
macrosetae;  long-winged  specimens  have 
been  observed.  This  species  has  been  col- 
lected in  Colorado  and  Wyoming. 

Athysanella  robusta  Baker 

Figs.  6-9 

Athysanella  robusta  Baker  1898:187. 
Athysanella  montana  Osborn  1930:700. 

Length  of  male  2.3  to  3. 1  mm,  female  3.4  to 
3.7  mm;  head  width  of  male  0.95  to  1. 15  mm, 
female  1.05  to  1.15  mm;  pronotal  width  of 
male  0.9  to  1. 15  mm,  female  1.05  to  1. 15  mm; 
interocular  width  of  male  0.4  to  0.55  mm, 
female  0.5  to  0.55  mm;  vertex  length  of  male 
0.35  to  0.5  mm,  female  0.4  to  0.5  mm;  prono- 
tal length  of  male  0.3  to  0.4  mm,  female  0.35 
to  0.4  mm.  Vertex  length  0.8  to  0.9  interocu- 
lar width;  pronotal  length  0.76  to  0.88  vertex 
width. 

Character  code:  0-0-1-1-2-1-1-0-2-0-2-0-1- 
0-0-1-0-0. 

The  type  of  this  species  has  not  been  stud- 
ied; identification  is  based  on  determined  ma- 
terial from  various  collections  and  the  litera- 
ture. 

Athysanella  robusta  is  related  to  pastora 
but  can  be  separated  by  the  unique  shape  of 
the  aedeagus  (Fig.  6),  the  pronotum  nearly  as 
wide  as  the  head,  and  the  connective,  which  is 
as  long  as  the  style.  There  are  no  fuscous  spots 
on  the  vertex  of  this  species.  It  has  been  col- 
lected in  Colorado,  Montana,  Nebraska, 
North  Dakota,  Wyoming,  and  Canada. 

Athysanella  pastora  Blocker,  n.  sp. 

Figs.  10-13 

Length  of  male  2.5  to  2.7  mm,  female  3.6  to 


22 


Great  Basin  Naturalist  Memoirs 


No.  12 


V^^ 


Figs.  1-1.3.  Figs.  1-5.  Athysanella  gardenia:  1,  aedeagus,  lateral  view;  2,  style,  broad  aspect;  3,  pygofer,  lateral  view; 
4,  plate,  ventral  view;  5,  female  sternum  VII,  ventral  view.  Figs.  6-9.  Athysanella  rohusta.  6,  aedeagus,  lateral  view; 
7,  style,  broad  aspect;  8,  pygofer,  lateral  view;  9,  female  sternum  VII,  ventral  view.  Figs.  10-13.  Athysanella  pastora: 
10,  aedeagus,  lateral  view;  11,  style,  broad  aspect;  12,  pygofer,  lateral  view;  13,  female  sternum  VII,  ventral  view. 


3.9  mm;  head  width  of  male  1.05  to  1.15  mm, 
female  1.2  to  1.25  mm;  pronotal  width  of  male 
0.9  to  1.0  mm,  female  1.05  to  1. 15  mm;  inter- 
ocular  width  of  male  0.4  to  0.5  mm,  female  0.5 
to  0.6  mm;  vertex  length  of  male  0.35  to  0.45 
mm,  female  0. 45  to  0. 5  mm ;  pronotal  length  of 
male  0.3  to  0.4  mm,  female  0.35  to  0.4  mm. 
Vertex  length  0.84  to  0.95  interocular  width; 
pronotal  length  0.76  to  0.94  vertex  length. 

Character  code:  1-1-0-1-1-1-0-1-1-2-2-0-0- 
0-0-0-0-0. 

Color   stramineous;    vertex   with    fuscous 


spots;  vertex,  pronotum,  and  abdomen  with 
additional,  conspicuous  brown  pattern;  fore- 
wings  commonly  with  darkened  stripes;  face 
with  lateral  fuscous  marks;  legs  with  various 
amounts  of  fuscous  coloring. 

Forewings  brachypterous,  exposing  3  to  4 
pregenital  abdominal  terga;  ocellus  approxi- 
mately 1.5  its  diameter  from  eye;  hind  tibial 
spur  3/4  length  of  1st  tarsomere;  female  ab- 
dominal sternum  VII  (Fig.  13)  variable  but 
hind  margin  usually  with  conspicuous  lateral 
lobes,  medial  third  truncate  or  rounded,  vari- 
ously embrowned. 


1988 


Blocker,  Johnson:  Subgenus  Athysanella 


23 


Pygofer  with  caudal  margin  extended, 
slightly  embrowned  on  caudoventral  border, 
densely  set  with  macrosetae  (25+);  anal  tube 
attains  apex  of  pygofer;  valve  with  caudal  mar- 
gin rounded;  plates  rounded  and  conspicu- 
ously embrowned  apically,  slightly  separated 
at  base;  connective  2/3  or  less  length  of  style; 
styles  rounded  apically,  with  conspicuous 
preapical  lobe,  apex  slightly  exceeding  apex  of 
plates;  aedeagus  with  shaft  simple,  slightly 
enlarged  apically,  elongate,  with  dorsal 
apodeme  slightly  overlapping  base  of  shaft. 

Holotype,  male,  from  Chaves  Co.,  New 
Mexico,  Bitter  Lake  NWR,  14  Aug  1984  (R.  F. 
Whitcomb);  4  male  and  8  female  paratypes, 
same  data.  Holotype  and  paratypes  in  USNM; 
paratypes  in  IPL  and  KSU. 

Athysanella  pastora  is  related  to  robusta 
and  glohosa  but  can  be  separated  from  the 
former  by  the  absence  of  an  aedeagal  process. 
It  can  be  separated  from  glohosa  by  the  shape 
of  the  apex  of  the  style,  which  is  not  widened 
(Fig.  11),  and  by  the  shaft  of  the  aedeagus 
(Fig.  10),  which  is  elongate  and  not  serrated. 
This  species  has  been  collected  only  on  gyp 
grama,  Bouteloua  hreviseta,  in  gypsum  flats  of 
southeastern  New  Mexico. 

Athysanella  glohosa  Ball  &  Beamer 

Figs.  14-15 

Athysanella  glohosa  Ball  &  Beamer  1940:18. 

Length  of  male  2.9  mm;  head  width  1.2 
mm;  pronotal  width  1.1  mm;  interocular 
width  0.45  mm;  vertex  length  0.35  mm; 
pronotal  length  0.4  mm.  Vertex  length  0.78 
interocular  width;  pronotal  length  1.07  vertex 
length.  Female  unknown. 

Character  code:  1-0-1-1-2-1-1-0-1-1-1-0-0- 
1-0-0-1-0. 

Holotype,  male,  from  Grand  Canyon,  Ari- 
zona, 4  Aug  1930  (E.  D.  Ball)  in  USNM.  Male 
and  female  paratypes,  same  data,  at  KU. 

Athysanella  glohosa  is  related  to  gardenia 
but  can  be  separated  by  the  shape  of  the  shaft 
of  the  aedeagus,  which  is  serrate  at  midlength 
(Fig.  14).  Female  paratypes  examined  are 
doubtfully  conspecific  with  the  male.  This 
species  is  known  only  from  its  type  locality  in 
Arizona;  a  specimen  from  Utah  is  assigned  to 
this  species  but  may  represent  a  new  species. 

Athysanella  planata  Ball  &  Beamer 

Figs.  16-18 

Athysanella  planata  Ball  &  Beamer  1940:24. 


Length  of  male  2.6  to  2.7  mm,  female  3.4  to 
4.0  mm;  head  width  of  male  1.0  to  1.05  mm, 
female  1. 1  to  1. 15  mm;  pronotal  width  of  male 
0.9  to  1.0  mm,  female  1.0  to  1.1  mm;  interocu- 
lar width  of  male  0.35  to  0.4  mm,  female  0.4  to 
0.5  mm;  vertex  length  of  male  0.35  mm  to  0.4 
mm,  female  0.4  to  0.45  mm;  pronotal  length  of 
male  0.3  to  0.35  mm,  female  0.3  to  0.35  mm. 
Vertex  length  0.87  to  1.0  interocular  width; 
pronotal  length  0.8  to  0.86  vertex  length. 

Character  code:  0-1-0-0-1-0-0-0-1-1-1-2-0- 
0-0-0-0-0. 

Holotype,  male,  and  allotype,  female,  from 
Newberry  Springs,  California,  30  July  1936 
(D.  R.  Lindsay  and  R.  H.  Beamer)  in  KU. 

Athysanella  planata  is  related  to  krameri 
but  can  be  separated  by  the  presence  of  pro- 
cesses on  the  base  of  the  shaft  of  the  aedeagus 
(Fig.  16),  a  more  slender  apical  arm  of  the 
style  (Fig.  17),  and  the  absence  of  fuscous 
spots  on  the  vertex.  This  species  has  been 
collected  only  from  the  type  locality  in  Cali- 
fornia. 

Athysanella  krameri  Blocker,  n.  sp. 

Figs.  19-21 

Length  of  male  3.0  to  3. 1  mm,  female  4.4  to 
4.7  mm;  head  width  of  male  1.05  to  1.15  mm, 
female  1.2  to  1.25  mm;  pronotal  width  of  male 
1.0  to  1.1  mm,  female  1.1  to  1.2  mm;  interocu- 
lar width  of  male  0.45  to  0.5  mm,  female  0.55 
to  0.6  mm;  vertex  length  of  male  0.35  to  0.45 
mm,  female  0.45  to  0.5  mm;  pronotal  length  of 
male  0.3  to  0.35  mm,  female  0.35  to  0.4  mm. 
Vertex  length  0.83  to  0.95  interocular  width; 
pronotal  length  0.77  to  0.87  vertex  length. 

Character  code:  1-1-0-0-1-1-0-0-2-1-2-0-0- 
0-0-0-0-0 . 

Color  stramineous;  vertex  commonly  with 
fuscous  spots,  often  with  orange  dashes;  male 
pygofer  commonly  with  conspicuous  fuscous 
spot;  remainder  of  body  mostly  unicolorous; 
female  without  fuscous  spots  on  vertex. 

Forewings  brachypterous,  exposing  3.5  to 
4.5  pregenital  abdominal  terga;  ocellus  1.5  its 
diameter  (or  less)  from  eye;  hind  tibial  spur 
approximately  3/4  length  of  1st  tarsomere;  fe- 
male abdominal  sternum  VII  (Fig.  21)  with 
elongate  lateral  lobes  and  rounded  medial 
lobe  on  hind  margin. 

Pygofer  rounded  apically,  with  about  20 
macrosetae;  valve  rounded  on  apical  margin; 
plates  rounded  apically,  slightly  separated  or 
touching  at  base;  connective  less  than  3/4 


24 


Great  Basin  Naturalist  Memoirs 


No.  12 


Figs.  14-27.  Fig.s.  14-15.  Athijsanella  globosa:  14,  aedeagus,  lateral  view;  15,  style,  broad  aspect.  Figs.  16-18. 
Athysanella  planata.  16,  aedeagus,  lateral  view;  17,  style,  broad  aspect;  18,  female  sternum  Vll,  ventral  view.  Figs. 
19-21.  Athysanella  krameri:  19,  aedeagus,  lateral  aspect;  20,  style,  broad  aspect;  21,  female  sternum  Vll,  ventral 
aspect.  Figs.  22-23.  Athysanella  whitcombi:  22,  style,  broad  aspect;  23,  female  sternum  Vll,  ventral  view.  Figs.  24-27. 
Athysanella  plana:  24,  aedeagus,  lateral  view;  25,  aedeagus,  anterodorsal  view;  26,  style,  broad  aspect;  27,  female 
sternum  VII,  ventral  view. 


length  of  style;  styles  (Fig.  20)  bifid  apically, 
dorsal  arm  short,  robust,  no  preapical  lobe, 
exceeding  apex  of  plates,  equal  to  apex  of 
pygofer;  aedeagus  (Fig.  19)  with  shaft  short, 
widest  basally,  and  tapered  to  acute  apical 
hook. 

Holotype,  male,  from  Little  Lake,  Inyo 
Co.,  California,  7  June  1929  (E.  P.  Van 
Duzee);  5  male  and  6  female  paratypes,  same 


data;  3  males  from  Olancha,  Inyo  Co.,  8  June 
1929;  1  male  from  Lone  Pine,  Inyo  Co.,  10 
June  1929.  Holotype  and  paratypes  in  CAS, 
paratypes  in  KSU. 

Athysanella  krameri  is  related  to  planata 
but  can  be  separated  by  the  absence  of  projec- 
tions at  the  base  of  the  aedeagus  and  the  dorsal 
arm  of  the  style,  which  is  shorter  and  more 
robust.  An  undeveloped  male  specimen  has 


1988 


Blocker,  Johnson:  Subgenus  Athysanella 


25 


been  observed.  This  species  is  known  only 
from  the  above  locahties  in  CaHfornia. 

Athysanella  whitcombi  Blocker,  n.  sp. 

Figs.  22-23 

Length  of  male  2.6  mm,  female  4.0  mm; 
head  width  of  male  1.0  mm,  female  1.15  to  1.2 
mm;  pronotal  width  of  male  0.9  mm,  female 
1.0  to  1.05  mm;  interocular  width  of  male  0.4 
mm,  female  0.55  to  0.6  mm;  vertex  length  of 
male  0.35  to  0.4  mm,  female  0.5  mm;  pronotal 
length  of  male  0.25  to  0.3  mm,  female  0.45 
mm.  Vertex  length  0.93  to  0.94  interocular 
width;  pronotal  length  0.73  to  0.74  vertex 
length. 

Character  code:  0-1-1-0-1-0-0-0-1-1-1-1-0- 
0-0-0-0-1. 

Color  stramineous;  vertex  without  fuscous 
spots;  body  virtually  unicolorous. 

Forewings  of  male  brachypterous,  exposing 
4.5  pregenital  abdominal  terga;  ocellus  ap- 
proximately 2X  its  diameter  from  eye;  hind 
tibial  spur  about  1/2  length  of  1st  tarsomere; 
female  abdominal  sternum  VII  with  lateral 
lobes,  without  medial  lobe  on  caudal  margin. 

Pygofer  rounded,  with  about  20  macrose- 
tae;  valve  rounded  on  caudal  margin;  plates 
rounded  apically,  touching  at  base  then  diver- 
gent apically;  connective  3/4  length  of  style; 
styles  bifid  apically,  dorsal  arm  slender,  no 
preapical  lobe,  exceeding  apex  of  plates, 
equal  to  apex  of  pygofer;  aedeagus  with  shaft 
and  dorsal  apodeme  as  plana. 

Holotype,  male,  from  Austin,  Nevada,  12 
Aug  1940  (D.  E.  Hardy)  and  a  long- winged 
female  paratype,  same  data,  in  KU. 

Athysanella  whitcombi  is  related  to  plana 
but  can  be  separated  by  its  smaller  size,  the 
absence  of  fuscous  spots  on  the  vertex,  and  the 
more  slender  dorsal  apical  arm  of  the  style. 
This  species  is  named  for  R.  F.  Whitcomb, 
who  has  made  numerous  contributions  to  this 
research;  it  has  been  collected  in  Nevada. 

Athysanella  plana  Ball  &  Beamer 

Figs.  24-27 

Athysanella  plana  Ball  &  Beamer  1940:13. 

Length  of  male  3.0  to  3.3  mm,  female  4.5  to 
4.7  mm;  head  width  of  male  1.05  to  1.15  mm, 
female  1.2  to  1.25  mm;  pronotal  width  of  male 
1.0  to  1.05  mm,  female  1.15  to  1.2  mm;  intero- 
cular width  of  male  0.45  to  0.5  mm,  female 
0.55  to  0.6  mm;  vertex  length  of  male  0.4  to 
0.45  mm,  female  0.45  to  0.5  mm;  pronotal 


length  of  male  0.3  to  0.4  mm,  female  0.35  to 
0.4  mm.  Vertex  length  0.89  to  0.95  interocu- 
lar width;  pronotal  length  0.77  to  0.82  vertex 
length. 

Character  code:  1-1-1-0-2-0-0-0-1-1-1-1-0- 
0-0-0-0-0. 

Holotype,  male,  and  allotype,  female,  from 
Califa,  California,  12  June  1935  (P.  W.  Oman) 
in  USNM;  paratypes  in  USNM. 

Athysanella  plana  is  related  to  aspera  but 
can  be  separated  by  the  expanded  base  of  the 
dorsal  apodeme  of  the  aedeagus  (Fig.  25)  and 
the  longer  dorsal  apical  arm  of  the  style  (Fig. 
26).  This  species  has  been  collected  only  from 
the  Central  Valley  (Fresno  Co.)  of  California. 

Athysanella  aspera  Ball  &  Beamer 
Figs.  28-30 

Athysanella  aspera  Ball  &  Beamer  1940:17. 

Length  of  male  2.5  to  3. 1  mm,  female  4.3  to 
5.0  mm;  head  width  of  male  0.95  to  1. 15  mm, 
female  1. 15  to  1.3  mm;  pronotal  width  of  male 
0.9  to  1.1  mm,  female  1.1  to  1.25  mm;  interoc- 
ular width  of  male  0.4  to  0.5  mm,  female  0.45 
to  0.5  mm;  vertex  length  of  male  0.35  to  0.45 
mm,  female  0.4  to  0.5  mm;  pronotal  length  of 
male  0.3  to  0.4  mm,  female  0.35  to  0.45  mm. 
Vertex  length  0.8  to  0.95  interocular  width; 
pronotal  length  0.76  to  0.93  vertex  length. 

Character  code:  1-1-1-0-2-1-0-0-2-1-2-1-0- 
0-0-0-0-0. 

Holotype,  male,  and  allotype,  female,  from 
Mojave,  California,  1  July  1931  (E.  D.  Ball)  in 
USNM. 

Athysanella  aspera  is  related  to  plana  but 
can  be  separated  by  the  apex  of  the  style, 
which  has  a  shorter  dorsal  arm  (Fig.  29),  and 
the  dorsal  apodeme  of  the  aedeagus,  which  is 
not  widened  apically.  In  one  specimen,  the 
vertex  length  equaled  the  interocular  width. 
The  shaft  of  the  aedeagus  has  an  indistinct  pair 
of  basal  tubercles  similar  to  planata.  A  male 
specimen  parasitized  by  Pipinculidae  and 
specimens  with  undeveloped  genitalia  have 
been  observed.  This  species  has  been  col- 
lected from  the  Mojave  Desert  in  California. 

Athysanella  aphoda  Blocker,  n.  sp. 

Figs.  31-32 

Length  of  male  3.5  mm;  head  width  1.2 
mm;  pronotal  width  1.1  mm;  interocular 
width  0.5  mm;  vertex  length  0.45  mm;  prono- 
tal length  0.4  mm;  female  unknown.  Vertex 
0.9  interocular  width;  pronotal  length  0.83  to 


26 


Great  Basin  Naturalist  Memoirs 


No.  12 


Figs.  28-39.  Figs.  28-30.  Athysanella  aspera:  28,  aedeagus,  lateral  view;  29,  style,  broad  aspect;  30,  female  sternum 
VII,  ventral  view.  Figs.  31-32.  Athysanella  aphoda:  31,  aedeagus,  lateral  view;  32,  style,  broad  aspect.  Figs.  33-36. 
Athysanella  strobila:  33,  aedeagus,  lateral  view;  .34,  style,  broad  aspect;  35,  pygofer,  lateral  aspect;  36,  female  sternum 
VII,  ventral  view.  Figs.  37-.39.  Athysanella  supina:  37,  aedeagus,  lateral  view;  38,  style,  broad  aspect;  39,  female 
sternum  VII,  ventral  view. 


0.84  vertex  length.  Female  unknown. 

Character  code:  1-0-1-0-2-0-1-0-2-1-2-2-0- 
0-0-1-0-0. 

Color  stramineous;  vertex  and  pronotum 
with  distinct,  darker  pattern;  forewings  with 
dark  stripes;  face  with  distinct  lateral  arcs;  legs 
with  slight  fuscous  coloring. 

Forewings  brachypterous,  exposing  3.5 
pregenital  abdominal  terga;  ocellus  approxi- 


mately its  diameter  from  eye;  hind  tibial  spur 
slightly  shorter  than  1st  tarsomere. 

Pygofer  as  strobila  but  larger;  valve 
rounded  posteriorly;  plates  truncate  apically; 
styles  (Fig.  32)  with  dorsal  arm  bulbous  api- 
cally; aedeagus  (Fig.  31)  with  apical  hook 
more  distinct  than  strobila ,  shaft  with  indis- 
tinct, small  process  at  base. 

Holotype,  male,  from  Lander  Co. ,  Nevada, 


1988 


Blocker,  Johnson:  Subgenus  Athysanella 


27 


30  mi  N  Austin,  24  Julv  1986  (H.  D.  Blocker) 
in  KSU. 

Athysanella  aphoda  is  closely  related  to  and 
possibly  conspecific  with  strohila  but  can  be 
readily  separated  by  its  larger  size,  the  bul- 
bous apex  of  the  dorsal  arm  of  the  style,  and 
the  more  robust  aedeagus,  which  has  a  more 
pronounced  apical  hook.  This  species  is  repre- 
sented by  the  holotype  from  Nevada. 

Athysanella  strohila  Blocker,  n.  sp. 

Figs.  33-36 

Length  of  male  2. 6  to  3. 9  mm,  female  4. 2  to 
5.3  mm;  head  width  of  male  1.05  to  1.3  mm, 
female  1.15  to  1.35  mm;  pronotal  width  of 
male  0.95  to  1.25  mm,  female  1.05  to  1.3  mm; 
interocular  width  of  male  0.4  to  0.55  mm, 
female  0.45  to  0.65  mm;  vertex  length  of  male 
0.4  to  0.55  mm,  female  0.45  to  0.65  mm; 
pronotal  length  of  male  0.3  to  0.4  mm,  female 
0.3  to  0.45  mm.  Vertex  length  0.89  to  1.0 
interocular  width;  pronotal  length  0.7  to  0.82 
vertex  length. 

Character  code:  0-1-0-0-2-0-0-0-2-1-2-2-0- 
1-0-0-0-0. 

Color  stramineous,  occasionally  patterned; 
vertex  without  fuscous  spots;  abdomen  occa- 
sionally with  some  darker  pattern;  other  body 
areas  occasionally  with  irregular,  brown  pat- 
tern. 

Forewings  brachypterous,  exposing  3.0  to 
4.5  pregenital  abdominal  terga;  ocellus  ap- 
proximately its  diameter  from  eye;  hind  tibial 
spur  3/4  length  of  1st  tarsomere;  female  ab- 
dominal sternum  VII  (Fig.  36)  with  conspicu- 
ous lateral  lobes  on  posterior  margin,  with  a 
broadly  rounded  medial  lobe. 

Pygofer  rounded  on  caudoventral  margin, 
with  a  distinct  lobe  (Fig.  35),  with  15  or  fewer 
macrosetae;  valve  rounded  posteriorly;  plates 
truncate  apically,  nearly  touching  basally, 
then  diverging;  connective  3/4  length  of  style; 
styles  bifid  (Fig.  34)  apically,  with  ventral  arm 
thickened,  exceeding  apex  of  plates,  equal  to 
apex  of  pygofer;  aedeagus  with  shaft  short, 
widest  in  proximal  1/2,  slightly  tapered  in  api- 
cal 1/2  to  a  single  apical  hook. 

Holotype,  male,  from  Lone  Pine,  Califor- 
nia, 28  July  1940  (R.  H.  Beamer);  4  male  and  3 
female  paratypes,  same  data;  male  and  fe- 
male, Fallon,  Nevada,  12  Aug  1940  (L.  C. 
Kuitert);  2  males  and  8  females.  New  Hum- 
boldt Co.,  Nevada,  7  mi  W  Denio,  23  June 
1971  (Oman);  6  males  and  3  females.  Eureka 


Co.,  Nevada,  W.  Eureka,  25  July,  1986  (H. 
D.  Blocker);  male  and  female,  Lander  Co., 
Nevada,  5  mi  W  Austin,  24  July  1986  (H.  D. 
Blocker);  male,  Owyhee  Co.,  Idaho,  10  mi  W 
Bruneau,  9  July  1981  (H.  D.  Blocker,  J.  W. 
Johnson);  numerous  specimens  from  Albert 
Lake,  Oregon,  5  mi  N  Valley  Falls,  18  May 
1969  (Oman),  and  6  mi  N  Silver  Lake,  Ore- 
gon, 17  May  1969  (Oman).  Holotype  and 
paratvpes  in  KU;  paratypes  in  KSU  and 
OrSU. 

Athysanella  strohila  is  closely  related  to  as- 
pera  but  can  be  separated  by  the  shape  of  the 
shaft  of  the  aedeagus,  which  is  more  robust 
and  shorter,  and  by  the  dorsal  apical  arm  of 
the  style,  which  is  longer  and  acute  apically.  It 
can  be  recognized  by  the  unique  lobe  on  the 
caudal  margin  of  the  pygofer.  An  undevel- 
oped female  specimen  has  been  examined. 
This  species  has  been  collected  in  California, 
Idaho,  Nevada,  and  Oregon. 

Athysanella  supina  Ball  &  Beamer 

Figs.  37-39 

Athysanella  supina  Ball  &  Beamer  1940:22. 

Length  of  male  2.8  to  3.2  mm,  female  4.3  to 
4.6  mm;  head  width  of  male  1.0  to  1.1  mm, 
female  1.15  to  1.25  mm;  pronotal  width  of 
male  1.0  to  1.05  mm,  female  1.1  to  1.2  mm; 
interocular  width  of  male  0.45  to  0.5  mm, 
female  0.55  to  0.6  mm;  vertex  length  of  male 
0.45  to  0.55  mm,  female  0.5  to  0.6  mm;  prono- 
tal length  of  male  0.35  to  0.4  mm,  female  0.35 
to  0.45  mm.  Vertex  length  1.0  to  1. 15  interoc- 
ular width;  pronotal  length  0.68  to  0.75  vertex 
length. 

Character  code:  1-1-1-0-1-0-0-0-2-1-2-1-0- 
1-0-0-1-0. 

Holotype,  male,  and  allotype,  female,  from 
Boca  Chica,  Texas,  30  May  1933  (P.  W. 
Oman)  in  USNM;  paratypes  in  USNM  and 
KU. 

Athysanella  supina  is  related  to  fredonia 
but  can  be  separated  by  the  shaft  of  the  aedea- 
gus, which  has  a  shorter  crest  on  the  ventral 
margin  (Fig.  37),  and  the  shape  of  the  apex  of 
the  style,  which  is  more  bifid  (Fig.  38).  This 
species,  described  from  Texas,  has  recently 
been  collected  there  (A.  L.  Hicks)  from  shore- 
grass,  Monathochloe  littoralis. 

Athysanella  foeda  Ball  &  Beamer 

Figs.  40-43 

Athysanella  foeda  Ball  &  Beamer  1940:19. 


28 


Great  Basin  Naturalist  Memoirs 


No.  12 


Figs.  40-.52.  Figs.  40-43.  Athysanella  foeda:  40,  head  and  prothorax,  dorsal  \'ievv;  41,  aedeagus,  lateral  view;  42, 
style,  broad  aspect;  4.3,  female  sternum  VII,  ventral  view.  Figs.  44-46.  Athysanella  parca:  44,  aedeagus,  lateral  view; 
45,  style,  broad  aspect;  46,  female  sternum  VII,  ventral  view.  Figs.  47-49.  Athysanella  rostrata:  47,  aedeagus,  lateral 
view;  48,  style,  broad  aspect;  49,  female  sternum  VII,  ventral  aspect.  Figs.  50-52.  Athysanella  fredonia:  50,  aedeagus, 
lateral  view;  51,  style,  broad  aspect;  52,  female  sternum  VII,  ventral  view. 


Length  of  male  3.4  to  3.6  mm,  female  4.9  to 

5.1  mm;  head  width  of  male  1.2  to  1.3  mm, 
female  1.4  to  1.5  mm;  pronotal  width  of  male 

1.2  to  1.3  mm,  female  1.35  to  1.4  mm;  interoc- 
ular  width  of  male  0.55  to  0.6  mm,  female  0.65 
to  0.75  mm;  vertex  length  of  male  0.5  to  0.55 
mm,  female  0.55  to  0.6  mm;  pronotal  length  of 


male  0.4  to  0.45  mm,  female  0.4  to  0.45  mm. 
Vertex  length  0.83  to  0.88  interocular  width; 
pronotal  length  0.76  to  0.85  vertex  length. 

Character  code:  0-1-1-0-1-0-0-0-1-2-2-1-0- 
1-0-0-0-1. 

Holotype,  male,  and  allotype,  female,  from 
Laramie,  Wyoming,  23  June  1935  (R.  H.  and 


1988 


Blocker,  Johnson:  Subgenus  Athysanella 


29 


J.  Beamer)  in  KU;  paratypes  in  KU. 

Athysanella  foeda  is  related  to  parca  but 
can  be  separated  by  its  larger  size  and  the 
presence  of  a  conspicuous  lateral  lobe  at  the 
apex  of  the  style  (Fig.  42).  It  can  be  separated 
from  fredonia  by  the  shape  of  the  apex  of  the 
style,  which  is  distinctly  bifid.  This  species 
has  been  collected  in  Wyoming. 

Athysanella  parca  Ball  &  Beamer 

Figs.  44-46 

Athysanella  parca  Ball  &  Beamer  1940:15. 

Length  of  male  2.7  to  3.4  mm,  female  3.9  to 
4.4  mm;  head  width  of  male  1.0  to  1.15  mm, 
female  1. 15  to  1.4  mm;  pronotal  width  of  male 
0.9  to  1.15  mm,  female  1.05  to  1.25  mm; 
interocular  width  of  male  0.4  to  0.5  mm,  fe- 
male 0.5  to  0.65  mm;  vertex  length  of  male  0.4 
to  0.45  mm,  female  0.45  to  0.55  mm;  pronotal 
length  of  male  0.3  to  0.4  mm,  female  0.4  to 
0.45  mm.  Vertex  length  0.89  to  0.95  interocu- 
lar width;  pronotal  length  0.82  to  0.88  vertex 
length. 

Character  code:  1-1-0-0-1-0-0-0-2-1-2-2-0- 
0-0-0-0-0. 

Holotype,  male,  and  allotype,  female,  from 
Belen,  New  Mexico,  20  July  1936  (R.  H. 
Beamer  and  D.  R.  Lindsay)  in  KU;  paratypes 
inKU. 

Athysanella  parca  is  related  to  foeda  but 
can  be  separated  by  the  absence  of  a  lateral 
flange  on  the  style  and  the  usual  presence  of 
fuscous  spots  on  the  vertex.  It  can  be  sepa- 
rated from  incerta  by  the  shape  of  the  apex  of 
the  shaft  of  the  aedeagus,  which  has  a  single 
acute  apical  hook  (Fig.  44).  A  long-winged 
female  has  been  examined.  This  species  has 
been  collected  in  Colorado  and  New  Mexico; 
in  New  Mexico,  the  host  is  saltgrass,  Distichlis 
stricta. 

Athysanella  rostrata  Ball  &  Beamer 

Figs.  47-49 

Athysanella  rostrata  Ball  &  Beamer  1940:21. 

Length  of  male  3.5  mm,  female  5.0  to  5.1 
mm;  head  width  of  male  1.15  to  1.25  mm, 
female  1.3  to  1.35  mm;  pronotal  width  of  male 
1.1  to  1.15  mm,  female  1.2  to  1.25  mm;  intero- 
cular width  of  male  0.5  to  0.55  mm,  female  0.6 
to  0.65  mm;  vertex  length  of  male  0.45  to  0.5 
mm,  female  0.5  to  0.55  mm;  pronotal  length  of 
male  0.35  to  0.4  mm,  female  0.4  to  0.45  mm. 
Vertex  length  0.86  to  0.91  interocular  width; 


pronotal  length  0.75  to  0.78  vertex  length. 

Character  code:  0-1-0-0-2-0-0-0-2-2-2-2-0- 
1-0-0-0-0. 

Holotype,  male,  and  allotype,  female,  from 
Perris,  California,  5  June  1935  (P.  W.  Oman) 
in  USNM;  paratypes  in  USNM. 

Athysanella  rostrata  is  related  to  foeda  but 
can  be  separated  by  the  distinctly  elongate 
dorsal  arm  of  the  style  (Fig.  48).  A  paratype 
with  undeveloped  genitalia  has  been  exam- 
ined. This  species  has  been  collected  only 
from  its  type  locality  in  southern  California. 

Athysanella  fredonia  Ball  &  Beamer 

Figs.  50-52 

Athtjsanella  fredonia  Ball  and  Beamer,  1940:12  (in  part). 

Length  of  male  2.6  to  2.9  mm,  female  3.9  to 
4.3  mm;  head  width  of  male  1.0  to  1.15  mm, 
female  1. 15  to  1.4  mm;  pronotal  width  of  male 
0.95  to  1.1  mm,  female  1.0  to  1.25  mm;  intero- 
cular width  of  male  0.4  to  0.45  mm,  female 
0.45  to  0.6  mm;  vertex  length  of  male  0.35  to 
0.4  mm,  female  0.4  to  0.5  mm;  pronotal 
length  of  male  0.3  to  0.35  mm,  female  0.35  to 
0.4  mm.  Vertex  length  0.83  to  0.94  interocu- 
lar width;  pronotal  length  0.75  to  0.87  vertex 
length. 

Character  code:  1-1-0-0-1-0-0-0-1-1-1-0-0- 
1-0-0-1-0. 

Color  stramineous;  vertex  with  fuscous 
spots  anteriorly;  pronotum  commonly  with 
small  brown  spots;  other  dorsal  body  pattern 
variable;  face  with  lateral  fuscous  marks;  legs 
with  various  amounts  of  fuscous  coloring. 

Forewings  brachypterous,  exposing  3.5  to 
4.0  pregenital  abdominal  terga;  ocellus  ap- 
proximately its  diameter  from  eye;  hind  tibial 
spur  over  3/4  length  of  1st  tarsomere;  female 
abdominal  sternum  VII  (Fig.  52)  with  lateral 
lobes  extending  posteriorly  slightly  more  than 
median  lobe. 

Pygofer  rounded  apically,  densely  set  with 
short  setae;  valve  rounded  on  apical  margin; 
plates  rounded  apically,  slightly  separated  at 
base;  connective  shorter  than  style;  styles 
(Fig.  51)  widened  apically,  nearly  truncate, 
dorsal  arm  acute  apically,  curved,  usually 
with  conspicuous  lateral  flange,  equal  to  apex 
of  plates  and  pygofer;  aedeagus  (Fig.  50)  with 
shaft  not  widened  apically,  finely  serrate  on 
ventral  margin. 

Holotype,  male,  from  Fredonia,  Arizona,  6 
Aug  1930  (E.  D.  Ball)  in  USNM. 


30 


Great  Basin  Naturalist  Memoirs 


No.  12 


Athysanella  fredonia  is  related  to  cursa  but 
can  be  separated  by  the  longer  shaft  of  the 
aedeagus,  which  is  not  widened  apically.  Ex- 
amination of  the  holotype  has  revealed  that 
the  male  genitalia  differ  from  the  genitalia 
described  by  Ball  and  Beamer  (1940).  For  this 
reason,  fredonia  is  redescribed  and  a  new 
species,  cursa,  which  agrees  with  the  pub- 
lished illustration,  is  described.  It  has  been 
collected  in  Arizona,  Colorado,  New  Mexico, 
Nevada,  and  Utah,  where  it  occurs  on  galleta, 
Hilariajamesii.  At  the  type  locality,  it  occurs 
with  cursa  on  the  same  host  (R.  F.  Whit- 
comb). 

Athysanella  cursa  Blocker,  n.  sp. 

Figs.  53-55 

Athysanella  fredonia  Ball  &  Beamer  (in  part). 

Length  of  male  2.5  to  3.0  mm,  female  4.0  to 

4.4  mm;  head  width  of  male  1.05  to  1. 15  mm, 
female  1.2  to  1.3  mm;  pronotal  width  of  male 
0.95  to  1.05  mm,  female  1.1  to  1.2  mm;  intero- 
cular  width  of  male  0.4  to  0.45  mm,  female  0.5 
to  0.6  mm;  vertex  length  of  male  0.35  to  0.4 
mm,  female  0.45  to  0.5  mm;  pronotal  length  of 
male  0.3  to  0.35  mm,  female  0.35  to  0.4  mm. 
Vertex  length  0.88  to  0.94  interocular  width; 
pronotal  length  0.8  to  0.88  vertex  length. 

Character  code:  0-1-0-0-1-0-0-0-1-1-1-1-0- 
1-0-0-1-0. 

Color  stramineous  with  fuscous  spots  on 
vertex;  vertex,  pronotum,  and  abdomen  vari- 
ously patterned  with  brown  markings; 
forewings  commonly  with  brown  stripes;  face 
with  lateral  fuscous  marks,  legs  with  various 
amounts  of  fuscous  coloring. 

Forewings  brachypterous,  exposing  3.5  to 

4.5  pregenital  abdominal  sterna;  ocellus  ap- 
proximately 1.5X  its  diameter  from  eye;  hind 
tibial  spur  usually  about  1/2  length  of  1st  tar- 
somere;  female  abdominal  sternum  VII  (Fig. 
55)  with  lateral  lobes  slightly  longer  than  me- 
dian lobe. 

Pygofer  rounded  but  slightly  extended  api- 
cally; densely  set  with  short  setae;  valve  with 
caudal  margin  rounded;  plates  rounded  api- 
cally; connective  shorter  than  styles;  styles 
(Fig.  54)  widened  apically  and  shallowly  bifid, 
dorsal  arm  narrowed  and  acute  apically,  usu- 
ally with  a  conspicuous  lateral  flange,  equal  to 
apex  of  plates  and  pygofer;  aedeagus  (Fig.  53) 
with  shaft  short,  lightly  serrate  on  ventral 
margin,  with  pronounced  flanges  on  ventral 
margin,  slightly  widest  in  apical  1/2. 


Holotype,  male,  from  Whites  City,  New 
Mexico,  13  July  1936  (R.  H.  Beamer).  One 
male  and  two  female  paratypes  from  Flagstaff, 
Arizona,  27  July  1936  (R.  H.  Beamer);  two 
males  from  Flagstaff,  Arizona,  21  July  1936 
(D.  R.  Lindsey);  female  from  St.  Johns,  Ari- 
zona, 26  July  1936  (R.  H.  Beamer);  two  males 
from  Estancia,  New  Mexico,  24  June  1940  (R. 
H.  Beamer);  five  males  and  one  female  from 
Cuervo,  New  Mexico,  23  June  1940  (R.  H. 
Beamer).  Holotype  in  KU;  paratypes  in  KU 
and  KSU. 

Athysanella  cursa  is  closely  related  to  hemi- 
jona  but  can  be  separated  by  the  shorter 
aedeagus  with  pronounced  flanges  on  the  ven- 
tral margin,  the  shorter  tibial  spurs,  and  the 
pygofer,  which  is  slightly  extended  apically. 
The  plates  occasionally  have  a  macroseta. 
Long-winged  specimens  and  specimens  para- 
sitized by  Dryinidae  have  been  observed. 
This  species  is  illustrated  as  fredonia  in  Ball 
and  Beamer  (1940),  but  a  study  of  the  holo- 
type of  fredonia  reveals  that  this  is  a  different 
species.  It  has  been  reported  from  the  Desert 
Plains  region  of  Arizona,  Colorado,  New  Mex- 
ico, and  Utah,  where  it  occurs  on  galleta, 
Hilariajamesii. 


Athysanella  he?nijona  Blocker,  n. 

Figs.  ,56-58 


sp. 


Length  of  male  3.2  to  3.3  mm,  female  4.8  to 
4.9  mm;  head  width  of  male  1.2  to  1.25  mm, 
female  1.3  to  1.4  mm;  pronotal  width  of  male 
1. 1  to  1. 15  mm,  female  1.25  to  1.3  mm;  intero- 
cular width  of  male  0.45  to  0.55  mm,  female 
0.5  to  0.55  mm;  vertex  length  of  male  0.4  to 
0.45  mm,  female  0.5  to  0.55  mm;  pronotal 
length  of  male  0.35  to  0.4  mm,  female  0.4  to 
0.45  mm.  Vertex  length  0.85  to  0.95  interocu- 
lar width;  pronotal  length  0.73  to  0.83  vertex 
length. 

Character  code:  1-1-0-0-1-0-0-0-1-1-1-1-0- 
1-0-0-1-1. 

Color  stramineous;  vertex  with  fuscous 
spots;  vertex,  pronotum,  and  abdomen  with 
conspicuous,  dark  pattern;  forewings  with 
darkened  stripes;  face  with  lateral  fuscous 
marks;  legs  with  various  amounts  of  fuscous 
coloring. 

Forewings  brachypterous,  exposing  3.5  to 
4.5  pregenital  abdominal  terga;  ocellus  ap- 
proximately 1.5X  its  diameter  from  eye;  hind 
tibial  spur  over  3/4  length  of  1st  tarsomere; 
female  abdominal  sternum  VII  (Fig.  58)  with 


1988 


Blocker,  Johnson:  Subgenus  Athysanella 


31 


Figs.  53-70.  Figs.  53-55.  Athysanella  cursa:  53,  aedeagus,  lateral  view;  54,  style,  broad  aspect;  55,  female  sternum 
VII,  ventral  view.  Figs.  56-58.  Athysanella  hemijona.  56,  aedeagus,  lateral  view;  57,  style,  broad  aspect;  58,  female 
sternum  VII,  ventral  view.  Figs.  59-61.  Athysanella  incerta:  59,  aedeagus,  lateral  view;  60,  style,  broad  aspect;  61, 
female  sternum  VII,  ventral  view.  Figs.  62-65.  Athysanella  kadokana:  62,  aedeagus,  lateral  view;  63,  style,  broad 
aspect;  64,  65,  female  sternum  VII,  ventral  view.  Figs.  66-70.  Athysanella  salsa:  66,  head  and  pronotum,  dorsal  view; 
67,  aedeagus,  lateral  view;  68,  style,  broad  aspect;  69,  male  plate,  ventral  view;  70,  female  sternum  VII,  ventral  view. 


median  lobe  approximately  same  length  as 
lateral  lobes. 

Pygofer  rounded  apically,  densely  set  with 
short,  stubby  setae;  valve  with  caudal  margin 
broadly  rounded;  plates  broadly  rounded, 
slightly  separated  at  base;  connective  shorter 
than  style;  styles  (Fig.  57)  widened  apically, 
shallowly  bifid,  dorsal  arm  constricted  and 


curved;  preapical  lobe  inconspicuous,  with 
conspicuous  apical  lateral  lobe,  extending  to 
apex  of  plates  and  pygofer;  aedeagus  (Fig.  56) 
with  shaft  short,  slightly  wider  apically,  ser- 
rated on  ventral  margin. 

Holotype,  male,  from  Inyo  Co.,  California, 
Westgard  Pass,  14  June  1983,  sweeping 
grasses  (R.  J.  Gill  and  D.  Blocker).  Two  male 


32 


Great  Basin  Naturalist  Memoirs 


No.  12 


and  three  female  paratypes,  same  data.  Holo- 
type  and  paratype  in  KSU;  paratypes  in  CDA 
and  KU. 

Athysanella  hemijona  is  related  to  ctirsa  but 
can  be  separated  by  the  shaft  of  the  aedeagus, 
which  is  longer,  and  with  the  ventral  margin 
not  as  flared.  This  species  is  known  only  from 
its  type  locality  in  California. 

Athysanella  mcerta  Ball  &  Beamer 

Figs.  59-61 

Athysanella  incerta  Ball  &  Beamer  1940:15. 

Length  of  male  2.7  to  3.3  mm,  female  3.9  to 
4.1  mm;  head  width  of  male  1.05  to  1.1  mm, 
female  1. 1  to  1.25  mm;  pronotal  width  of  male 
1.0  to  1. 1  mm;  female  1.0  to  1. 15  mm;  interoc- 
ular  width  of  male  0.45  to  0.5  mm,  female  0.5 
to  0.55  mm;  vertex  length  of  male  0.35  to  0.45 
mm,  female  0.45  to  0.5  mm;  pronotal  length  of 
male  0.3  to  0.35  mm,  female  0.35  to  0.4  mm. 
Vertex  length  0.84  to  0.95  interocular  width; 
pronotal  length  0.75  to  0.82  vertex  length. 

Character  code:  1-1-0-0-1-0-0-0-2-1-2-2-0- 
0-0-0-0-0. 

Holotype,  male,  and  allotype,  female,  from 
Lamar,  Colorado,  20  Aug  1936  (R.  H.  Beam- 
er) in  KU;  paratypes  in  KU,  USNM,  and 
CSU. 

Athysanella  incerta  is  very  closely  related 
to  kadokana  but  can  be  separated  by  the 
shape  of  the  apex  of  the  style  (Fig.  60),  which 
is  not  excavated  as  deeply,  and  with  the  dorsal 
arm  neither  expanded  apically  nor  with  an 
acute  projection.  Fuscous  spots  are  some- 
times present  on  the  vertex  of  incerta .  Long- 
winged  specimens  have  been  examined.  This 
species  has  been  collected  in  Colorado,  Kan- 
sas, Montana,  Nebraska,  New  Mexico,  and 
Wyoming,  and  appears  to  be  a  specialist  on 
Distichlis  spicata. 

Athysanella  kadokana  Knull 

Figs.  62-65 

Athysanella  kadokana  Knull  1951:180. 

Length  of  male  2.7  to  3.4  mm,  female  4.0  to 
4.4  mm;  head  width  of  male  0.95  to  1.1  mm, 
female  1. 1  to  1.25  mm;  pronotal  width  of  male 
0.9  to  1.05  mm,  female  1.05  to  1.15  mm; 
interocular  width  of  male  0.4  to  0.5  mm,  fe- 
male 0.5  to  0.6  mm;  vertex  length  of  male  0.35 
to  0.45  mm,  female  0.45  to  0.55  mm;  pronotal 
length  of  male  0.3  to  0.35  nmi,  female  0.35  to 
0.4  mm.  Vertex  length  0.84  to  1.0  interocular 


width:  pronotal  length  0.72  to  0.82  vertex 
length. 

Character  code:  0-1-1-0-1-0-0-0-1-2-2-2-0- 
1-0-0-0-1. 

Holotvpe,  male,  from  Kadoka,  South  Da- 
kota, Badlands,  19  July  1950  (H.  C.  Severin) 
atOhSU. 

This  species  is  closely  related  to  salsa  but 
can  be  separated  by  the  shape  of  the  male 
plate,  which  is  rounded,  and  by  the  apex  of 
the  style,  which  has  a  more  conspicuous  ven- 
tral arm  (Fig.  63).  The  shape  of  the  posterior 
margin  of  the  female  sternum  VII  is  variable 
in  this  species  (Figs.  64,  65).  Specimens  para- 
sitized with  Pipinculidae  and  Dryinidae  and 
specimens  with  undeveloped  genitalia  have 
been  examined;  occasionally  a  specimen  with 
faint  spots  on  the  vertex  is  seen.  This  species, 
collected  in  Colorado,  South  Dakota,  Mon- 
tana, Nebraska,  Wyoming,  and  Canada,  ap- 
pears to  be  a  specialist  on  Distichlis  spicata. 

Athysanella  salsa  Ball  &  Beamer 

Figs.  66-70 

Athysanella  salsa  Ball  &  Beamer  1940:20. 

Length  of  male  2.9  to  3.2  mm,  female  3.6  to 
4.4  mm;  head  width  of  male  1.0  to  1.15  mm, 
female  1.0  to  1.2  mm;  pronotal  width  of  male 
0.9  to  1.05  mm,  female  0.95  to  1.15  mm; 
interocular  width  of  male  0.45  to  0.5  mm, 
female  0.45  to  0.55  mm;  vertex  length  of  male 
0.4  to  0.5  mm,  female  0.45  to  0.55  mm;  prono- 
tal length  of  male  0.3  to  0.4  mm,  female  0.3  to 
0.4  mm.  Vertex  length  0.94  to  1. 1  interocular 
width;  pronotal  length  0.72  to  0.88  vertex 
length. 

Character  code:  0-1-1-0-2-0-0-0-2-1-2-2-0- 
1-0-0-0-0. 

Holotvpe,  male,  and  allotype,  female,  from 
St.  John,'  Kansas,  11  Sept  1936  (R.  H.  Beamer) 
in  KU. 

Athysanella  salsa  is  closely  related  to 
kadokana  but  can  be  separated  by  the  trun- 
cate apices  of  the  male  plates  and  the  less 
conspicuous  ventral  apical  arm  of  the  style 
(Fig.  68).  Specimens  studied  from  Mexico 
have  fuscous  spots  on  the  vertex  and  dark 
stripes  on  the  forewings.  Long-winged  fe- 
males have  been  examined.  Males  and  fe- 
males parasitized  by  Dryinidae  and  undevel- 
oped specimens  have  been  examined.  This 
species  has  been  collected  in  Kansas,  Texas, 
and  Mexico,  and  appears  to  be  a  specialist  on 
Distichlis  spicata. 


1988 


Blocker,  Johnson:  Subgenus  Atmysanella 


33 


^-m^ 


\r^r^ 


Figs.  71-86.  Figs.  71-73.  Athysanella  kanabana:  71,  aedeagus,  lateral  view;  72,  style,  broad  aspect;  73,  female 
sternum  VII,  ventral  view.  Figs.  74-76.  Athysanella  furnaca:  74,  aedeagus,  lateral  aspect;  75,  style,  broad  aspect;  76, 
female  sternum  VII,  ventral  view.  Figs.  77-79.  Athysanella  marthae:  11,  aedeagus,  lateral  view;  78,  style,  broad 
aspect;  79,  female  sternum  VII,  ventral  view.  Figs.  80-83.  Athysanella  bifida:  80,  aedeagus,  lateral  view;  81,  82,  style, 
broad  aspect;  83,  female  sternum  VII,  ventral  view.  Figs.  84-86.  Athysanella  utahna:  84,  aedeagus,  lateral  view;  85, 
style,  broad  aspect;  86,  female  sternum  VII,  ventral  aspect. 


Athysanella  kanabana  Ball  &  Beamer 

Figs.  71-73 

Athysanella  kanabana  Ball  &  Beamer  1940:20. 

Length  of  male  3. 1  to  3.6  mm,  female  4.4  to 
4.7  mm;  head  width  of  male  1.1  to  1.2  mm, 
female  1.25  to  1.4  mm;  pronotal  width  of  male 
1.05  to  1.15  mm,  female  1.2  to  1.3  mm;  inter- 


ocular  width  of  male  0.5  to  0.55  mm,  female 
0.6  to  0.7  mm;  vertex  length  of  male  0.4  to  0.5 
mm,  female  0.5  to  0.55  mm;  pronotal  length  of 
male  0.35  to  0.4  mm,  female  0.35  to  0.45  mm. 
Vertex  length  0.8  to  0.95  interocular  width; 
pronotal  length  0.76  to  0.88  vertex  length. 

Character  code:  0-1-1-1-1-0-0-0-2-1-2-2-0- 
1-0-1-1-0. 


34 


Great  Basin  Naturalist  Memoirs 


No.  12 


Holotype,  male,  and  allotype,  female,  from 
Kanab,  Utah,  9  Aug  1936  (R.  H.  Beamer)  in 
KU;  paratypes  in  KU  and  USNM. 

Athysanella  kanabana  is  related  to  tenera 
but  can  be  separated  by  the  embrowned  area 
on  the  plates,  the  aedeagus  with  apical  crests 
on  the  shaft  (Fig.  71),  and  the  unique  shape  of 
the  apex  of  the  style  (Fig.  72).  Occasionally, 
there  are  fuscous  spots  on  the  vertex.  Unde- 
veloped specimens  are  represented  in  the 
paratype  series.  This  species  has  been  col- 
lected on  Sporobolus  airoides  in  Arizona, 
Kansas,  New  Mexico,  and  Utah. 

Athysanella  furnaca  Blocker,  n.  sp. 

Figs.  74-76 

Length  of  male  3.1  mm,  female  4.4  to  4.7 
mm;  head  width  of  male  1. 15  mm,  female  1.25 
to  1.3  mm;  pronotal  width  of  male  1.1  mm, 
female  1.2  to  1.25  mm;  interocular  width  of 
male  0.5  mm,  female  0.5  to  0.55  mm;  vertex 
length  of  male  0. 5  mm,  female  0.5  to  0. 55  mm; 
pronotal  length  of  male  0.4  mm,  female  0.4  to 
0.45  mm.  Vertex  length  equal  to  interocular 
wddth;  pronotal  length  0.78  to  0.79  vertex 
length. 

Character  code:  0-0-1-0-2-0-0-0-2-1-2-2-0- 
1-0-0-1-1. 

Color  stramineous,  except  for  dark  dots  on 
abdomen  and  slight  fuscous  on  legs. 

Forewings  brachypterous,  exposing  3.5 
pregenital  abdominal  terga;  ocellus  approxi- 
mately 2X  its  diameter  from  eye;  hind  tibial 
spur  1/2  length  (or  less)  of  1st  tarsomere;  fe- 
male abdominal  sternum  VII  (Fig.  76)  with 
short  lateral  lobes  and  slight  median  projec- 
tion. 

Pygofer  with  caudal  margin  extended,  ex- 
ceeding anal  tube,  with  approximately  25  se- 
tae; valve  with  caudal  margin  rounded;  plates 
truncate  apically,  barely  exceeding  apex  of 
valve;  style  with  ventral  arm  short  (Fig.  75), 
conspicuously  flattened,  dorsal  arm  elongate; 
connective  3/4  length  of  pygofer;  aedeagus 
(Fig.  74)  with  conspicuous  keels  on  shaft. 

Holotype,  male,  from  Death  Valley,  Cali- 
fornia, 5.7  mi  N  Furnace  Cr.,  19  March  1971 
(Oman);  two  female  paratypes,  same  data. 
Holotype  and  paratype  in  OrSU;  paratype  in 
KSU. 

Athysanella  furnaca  is  related  to  kanabana 
but  can  be  distinguished  by  the  medial  crests 
on  the  shaft  of  the  aedeagus  and  the  connec- 
tive, which  is  shorter  than  the  style.  This  spe- 


cies is  known  only  from  the  type  locality  in  the 
Mojave  Desert  of  California. 

Athysanella  marthae  Blocker,  n.  sp. 

Figs.  77-79 

Length  of  male  2.9  to  3.3  mm,  female  4. 1  to 
4.6  mm;  head  width  of  male  1.15  to  1.25  mm, 
female  1.25  to  1.35  mm;  pronotal  width  of 
male  1.1  to  1.2  mm,  female  1.1  to  1.2  mm; 
interocular  width  of  male  0.45  to  0.5  mm, 
female  0.5  to  0.55  mm;  vertex  length  of  male 
0.4  to  0.45  mm,  female  0. 45  to  0. 5  mm;  prono- 
tal length  of  male  0.35  to  0.4  mm,  female  0.4 
to  0.45  mm.  Vertex  length  0.89  to  0.95  intero- 
cular width;  pronotal  length  0.82  to  0.94  ver- 
tex length. 

Character  code:  0-1-1-0-2-0-0-0-2-2-2-1-0- 
1-0-0-1-1. 

Color  uniformly  stramineous,  except  for  ir- 
regular, fuscous  markings  on  venter  and  legs. 

Forewings  brachypterous,  exposing  3  to  4.5 
pregenital  abdominal  terga;  ocellus  approxi- 
mately 2X  its  diameter  from  eye;  hind  tibial 
spur  2/3  length  of  1st  tarsomere;  female  ab- 
dominal sternum  VII  (Fig.  79)  with  conspicu- 
ous lateral  lobes  on  hind  margin. 

Pygofer  broadly  rounded  apically,  with 
fewer  than  25  macrosetae,  exceeding  apex  of 
plates;  anal  tube  exceeding  the  apex  of 
pygofer;  valve  with  caudal  margin  rounded; 
plates  short,  truncate,  with  conspicuous  mi- 
crosetae;  style  (Fig.  78)  with  conspicuous  lat- 
eral flange  and  preapical  lobe,  deeply  bifid 
apically,  dorsal  arm  truncate,  ventral  arm 
rounded,  exceeding  apex  of  plates  and 
pygofer;  connective  3/4  length  of  style;  aedea- 
gus (Fig.  77)  with  caudal  margin  of  shaft  ser- 
rate, troughlike  in  caudal  view,  with  incon- 
spicuous process  apically,  dorsal  apodeme 
simple. 

Holotype,  male,  from  Cuatro  Cienegas, 
Coahuila,  Mexico,  9  June  1985,  (A.  L.  Hicks) 
1266B;  five  male  and  five  females  paratypes, 
same  data.  Holotype  and  paratypes  in  KSU; 
paratypes  in  IPL. 

Athysanella  nuirthae  is  related  to  bifida  but 
can  be  distinguished  by  the  conspicuous  lat- 
eral flange  of  the  style  and  the  short,  truncate 
plates.  This  species  has  been  collected  only  at 
the  type  locality  in  Mexico  on  Monanthochloe 
littoralis.  I  name  this  species  for  my  wife. 

Athysanella  bifida  Ball  &  Beamer 

Figs.  80-83 

Athysanella  bifida  Ball  fit  Beamer  1940:23. 


1988 


Blocker,  Johnson;  Subgenus  Athysanella 


35 


Length  of  male  2.4  to  3.0  mm,  female  3. 1  to 
3.7  mm;  head  width  of  male  0.9  to  1.05  mm, 
female  0.95  to  1. 1  mm;  pronotal  width  of  male 
0.8  to  1.0mm,  female 0.9  to  1.0mm;interocu- 
lar  width  of  male  0.4  to  0.45  mm,  female  0.4  to 
0.5  mm;  vertex  length  of  male  0.35  to  0.45 
mm,  female  0.4  to  0.5  mm;  pronotal  length  of 
male  0.25  to  0.35  mm,  female  0.3  to  0.35  mm. 
Vertex  length  0.87  to  1.0  interocular  width; 
pronotal  length  0.7  to  0.82  vertex  length. 

Character  code:  0-1-1-0-1-0-0-0-1-2-2-1-0- 
1-0-0-1-0. 

Holotype,  male,  and  allotype,  female,  from 
Monument,  Colorado,  19  Aug  1936  (R.  H. 
Beamer)  in  KU;  paratypes  in  KU  and  USNM. 

Athysanella  bifida  is  related  to  utahna  and 
ijwnana  but  can  be  separated  by  the  elongate 
shaft  of  the  aedeagus  (Fig.  80),  which  is  not 
widened  at  midlength.  Specimens  parasitized 
by  Dryinidae  and  Strepsiptera,  undeveloped 
specimens,  and  long-winged  specimens  have 
been  examined.  This  species  has  been  col- 
lected in  Arizona,  Colorado,  Montana,  New 
Mexico,  North  Dakota,  South  Dakota,  Texas, 
Wyoming,  and  Canada.  In  the  southern  part 
of  its  range,  at  least,  the  host  is  blue  grama, 
Bouteloua  gracilis  (Hicks  et  al.  1988). 

Athysanella  utahna  Osborn 

Figs.  84-86 

Athysanella  utahna  Osborn  1930:705. 

Length  of  male  2.6  to  3.3  mm,  female  3.9  to 
4.9  mm;  head  width  of  male  0.9  to  1.05  mm, 
female  1.05  to  1.25  mm;  pronotal  width  of 
male  0.85  to  1.0  mm,  female  1.0  to  1.1  mm; 
interocular  width  of  male  0.35  to  0.45  mm, 
female  0.45  to  0.55  mm;  vertex  length  of  male 
0.4  to  0.45  mm,  female  0.45  to  0.5  mm;  prono- 
tal length  of  male  0.3  to  0.35  mm,  female  0.3 
to  0.4  mm.  Vertex  length  0.94  to  1. 13  interoc- 
ular width;  pronotal  length  0.7  to  0.82  vertex 
length. 

Character  code:  0-1-0-0-1-0-0-0-2-1-2-0-0- 
1-0-0-1-1. 

Lectotype,  male,  and  lectoallotype,  fe- 
male, from  Ephraim,  Utah,  20  July  1914  (E. 
D.  Ball)  in  USNM. 

Athysanella  utahna  is  related  to  yumana 
but  can  be  separated  by  the  more  slender 
apical  dorsal  arm  and  less  robust  ventral  arm 
of  the  style  (Fig.  85).  It  can  be  separated  from 
bifida  by  the  shorter  shaft  of  the  aedeagus 
(Fig.  84).  Rarely  a  specimen  is  seen  with  fus- 
cous spots  on  the  vertex;  the  styles  occasion- 


ally have  a  macroseta.  Specimens  with  long 
wings  and  undeveloped  genitalia  and  speci- 
mens parasitized  by  Dryinidae  and  Pipinculi- 
dae  have  been  examined.  This  species  has 
been  collected  in  Arizona,  California,  Colo- 
rado, Idaho,  Nevada,  Utah,  Oregon,  Wash- 
ington, Canada,  and  Mexico.  Reported  from 
saltgrass,  Distichlis  spicata,  in  Mexico  and 
Utah. 

Athysanella  yumana  Osborn 

Figs.  87-89 

Athysanella  yumana  Osborn  1930:704. 

Length  ofmale  2.8  to  3.1  mm,  female  3.8  to 

4.7  mm;  head  width  ofmale  1.05  to  1. 15  mm, 
female  1. 1  to  1.25  mm;  pronotal  width  ofmale 
0.95  to  1.1  mm,  female  1.0  to  1.2  mm;  interoc- 
ular width  ofmale  0.4  to  0.45  mm,  female  0.45 
to  0.55  mm;  vertex  length  ofmale  0.4  to  0.45 
mm,  female  0.45  to  0.50  mm;  pronotal  length 
ofmale  0.3  to  0.35  mm,  female  0.3  to  0.4  mm. 
Vertex  length  0.94  to  1.0  interocular  width; 
pronotal  length  0.76  to  0.82  vertex  length. 

Character  code:  1-1-1-0-2-0-0-0-2-1-2-2-0- 
1-0-0-1-0. 

Lectotype,  male,  and  lectoallotype,  fe- 
male, from  Yuma,  Arizona  (H.  Osborn)  in 
USNM;  paratypes  in  USNM  and  KU. 

Athysanella  yumana  is  related  to  utahna 
but  can  be  separated  by  the  shape  of  the  apex 
of  the  style  (Fig.  88),  which  has  more  robust 
ventral  and  dorsal  arms.  Occasionally,  there 
are  fuscous  spots  on  the  vertex.  The  style  may 
have  an  occasional  macroseta.  This  species  has 
been  reported  from  saltgrass,  Distichlis  spi- 
cata; it  has  been  collected  in  Arizona  and  Cali- 
fornia. 

Athysanella  deserta  Blocker,  n.  sp. 

Figs.  90-92 

Length  ofmale  3.0  to  3.3  mm,  female  4.3  to 

4.8  mm;  head  width  ofmale  1.1  to  1.2  mm, 
female  1.2  to  1.3  mm;  pronotal  width  ofmale 
1.05  to  1.15  mm,  female  1.15  to  1.25  mm; 
interocular  width  of  male  0.45  to  0.5  mm, 
female  0.5  to  0.6  mm;  head  width  ofmale  0.45 
to  0.5  mm,  female  0.5  to  0.6  mm;  pronotal 
length  ofmale  0.35  to  0.4  mm,  female  0.35  to 
0.45  mm.  Vertex  length  0.9  to  1.0  interocular 
width;  pronotal  length  0.75  to  0.83  vertex 
length. 

Character  code:  0-0-1-0-2-0-1-0-2-2-2-2-0- 
1-0-0-1-1. 


36 


Great  Basin  Naturalist  Memoirs 


No.  12 


Figs.  87-103. Figs.  87-89.  Athijsanella  ijmnana:  87,  aedeagiis,  lateral  view;  88,  style,  broad  aspect;  89,  female 
sternum  VII,  ventral  view.  Figs.  90-92.  Athijsanella  deserta:  90,  aedeagus,  lateral  view;  91,  style,  broad  aspect;  92, 
female  sternum  VII,  ventral  view.  Figs.  93-97.  Athijsanella  magdalena:  93,  head  and  pronotum,  dorsal  view;  94, 
aedeagus,  lateral  view;  95,  style,  broad  aspect;  96,  male  plate,  ventral  view;  97,  female  sternum  VII,  ventral  view.  Figs. 
98-100.  Athijsanella  laeta.  98,  aedeagus,  lateral  view;  99,  style,  broad  aspect;  100,  female  sternum  VII,  ventral  view. 
Figs.  101-103.  Athijsanella  stijlosa:  101,  aedeagus,  lateral  view;  102,  style,  broad  aspect;  103,  female  sternum  VII, 
ventral  view. 


Color  stramineous;  faint  orange  pattern  of- 
ten seen  on  vertex;  faint  fuscous  dots  may  be 
present  on  abdominal  terga;  face  usually  with 
faint  lateral  lines;  venter  with  various  amounts 
of  fuscous  coloring. 

Forewings  brachypterous,  exposing  3.5  to 
4.0  pregenital  abdominal  terga;  ocellus  ap- 
proximately 1.5  its  diameter  from  eye;  hind 
tibial  spur  1/3  to  1/2  length  of  1st  tarsomere; 
female  abdominal  sternum  VII  (Fig.  92)  with 


conspicuous  lateral  lobes,  inconspicuous  me- 
dial lobe  rounded,  with  various  amounts  of 
fuscous  coloring. 

Pygofer  with  caudal  margin  rounded,  with 
25 -h  microsetae;  plates  separated  at  base, 
truncate  apically,  not  reaching  apex  of  pygo- 
fer; anal  tube  exceeding  apex  of  pygofer;  valve 
with  caudal  margin  rounded;  connective 
2/3  length  of  style;  styles  (Fig.  91)  slightly 
exceeding  apex  of  pygofer  with  apical  arm 


1988 


Blocker,  Johnson:  Subgenus  Athysanella 


37 


enlarged  at  apex,  ventral  arm  with  small, 
acute  process;  aedeagus  (Fig.  90)  with  shaft 
narrowed  apically,  finely  serrate  on  ventral 
margin,  approximately  2X  length  of  dorsal 
apodeme. 

Holotype,  male,  from  Death  Valley,  Cali- 
fornia, 36  mi  S  Furnace  Cr.,  19  March  1971 
(Oman);  18  male  and  15  female  paratypes, 
same  data.  Holotypes  and  paratypes  in  OrSU; 
paratypes  in  KSU. 

Athysanella  deserta  is  related  to  ijiwiana 
but  can  be  separated  by  the  shape  of  the  apical 
arms  of  the  style  with  the  dorsal  arm  enlarged 
at  the  apex  and  the  ventral  arm  with  a  small, 
acute  process.  This  species  has  been  collected 
only  at  the  type  locality  in  the  Mojave  Desert 
of  California. 

Athysanella  magdalena  Baker 

Figs.  93-97 

Athysanella  magdalena  Baker  1898:185. 

Length  ofmale  2.2  to  2.7  mm,  female  3.0  to 
3.6  mm;  head  width  ofmale  0.85  to  0.95  mm, 
female  0.95  to  1. 1  mm;  pronotal  width  ofmale 
0.75  to  0.9  mm,  female  0.9  to  1.1  mm;  interoc- 
ular  width  ofmale  0.3  to  0. 4  mm,  female  0. 4  to 
0.5  mm;  vertex  length  ofmale  0.3  to  0.4  mm, 
female  0.35  to  0.45  mm;  pronotal  length  of 
male  0.25  to  0.35  mm,  female  0.3  to  0.35  mm. 
Vertex  length  0.93  to  1.1  interocular  width; 
pronotal  length  0.8  to  0.91  vertex  length. 

Character  code:  1-1-0-0-0-0-0-0-0-2-0-1-0- 
1-0-0-1-0. 

Lectotype,  female,  from  Magdalena  Mts., 
New  Mexico,  Aug  1894  (F.  H.  Snow),  and 
lectoallotype,  male,  from  Forresters  Ranch, 
Laramie  Co. ,  Colorado,  3  Aug  1896  (No.  2013 
ofBaker)inUSNM. 

Athysanella  magdalena  is  related  to  tenera 
but  can  be  separated  by  the  elongate  plates 
and  the  shape  of  the  apex  of  the  style  (Fig.  95), 
which  is  more  clearly  bifid.  Plates  occasionally 
have  one  or  more  macrosetae,  and  occasion- 
ally the  male  hind  tibial  spur  is  undeveloped. 
Numerous  undeveloped  specimens,  speci- 
mens parasitized  by  Dryinidae  and  Strep- 
siptera,  and  long-winged  males  and  females 
have  been  examined.  This  species,  collected 
in  Arizona,  California,  Colorado,  Montana, 
New  Mexico,  Nevada,  Oklahoma,  Texas, 
Utah,  Wyoming,  and  Mexico,  is  a  common 
specialist  of  blue  grama,  Bouteloua  gracilis 
(Whitcomb  et  al.  1987,  Hicks  et  al.  1988). 


Athysanella  laeta  Ball  &  Beamer 

Figs.  98-100 

Athysanella  laeta  Ball  and  Beamer,  1940:16. 

Length  ofmale  2.7  to  2.9  mm,  female  3. 8  to 
4.1  mm;  head  width  ofmale  1.0  to  1.1  mm, 
female  1.1  to  1.2  mm;  pronotal  width  ofmale 
0.95  to  1.05  mm,  female  1.05  to  1.15  mm; 
interocular  width  of  male  0.45  to  0.5  mm, 
female  0.5  to  0.55  mm;  vertex  length  ofmale 
0.35  to  0.4  mm,  female  0.4  to  0.45  mm;  prono- 
tal length  ofmale  0.35  to  0.4  mm,  female  0.35 
to  0.45  mm.  Vertex  length  0.75  to  0.89  intero- 
cular width;  pronotal  length  0.87  to  1.0  vertex 
length. 

Character  code:  1-1-1-0-2-0-0-0-2-1-2-0-0- 
1-0-0-1-1. 

Holotype,  male,  from  Huachuca  Mts.,  Ari- 
zona, 15  July  1934  (E.  D.  Ball);  allotype,  fe- 
male, same  data  except  14  July,  in  USNM; 
paratypes  in  USNM  and  KU. 

Athysanella  laeta  is  related  to  incongrua 
but  can  be  separated  by  its  usually  smaller  size 
and  the  shape  of  the  shaft  of  the  aedeagus  (Fig. 
98),  which  is  shorter  and  not  as  avicephaliform 
apically.  Some  of  the  male  paratypes  have 
undeveloped  genitalia.  This  species  has  been 
collected  in  New  Mexico  and  Arizona. 

Athysanella  stylosa  Blocker,  n.  sp. 

Figs.  101-103 

Length  ofmale  2.4  to  2.8  mm,  female  3.8  to 
4.1  mm;  head  width  ofmale  0.95  to  1.1  mm, 
female  1. 15  to  1.2  mm;  pronotal  width  ofmale 
0.9  to  1.0  mm,  female  1.05  to  1.1  mm;  interoc- 
ular width  ofmale  0.35  to  0.45  mm,  female 
0.45  to  0.5  mm;  vertex  length  ofmale  0.35  to 
0.45  mm,  female  0.4  to  0.5  mm;  pronotal 
length  ofmale  0.3  to  0.35  mm,  female  0.3  to 
0.4  mm.  Vertex  length  0.93  to  1.06  interocu- 
lar width;  pronotal  length  0.75  to  0.82  vertex 
length. 

Color  stramineous  with  brown  pattern  on 
vertex  and  pronotum,  vertex  with  fuscous 
spots,  wings  with  brown  stripes,  face  with 
brown  arcs. 

Forewings  brachypterous,  exposing  3  to  4.5 
pregenital  abdominal  terga;  ocellus  approxi- 
mately its  diameter  from  eye;  hind  tibial  spur 
3/4  1st  tarsomere;  female  abdominal  sternum 
VII  (Fig.  103)  with  well-developed  lateral 
lobes,  slightly  concave  medially. 

Pygofer  broadly  rounded  apically,  with 
fewer  than  25  setae;  anal  tube  exceeds  apex  of 


38 


Great  Basin  Naturalist  Memoirs 


No.  12 


pygofer;  valve  with  caudal  margin  rounded; 
plates  rounded  apically;  styles  (Fig.  102)  bifid, 
dorsal  arm  acute,  ventral  arm  truncate,  with- 
out lateral  flange  or  preapical  lobe,  exceeding 
apex  of  plates  and  pygofer;  connective  3/4 
length  of  style;  aedeagus  (Fig.  101)  with  shaft 
serrate  on  caudal  margin,  troughlike  in  caudal 
view,  with  inconspicuous  process  apically. 

Holotype,  male,  from  Roswell,  Chaves 
Co.,  New  Mexico,  Bitter  Lake  Natl.  WLR,  14 
Aug  1984  (R.  F.  Whitcomb)  0868A;  17  male 
and  7  female  paratypes,  same  data;  1  male 
paratype,  same  data  except  21  Aug  1985, 
01931.  Holotype  and  paratypes  in  KSU; 
paratypes  in  USNM  and  IPL. 

Athysanella  stylosa  is  related  to  supina  but 
can  be  distinguished  by  the  fuscous  spots  on 
the  vertex  and  by  the  shaft  of  the  aedeagus, 
which  is  not  as  widened  medially.  This  species 
is  different  in  that  the  vertex  length  is  equal  to 
the  interocular  width  and  fuscous  spots  are 
present.  It  has  been  collected  on  gyp  drop- 
seed,  Sporobolus  nealleiji,  in  the  gypsum  flats 
of  southeastern  New  Mexico. 

Athysanella  incongrua  Baker 

Figs.  104-106 

Athysanella  incongrua  Baker  1898:188. 

Length  of  male  3.0  to  3.4  mm,  female  4.2  to 
4.8  mm;  head  width  of  male  1.1  to  1.2  mm, 
female  1.15  to  1.35  mm;  pronotal  width  of 
male  1.05  to  1. 15  mm,  female  1. 1  to  1.25  mm, 
interocular  width  of  male  0.45  to  0.5  mm, 
female  0.55  to  0.65  mm;  vertex  length  of  male 
0.35  to  0.45  mm,  female  0.45  to  0.5  mm; 
pronotal  length  of  male  0.35  to  0.4  mm,  fe- 
male 0.35  to  0.45  mm.  Vertex  length  0.76  to 
0.85  interocular  width;  pronotal  length  0.88  to 
1.0  vertex  length. 

Character  code:  1-1-0-0-2-1-0-0-2-1-2-2-0- 
1-0-0-0-0. 

Lectotype,  male,  from  Fort  Collins,  Colo- 
rado, 21  July  (C.  F.  Baker)  in  USNM. 

Athysanella  incongrua  is  related  to  laeta 
and  terebrans  but  can  be  separated  by  the 
length  and  shape  of  the  apex  of  the  shaft  of  the 
aedeagus  (Fig.  104).  The  fuscous  spots  on  the 
vertex  are  commonly  missing.  Long-winged 
males  and  females  have  been  examined.  Spec- 
imens, collected  in  Colorado,  Iowa,  Kansas, 
Nebraska,  New  Hampshire,  New  Mexico, 
Oklahoma,  South  Dakota,  Texas,  and  Wyo- 
ming,   appear   to   be   associated   with   little 


bluestem,  Schizachyrium  scroparium.  The 
specimen  from  New  Hampshire  was  collected 
by  P.  W.  Oman  at  Wiley  House,  20  Aug  1934. 

Athysanella  tenera  Ball  &  Beamer 

Figs.  107-109 

Athysanella  tenera  Ball  &  Beamer  1940:17. 

Length  of  male  2.5  to  3.0  mm,  female  3.9  to 
4.0  mm;  head  width  of  male  1.0  to  1.1  mm, 
female  1.1  to  1.2  mm;  pronotal  width  of  male 
0.9  to  1.0  mm,  female  1.0  to  1.15  mm;  interoc- 
ular width  of  male  0.4  to  0.45  mm,  female  0.45 
to  0.5  mm;  vertex  length  of  male  0.35  to  0.45 
mm,  female  0.4  to  0.45  mm;  pronotal  length  of 
male  0.3  to  0.35  mm,  female  0.35  to  0.4  mm. 
Vertex  length  0.94  to  1.0  interocular  width; 
pronotal  length  0.76  to  0.8  vertex  length. 

Character  code:  1-1-0-0-1-0-0-0-1-0-2-2-0- 
1-0-0-1-0. 

Holotype,  male,  and  allotype,  female,  from 
Las  Vegas,  Nevada,  8  Aug  1936  (R.  H. 
Beamer)  in  KU;  paratypes  in  KU. 

Athysanella  tenera  is  related  to  magdalena 
but  can  be  distinguished  by  the  shorter  plates 
and  the  irregular  shape  of  the  outer  apical  arm 
of  the  style  (Fig.  108),  which  is  widened  and 
appears  to  be  trifid.  This  species  is  known  only 
from  the  type  locality  in  Nevada. 

Athysanella  vativala  Blocker,  n.  sp. 

Figs.  110-112 

Length  of  male  2.9  to  3.2  mm,  female  4.2  to 
4.6  mm;  head  width  of  male  1.05  to  1.15  mm, 
female  1.25  to  1.35  mm;  pronotal  width  of 
male  1.0  to  1.15  mm,  female  1.15  to  1.3  mm; 
interocular  width  of  male  0.4  to  0.55  mm, 
female  0.55  to  0.65  mm;  vertex  length  of  male 
0.35  to  0.5  mm,  female  0.45  to  0.55  mm; 
pronotal  length  of  male  0.35  to  0.4  mm,  fe- 
male 0.35  to  0.45  mm.  Vertex  length  0.78  to 
0.9  interocular  width;  pronotal  length  0.74  to 
0.94  vertex  length. 

Character  code:  1-0-0-0-1-0-0-0-1-1-1-0-0- 
1-0-0-1-0. 

Color  stramineous;  vertex  with  fuscous 
spots;  vertex,  pronotum,  and  abdomen  with 
additional  brown  pattern;  fore  wings  with 
darkened  stripes;  face  with  lateral  fuscous 
marks;  legs  with  fuscous  areas. 

Forewingsbrachypterous,  exposing  4  to  4. 5 
pregenital  abdominal  terga;  ocellus  2X  its  di- 
ameter from  eye  or  less;  hind  tibial  spur  1/2 
length  of  1st  tarsomere;  female  abdominal 


1988 


Blocker,  Johnson:  Subgenus  Athysanella 


39 


V^ 


113 


114 


Figs.  104-117.  Figs.  104-106.  Athysanella  incongrua:  104,  aedeagus,  lateral  view;  105,  style,  broad  aspect;  106, 
female  sternum  VII,  ventral  view.  Figs.  107-109.  Athysanella  tenera:  107,  aedeagus,  lateral  view;  108,  style,  broad 
aspect;  109,  female  sternum  VII,  ventral  view.  Figs.  110-112.  Athysanella  vativala:  110,  aedeagus,  lateral  view;  111, 
style,  broad  aspect;  112,  female  sternum  VII,  ventral  view.  Figs.  113-117.  Athysanella  terebrans:  113,  head  and 
pronotum,  dorsal  view;  114,  aedeagus,  lateral  view;  115,  116,  style,  broad  aspect;  117,  female  sternum  VII,  ventral 


sternum  VII  (Fig.  1 12)  with  lateral  lobes  much 
longer  than  medial  lobe. 

Pygofer  rounded  apically,  fewer  than  25 
macrosetae;  valve  with  caudal  margin  angu- 
late;  plates  separated  at  base,  rounded  api- 
cally; connective  3/4  length  of  style;  styles 
(Fig.  Ill)  deeply  bifid,  dorsal  arm  elongate 
and  slender,  ventral  arm  thickened,  no 
preapical  lobe  present;  aedeagus,  in  lateral 
view,  with  shaft  slightly  widened  in  apical  half 
(Fig.  110),  troughlike  on  ventral  margin,  and 
slightly  serrate. 


Holotype,  male,  from  Sheridan  Co.,  Ne- 
braska, 7  mi  N  Rushville,  Hwy  87,  9  Aug  1979 
(H.  D.  Blocker  and  R.  A.  Sweet);  female 
paratype,  same  data;  4  male  and  7  female 
paratypes,  Cherry  Co.,  Nebraska,  S  Valen- 
tine, Hwy  83,  mi  202,  10  Aug  1979  (H.  D. 
Blocker  and  R.  A.  Sweet);  2  males.  Cherry 
Co.,  Nebraska,  near  Manker,  Hwy  20,  9  Aug 
1979  (H.  D.  Blocker  and  R.  A.  Sweet);  1  male. 
Bowman,  North  Dakota,  5  July  1968,  GL  819 
(Harris  and  Cooley);  2  males  and  10  females, 
Tryon,    McPherson  Co.,    Nebraska,   7  Aug 


40 


Great  Basin  Naturalist  Memoirs 


No.  12 


1977,  (R.  F.  Whitcomb)  0480.  Holotype  and 
paratypes  in  KSU;  paratypes  in  CNC,  IPL, 
USNM. 

Athysanella  vativala  is  very  closely  related 
and  possibly  conspecific  with  terebrans  but 
can  be  separated  by  the  shape  of  the  apex  of 
the  style,  which  has  a  more  slender  dorsal 
arm,  a  more  patterned  and  longer  vertex,  and 
the  ocelli  usually  more  distant  from  the  eye. 
Undeveloped  specimens  and  specimens  para- 
sitized with  Strepsiptera  have  been  exam- 
ined. This  species  has  been  collected  in  Ne- 
braska and  North  Dakota  on  Calanovilfa 
longifolia. 

Athysanella  terebrans  (Gillette  &  Baker) 

Figs.  113-117 

Euttetix  terebrans  Gillette  &  Baker  1895:102. 

Length  of  male  2.7  to  3. 1  mm,  female  4.2  to 
4.6  mm;  head  width  of  male  1.0  to  1.15  mm, 
female  1.2  to  1.35  mm;  pronotal  width  of  male 
0.95  to  1.05  mm,  female  1.15  to  1.3  mm; 
interocular  width  of  male  0.45  to  0.5  mm, 
female  0.55  to  0.6  mm;  vertex  length  of  male 
0.35  to  0.45  mm,  female  0.45  to  0.5  mm; 
pronotal  length  of  male  0.3  to  0.4  mm,  female 
0.4  to  0.45  mm.  Vertex  length  0.75  to  0.89 
interocular  width;  pronotal  length  0.82  to  1.0 
vertex  length. 

Character  code:  1-0-0-0-1-0-0-0-1-1-1-0-0- 
1-0-0-1-0. 

Holotype,  female,  from  North  Park,  Colo- 
rado, 30  July  (Gillette)  in  USNM;  allotype, 
male,  from  Wray,  Colorado,  13  July  1899,  at 
Colorado  State  University. 

Athysanella  terebrans  is  related  to  incon- 
grua  but  can  be  distinguished  by  the  dorsal 
apical  arm  of  the  style  (Figs.  115,  116),  which 
is  rounded,  and  by  the  shaft  of  the  aedeagus 
(Fig.  114),  which  is  not  conspicuously 
widened  apically;  the  female  abdominal  ster- 
num VII  (Fig.  117)  may  be  variable  in  shape. 
Long-winged  and  undeveloped  specimens 
were  commonly  examined.  This  species  has 
been  collected  in  Colorado,  Montana,  Ne- 
braska, North  Dakota,  South  Dakota,  Utah, 
Wyoming,  and  Canada.  It  is  reported  from 
Dystichlis  striata  in  Manitoba. 

Phylogeny 

Phylogeny  of  the  subgenera  oi  Athysanella 
is  discussed  in  Blocker  and  Johnson  (1988). 
The  subgenus  Athysanella  is  designated  a  sis- 


ter o{ Gladionura  and  can  be  separated  by  the 
absence  of  a  pygofer  process  in  the  former. 
The  presence  of  a  pygofer  process  is  a  conver- 
gent character  that  occurs  in  two  other  less 
closely  related  subgenera.  We  selected  40 
characters  that  show  differences  within  the 
subgenus  for  analysis.  An  intuitive  phylogeny 
based  on  the  proposed  sequence  of  occur- 
rence of  major  derived  characters  was  gener- 
ated, as  well  as  an  analysis  by  PAUP  (Swofford 
1986).  Results  of  these  analyses  are  shown  in 
Figures  118  and  119.  PAUP  generated  five 
equally  most  parsimonious  trees;  the  tree 
with  no  unresolved  trifurcations  is  presented 
here.  A  hypothetical  ancestor  consisting  of  a 
uniform  plesiomorphic  character  set  was  uti- 
lized. Character  sequences  are  contained  in 
the  preceding  descriptions;  the  entire  PAUP 
data  set  is  available  on  request.  Overall,  the 
PAUP  cladogram  and  the  intuitive  phylogeny 
are  in  good  agreement.  Species  groups  1,  3,  4, 
and  7  are  consistently  grouped  together  (Fig. 
119),  whereas  the  rostrata  and  incongrua- 
planata  groups  are  widely  fragmented.  It  is 
possible  that  this  fragmentation  of  the  groups 
is  indicative  of  the  high  degree  of  homoplasy 
in  the  genus. 

The  robusta  group  is  characterized  by  re- 
tention of  the  rounded  shape  of  the  style  apex; 
the  male  plates  are  embrowned  in  all  species 
(2');  the  styles  are  widened  and  often  bifid 
apically  in  all  other  species  (1').  This  group  is 
recognized  in  the  PAUP  output. 

The  rostrata  group  has  an  extended  apex  of 
the  male  style  (1");  PAUP  has  the  species  in 
this  group  occurring  close  together  but  none 
as  sisters. 

The  incerta  group  is  characterized  by  the 
presence  of  a  semicircular  apex  of  the  shaft  of 
the  aedeagus  (3').  There  is  fairly  good  congru- 
ence in  the  cladogram,  which  includes 
planata  as  a  sister  o( incerta  . 

The  terebrans-bifida  group  is  characterized 
by  an  aedeagal  shaft  that  is  serrate  on  the 
caudal  margin  (4').  The  terebrans  group  has  a 
shortened  vertex  (5');  five  of  the  seven  species 
are  recognized  by  PAUP;  the  other  two  (laeta 
and  marthae)  split  out  sooner  but  are  close. 
The  bifida  group  retains  an  elongate  vertex; 
they  are  represented  in  the  last  half  of  the 
cladogram,  close  together,  but  only  bifida  and 
magdalena  arc  sisters. 

The  incongrua-planata  group  retains  an 
aedeagus  with  a  simple  shaft.  PAUP  separates 


1988 


Blocker.  Johnson:  Subgenus  Athysanella 


41 


ro 


4- 


CO 


rf 


4^ 


Q 


^ 


O) 


pastora 

robust  a 

glob  OS  a 

gardenia 

kadokana 

salsa 

incerta 

vativala 

terebrans 

lacta 

marthae 

fredonla 

cursa 

hemijona 

bifida 

utahna 

■  yumana 

■  deserta 
•stylos a 

■  tenera 

■  magdalena 
•  supina 

foeda 

parca 

incongrua 

aphoda 

strobila 

aspera 

plana 

whitcombi 

krameri 

planata 

rostrata 

kanabana 

furnaca 


\ 


I 


q 


o~-  en  4^  OJ  t^ 


Si 


CQ  CQ  OQ  OQ  CO  Ua 

-!  '-J  "1  "1  ^  ■-! 

o  o  o  o  o  o 

c  c;  c  c  c  c 

"X3  "1:3  13  'T3  "O  "O 


•^  - 

■^  - 

^  ■ 
CO  - 
^   - 
CO 
CO 


I  to  — 


tf 


ro 

O) 

ro 

ro 
en 
c;i 

O)  - 


Lcs: 


4^ 

cn 

^  - 


rC 


cn 

Oi 
4^- 


Oulgroup 
robusla 
pastora 
globosa 
gardenia 
krameri 
aspera 
wbitcoinbi 
plana 
parca 
aphoda 
incerta 
planata 
salsa 
kadokana 
strobila 
rostrata 
incongrua 
kanabana 
deserta 
furnaca 
yumana 
tenera 
supina 
foeda 
marthae 
bifida 
magdalena 
laeta 
utahna 
•stylata 
•  vativala 
terebrans 
fredonia 
cursa 
hemijona 


Fig.  118.  Intuitive  cladogram. 

the  incongrua  group  to  a  greater  degree  than 
any  other  group  in  the  intuitive  cladogram. 
The  planata  group  species  are  characterized 
by  an  ocellus  that  is  more  remote  from  the  eye 
than  other  groups  (6')  and  all  split  out  in  the 
first  half  of  the  PAUP  cladogram.  Four  species 
{krameri,  aspera,  whitcombi,  and  p/ona)  arise 
from  the  cladogram  in  sequence. 

Acknowledgments 

Paul  Oman  has  always  been  gracious  and 
very  helpful  in  his  support  of  leafhopper  sys- 
tematics;  it  is  a  high  honor  to  contribute  to  a 


Fig.  119.  PAUP  cladogram. 

volume  dedicated  to  his  many  contributions 
to  our  science.  Merv  Nielson  collected  with 
the  senior  author  in  Mexico  and  elsewhere 
and  has  offered  assistance  throughout  this 
study.  Likewise,  the  assistance  of  Bob  Whit- 
comb  and  Andy  Hicks  has  added  significantly, 
particularly  with  host  information  and  species 
distribution.  Bob  Brooks  frequently  furnished 
work  space  in  the  Snow  Entomological  Mu- 
seum. Paul  Cwikla  has  read  drafts  of  this  and 
related  manuscripts,  and  JefiFry  Ediger,  a  stu- 
dent at  KSU,  inked  the  illustrations  of  the 
genitalia. 


42 


Great  Basin  Naturalist  Memoirs 


No.  12 


Literature  Cited 

Ball,  E.  D  ,  and  R  H  Beamer  1940.  A  revision  of  the 
genus  Athysanella  and  some  related  genera  (Ho- 
moptera:  Cicadellidae).  Univ.  Kansas  Sci.  Bull. 
26:  5-82. 

Blocker,  H.  D.  1984.  Morphological  irregularities  in  the 
external  genital  structures  of  Athysanella  (Ho- 
moptera:  Cicadellidae:  Deltocephalinae).  Bull. 
Soc.  Entomol.  Suisse  57:  412. 

Blocker,  H.  D.,  and  J.  W  Johnson.  1988.  Subgenera  of 
the  genus  Athysanella  Baker  (Homoptera:  Ci- 
cadellidae: Deltocephalinae)  and  a  proposed  phy- 
logeny.  Proc.  VI  Auchenorrhyncha  Mtg.,  Torino, 
Italy,  September  1987. 

Blocker,  H.  D.,  andC  S.  Wesley  1985.  Distribution  of 
Athysanella  (Homoptera:  Cicadellidae:  Delto- 
cephalinae) in  Canada  and  Alaska  with  descrip- 
tions of  three  new  species.  J.  Kansas  Entomol. 


Soc.  52:  377-386. 

Hicks,  A  L  ,  R  F  Whitcomb,  H  D  Blocker,  and  K.  A 
Allred   1988.  Why  are  there  so  many  species  of 
Athysanella?  Proc.   VI  Auchenorrhyncha  Mtg., 
Torino,  Italy,  September  1987. 

Swofford,  D  L.  1986.  PAUP,  phylogenetic  analysis  us- 
ing parsimony.  Private  publication. 

Wesley,  C  S  ,  and  H  D  Blocker  1985.  Athysanella  of 
Mexico  including  descriptions  of  a  new  subgenus 
and  six  new  species  (Homoptera:  Cicadellidae). 
Entomography  3:  163-180. 

Whitcomb,  R  F  ,  A  L  Hicks,  D  E  Lynn,  K  A  Allred, 
andH  D  Blocker.  1987.  Geographic  variation  in 
host  relationships  of  leafhoppers  (Homoptera:  Ci- 
cadellidae) in  North  American  Grasslands.  In:  M. 
R.  Wilson  and  L.  R,  Nault,  eds.,  Proc.  2nd  Int. 
Workshop  on  Leafhoppers  and  Planthoppers  of 
Economic  Importance.  Commonwealth  Inst,  of 
Entomol.,  London. 


A  NEW  GENUS,  ILEOPELTUS,  RELATED  TO  CHLOROTETTIX 
(HOMOPTERA:  CICADELLIDAE) 

Paul  S.  Cwikla' 

Abstract — On  the  basis  of  their  unique  genitahc  characters,  the  Neotropical  leafhopper  species  previously  treated 
in  the  genus  Doleranus  Ball  are  placed  in  a  new  genus,  Ileopeltus.  Twelve  species  are  treated,  including  six  described 
as  new.  A  key  to  the  species  and  a  cladistic  hypothesis  using  Chlorotettix  Van  Duzee  as  the  outgroup  are  presented.  The 
new  species  of  Ileopeltus  include:  nanocanthus  (Panama),  dorsalus  and  clavatus  (Brazil  and  Venezuela),  ventriculus 
and  haplus  (Brazil),  and  blockeri  (Venezuela). 


This  paper  is  the  first  in  a  series  of  revisions 
deahng  with  the  New  World  deltocephahne 
genera,  which  have  their  crown  completely 
microsculptured.  These  leaflioppers,  com- 
monly called  the  broad-headed  leaflioppers, 
are  best  known  from  the  genus  Chlorotettix 
Van  Duzee.  Species  of  this  genus  are  com- 
monly found  in  low-lying,  grassy  habitats  of 
North  and  South  America. 

Traditionally,  those  leaflioppers  with  a 
completely  microsculptured  crown  have  been 
restricted  to  two  genera,  Chlorotettix  Van 
Duzee  and  Doleranus  Ball  (Oman  1949).  Lin- 
navuori  (1959)  mentioned  that  some  species  of 
Paratanus  Young  and  Stirellus  Osborn  &  Ball 
also  have  their  crown  microsculptured,  al- 
though no  degree  of  phylogenetic  relationship 
was  implied. 

More  recently,  I  (Cwikla  1988)  examined 
the  North  American  species  o{ Doleranus  and 
found  that  they  fell  within  an  acceptable  range 
of  variation  for  Chlorotettix.  I  did  not,  how- 
ever, consider  the  Neotropical  species  of  Dol- 
eranus congeneric  with  Chlorotettix  because 
of  the  following  synapomorphies:  the  asym- 
metrical aedeagus,  the  aedeagus  without  pro- 
cesses, and,  with  the  exception  of  only  two 
species,  the  male  plates  fused  with  the  valve. 
Because  of  these  unique  attributes,  the 
Neotropical  species  of  Doleranus  treated  by 
Linnavuori  (1959)  and  six  species  described  as 
new  are  designated  a  new  genus,  Ileopeltus. 

Virtually  nothing  is  known  about  the  biol- 
ogy of  Ileopeltus  species.  Ileopeltus  tethys  has 
been  collected  from  sugar  cane,  weeds,  sweet 
potato  (Wolcott  1923),  and  grassy  pastures 
(Caldwell  and  Martorell  1950)  in  Puerto  Rico. 


Hosts  for  the  other  species  of  Ileopeltus  are 
probably  grasses. 

With  the  exception  of/,  tethys,  specimens 
of  Ileopeltus  species  are  rare  in  collections. 
For  the  most  part  they  appear  restricted  be- 
tween the  Tropic  of  Cancer  and  the  Tropic  of 
Capricorn.  Specimens  have  been  collected  in 
low-  to  mid-altitude  areas. 

Twelve  species  are  included  in  this  new 
genus,  of  which  six  are  described  as  new.  In 
addition,  this  paper  provides  a  key  to  the  spe- 
cies and  a  discussion  of  the  phylogenetic  rela- 
tionships among  the  species.  Specimens  were 
prepared  for  SEM  study  as  described  in 
Cwikla  and  Freytag  (1983).  Names  of  institu- 
tions associated  with  abbreviations  used  in  the 
text  are  in  the  acknowledgments  section. 

Ileopeltus,  n.  gen. 

Diagnosis. — Ileopeltus  species  can  be  sep- 
arated from  other  deltocephaline  genera  by 
the  asymmetrical  aedeagus  without  processes 
and  the  crown  completely  microsculptured. 

Color. — Yellowish  green  or  ochraceous. 
Markings  occasionally  present  on  crown  and 
forewing. 

Structural  features. — Small,  delto- 
cephaline leaflioppers.  Crown  roundly  pro- 
duced, median  length  slightly  longer  than 
length  next  to  eye.  Clypellus  not  constructed 
proximally.  Forewing  with  cross-vein  in  claval 
area. 

Male  genitalia. — Pygofer  roundly  pro- 
duced or  truncated,  process  present  or  ab- 
sent. Anal  tube  weakly  sclerotized  dorsally. 
Valve  acutely  triangular,  usually  fused  to  plate 
posteriorly.  Plate  short  or  slightly  longer  than 


'Department  of  Entomology,  University  of  Missouri,  Columbia.  Missouri  65211,  Present  address:  14560  Josephine  Street,  Irwin,  Pennsylvania  15642. 


43 


44 


Great  Basin  Naturalist  Memoirs 
I 


No.  12 


Figs.  1-4.  Ileopeltus  tethijs  Van  Duzee;  1,  pygofer,  left  lateral  aspect;  2,  right  style,  dorsal  aspect;  3,  aedeagus, 
ventral  aspect;  4,  aedeagus  and  apex  of  connective,  left  lateral  aspect. 


pygofer,  rounded  apically,  macrosetae  unise- 
riate,  second  short  row  of  microsetaelike 
structures  occasionally  present,  lateral  mar- 
gin sinuate,  straight,  or  slightly  convex.  Stylar 
apex  linear,  preapical  angle  usually  absent. 
Connective  as  long  as  or  shorter  than  aedea- 
gus. Aedeagus  without  processes,  asymmetri- 
cal, apex  acute  in  posterior  aspect.  Gonopore 
on  right  side  at  base  of  shaft. 

Type  species. — Chlorotettix  tethijs  Van 
Duzee  1907:71. 

Distribution. — Neotropical,  usually  re- 
stricted between  the  Tropic  of  Cancer  and  the 
Tropic  of  Capricorn. 

Notes. — This  genus  has  been  previously 
described  as  Doleranus  by  Linnavuori  (1959). 
The  Neotropical  species  are  not  considered 
congeneric  with  the  Nearctic  forms  and  have 
been  treated  as  synonyms  oi  Chlorotettix  (in- 
cluding the  type  species  of  Doleranus,  Tham- 
notettix  longulus  Gillette  &  Baker)  in  a  disser- 
tation by  Cwikla  (1988). 

Key  to  the  Male  Ileopeltus  Species 

1.  Pygofer   without    a   process    (Fig.    1),    or    if 

present  then  e.xtremely  small  (Fig.  7) 2 

—  Pygofer  with  long,  distinct  processes  (Fig.  43)    3 

2(1).        Pygofer  without  a  process  (Fig.  1) 

tethys  (Van  Duzee) 


—  Pygofer  with  short  process  on  posterodorsal  mar- 
gin in  lateral  aspect nanocanthus ,  n.  sp. 

.3(1).        Stylar  apex  curved  laterally  (Fig.  52) 

Cyclops  (Linnavuori) 

—  Stylar  apex  linear  (Figs.  12,  30) 4 

4(3).  Pygofer  with  process  directed  ventrad  in  lat- 
eral aspect  ventriculus.  n.  sp. 

—  Pygofer  with  process  directed  mesad  or  dor- 
sad in  lateral  aspect 5 

5(4).  Pygofer  with  process  heavily  sclerotized,  in- 
serted on  medial  side  in  lateral  aspect  (Fig. 
27) 8 

—  Pygofer  with  process  not  heavily  sclerotized, 
not  inserted  on  medial  side  in  lateral  aspect 

(Fig.  11) 6 

6(5).        Aedeagal  shaft  wide  in  ventral  aspect  (Fig.  25) 

hapliis,  n.  sp. 

—  Aedeagal  shaft  narrow  in  ventral  aspect  (Fig. 
22) .  .  .' 7 

7(6).        Stylar  apex  truncate  (Fig.  12)  .  .    dorsalus,  n.  sp. 

—  Stylar  apex  narrow  (Fig.  20)  .  spinosus  (DeLong) 

8(5).        Stylar  apex  truncate  and  constricted  subapi- 

cally  (Fig.  40) clavaUis,  n.  sp. 

—  Stylar  apex  rounded  or  if  truncate,  then  not 
constricted  subapically 9 

9(8).        Pygofer  with  process  inserted  on  middle  of 

ventral  margin  (Fig.  33) blockeri,  n.  sp. 

—  Pygofer  with  process  inserted  on  posteroven- 
tral  margin  (Figs.  27,  49)    10 

10(9).      Aedeagus  wide  and  highly  asymmetrical  in 


1988 


CwiKLA:  The  Genus  Ileopeltus 


45 


Fig.  5.  Ileopeltus  tethys  Van  Duzee,  dorsal  habitus. 

ventral  aspect  (Fig.  49) 

cuneus  (DeLong  &  Martinson) 

—  Aedeagus  narrow  in  ventral  aspect 11 

11(10).  Aedeagus  with  large  gonopore  in  lateral  as- 
pect (Fig.  32);  plate  fused  with  valve 

hasttilus  (DeLong  &  Linnavuori) 

—  Aedeagus  with  small  gonopore  in  lateral  as- 
pect (Fig.  46);  plate  not  appearing  fused  with 
valve aberrans  (Osborn) 

Ileopeltus  tethys  (Van  Duzee),  n.  comb. 
Figs.  1-5,  57,  59;  Map  1 

Chlorotettix  tethys  Van  Duzee  1907:71. 

Chlorotettix  hidentatus  DeLong  1923:264,  Wolcott  1936: 

86. 
Chlorotettix  dilutus  Oshorn  1923:73,  Osborn  1935:118. 
Doleranus  kinonanus  Ball  1936:432,   Linnavuori  1959: 

274-275. 

Diagnosis. — Ileopeltus  tethys  is  near  /. 
nanocantus  and  can  be  separated  from  it  and 
other  Ileopeltus  species  by  the  lack  of  a 
pygoferal  spine. 

Length. — Male  3.8-4.8  mm,  female  4.1- 
5.4  mm. 


Color, — Yellowish  green.  Eye  reddish 
brown.  Forewing  yellowish  subhyaline,  small 
brown  patch  occasionally  present  on  middle  of 
wing  and/or  on  claval  area. 

Male  genitalia. — Pygofer  with  posterior 
margin  roundly  produced,  processes  absent. 
Plate  short,  approximately  half  the  length  of 
pygofer,  lateral  margin  straight  or  slightly 
concave,  not  fused  to  valve.  Stylar  apex  stout, 
linear,  preapical  angle  absent.  Aedeagus 
asymmetrical  in  posterior  aspect,  left  subapi- 
cal  margin  sinuate  in  posterior  aspect,  right 
subapical  margin  straight  or  somewhat  convex 
in  posterior  aspect.  Gonopore  at  base  on  right 
side. 

Female  seventh  sternum. — Posterior 
margin  shallowly  excavated,  median  of  exca- 
vation with  short,  produced  projection  bear- 
ing two  short  teeth  (see  Linnavuori  1959:  Fig. 
113g).  Depth  of  excavation  varies  from  only 
slightly  to  one-third  width  of  segment. 

Type. — A  single  female  cotype  bearing  the 
labels  "Martinique  W.I.,  VII-26"  and  "Aug. 
Busck  collector"  was  examined  from  the 
USNM. 

Distribution. — ^Many  specimens  from  the 
following  localities  were  examined:  Central 
America:  Honduras,  Mexico  (Chiapas,  Guer- 
rero, Michoacan,  Oaxaca,  Tamaulipas,  Vera- 
cruz), Nicaragua,  Panama.  West  Indies:  An- 
tigua, Cayman  Islands,  Dominican  Republic, 
Guadeloupe,  Haiti,  Jamaica,  Montserrat, 
Nevis,  Puerto  Rico,  Trinidad,  and  Tobago. 
South  America:  Venezuela  (Guarico). 

Specimens  were  collected  from  May 
through  December  and  are  deposited  in  the 
UPB,  UCV,  UKC,  USNM,  OSUC,  and 
BMNH  collections. 

Note. — The  types  of  hidentatus,  dilutus, 
and  kinonanus  were  not  available  for  study; 
consequently,  previous  synonymies  of  these 
names  under  /.  tethys  were  not  verified.  Be- 
cause the  types  are  females,  it  is  doubtful  that 
they  can  be  correctly  associated  with  males  at 
this  time. 

Ileopeltus  nanocanthus,  n.  sp. 

Figs.  6-10,  58,  60;  Map  2 

Diagnosis. — Ileopeltus  nanocanthus  is 
near  /.  tethys  and  can  be  separated  from  it  and 
other  Ileopeltus  species  by  the  short  pygoferal 
spine. 

Length. — Male  4.6-5.4  mm,  female  4.8- 
5.3  mm. 


46 


Great  Basin  Naturalist  Memoirs 


No.  12 


Map  1.  Distribution  of  Ileopeltiis  tethijs. 


Color. — Ochraceous  or  greenish  yellow. 
Eye  reddish  brown.  Crown  occasionally  with 
faint  orange,  median  band.  Pronotum  with 
four  longitudinal,  faint  orange  bands.  Fore- 
wing  yellowish  subhyaline,  brown  spots 
present  at  bases  of  anteapical  cells  and  on 
claval  area. 

Male  genitalia. — Pygofer  with  ventral 
margin  roundly  produced  into  short  spine  on 
dorsoposterior  margin,  spine  directed  medi- 
ally in  ventral  aspect.  Plate  short,  anterior 
margin  convex.  Stylar  apex  narrowed,  di- 
rected posteriorly,  preapical  angle  absent. 
Aedeagus  linear,  asymmetrical  in  posterior 
aspect,  apex  with  opposite  curve  compared  to 
/.  tethys.  Gonopore  at  base  on  right  side. 

Female  seventh  sternum. — Posterior 
margin  with  U-shaped  excavation  extending 
half  length  of  segment,  base  of  excavation  with 
small  V-shaped  notch,  notch  surrounded  by 
light  brown  coloration,  lateral  angle  rounded. 

Type. — Male  holotype,  Panama,  Canal 
Zone,  Curundu,  26-3'()-XI-1986,  B.  Sieber- 
glied  collector.  Four  female  paratypes,  same 


data  as  holotype;  one  male  paratype,  Panama, 
Chiriqui,  Rio  Colorado,  1,200  m,  8°5'N, 
82°43'W,  26-XII-1974,  H.  Wolda  collector; 
male  paratvpe,  Panama,  Panama,  Las  Cum- 
bres,  17-23-11-1982,  H.  Wolda  collector. 
Holotype,  two  paratypes  with  same  data  as 
holotype,  and  one  paratype  from  Las  Cum- 
bres  deposited  in  OSUC,  remainder  in  HWC. 
Etymology. — Greek,  nano  (dwarf)  and 
acantha  (spine)  refer  to  the  small  pygofer  pro- 


Ileopeltiis  dorsalus,  n.  sp. 

Fig.s.  11-14;  Map  2 

Diagnosis. — Ileopeltus  dorsalus  is  near  7. 
ventriculus  and  can  be  separated  from  it  by 
the  acute,  dorsally  directed  pygoferal  spine. 

LENCiTH. — Male  4.4  mm,  female  unknown. 

Color. — Ochraceous,  either  without  dark 
markings  or  crown  with  light  brown  surround- 
ing coronal  sulcus.  Eye  red.  Pronotum  with 
six  light  brown,  longitudinal  bands.  Scutel- 
lum  with  brown  lateral  angles.  Forewing  sub- 
hyaline  with  brown  patches  at  base  of  anal 


1988 


CwiKLA:  The  Genus  Ileopeltus 


47 


Figs.  6-10.  Ileopeltus  nanocanthus,  n.  sp.:  6,  pygofer,  left  lateral  aspect;  7,  pygofer,  ventral  aspect;  8,  right  style, 
dorsal  aspect;  9,  aedeagus,  ventral  aspect;  10,  aedeagus  and  apex  of  connective,  left  lateral  aspect. 


Map  2.  Distribution  of  Ileopeltus  nanocanthus  (squares),  /.  dorsalus  (circles),  and  I.  ventriculus  (triangles) 


48 


Great  Basin  Naturalist  Memoirs 


No.  12 


Figs.  11-14.  Ileopeltus  dorsalus,  n.  sp.:  11,  pygofer,  left  lateral  aspect;  12,  right  style,  dorsal  aspect;  1.3,  aedeagus, 
ventral  aspect;  14,  aedeagus  and  apex  of  connective,  left  lateral  aspect. 


veins,  proximal  and  distal  ends  of  inner  ante- 
apical  cell,  and  proximal  end  of  central  ante- 
apical  cell. 

Male  genitalia. — Pygofer  with  posterior 
margin  produced  into  dorsally  directed  spine, 
acute  apically,  not  heavily  sclerotized,  small 
group  of  microsetaelike  structures  near  mid- 
dle of  posterior  margin.  Plate  triangular,  lat- 
eral margin  straight.  Stylar  apex  truncate,  di- 
rected posteriorly,  preapical  angle  small. 
Aedeagus  only  slightly  asymmetrical. 

Type. — Male  holotype,  Venezuela,  Aragua, 
El  Limon,  22-11-1973,  Malaise  trap,  C.  J. 
Resales  collector.  Male  paratype,  Brazil, 
Ceara,  Barbalha,  V-1969,  M.  Alvarenga,  B.M. 
1971-165.  Holotype  deposited  in  OSUC, 
paratype  in  BMNH. 

Etymology, — Latin,  dorso  (back)  refers  to 
the  dorsally  directed  pygoferal  spine. 

Ileopeltus  ventriciilus ,  n.  sp. 

Figs.  1.5-18;  Map  2 

Diagnosis. — Ileopeltus  ventriculus  is  near 
spinosus  and  can  be  separated  from  it  and 
other  Ileopeltus  species  by  the  pygoferal  pro- 
cess directed  ventrally. 

Length. — Male  4.4  mm,  female  unknown. 

Color.— Ochraceous,  without  distinct  dark 


markings.  Eye  reddish  brown. 

Male  genitalia. — Pygofer  with  short,  ven- 
trally directed  process  inserted  on  posterior 
margin.  Plate  elongate,  lateral  margin  sinu- 
ate, apex  curved  dorsally,  bluntly  rounded. 
Style  elongate,  apex  linear,  slightly  curved 
laterally.  Aedeagus  elongate,  broadly  curved 
dorsally  in  lateral  aspect,  apex  acute  in  poste- 
rior aspect. 

Type. — Male  holotype,  Brazil,  Ceara, 
Crato,  May  1969,  M.  Alvarenga  collector. 
Type  deposited  in  BMNH. 

Etymology. — Latin,  ventricul  (belly) 
refers  to  the  ventrally  directed  pygofer  pro- 
cess. 

Ileopeltus  spinosus  (Dehong),  n.  comb. 

Figs.  19-22;  Map  3 

Chlorotettix  spinosus  DeLong  1945:10-11. 

Diagnosis. — Ileopeltus  spiiiosus  is  near  /. 
dorsalus  and  can  be  separated  from  it  and 
other  Ileopeltus  species  by  the  presence  of  a 
preapical  angle  on  the  style  and  the  unique 
pygoferal  process. 

Length. — Male  5.2-5.5  mm,  female  un- 
known. 

Color. — Greenish  yellow,  without  dark- 
markings.  Eye  gray  or  greenish  yellow. 


1988 


CwiKLA:  The  Genus  Ileopeltus 


49 


Figs.  15-18.  Ileopeltus  ventricuhis ,  n.  sp.:  15,  pygofer,  left  lateral  aspect;  16,  right  style,  dorsal  aspect;  17,  aedeagus, 
ventral  aspect;  18,  aedeagus  and  apex  of  connective,  left  lateral  aspect. 


Male  genitalia. — Pygofer  with  posterior 
margin  broadly  rounded  apically,  heavily 
sclerotized  process  inserted  on  posteroventral 
margin,  directed  posteriorly  then  curved  me- 
dially. Plate  with  lateral  margin  slightly  sinu- 
ate, apex  elongate,  pointed.  Stylar  apex  nar- 
rowly elongate,  directed  slightly  laterally, 
preapical  angle  roundly  triangular.  Aedeagus 
elongate,  not  strongly  asymmetrical,  narrow 
in  ventral  aspect.  Gonopore  on  right  side  near 
base. 

Type. — Male  holotype  bearing  the  labels 
"Buena  Vista,  Gro.  [Guerrero,  Mexico],  23-X- 
1941,  3,400  ft."  and  "DeLong,  Good,  Cald- 
well and  Plummer"  was  examined  from 
OSUC. 

Distribution. — Besides  the  type,  three 
paratypes  from  Mazaclan,  Guerrero,  Mexico, 
3-X-1945,  were  examined  from  OSUC. 

Note. — DeLong  (1945)  reported  this  spe- 
cies from  meadow  grasses. 

Ileopeltus  haplus,  n.  sp. 

Figs  23-26;  Map  3 

Diagnosis. — Ileopeltus  haplus  is  near  /. 
spinosus  and  can  be  separated  from  it  and 
other  Ileopeltus  species  by  the  unique  pygofer 
process  and  the  wide  aedeagus  in  ventral  as- 
pect. 


Length. — Male  5.3  mm,  female  unknown. 

Color. — Yellowish  green,  without  dark 
markings.  Eye  yellowish  green. 

Male  genitalia. — Pygofer  with  posterior 
margin  produced  into  dorsally  directed  spine, 
spine  not  heavily  sclerotized.  Plate  triangular, 
lateral  margin  straight,  apex  rounded.  Stylar 
apex  linear,  preapical  angle  absent.  Aedeagus 
asymmetrical,  left  margin  forming  carina  in 
posterior  aspect.  Gonopore  on  right  side  near 
base. 

Type. — Male  holotype,  [Brazil]  MS  [Matto 
Grosso  du  Sul],  Campo  Grande,  l-X-1982, 
W.  KoUer  collector.  Type  deposited  in  UPB. 

Eri'MOLOGY. — Greek,  haplo  (simple)  refers 
to  the  color  of  this  species. 

Ileopeltus  hastulus  (DeLong  & 
Linnavuori),  n.  comb. 

Figs.  27-32;  Map  3 

Chlorotettix  hastulus  DeLong  &  Linnavuori  1978:121- 
122. 

Diagnosis. — This  species  can  be  separated 
from  other  members  of  this  genus  by  the  long 
pygofer  process  and  the  aedeagus  with  a  large 
gonopore. 

Length. — Male  5.0-5.2  mm,  female  un- 
known. 


50 


Great  Basin  Naturalist  Memoirs 


No.  12 


Figs.   19-22.  Ileopeltus  spinosus  (DeLong):  19,  pygofer,  left  lateral  aspect;  20,  right  style,  dorsal  aspect;  21, 
aedeagus,  ventral  aspect;  22,  aedeagus  and  apex  of  connective,  left  lateral  aspect. 


Map  3.  Distribution  of //eo/«'/<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 

Literature  Cited 

Ball.  E  G  1936.  Some  new  leafhoppers  related  to  Tham- 
notettix.  J.  Washington  Acad.  Sci.  26:  430-434. 

Caldwell,  J  S  ,  and  L  F  Martorell  1950.  Review  of 
the  Auchenorynchous  Homoptera  of  Puerto  Rico. 
Part  1.  Cicadellidae.  Puerto  Rico  Univ.  J.  Agric. 
34:  1-132. 

CwiKLA,  P.  S.  1988.  A  systematic  study  of  the  leafhopper 
genus  Chlorotettix  (Homoptera:  Cicadellidae). 
Unpublished  dissertation,  Ohio  State  University, 
Columbus. 

CwiKLA,  P  S  ,  AND  P.  H  Freytag  1983.  External  mor- 
phology of  Xestocephalus  subtessellatus  (Ho- 
moptera: Cicadellidae:  Xestocephalinae).  Ann. 
Entomol.  Soc.  Amer.  76:  641-652. 

DeLong.  D  M.  1923.  New  species.  Pages  257-270  in 
G.  N.  Wolcott,  Insectae  Portoricensis.  Puerto 
Rico  Univ.  J.  Agric.  Vol.  7. 

1945.  The  genus  Chlorotettix  (Homoptera:  Ci- 
cadellidae) in  Mexico.  Lloydia8:  1-30. 

DeLong,  DM,  and  P  S.  Cwikla  1984.  New  Neotropi- 
cal Deltocephalinae  (Homoptera:  Cicadellidae).  J. 
Kansas  Entomol.  Soc.  57:  725-728. 

DeLong,  D  M  ,  andC  Martinson  1974.  New  species  of 
Chlorotettix  from  Argentina,  Bolivia,  Brazil,  and 
Mexico.  J.  Kansas  Entomol.  Soc.  47:  261-267. 

DeLong,  D  M  ,  and  R  E  Linnavuorl  1978.  Studies  on 
Neotropical  leafhoppers  (Homoptera:  Cicadelli- 
dae). Entomol.  Scand.  9:  111-123. 

Hennig,  W  1966.  Phylogenetic  systematics.  University 
of  Illinois,  Urbana. 

Linnavuorl  R  E  1959.  Revision  of  the  Neotropical  Del- 
tocephalinae and  some  related  subfamilies  (Ho- 
moptera: Cicadellidae).  Ann.  Zool.  Soc.  Vanamo 


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 

0 

0 

0 

0 

0 

0 

0 

0 

0 

0 

0 

0 

1.  cuneus 

1 

1 

0 

1 

1 

1 

0 

1 

1 

I.  Cyclops 

1 

1 

0 

0 

1 

1 

1 

0 

0 

I.  aherrans 

1 

1 

0 

I 

0 

0 

1 

0 

0 

0 

1.  blockeri 

0 

1 

0 

1 

0 

0 

0 

0 

0 

I.  hastulus 

0 

1 

0 

1 

0 

0 

0 

0 

1 

I.  tethijs 

0 

0 

0 

0 

0 

0 

1 

1 

0 

0 

0 

I.  nanocanthus 

0 

0 

0 

0 

0 

1 

1 

0 

0 

0 

I.  haplus 

0 

1 

1 

0 

0 

0 

1 

0 

1 

0 

I.  dorsahis 

1 

1 

1 

0 

0 

0 

0 

0 

0 

0 

1.  clavatus 

0 

1 

0 

1 

1 

0 

1 

0 

0 

0 

I.  sphwsus 

0 

1 

1 

0 

0 

0 

1 

0 

1 

0 

I.  ventriculus 

0 

1 

1 

0 

0 

0 

1 

0 

0 

0 

20:  1-370. 

OSBORN.  H  1923.  Neotropical  Homoptera  of  the 
Carnegie  Museum.  Part  2.  Records  and  descrip- 
tions of  five  new  genera  and  sixty-five  new  species 
of  the  subfamily  Jassinae.  Carnegie  Museum  Ann. 
1.5:  27-79. 

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- 
nary annotated  check-list  of  the  insects  of  Puerto 
Rico,  with  descriptions  of  new  species.  Puerto 
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 


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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. 


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Great  Basin  Naturalist  Memoirs 


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 


Great  Basin  Naturalist  Memoirs 


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, 


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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, 


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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. 


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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 


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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. 


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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. 

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KiRKALDY,  G  W  1900.  Bibliographical  and  nomenclatu- 
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1910.  Fauna  Hawaiiensis  2:  531-700. 

KiRSCHBAUM,  C  L  1868.  Die  Cicadinen  der  Gegend  von 
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zahl  neuer  oder  schwer  zu  unterscheidender 
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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- 
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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- 
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OSSIANNILSSON,  F  1983.  The  Auchenorrhyncha  (Ho- 
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the  family  Cicadellidae:  Deltocephalinae,  cata- 
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Simon,  C.  1987.  Hawaiian  evolutionary  biology:  an  intro- 
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SoUTHWOOD,  T  R  E  1978.  Ecological  methods.  Chap- 
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TiMBERLAKE,  P  H  1918.  Note  on  the  non-identity  of  a 
common  Hawaiian  jassid  with  Nesosteles  hebe 
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tettix  (Homoptera:  Auchenorrhyncha).  Journal  of 
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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 


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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 


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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 


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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; 


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107 


connective  long,  extending  to  about  apex  of 
style  (Fig.  2);  style  with  distal  shaft  broad, 
lateral  margins  smooth,  stylar  spine  broad 
and  bluntly  pointed,  projecting  laterodistallv 
(Fig.  4). 

Female. — Seventh  sternum  produced  dis- 
tally  to  form  broad  medial  lobe,  with  median, 
shallow  notch  on  its  caudal  margin,  configura- 
tion of  lobe  variable,  sometimes  with  small 
lobe  at  base  of  notch. 

Distribution. — This  species  has  been  col- 
lected in  the  Mexican  states  of  Distrito  Fed- 
eral, Michoacan,  Morelos,  and  Hidalgo. 

Host. — Some  specimens  were  collected  in 
pine  forests  at  elevations  of  7,500  feet  from 
July  to  October  (DeLong  1946). 

Remarks. — The  configuration  of  the  female 
seventh  sterna  of  the  holotypes  of  nigridens 
and  sexpunctatiis  is  similar,  with  some  intra- 
and  interpopulation  variation  in  the  angle  of 
the  posteriolateral  margins.  The  abdomen  of 
the  female  holotype  oi  albocinctus  is  missing, 
but  comparisons  were  made  with  paratype 
female  specimens  after  they  were  associated 
with  holotype  and  with  DeLong's  illustration 
of  the  seventh  sternum.  Although  the  male 
genital  structures  of  the  allotypes  of  albocinc- 
tus  and  nigridens  are  nearly  identical,  the 
latter  is  more  deeply  sclerotized  than  the  for- 
mer. The  sexes  of  both  species  appear  to  be 
properly  associated. 

The  female  holotype  of  sexpunctatus  is 
identical  in  general  habitus  with  the  female 
holotype  oi  nigridens,  but  both  species  differ 
markedly  in  color  patterns  from  albocinctus. 
The  latter  species  has  a  distinctive  pronotal 
band  that  is  absent  in  the  specimens  of  the  two 
former  species.  The  male  genital  structures, 
however,  are  identical  among  these  popula- 
tions, which  suggests  that  two  infraspecific 
entities  may  be  involved.  Such  action  is  de- 
ferred until  more  material  is  collected  and 
studied. 

Colladonus  albocinctus  is  distinguished 
homfasciaticollis  by  the  position  of  the  pygo- 
fer  spine,  which  is  distinctly  basad  of  the  mid- 
dle of  the  caudal  margin  of  the  pygofer. 

Colladonus  belli  (Uhler) 
Figs.  6-10 

Jassus  belli  Uhler  1877:471  [Holotype  9  (USNM)  (exam- 
ined).] 
Thamnotettix  belli:  Van  Duzee  1892:306 
Colladonus  belli:  Ball  1936:58 
Idiodonus  belli:  Medler  1943:18 


Colladonus  belli:  Nielson  1957:38 
Colladonus  belli:  Metcalf  1967: 1245 
Thamnotettix  semipullatus  Van  Duzee  1892:306 
Colladonus  semipullatus:  Oman  1949:125 
Thamnotettix  fiilletti  Van  Duzee  1892:306 
Colladonus  gilk'tti:  Oman  1949:125 
Thamnotettix  sonorae  (;illette  &  Baker  1895:100 
Colladonus  sonorae:  Oman  1949:125 

Length. — Male,  4.0-4.4  mm;  female,  4.4- 
5.2  mm. 

A  well-marked  species  from  pale  yellow  to 
deep  yellow  with  two  black  spots  on  anterior 
margin  of  crown  and  sometimes  with  a  black, 
transverse  band  on  disc  of  crown;  pronotum 
with  pale  yellow  to  deep  yellow,  transverse 
band;  forewings  with  veins  pale  to  yellow; 
color  intensity  highly  variable.  Similar  to  fas- 
ciaticollis  in  general  habitus  but  with  distinc- 
tive male  genitalia. 

Head  with  anterior  margin  not  produced, 
apex  rounded. 

Male. — Pygofer  in  lateral  view  with  short, 
stout  spine  near  or  above  middle  of  caudal 
margin,  spine  projecting  dorsally  (Fig.  6); 
aedeagus  in  lateral  view  with  long,  bifurcated 
distal  processes  extending  beyond  midlength 
of  shaft  (Fig.  8),  crossing  over  in  dorsal  view 
(Fig.  10),  gonopore  about  middle  of  shaft;  con- 
nective short,  not  reaching  apex  of  style  (Fig. 
7);  style  with  moderately  broad  stylar  shaft, 
inner  lateral  margin  serrate,  outer  one 
smooth,  stylar  spine  moderately  long,  pro- 
jecting laterally  (Fig.  9). 

Female. — Seventh  sternum  with  truncate 
caudal  margin,  shallow,  U-shaped  excision 
medially. 

Distribution. — This  species  is  widespread 
in  mountainous  areas  from  Mexico  north  to 
British  Columbia.  In  Mexico  the  author  col- 
lected specimens  in  the  state  of  Chihuahua.  It 
has  been  reported  in  seven  western  states  in 
the  United  States  and  in  the  province  of 
British  Columbia  (Nielson  1957). 

Host. — Adults  have  been  swept  from  a  va- 
riety of  trees  and  shrubs  by  the  author.  Sev- 
eral specimens  were  taken  from  Monarda  sp. 
near  citriodora  Cerv.  in  Chihuahua  state  in 
Mexico  by  the  author. 

Remarks. — Colladonus  belli  may  repre- 
sent a  species  complex  that  occupies  a  rather 
narrow,  longitudinal  range  in  western  North 
America.  Specimens  north  of  Utah  and 
Colorado  are  almost  uniformly  yellow-green 
without  a  dark  band  across  the  crown, 
whereas    southern    populations    are    more 


108 


Great  Basin  Naturalist  Memoirs 


No.  12 


Figs.  6-10.  CoUadomts  belli  (Uhler):  6,  male  pygofer,  lateral  view;  7,  connective  and  right  style,  dorsal  view;  8, 
aedeagus,  lateral  view;  9,  apex  of  right  style,  enlarged  dorsal  view;  10,  apex  of  aedeagus  and  distal  processes,  dorsal 
view. 


deeply  marked  with  yellow  and  black  mark- 
ings. The  position  of  the  pygofer  spine,  al- 
though similar  in  configuration,  varies  from 
distad  of  the  middle  of  the  caudal  margin  in 
northern  populations  to  the  middle  of  the  cau- 
dal margin  in  southern  populations.  No  varia- 
tion in  the  configuration  of  the  female  seventh 
sternum  was  evident,  however. 

Much  additional  collecting  over  the  entire 
range  of  this  species  complex  is  needed.  Pop- 
ulations are  not  commonly  collected,  and 
when  found,  specimens  are  usually  few  in 
number.  A  reexamination  of  the  type  speci- 
mens of  belli,  semipullatus,  ^illettei,  and 
sonorae  is  recommended  for  any  future  study. 

From  fasciaticollis,  to  which  it  is  similar, 
belli  can  be  distinguished  by  the  short  pygofer 
species  has  been  made,  nor  has  any  type  of 


die  to  upper  middle  portion  of  the  caudal 
margin. 

Colladonus  serratiis,  n.  sp. 
Figs.  11-14 

Length. — Male,  4.8-5.0  mm;  female,  5.4 
mm. 

Color  dark  brown  to  blackish;  crown  yellow 
with  two  large,  (juadrate,  black  spots  on  ante- 
rior margin,  disc  with  broad,  black,  transverse 
band;  pronotum  with  yellow,  narrow  to 
broad,  transverse  band;  forewings  black  ex- 
cept for  transculent  area  along  middle  of 
costa,  veins  yellow;  face  yellow  with  black 
markings. 

Head  with  anterior  margin  produced,  apex 
obtusely  angled. 

Male. — Pygofer  in  lateral  view  with 
broadly  roimded  caudal  margin,  caudoventral 


1988 


NiELSON:  COLLADONUS  OF  MEXICO  AND  CENTRAL  AMERICA 


109 


Figs.  11-14.  CoUadonus  serratus,  n.  sp.:  11,  male  pygofer,  lateral  view;  12,  connective,  right  style,  and  aedeagus, 
dorsal  view;  13,  aedeagus,  lateral  view;  14,  apex  of  right  style,  enlarged  dorsal  view. 


margin  toothed  or  serrate,  short  spine  above 
middle  of  caudal  margin,  directed  dorsad 
(Fig.  11);  aedeagus  in  lateral  view  narrow 
along  major  portion  of  shaft  with  moderately 
long,  distal,  bifurcated  processes  reaching  to 
about  middle  of  shaft  (Fig.  13),  processes 
broad  medially,  gradually  tapered  distally, 
crossing  over  in  dorsal  view  (Fig.  12),  gono- 
pore  basad  of  middle  of  shaft;  connective 
short,  not  reaching  apex  of  style  (Fig.  12); 
style  with  broad  stylar  shaft,  stylar  spine  sub- 
apical,  small,  sharply  pointed  (Fig.  14). 

Female. — Seventh  sternum  with  truncate 
caudal  margin,  small  tooth  medially. 

HoLOTYPE  (male). — Mexico:  Mex.,  7  miles 
east  of  Amecameca,  9,300  ft,  31. VI.  1974,  C. 
W.  and  L.  O'Brien  and  Marshal  (USNM). 
Allotype  female,  same  locality  as  holotype, 
10,000  ft,  17.  VIII.  1982,  C.  W.  and  L.  O'Brien 
and  G.  Wibner  (USNM).  Paratypes:  one 
male,  same  data  as  female  allotype  (in  au- 
thor's collection);  one  male,  Mexico,  #16,  7 
miles  southeast  of  Amecameca,  9,000  ft, 
17.  VIII.  1969,  George  W.  Byers  (UK). 

Remarks. — ^This  is  a  rather  unique  species 
that  is  not  related  to  other  species  oi  CoUa- 


donus in  Mexico.  The  combination  of  the  sub- 
apical  stylar  spine  and  toothed  caudovcntral 
margin  of  the  pygofer  will  readily  distinguish 
it  from  all  other  known  species  in  the  region. 

CoUadonus  fasciaticollis  (Stal) 
Figs.  15-22 

J assiis  fasciaticollis  Stal  1864:86  [Holotype  9  (NM)  (ex- 
amined).] 

Thamnotettix  fasciaticollis:  DeLong  1946:13 

CoUadonus  fasciaticollis:  Nielson  1957:37 

Thamnotettix  fasciaticollis:  Metcalf  1967:756 

Idiodonus  tubtdus  DeLong  1946:22  [Holotype  9  (OSU) 
(examined).] 

CoUadonus  tubulus:  Nielson  1957:37 

Idiodonus  tubulus:  Metcalf  1967:1303 

Idiodonus  tubulus:  DeLong  1984:10 

Idiodonus  diserus  DeLong  1946:24  [Holotype  9  (OSU) 
(examined).] 

CoUadonus  diserus:  Nielson  1957:37 

Idiodonus  diserus:  Metcalf  1967:1297 

Idiodonus  diserus:  DeLong  1984:11 

Idiodonus  pravus  DeLong  1946:24  [Holotype  9  (OSU) 
(examined).]  New  synonymy 

Idiodonus  pravus:  Metcalf  1967:1301 

Idiodonus  pravus:  DeLong  1984:11 

Idiodonus  dampfi  DeLong  1946:20  [In  part,  allotype  6 
and  8  paratypes  (OSU)] 

Idiodonus  dampfi:  Linnavuori  1959:280  [Misdetermined 
specimens.] 


no 


Great  Basin  Naturalist  Memoirs 


No.  12 


Figs.  15-22.  Colladonus  fasciaticollis  (Stal):  15,  male  pygofer  of  specimen  from  Costa  Rica,  lateral  view;  16,  male 
pygofer  of  allotype  oi  tubulus ,  lateral  view;  17,  male  pygofer  of  allotype  oi  disertis,  lateral  view;  18,  male  pygofer  of 
allotype  of  dampfi  (=  fasciaticollis),  lateral  view;  19,  connective  and  right  style,  dorsal  view;  20,  aedeagus,  lateral  view; 
21,  apex  of  right  style,  enlarged  dorsal  view;  22,  apex  of  aedeagus  and  distal  processes,  enlarged  dorsal  view. 


Length. — Male,  4.4-4.6  mm;  female, 
4.8-5.0  mm. 

A  well-marked  black  and  yellow  species 
usually,  but  sometimes  paler  in  some  speci- 
mens. Similar  in  general  habitus  to  belli  and  to 
albocinctus  in  male  genitalic  characters  but 
distinct  from  both  species. 

Head  with  anterior  margin  not  produced, 
apex  nearly  rounded. 

Male. — Pygofer  in  lateral  view  with  short, 
bluntly  pointed  spine  arising  below  middle  of 
caudal  margin  (some  variation  in  origin  of 
spine  on  caudoventral  margin),  basal  3/4  of 
spine  sclerotized  and  fused  to  pygofer  wall 
(Figs.  15,  16,  17,  18);  aedeagus  in  lateral  view 
with  moderately  long  and  narrow  (sometimes 
broad),  distal,  bifurcated  processes  extending 
to  about  middle  of  shaft  (Fig.  20),  crossing 
over  in  dorsal  view  (Fig.   22),   gonopore  at 


about  middle  of  shaft;  connective  short,  apex 
not  reaching  to  apex  of  style  (Fig.  19);  style 
with  tapered  stylar  shaft,  lateral  margins 
smooth,  stylar  spine  slanted  laterodistally 
(Fig.  22). 

Female. — Seventh  sternum  with  caudal 
margin  obtusely  angled  with  shallow,  U- 
shaped,  median  emargination. 

Distribution. — Colladonus  fasciaticollis  is 
widely  distributed  from  southern  California  to 
Costa  Rica.  Its  northern  range  into  the  United 
States  is  restricted  to  the  southern  border  of 
California.  It  is  common  in  southern  Mexico 
in  the  states  of  Distrito  Federal,  Veracruz, 
Michoacan,  and  Puebla. 

Host. — This  species  presumably  occurs  on 
unspecified  trees  or  shrubs  in  Mexico  from 
September  to  November  at  elevations  from 


1988 


NiELSON:CoLLADONus  OF  Mexico  AND  Central  America 


111 


5,000  to  10,000  feet  (DeLong  1946).  In  Costa 
Rica  it  was  taken  on  Lippia  herlandieri  in 
February  (Nielson  1957). 

Remarks. — In  my  earlier  work  (Nielson 
1957),  tubulus  and  diserus  were  transferred  to 
Colladonus  and  suppressed  as  synonyms  of 
fasciaticollis .  Reexamination  of  the  female 
holotypes  and  comparison  of  them  with  au- 
thentically determined  female  specimens  of 
fasciaticollis  confirm  that  action. 

The  female  holotype  of  pravus  is  identical 
to  its  counteipart  holotype  specimens  of  the 
above  species.  The  male  allotype  of  dampfi 
from  Bella  Vista,  Mexico  (MB  366),  which 
apparently  had  been  misidentified  and  im- 
properly associated  with  the  female  holotype, 
was  associated  with  the  male  holotype  of 
pravus  from  Zitacuaro,  Michoacan.  Two  of 
five  male  paratype  specimens  and  three  fe- 
male paratype  specimens  of  dampfi  from 
Orizaba,  Veracruz,  were  also  associated  with 
pravus. 

From  albocinctus  and  belli,  to  which  it  has 
similar  genitalia,  fasciaticollis  can  be  distin- 
guished by  the  pygofer  spine  that  arises  from 
near  the  middle  of  the  caudal  margin,  the 
shorter  aedeagus,  and  the  stouter,  bifurcated 
distal  processes. 

Colladonus  idtimus,  n.  sp. 

Figs.  23-26 

Length. — Male,  3.9-4.2  mm;  female, 
4.7-4.9  mm. 

General  color  dark  brown  to  black;  crown 
yellow  with  two  black  spots  on  anterior  mar- 
gin and  narrow,  black,  transverse  band  be- 
tween eyes;  pronotum  black  with  narrow,  yel- 
low, transverse  band  at  middle;  forewings 
black  with  yellow  veins,  costal  area  hyaline; 
face  yellow  with  black  markings.  Similar  in 
general  habitus  to  dampfi  (DeLong)  but  with 
distinctive  male  genitalia. 

Head  with  anterior  margin  produced,  apex 
rounded. 

Male. — Pygofer  in  lateral  view  with  caudal 
margin  broadly  rounded,  pygofer  spine  short, 
slightly  longer  than  basal  width,  arising  just 
below  midline  of  pygofer  and  projecting  cau- 
dodorsally  (Fig.  23);  aedeagus  in  lateral  view 
with  narrow  shaft,  distal,  bifurcated  processes 
long,  more  than  1/2  length  of  shaft  and  extend- 
ing beyond  gonopore  (Fig.  25),  processes  of 
uniform  width  except  for  tapered  apex, 
gonopore  at  about  midlength  of  shaft;  connec- 


tive short,  apex  not  reaching  apex  of  style 
(Fig.  24);  style  with  broad  stylar  shaft,  nearly 
straight,  stylar  spine  short,  sharply  pointed 
distally  and  projecting  laterally  (Fig.  26). 

Female. — Seventh  sternum  with  caudal 
margin  nearly  truncate,  with  small  indenta- 
tion medially. 

Holotype  (male). — Panama:  Chiriqui, 
Bambito,  1,400  m,  10. VI.  1976,  Wolda  and 
Estribi(USNM). 

Remarks. — Numerous  specimens  of  this 
species  that  were  collected  in  Costa  Rica  and 
preserved  in  alcohol  were  too  badly  distorted 
and  discolored  to  be  used  in  the  type  series. 
They  were  all  taken  at  Cerro  de  la  Muerte, 
10,000  ft,  I.  VIII. 1966,  S.  L.  Wood,  and  are  in 
the  M.  L.  Bean  Museum  collection.  A  single 
female  specimen  of  this  species  from  Yepo- 
capa,  Guatemala,  XII.  1948,  H.  T.  Dalmat, 
was  also  examined. 

This  species  thus  far  represents  the  south- 
ernmost extremity  of  the  range  ofColladonus. 
It  can  be  distinguished  from  dampfi  by  the 
shorter  pygofer  spine,  by  the  gonopore, 
which  lies  midlength  of  the  aedeagal  shaft, 
and  by  the  shorter  connective. 

Colladonus  tolucensis,  n.  sp. 
Figs.  27-30 

Length. — Male,  4.5-4.6  mm;  female,  5.0- 
5.2  mm. 

Color  tan  with  black  markings;  crown  with 
two  spots  on  anterior  margin,  black,  trans- 
verse band  on  middle  of  crown;  pronotum 
with  broad,  pale  ivory,  transverse  band, 
sometimes  with  broken  black  stripe  on  either 
side  of  middle;  forewings  with  pale  veins, 
clavus  sometimes  dark  brown,  and  with  dark 
brown,  longitudinal  stripe  above  costal  cell; 
face  deep  tan  with  black  markings  on  clypeus. 
Similar  in  general  habitus  to  several  species 
possessing  broad  pronotal  band  and  in  male 
genital  characters  to  idtimus. 

Head  with  anterior  margin  produced,  apex 
obtusely  angled. 

Male. — Pygofer  in  lateral  view  with  broad, 
obtusely  rounded  caudal  margin,  very  short 
spine  arising  from  middle  of  margin  and  some- 
times hidden  from  lateral  view,  projecting 
distally  (Fig.  27);  aedeagus  in  lateral  view  with 
narrow  shaft,  distal,  bifurcated  processes 
long,  extending  slightly  beyond  midlength  of 
shaft  (Fig.  29),  processes  broad  medially, 
pointed  distally,  gonopore  at  about  midlength 


112 


Great  Basin  Naturalist  Memoirs 


No.  12 


Figs.  23-26.  CoUadonus  ultimus,  n.  sp. :  23,  male  pygofer,  lateral  view;  24,  connective,  right  style,  and  aedeagus, 
dorsal  view;  25,  aedeagus,  lateral  view;  26,  apex  of  right  style,  enlarged  dorsal  view. 


of  shaft;  connective  short,  not  reaching  apex  of 
style  (Fig.  28);  style  with  tapered  stylar  shaft, 
stylar  spine  small,  directed  laterodistallv  (Fig. 
30). 

Female. — Seventh  sternum  with  caudal 
margin  nearly  truncate,  with  broad  median 
excavation. 

HoLOTYPE  (male). — Mexico:  Distrito  Fed- 
eral, Toluca  road,  26.  IX.  1945,  DeLong, 
Hershberger,  and  Elliot  (OSU),  allotype  fe- 
male, Mexico  City,  D.F.,  13.IX.1939,  D.  M. 
DeLong  (OSU).  Paratypes:  one  male,  same 
data  as  holotype  (in  author's  collection);  one 
female,  same  data  as  allotype  (in  author's  col- 
lection); 29  females,  same  data  as  allotvpe 
(OSU). 

Remarks. — From  ultimus,  to  which  it  is 
similar,  tolucensis  can  be  distinguished  by  the 
unexposed  pygofer  spine,  by  the  tapered  sty- 
lar shaft,  and  by  the  color  patterns  on  the 
pronotum. 

CoUadonus  dampfi  (DeLong) 

Figs.  31-34 


kliodoniis  dampfi  DeLong  1946:20  [Holotype  9  (OSU) 

(examined).] 
CoUadonus  dampfi:  Nielson  1957:51 
Idiodonus  dampfi:  Metcalf  1967:1297 
Idiodomis  dampfi:  DeLong  1984:11 

Length. — Male,  3.9-4.0  mm;  female,  4.5 
mm. 

A  well-marked,  black  and  yellow  species 
with  two  distinct,  black  spots  on  anterior  mar- 
gin of  crown,  narrow,  transverse,  black  band 
on  disc;  pronotum  with  broad,  black  band 
along  anterior  margin,  broad,  yellow,  trans- 
verse band  medially,  narrow,  black  band  on 
posterior  margin;  forewings  with  pale  yellow 
to  deep  yellow  veins,  cells  brown  to  black. 
Similar  to  other  pronotal  banded  species  and 
to  bicinctus  (DeLong)  in  male  genital  charac- 
ters. 

Head  with  anterior  margin  not  produced, 
apex  obtusely  rounded. 

Male. — Pygofer  in  lateral  view  with  mod- 
erately long,  narrow  spine  arising  from  mid- 
dle of  caudal  margin  and  projecting  dorsally 


1988 


NiELSON:  COLLADONUS  OF  MEXICO  AND  CkNTHAI.  AMERICA 


113 


Figs.  27-30.  Colladonus  tohicensis,  n.  sp. :  27,  male  pygofer,  lateral  view;  28,  connective,  right  style,  and  aedeagus, 
dorsal  view;  29,  aedeagus,  lateral  view;  30,  apex  of  right  style,  enlarged  dorsal  view. 


(Fig.  31);  aedeagus  with  distal,  bifurcated  pro- 
cesses long,  extending  beyond  midlength  of 
shaft  (Fig.  33),  crossing  over  in  dorsal  view, 
gonopore  basad  of  middle  of  shaft;  connective 
long,  extending  to  about  apex  of  style  (Fig. 
32);  style  with  narrow  stylar  shaft,  inner  lat- 
eral margin  roughly  dentate,  stylar  spine  long 
and  projecting  laterally  (Fig.  34). 

Female. — Seventh  sternum  with  caudal 
margin  narrowed  distally,  shallow,  U-shaped 
indentation  medially. 

Distribution. — Colladonus  dampfi  is  known 
only  from  the  Mexican  states  of  Veracruz  and 
Morelos. 

Host. — Recorded  from  unspecified  trees 
and  shrubs  from  3,000  to  9,000  feet  during 
September  to  November  (DeLong  1946). 

Remarks. — The  type  series  of  dampfi  is 
mixed.  The  female  holotype  and  male  allotype 
were  not  properly  associated;  the  latter  is 
identical  to  pravus,  which  is  treated  as  a  ju- 
nior synonym  oi fasciaticollis  in  this  paper.  A 
male  paratype  oi  dampfi  from  Cordoba,  Vera- 
cruz (type  locality),  was  associated  with  the 
female  holotype  of  dampfi  and  used  in  illus- 


trating the  male  genitalia.  Five  additional 
male  paratypes,  three  from  Cuernavaca, 
Morelos,  and  two  from  unspecified  localities 
(MB  95,  Mexico,  and  MF  8523,  Mexico)  were 
identical  to  the  male  paratype  specimen  asso- 
ciated with  the  female  holotype.  The  remain- 
ing specimens  in  the  type  series  are  discussed 
under  fasciaticollis. 

Colladonus  darnpfi  can  be  distinguished 
from  bicinctus  by  the  short  and  more  rounded 
anterior  margin  of  the  head,  by  the  longer 
pygofer  spine,  and  by  the  longer,  distal,  bifur- 
cated aedeagal  processes. 

Colladonus  bicinctus  (DeLong) 
Figs.  35-39 

I diodomis  bicinctus  DeLong  1946:18  [Uo\otype  9  (OSU) 

(examined).] 
Colladonus  bicinctus:  Nielson  1957:51 
Idiodonus  bicinctus:  Metcalf  1967:1287 
Idiodonus  bicinctus:  DeLong  1984:10 

Length. — Male,  4.2  mm;  female,  4.5-4.8 
mm. 

A  well-marked  species  of  deep  yellow  with 
black  markings;  crown  with  two  triangular, 


114 


Great  Basin  Naturalist  Memoirs 


No.  12 


Figs.  31-34.  CoUadonus  dampfi  (DeLong):  31,  male  pygofer,  lateral  view;  32,  connective,  right  style,  and  aedeagus, 
dorsal  view;  33,  aedeagus,  lateral  view;  34,  apex  of  right  style,  enlarged  dorsal  view. 


black  spots  on  anterior  margin  and  narrow, 
black,  transverse  band  on  middle  of  disc; 
pronotum  with  narrow,  black  band  along 
anterior  margin  and  narrow,  black,  transverse 
band  on  middle  transecting  broad,  yellow, 
transverse  band;  forewings  with  veins  yellow, 
cells  black.  Similar  in  general  habitus  and 
male  genitalic  characters  to  claustrus  (De- 
Long). 

Head  with  anterior  margin  produced,  apex 
angled. 

Male. — Pygofer  in  lateral  view  with  mod- 
erately long  spine  on  middle  of  caudal  margin, 
spine  directed  caudodorsally  (Fig.  35);  aedea- 
gus in  lateral  view  with  stout,  distal,  bifur- 
cated processes  reaching  to  about  midlength 
of  shaft  (Fig.  37),  not  crossing  over  in  dorsal 
view  (Fig.  39),  gonopore  slightly  basad  of  mid- 
length  of  shaft;  connective  short,  not  reaching 
apex  of  style  (Fig.  36);  style  with  narrow  stylar 
shaft,  lateral  margins  smooth,  stylar  spine 
very  short  and  bluntly  pointed  (Fig.  38). 


Female — Seventh  sternum  with  caudal 
margin  shallowly  and  broadly  concave,  small 
indentation  medially. 

Distribution — This  species  is  known  only 
from  the  Mexican  state  of  Distrito  Federal. 

Host. — According  to  DeLong  (1946)  the 
species  was  collected  at  7,500  feet  from  un- 
specified shrubs  or  trees  during  a  June- 
September  period. 

Remarks, — ^This  species  was  described 
from  the  female  holotype  and  a  female  para- 
type  specimen.  However,  a  male  paratype 
specimen  (MB  352,  Mexico,  A.  Dampf,  coll.), 
which  was  without  a  name  label  and  not  cited 
in  the  original  description,  was  found  among 
the  type  series.  The  specimen  was  associated 
with  the  female  holotype  and  used  in  illustrat- 
ing the  genitalia. 

The  species  can  be  separated  from  claus- 
trus by  the  presence  of  a  narrow,  transverse, 
black  band  on  the  middle  of  the  pronotum,  by 
the  narrower  pygofer  spine,  by  the  shorter 


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NiELSON:  COLLADONUS  OF  MEXICO  AND  CENTRAL  AMERICA 


115 


Figs.  35-39.  Colladonus  bicinctus  (DeLong);  35,  male  pygofer,  lateral  view;  36,  connective  and  right  style,  dorsal 
view;  37,  aedeagus,  lateral  view;  38,  ape,\  of  right  style,  enlarged  dorsal  view;  39,  apex  of  aedeagus  and  distal  processes, 
enlarged  dorsal  view. 


connective,  and  by  the  smooth-margined  sty- 
lar  shaft. 

Colladonus  claustrus  (DeLong) 
Figs.  40-44 

Idiodonus  claustrus  DeLong  1946:18  [Holotype  9  (OSU) 

(examined).] 
Colladonus  claustrus:  Nielson  1957:51 
Idiodonus  claustrus:  Metcalf  1967:1289 
Idiodonus  claustrus:  DeLong  1984:11 

Length. — Male,  4.4  mm;  female,  4.8-5.0 
mm. 

A  well-marked  species  with  color  patterns 
nearly  identical  to  bicinctus  except  paler  and 
without  median,  transverse,  black  band  on 
pronotum.  Similar  in  male  genital  characters 
to  bicinctus. 


Head  with  anterior  margin  produced,  apex 
angled. 

Male. — Pygofer  in  lateral  aspect  with  mod- 
erately long,  stout  spine  on  middle  of  caudal 
margin,  spine  directed  caudodorsally  (Fig. 
40):  aedeagus  with  slender,  bifurcated,  distal 
processes  reaching  to  about  middle  of  shaft, 
apices  not  reaching  gonopore  (Fig.  42),  cross- 
ing over  in  dorsal  view  (Fig.  44),  gonopore 
basad  of  midlength  of  shaft;  connective  short, 
not  reaching  apex  of  style  (Fig.  41);  style  with 
inner  lateral  margin  serrate,  outer  one 
smooth,  stylar  spine  short,  bluntly  pointed 
and  directed  laterad  (Fig.  43). 

Female. — Seventh  sternum  with  caudal 
margin  shallowly  and  broadly  concave,  small 


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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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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 


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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 


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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. 


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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 


1988 


NiELSON:  COLLADONUS  OF  MF.XICX)  AND  CENTRAL  AMERICA 


125 


Figs.  80-84.  Colladonus  anademus  (DeLong):  82,  male  pygofer,  lateral  view;  81,  connective  and  right  style,  dorsal 
view;  82,  aedeagus,  lateral  view;  83,  apex  of  right  style,  enlarged  dorsal  view;  84,  apex  of  aedeagus  and  distal  processes, 
enlarged  dorsal  view. 


Ajusco,  Llano  de  Cantimplora,  3,340  m, 
4.IX.1982,  C.  W.  and  L.  B.  O'Brien  and  G.  J. 
Wibmer  (CAS). 

Remarks. — From  trabilis,  to  which  it  is 
similar  in  certain  male  genitalic  features,  tes- 
sellatus  can  be  separated  by  the  gonopore, 
which  is  basad  of  midlength  of  the  aedeagal 
shaft,  by  the  short  connective,  and  by  the 
slender  stylar  shaft,  which  is  serrate  on  the 
inner  lateral  margin. 

Colladonus  anademus  (DeLong) 
Figs.  80-84 

Idiodonus   anademus    DeLong    1946;29    [Holotype    9 

(OSU)  (examined).] 
Colladonus  anademus:  Nielson  1957:51 
Idiodonus  anademus:  Metcalf  1967:1286 
Idiodonus  anademus:  DeLong  1983:90 
Idiodonus  anademus:  DeLong  1984:10 

Length. — Male,  4.8  mm;  female,  4.9-5.1 
mm. 


General  color  tawny  throughout  with  two 
distinct,  black,  elongate,  triangular  spots  on 
anterior  margin  of  crown,  small  black  mark- 
ings on  disc  of  crown  and  pronotum.  Similar  in 
general  habitus  to  incidus  (DeLong)  but  with 
distinctive  male  genitalia. 

Head  with  anterior  margin  not  produced, 
apex  rounded. 

Male, — Pygofer  in  lateral  view  with  cau- 
doventral  margin  produced  distally,  subtrun- 
cate  and  serrate  (Fig.  80),  pygofer  spine  very 
short,  basal  3/4  sclerotized  and  fused  to  inner 
mesal  margin  of  pygofer,  spine  projecting  dor- 
sad (Fig.  80);  aedeagus  in  lateral  view  with 
slender,  distal,  bifurcated  processes  reaching 
to  midlength  of  shaft  (Fig.  82),  crossing  over 
in  dorsal  view  (Fig.  84),  gonopore  at  about 
middle  of  shaft;  connective  long,  nearly 
reaching  to  apex  of  style  (Fig.  81);  style  with 
long,  narrow  stylar  shaft,  inner  lateral  margin 


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Great  Basin  Naturalist  Memoirs 


No.  12 


89 


Figs.  85-89.  Colladonus  incidus:  85,  male  pygofer,  lateral  view;  86,  connective  and  right  style,  dorsal  view;  87, 
aedeagus,  lateral  view;  88,  apex  of  right  style,  enlarged  lateral  view;  89,  apex  of  aedeagus  and  distal  processes,  enlarged 
dorsal  view. 


serrate,  outer  one  smooth,  stylar  spine  short, 
blunt,  and  directed  laterad. 

Female. — Seventh  sternum  with  caudal 
margin  convex,  shallow,  narrow,  V-shaped 
excision  medially. 

Host. — Specimens  were  collected  on  pine 
by  DeLong  (1946). 

Distribution. — This  species  is  known  only 
from  the  Mexican  state  of  Distrito  Federal. 

Remarks. — The  female  holotype  and  male 
allotype  appear  to  be  correctly  associated. 
This  species,  similar  to  incidus,  can  be  distin- 
guished by  the  produced,  subtruncate  cau- 
doventral  margin  of  the  pygofer,  by  the  short, 
dorsally  directed  pygofer  spine,  and  by  the 
long,  narrow  stylar  shaft. 

Colladonus  incidus  (DeLong),  n.  comb. 
Figs.  85-89 

Idiodonus  incidus  DeLong  1946:29  [Holotype  9  (OSU) 


(examined).] 
Idiodonus  iticidus:  Metcalf  1967:1299 
Idiodonus  incidus  DeLong  1984:11 
Idiodonus  pallidas:  DeLong  1983:90  [Holotype  6  (OSU) 

(examined).]  New  synonymy 
Idiodonus  pallidus:  DeLong  1984:10 

Length. — Male,  4.8  mm;  female,  4.9-5.1 


General  color  tawny  throughout  with  two 
small,  black  spots  on  anterior  margin  of 
crown.  Similar  in  general  habitus  to  anade- 
ynus  but  with  distinctive  male  genitalia. 

Head  with  anterior  margin  produced,  apex 
angled. 

Male. — Pygofer  in  lateral  view  with 
broadly  angled,  serrate  caudal  margin,  very 
small  spine  at  middle  of  margin,  spine  di- 
rected distad  (Fig.  85);  aedeagus  with  slender, 
distal,  bifurcated  processes  extending  beyond 
midlength  of  shaft  (Fig.  87),  crossing  over  in 


1988 


NiELSON:  COLLADONUS  OF  MEXICO  AND  CENTRAL  AMERICA 


127 


dorsal  view  (Fig.  89),  gonopore  at  midlength 
of  shaft;  connective  short,  not  reaching  to  apex 
of  style  (Fig.  86);  style  with  slender  stylar 
shaft,  lateral  margins  smooth,  stylar  spine 
long,  directed  laterad  (Fig.  88). 

Female. — Seventh  sternum  with  broadly 
convex  caudal  margin,  very  small  indentation 
medially. 

Distribution — This  species  is  known  only 
from  the  Mexican  state  of  Distrito  Federal. 

Host. — Specimens  were  collected  from 
unspecified  trees  or  shrubs  at  8,500  feet  dur- 
ing September  and  October  in  Mexico  (De- 
Long  1946). 

Remarks, — No  males  of  incidus  were  de- 
scribed in  the  type  series.  The  female  holo- 
type  of  incidus  was  associated  with  the  male 
holotype  of  paUidus  and  is  identical  with  the 
female  allotype  ofpaUidiis .  The  latter  name  is 
the  more  recent  and  is,  therefore,  suppressed 
as  a  junior  synonym  oi  incidus. 

From  anademus,  to  which  it  is  similar,  in- 
cidus can  be  easily  separated  by  the  serrated, 
broadly  angled  caudal  margin  of  the  pygofer 
with  its  very  small,  distally  directed  spine,  by 
the  shorter  connective,  and  by  the  longer, 
distal,  bifurcated  processes  of  the  aedeagus. 

Paracolladonus ,  new  genus 

Type-species :  Idiodonus  insculptus  De- 
Long  1946:25. 

Similar  to  Colladonus  Ball  and  Dolyobius 
Linnavuori  in  general  habitus  but  with  dis- 
tinctive male  genitalia.  General  color  tannish 
brown  in  female  to  black  in  male  with  dark 
markings  on  crown,  pronotum,  and  scutel- 
lum. 

Head  slightly  wider  than  pronotum,  ante- 
rior margin  broadly  rounded  and  slightly  pro- 
duced medially;  pronotum  with  posterior 
margin  broadly  concave;  scutellum  as  in  Col- 
ladonus ;  clypeus  broad  anteriorly;  venation  of 
forewing  similar  to  Colladonus,  inner  an- 
teapical  open  basally. 

Male  pygofer  with  large  spine  on  caudo- 
ventral  margin,  setose  caudodorsally;  aedea- 
gus long,  broadly  recurved,  somewhat 
compressed  dorsoventrally,  with  paired,  re- 
curved, short,  distal  processes,  processes 
sometimes  with  secondary  process  medially, 
gonopore  subapical  on  dorsal  surface;  connec- 
tive long  and  narrow;  style  broad  at  basal  half, 
with  foot-shaped  apex. 


Paracolladonus  insculptus 

(DeLong),  n.  comb. 

Fi^s.  90-94 

Idiodonus   in.sctdptiis:    DeLong    1946:25   [Holotype    $ 

(OSU)  (examined).] 
Idiodonus  insculptus:  Metealf  1967:1299 
Idiodonus  insculptus:  DeLong  1984:11 

Length. — Male,  5.3  mm;  female,  5.7-5.9 
mm. 

Long,  slender  species  with  dark  markings 
on  crown,  pronotum,  and  scutellum;  prono- 
tum black  in  male,  light  brown  in  female  with 
dark  markings  on  anterior  margin;  forewing 
light  to  dark  brown  with  yellowish  veins. 

Male  pygofer  with  very  long,  stout  spine  on 
caudoventral  margin,  spine  projecting  cau- 
dodorsad,  numerous  setae  confined  to  cau- 
dodorsal  area  (Fig.  90),  aedeagus  in  lateral 
view  long,  recurved,  with  distal  processes 
short,  broad  medially  (Fig.  92),  sometimes 
with  secondary  process  medially  (Fig.  94)  (if 
absent,  basal  attachment  apparent  in  dorsal 
view),  curved  dorsad  in  lateral  view,  slightly 
flared  laterally  in  dorsal  view  (Fig.  93),  shaft 
somewhat  flattened  dorsoventrally  (Fig.  94), 
gonopore  subapical  on  dorsal  surface;  connec- 
tive long  and  narrow,  reaching  to  or  beyond 
apex  of  style  (Fig.  91);  style  with  narrow  ridge 
or  flange  from  inner  lateral  margin  above 
preapical  lobe  to  apex  of  style  in  dorsal  view 
(Fig.  92). 

Remarks. — ^This  is  the  only  known  species 
assigned  to  the  genus.  It  is  known  from  the 
states  of  Distrito  Federal  and  Morelos  in  Mex- 
ico. DeLong  (1946)  collected  specimens  from 
pine  at  10,000  feet  in  September  and  Octo- 
ber. A  pair  of  specimens  were  collected  on  the 
Cuernavaca  road  in  Mexico  by  Ball  and  Stone. 

Paranurenus,  new  genus 

Type-species:  Idiodonus  latidens  DeLong 
1946:26 

Similar  to  Colladonus  in  general  habitus 
and  Nurenus  in  some  male  genital  characters. 
General  color  tawny  with  black  markings  on 
head.  Pronotum  with  ivory,  transverse  band, 
fore  wings  with  ivory  stripe  on  claval  suture. 

Head  about  as  wide  as  pronotum;  crown 
produced,  longer  medially  than  next  to  eye; 
pronotum  and  scutellum  as  in  Colladonus; 
forewing  as  in  Colladonus;  venation  partially 
obscured;  clypeus  long;  clypellus  long  and 
narrow. 


128 


Great  Basin  Naturalist  Memoirs 


No.  12 


Figs.  90-94.  Paracolladonufi  insculptus  (DeLong);  90,  male  pygofer,  lateral  view;  91,  connective  and  right  style, 
dorsal  view;  92,  aedeagus,  lateral  view  (allotype);  93,  aedeagus,  enlarged  dorsal  view  (allotype);  94,  aedeagus,  dorsal 
view  (specimen  from  Cuernavaca  Road,  Mex.). 


Male  pygofer  without  caudal  spine,  margin 
obliquely  truncate  as  in  Niirenus;  aedeagus 
small,  recurved,  flattened  dorsoventrally  with 
pair  of  short,  acuminate  distal  processes; 
gonopore  subapical  on  dorsal  surface;  connec- 
tive and  style  as  in  Colladonus. 

Paranurenus  is  represented  by  a  single 
known  species  from  Mexico. 

From  Colladonus,  to  which  it  is  similar  in 
general  habitus,  Paranurenus  can  easily  be 
differentiated  by  the  flattened  aedeagus  and 
by  the  absence  of  a  pygofer  spine.  The  head 
characters  and  the  flattened  shaft  of  the  aedea- 
gus will  differentiate  the  genus  from  Nurenus. 

Paranurenus  latidens  (DeLong),  n.  comb. 
Figs.  95-98 

Idiodonus  latidens:  DeLong  1945:26  [Holotype  9  (OSU) 
(examined).] 


Idiodonus  latidens:  Metcalf  1967:1300 
Idiodonus  latidens  DeLong  1984:11 

Length. — Male,  4.4-4.5  mm;  female,  4.8- 
5.1  mm. 

Color  tawny,  crown  ivory  with  two  black 
spots  on  anterior  margin  and  narrow,  black, 
transverse  band  on  disc  between  eyes;  prono- 
tum  tawny  with  narrow,  ivory,  transverse 
band  above  middle;  scutellum  tawny;  fore- 
wing  tawny  with  ivory  on  claval  suture  ex- 
tending to  pronotal  band,  fuscous  band  above 
costal  area;  face  ivory  to  tawny  with  black 
markings. 

Male. — Pygofer  in  lateral  view  with 
obliciuely  truncate  caudal  margin,  pygofer 
spine  absent,  macrosetae  on  caudodorsal 
quadrant  (Fig.  95);  aedeagus  in  lateral  view 
recurved,  socle  large,  shaft  slender,  flattened 


1988 


NiELSON;  COLLADONUS  OF  MEXICO  AND  CeNTKAL  AMERICA 


129 


95 


97 


98 


Figs.  95-98.  Paranurenus  latidens  (DeLong):  95,  male  pygofer,  lateral  view;  96,  connective  and  right  style,  dorsal 
view;  97,  aedeagus,  lateral  view;  98,  aedeagus,  ventral  view. 


dorsoventrally  (Fig.  97)  with  pair  of  short, 
acuminate  distal  processes,  processes  di- 
rected laterad  in  ventral  view  (Fig.  98);  gono- 
pore  subapical  on  dorsal  surface  of  shaft;  con- 
nective long  and  narrow,  extending  just  be- 
yond apex  of  style  (Fig.  96);  style  with  stylar 
shaft  short  and  enlarged  distally,  stylar  spine 
prominent,  subapical,  and  projecting  laterally 
(Fig.  96). 

Female. — Seventh  sternum  with  short, 
median,  spatulate  process  on  caudal  margin. 

Remarks. — This  species  has  no  known 
close  relatives.  The  styles  are  typical  of  many 
species  oi  Colladonus,  and  the  aedeagus,  ex- 
cept for  its  flattened  condition,  is  similar  in 
configuration  to  Nurenus.  The  head  charac- 
ters and  general  habitus  are  similar  to  species 
oiColladonus. 

The  species  is  known  from  the  states  of 
Michoacan  and  Distrito  Federal  in  Mexico, 


and  occur  at  elevations  from  5,000  to  8,000 
feet  in  February  and  September  on  unspeci- 
fied shrubs  or  trees  (DeLong  1946). 

Jaacunga,  new  genus 

Type-species:  Idiodonus  vinculus  DeLong 
1945:15 

Similar  to  Paracolladonus  in  general 
habitus  but  more  robust  with  slightly  wider 
head  and  distinctive  male  genitalia.  General 
color  tawny  with  markings  on  crown,  narrow 
band  on  pronotum  and  iridescent  forewings. 

Head  wider  than  pronotum,  anterior  mar- 
gin broadly  rounded  and  slightly  produced  as 
in  Paracolladonus;  pronotum  large,  posterior 
margin  broadly  and  shallowly  concave;  fore- 
wing  venation  as  in  Paracolladonus. 

Male  pygofer  without  caudal  spine,  setose 
submarginally;  aedeagus  stout  with  enlarged 
socle,  recurved  at  apical  third  with  terminal 


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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. 


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Great  Basin  Naturalist  Memoirs 


No.  12 


Remarks — Jaacunga  spatulatus  is  known 
from  the  type  series  collected  from  Mexico 
City  and  from  the  female  holotype  o^nibeUus, 
which  has  the  same  type  locality  as  spatulatus. 
The  specimens  were  collected  from  unspeci- 
fied shrubs  at  elevations  of  7,500  to  8,500  feet 
in  September  (DeLong  1946).  The  species  can 
be  distinguished  from  vinculus  by  the  shape  of 
four  terminal  spines  on  the  aedeagus  and  by 
the  shape  of  the  stylar  apex. 

Paracrassana,  new  genus 

Type-species:  Idiodonus  nigrifrons  De- 
Long  1983:91 

Near  Crassana  DeLong  &  Heishberger  but 
with  distinctive  head,  facial,  and  genitalic  fea- 
tures. General  color  tawny  with  black  mark- 
ings on  head  and  thorax,  face  black. 

Head  slightly  wider  than  pronotum,  ante- 
rior margin  slightly  angled  and  produced  me- 
dially, not  rounded  to  face,  clearly  demar- 
cated but  not  carinate,  narrow,  transverse 
impression  submarginally  between  ocelli, 
surface  shiny,  striate  longitudinally;  ocellus 
remote  from  eye;  forewing  as  in  Crassana; 
clypeus  broad  anteriorly,  about  as  wide  as 
long;  clypellus  long,  lateral  margins  expanded 
distally. 

Male  pygofer  long,  with  deep  incision 
basally  to  near  middle,  dorsal  margin  broadly 
excavated  near  middle,  row  of  fine  teeth  on 
ventral  margin;  aedeagus  symmetrical,  long, 
narrow,  nearly  tubelike,  socle  basal,  gono- 
pore  terminal,  sclerotized  endotheca  basad  of 
aedeagal  shaft;  connective  very  short  as  in 
Crassana  and  Neocrassana  Linnavuori;  plate 
long,  triangular,  tenth  segment  sclerotized 
laterally. 

Paracrassana  is  represented  by  a  single 
known  species  from  Mexico.  It  can  be  distin- 
guished from  Crassana  by  the  narrow  demar- 
cation between  the  crown  and  face,  the  nar- 
row impression  submarginally  on  the  crown, 
the  long  pygofer  with  basal  incision  and  tubu- 
lar aedeagus.  From  Neocrassana,  it  can  be 
distinguished  by  the  remote  position  of  the 
ocelli  from  eyes,  the  position  of  the  socle 
on  the  aedeagus,  and  characters  described 
above. 

Paracrassana  nigrifrons  (DeLong),  n.  comb. 
Figs.  105-107 

Idiodonus    nigrifrons    DeLong    1983;91    [Holotype    6 
(OSU)  (examined).] 


Length. — Male,  5.4  mm. 

Long,  robust  species,  crown  with  irregular, 
narrow,  black,  transverse  band  between 
ocelli;  pronotum  with  black  markings  on  ante- 
rior lateral  angles;  face  black  except  for  nar- 
row, yellow  anterior  margin;  forewing  iri- 
descent, veins  fuscous. 

Male  pygofer  in  lateral  view  long  and  nar- 
row, macrosetae  on  distal  third  except  caudal 
margin,  ventral  margin  with  fine  spines  (Fig. 
105);  aedeagus  long,  tubular,  curved  dorsally 
at  distal  third,  with  subdistal  flange  laterally 
on  dorsal  margin  (Fig.  106),  gonopore  apical, 
endotheca  not  forming  a  loop;  connective 
short,  broad;  style  narrow,  with  narrowed 
curved  apices. 

Female. — Unknown. 

Remarks. — This  species  is  known  only  from 
the  male  holotype  from  Huanchinango,  Pue- 
bla,  Mexico.  The  host  is  unknown. 

The  nomenclature  of  a  few  species  de- 
scribed in  Idiodonus  and  Colladonus  are 
treated  below: 

Bonneyana  caldwelli  (DeLong),  n.  comb. 

/(/io(/onusca/duJe//i  DeLong  1946:16  [Holotype  9  (OSU) 

(examined).] 
Idiodonus  caldwelli:  Metcalf  1967:1288 
Idiodonus  caldwelli  DeLong,  1984:10 
Idiodonus  apertus  DeLong  1946:14  [Holotype  9  (OSU) 

(examined).]  New  synonymy. 
Idiodonus  apertus:  Metcalf  1967:1287 
Idiodonus  apertus  Delong  1984:10 

I  have  compared  dissected  specimens  of 
caldwelli  and  schwartzi  (Ball)  and  found  them 
to  be  distinct.  Two  remaining  known  species 
in  the  genus,  osborni  (Ball)  and  terminalis 
(Ball),  have  not  been  examined;  but,  based  on 
the  original  descriptions,  they  appear  to  be 
distinct  from  caldwelli.  Further  study  of  the 
group  is  warranted,  however,  including  illus- 
trations and  descriptions  of  the  male  diagnos- 
tic characters. 

Ollarianus  tripartitus  DeLong 

Ollarianus  tripartitus  DeLong  1944:398 
/r/tO(/oni/sfl/fo(frons  DeLong  1983:89  [Holotype  9  (OSU) 

(examined).]  New  synonymy 
Idiodonus  alhifrons  DeLong  1984:10 

Examination  of  the  female  holotype  of  /. 
alhifrons  and  comparison  with  specimens  of 
several  species  o{ Ollarianus  show  that  it  be- 
longs in  that  genus  and  is  identical  with  O. 
tripartitus.  The  genus  Ollarianus  has  wide 


1988 


NiELsoN:  CoLLADONus  OF  Mexico  AND  Centiul  America 


133 


107 

Figs.  105-107.  Paracrassana  nigrifrons  (DeLong):  105,  male  pygofer,  lateral  view;  106,  connective,  right  style,  and 
aedeagus,  dorsal  view;  107,  aedeagus,  lateral  view. 


distribution  from  the  Nearctic  to  the  Neotrop- 
ical region  and  needs  revision. 

Paratanus  costatus  (DeLong),  n.  comb. 

Idiodonus  costatus  DeLong  1984:92  [Holotype  3  (OSU) 
(examined).] 

This  species  is  close  to  Paratanus  wygod- 
zinskyi  recurvatus  (Linnavuori)  but  differs 
from  it  in  color  characters.  It  may  be  a  distinct 
subspecies  of  the  nominate  form  or  a  separate 
species  pending  studies  of  the  genus  Para- 
tanus. 

Angulanus  DeLong,  n.  status 

Idiodonus  subgenus  Angulanus  DeLong  1946:30.  Type- 
species,  Idiodonus  incisurtis  DeLong  1946:30. 
[Holotype  9  (OSU)  (examined).] 

Idiodonus  (Angulanus):  Metcalf  1967:1304 


Idiodonus:  DeLong  1984:9.  [Inadvertent  synonymy  of 
Angulanus  DeLong  in  the  treatment  of  the  type- 
species,  incisurus  DeLong,  in  a  key  to  the  species 
oi  Idiodonus .] 

Examination  of  the  genital  structures  of  the 
male  allotype  clearly  showed  that  Angulanus 
deserves  generic-level  rank.  The  aedeagus  is 
asymmetrical,  and  the  pygofer  possesses  a 
prominent  spine  on  the  caudal  margin,  char- 
acters that  are  not  descriptive  of  the  genus 
Idiodonus. 

Angulanus  incisurus  (DeLong),  n.  comb. 

/diodonus  incisurus  DeLong  1946:30  [Holotype  9  (OSU) 

(examined).] 
Idiodonus  incisurus:  DeLong  1946:13,  14 
Idiodonus  (Angidanus)  incisurus:  Metcalf  1967:1304 
Idiodonus  (Angulanus)  incisurus:  DeLong  1984:10 


134 


Great  Basin  Naturalist  Memoirs 


No.  12 


Idiodonus  plummeri  DeLong 

Idiodonus    plummeri    DeLong    1946:25    [Holotype    9 

(OSU)  (examined).] 
Idiodonus  plummeri:  DeLong  1984:11 
Idiodonus  bakeri.  DeLong  1984:26  [Holotype  9  (OSU) 

(examined).]  New  synonymy 
Idiodonus  bakeri:  DeLong  1984:11 

The  generic  placement  of  this  species  ap- 
pears to  be  correct.  No  males  were  described 
in  the  type  series  oi plummeri  or  bakeri,  but  a 
male  paratype  specimen  labeled  "Idiodonus 
bakeri  DeLong,  MB  73,  Mexico,  A.  Dampf 
coll."  was  found  and  examined.  The  genitalic 
structures  clearly  show  that  it  belongs  in  Id- 
iodonus .  The  species  was  associated  with  the 
female  holotype  of  bakeri,  which  was  found  to 
be  identical  with  the  female  holotype  o( plum- 
meri. 

Species  incertae  sedis 

Three  species  of  Idiodonus  described  by 
DeLong,  edentulus,  excavatus,  and  copidus, 
are  treated  herein  as  incertae  sedis  until  males 
are  known  and  associated  with  females  of 
these  species.  Idiodonus  copidus  appears  to 
be  close  to  Jaacunga  and  may  belong  in  that 
genus. 

Acknowledgments 

I  thank  Dr.  Charles  A.  Triplehorn  and 
Dr.  Paul  Cwikla,  Ohio  State  University 
(OSU),  for  placing  on  loan  nearly  the  entire 
type  series  of  all  the  Idiodonus  species  from 
Mexico  and  Central  America  described  by  the 
late  Dr.  D.  M.  DeLong.  Without  this  material 
it  would  not  have  been  possible  to  unravel  the 
taxonomic  problems  involved  with  the  Id- 
iodonus and  Colladonus  fauna  in  that  part  of 
the  world.  I  also  thank  Dr.  James  P.  Kramer, 
U.S.  National  Museum,  for  the  loan  of  the 
type  of  Colladonus  delongi  Linnavuori  and 
material  of  genera  related  to  Colladonus. 

My  appreciation  is  extended  to  Jeanette 
Price  and  Pamela  Scott  for  the  illustrations 
and  to  Kaye  Thome  for  identifying  the  host 
plant  of  a  population  of  the  Colladonus  belli 
complex.  I  thank  Dr.  Leon  Hepner  for  read- 
ing the  manuscript  and  offering  helpful  sug- 
gestions to  improve  its  content. 


References  Cited 

Ball,  E  D  1937.  Some  new  species  of  leafhoppers  in 
groups  recently  segregated  from  Thamnotettix. 
Bull.  Brooklyn  Ent.  Soc.  32:  26-31. 

Bliven,  B  P  1954.  New  Hemiptera  from  redwood  (Miri- 
dae,  Cicadellidae).  Bull.  Brooklyn  Ent.  Soc.  49: 
119. 

DeLong,  D  M  1944.  The  genus  Ollarianus  (Homoptera: 
Cicadellidae)  in  North  America,  including  Mex- 
ico. J.  Wash.  Acad.  Sci.  34:  391-398. 

1946.  The  Mexican  species  oi Idiodonus  (Homop- 

tera-Cicadellidae).  Ohio  J.  Sci.  46:  13-30. 

1983.  New  species  and  distribution  notes  of  Mexi- 
can and  Bolivian  Idiodonus  (Homoptera:  Cicadell- 
idae) of  Mexico  and  Bolivia.  Entomol.  News  95(1): 
9-15. 

1984.  A  revised  key  to  the  Idiodonus  (Homoptera: 

Cicadellidae)  of  Mexico  and  Bolivia.  Entomol. 
News  95(1):  9-15. 

Gillette,  C.  P,  AND  C.  F.  Baker  1895.  A  preliminary  list 
of  the  Hemiptera  of  Colorado.  Colorado  Agric. 
Expt.  Sta.  Bull.  31.  1.37  pp. 

Medler,  J  T  1943.  The  leafhoppers  of  Minnesota.  Ho- 
moptera: Cicadellidae.  Minnesota  Agric.  Expt. 
Sta.  Tech.  Bull.  155.  196  pp. 

LiNNAVUORL  R  19.59.  Revision  of  the  Neotropical  Delto- 
cephalinae  and  some  related  subfamilies  (Ho- 
moptera). Suomal.  Elanja  Kasv.  Seuran  Vanamon 
Elaint.  Julk.  20:  1-370. 

Nielson,  M  W  19.57.  A  revision  of  the  genus  Colladonus 
(Homoptera,  Cicadellidae).  U.S.  Dept.  Agric. 
Tech.  Bull.  11,56:  1-.52, 

1962.   New  species  of  leafhoppers  in  the  genus 

Colladonus  (Homoptera,  Cicadellidae).  Ann.  En- 
tomol. Soc.  Amer.  .55:  143-147. 

1966.  A  synopsis  of  the  genus  Colladonus  (Ho- 
moptera: Cicadellidae).  J.  Kansas  Entomol.  Soc. 
.39:  .3.33-.336. 

Oman,  P  W  1949.  The  Nearctic  leallioppers  (Ho- 
moptera: Cicadellidae).  A  generic  classification 
and  check  list.  Washington  Entomol.  Soc.  Mem. 
3:  1-2,53. 

Stal,  C  1864.  Hemiptera  Mexicana  enumeravit  spe- 
ciesque  novas  descripsit.  Stettiner  Entomol.  Zeit. 
25:  49-86. 

Uhler,  P  R  1877.  Report  upon  the  insects  collected  by 
P.  R.  Uhler  during  the  explorations  of  1875,  in- 
cluding monographs  of  the  families  Cynidae  and 
Saldae  and  the  Hemiptera  collected  by  A.  S.  Pack- 
ard, Jr.  U.S.  Geol.  and  Geog.  Survey  of  the  Terr. 
Bull.  3(2):  355-475. 


NEW  WORLD  FULGORIDAE,  PART  I:  GENERA  WITH  ELONGATE 

HEAD  PROCESSES 

Lois  B.  OBi-it-n' 

Abstract. — Genera  new  to  science  described  below  include:  Amerzanna,  Sinuala,  and  Staluhra.  New  species 
include:  Amerzanna  peruana  (Peru),  Amijcle  hrevis  (Mexico),  A.  grandis  (Mexico),  A.  mankimi  (Honduras),  Aphro- 
disias  shaman  (Mexico),  Enchophora  maculata  and  E.  uniformis  (Peru),  Phrictus  delicatiis  (Brazil),  P.  dilinen.s 
(Colombia),  Sinuala  schmidti  (Belize),  S.  stall  (Honduras  and  El  Salvador),  S.  tuherculata  (Costa  Rica),  and  Staluhra 
brunnea  (Brazil  and  Guyana).  In  addition,  Artacie  dufourii  (Signoret)  is  removed  from  synonymy  with  A.  haemoptcra 
(Perty).  New  generic  synonymy  includes  C/H/oi>ifl  Stal(=  Ecuadoria)  Distant,  En/iyr/na  (Walker)  (=  UluhraStdl).  New 
synonymy  of  species  includes:  Copidocephala  merula  Distant  (=  Coanaco  melanoptera  Schmidt),  Copiducephala 
viridiguttata  Stal  (=  Coanaco  ornanda  Distant),  Diareusa  conspersa  Schmidt  (=  D.  dahli  Ossiannilsson),  Enchophora 
nigromactdata  Distant  (=  E.  nigrolimhata  Lallemand),  Enchophora  recurva  (Olivier)  (=  £.  hohemani  Stal),  Kri- 
chophora  sanguinea  Distant  (=  £. /Zorens  Distant  and  E.  longirostris  Distant),  Enchophora  tuberctdatu  (Ohvit'r)  (=  E. 
parvipennis  Walker),  Enchophora  viridipennis  Spinola  (=  E.  emincnta  Schmidt),  Enhydria  tessellata  (Walker)  (=  £. 
brachialis  Stal),  Fulgora  graciliceps  Blanchard  (=  Laternaria  orthocephahi  Fonseca),  Fulgora  laternaria  (Linnaeus) 
(=  F.  servillei  Spinola),  Phrictus  auromactdatus  Distant  (=  P.  notatus  Lellemand),  Phrictus  moebiusi  Schmidt  (=  P. 
sordidus  Caldwell).  The  two  species  of  Fulgora  are  synonymized  by  Ridout.  New  combinations  include  Chilobia 
dichopteroides  (Distant)  (Ecuadoria)  and  Stahtbra  ruftda  (Lallemand)  (Enhydria). 


Although  the  genera  of  New  World  Fulgori- 
dae  were  monographed  in  an  excellent  paper 
by  Stal  (1870b),  25  of  the  64  genera  have  been 
described  since  then.  Also, there  have  been 
many  subsequent  species  described,  many  in 
the  wrong  genus,  and  there  are  few  keys  to 
species.  Consequently,  even  though  they  are 
large  and  beautiful  insects,  it  is  very  difficult 
to  identify  most  species.  The  purpose  of  this 
paper  is  to  provide  a  guide  for  identification 
with  emphasis  on  the  use  of  external  charac- 
ters. 

Very  little  is  known  about  the  biology  of 
Fulgoridae.  Two  species  have  been  reported 
to  be  of  economic  importance  (Wilson  and 
O'Brien  1987),  Phrictus  diadema  (L.)  on  cocoa 
trees  {Theobroma  cacao  L.)  in  Brazil  (Silva 
1945)  and  Pyrops  candelaria  (L.)  on  longan 
and  mango  (Kershaw  and  Kirkaldy  1910). 
Host  records  for  species  contained  in  this  pa- 
per include  Amycle  pinyonae  Knull  &  Knull 
on  pinyon  pine,  Pinus  monophylla  Voss  (Knull 
and  Knull  1947),  Fulgora  laternaria  (L.)  on 
Hymenaea  courbaril  L.  or  quapinol  (Janzen 
1983),  Enchophora  pallidipunctata  Lalle- 
mand on  Copaifera  dourajeauni  (Legumi- 
nosae)  (label  data,  species  not  found  in  botani- 
cal references),  and  Rhabdocephala  brunnea 


Van  Duzee  on  Baccharis  sarothroides  Gray. 
Johnson  and  Foster  (1986)  explored  host 
specificity  in  a  tropical  forest,  where  plant 
diversity  might  make  host  finding  difficult, 
and  found  71%  of  Enchophora  longirostris 
Distant  (=  sanguinea  Distant)  on  Simarouba 
amara  Aubl.  (Sapindales)  and  84%  of  Phric- 
tus quinquepartitus  Distant  on  Terminalia 
oblonga  Steud.  (Combretaceae).  Adults  have 
been  reported  to  aggregate  on  tree  trunks  in 
ranks  and  to  move  in  conjunction  with  each 
other  (Johnson  and  Foster  1986,  O'Brien  and 
Wilson,  1985).  Further  host  data  will  be  given 
in  parts  to  be  published  later. 

Little  information  on  life  histories  is  avail- 
able. Kershaw  and  Kirkaldy  (1910)  noted  that 
the  eggs  o{  Pyrops  candelaria  were  laid  in  flat 
clumps  on  tree  trunks  and  covered  with  wax 
from  the  female  abdominal  segments;  nymphs 
dispersed  and  fed  on  branches;  and  adults  fed 
on  tree  trunks.  Hingston  (1932)  reported  rear- 
ing fulgorid  nymphs  from  what  he  described 
as  a  mantidlike  egg  case.  Other  descriptions  of 
immatures  are  those  of  Fulgora  phosphorea 
L.  (=  laternaria  L.)  (Hagmann  1928);  Pris- 
tiopsis  serratus  (F.)  (=  Cathedra  serrata  [¥.]) 
(da  Fonseca  1931),  and  Itzalana  subnuiculata 
Schmidt  (Wilson  and  O'Brien  1986).  As  far  as 


'Entomology,  Florida  A  &  M  University.  Tallahassee.  Florida,  USA. 


135 


136 


Great  Basin  Naturalist  Memoirs 


No.  12 


is  known,  fulgorids  feed  in  phloem  tissue  and 
are  univoltine.  No  accounts  of  other  typical 
fulgoroid  behaviors,  such  as  accoustical  com- 
munication and  association  with  ants,  have 
been  reported. 

Fulgoridae  are  subject  to  predation  by  fly- 
catchers (D.  Wechsler,  personal  communica- 
tion) and  parasitization  by  the  epipyropid 
moths  (Lepidoptera,  Epipyropidae).  The  col- 
ors of  the  closed  wings  would  appear  to  make 
them  inconspicuous  against  tree  trunks,  but 
they  can  be  found  fairly  easily  by  the  human 
eye  if  observed  in  profile.  I  assume  that  the 
red,  orange,  yellow,  and  white  coloration  so 
common  at  the  base  of  the  hind  wings  is  simi- 
lar in  function  to  that  of  the  underwing  moths 
(Noctuidae),  in  which  the  color  has  been 
shown  to  startle  birds  seeking  prey.  Eye  spots 
on  the  hind  wings  of  Cathedra  and  Fulgora 
would  seem  to  serve  a  startle  or  warning  func- 
tion as  well.  But  for  all  the  speculation  on  the 
bizarre  shapes  of  the  head  processes,  no  one 
has  reported  the  response  of  predators  to  the 
"alligator  head"  of  the  peanut  bug,  nor  to  any 
other  species. 

One  common  name  for  the  family  Fulgori- 
dae, "lantern-flies,"  was  given  because  Ful- 
gora were  reported  to  emit  light  (de  Merian 
1771),  but  recent  inquiries  and  enzyme  stud- 
ies have  failed  to  substantiate  this  phe- 
nomenon (Ridout  1983).  Many  Amerindians 
feared  Fulgora,  reporting  that  they  flew  a  zig- 
zag path  through  the  forest,  killing  everything 
they  touched  (Branner  1885).  Shamans  car- 
ried one  in  their  amulet  bags.  The  modern 
version  of  this  myth,  probably  invented  and 
surely  repeated  in  coed  entomology  classes  far 
out  in  the  bush,  is  that  if  a  human  is  bitten  by  a 
Fulgora,  death  is  certain  unless  one  is  saved 
by  the  antidote,  mating  within  24  hours;  non- 
sense, of  course. 

Lallemand  (1959,  1963)  revised  the  20  gen- 
era and  102  species  of  Fulgoridae  from  Africa 
and  the  28  genera  and  178  species  of  Asia  and 
Australia.  The  New  World  has  a  much  richer 
fauna,  composed  of  64  genera  and  250  species 
as  I  begin  this  monograph.  The  only  revisions 
of  New  World  genera  in  which  authors  ex- 
amined types  are  Cyrpoptus  (Kramer  1978) 
and  Fulgora  (Ridout,  personal  communica- 
tion). Other  generic  reviews  are  those  on  Ful- 
gora (da  Fonseca  1932),  Enchophora  (Metcalf 
1938),  Phrictus  (Caldwell  1945),  and  Poiocera 
(Gerstaecker  1860),  which  are  either  out  of 


date  or  relied  on  published  descriptions.  I 
have  examined  types  of  all  but  the  following 
species:  Artacie  dufourii  (Signoret)  (MZF?, 
letter  not  answered);  A.  haeriioptera  (Perty) 
(ZSBS,  not  found);  Cathedra  serrata  (Fabri- 
cius)  (not  listed  by  Zimsen  1964);  Copidoced- 
phala  melanoptera  Schmidt,  Diareusa  con- 
spersa  Schmidt,  and  Enchophora  etninenta 
Schmidt  (IZW);  Enchophora  ensifera  (Ger- 
mar)  and  E.  tuba  (Germar)  (lost  in  Lvov,  Rus- 
sia); E.  nigrolimbata  Lallemand  and  Enhydria 
rufula  Lallemand  (FSAG,  sent  but  not  yet 
arrived);  Olivier  s  species,  £.  recurva,  E.  tu- 
berculata,  and  F.  caerulescens,  and  the  types 
of  Fulgora,  which  were  studied  by  Ridout. 
Olivier's  (1791)  species  were  redescriptions 
with  Latin  binomials  of  Stoll's  (1781,  1788) 
descriptions  and  figures  in  French  and  Dutch 
or  Flemish,  using  names  in  those  languages. 
Olivier's  species  are  not  in  the  Paris  Museum 
(MHNP),  nor  in  his  personal  collection  de- 
posited there,  which  is  all  Coleoptera  (Bour- 
goin,  personal  communication).  Olivier  cites 
"Du  cabinet  de  M.  Holthuisen"  in  some  spe- 
cies; Horn  and  Kahle  (1935)  say  of  Holthuisen 
collection  "in  Hamburg  am  3. II.  1796  u. 
16. V.  1797  sowie  in  Stralsund  1800  verauk- 
tionert."  If  any  specimens  went  to  the  Ham- 
burg Museum  (ZMUH),  they  were  destroyed 
by  bombing  in  World  War  II,  except  for  a  few 
specimens  out  on  loan.  I  have  not  been  able  to 
trace  Olivier's  or  Stoll's  specimens  further. 
The  first  three  of  the  species  listed  here  were 
well  illustrated  when  described;  Schmidt's 
descriptions  clearly  fit  the  specimens  I  have; 
only  Lallemands,  Germar's,  and  Olivier's 
species  are  in  doubt. 

Fulgoridae  may  be  distinguished  from 
other  Fulgoroidea  by  the  presence  of  numer- 
ous cross-veins  in  the  hind  wings,  a  feature 
lacking  in  all  other  families.  Stal  also  noted 
that  the  carina  between  the  frons  and  gena 
continues  onto  the  clypeus,  but  this  feature 
also  occurs  in  some  Derbidae,  Dictyophari- 
dae,  and  Lophopidae.  Fulgoridae  share  with 
the  Dictyopharidae  similar  bilaterally  sym- 
metrical, usually  trilobed,  inflatable  male 
genitalia.  Emelyanov  (1979)  transferred  some 
Asian  genera  from  Dictyopharidae  to  Ful- 
goridae, citing  other  characters,  but  these 
characters  are  not  diagnostic  for  New  World 
families. 

The  stable  characters  easiest  to  use  are 
color,  especially  of  the  hind  wing;  the  shape 


1988 


O'BRIEN;  New  World  Fulgoridae 


137 


and  length  of  the  head  process  when  present; 
and  the  shape  and  carinae  of  the  frons,  vertex, 
and  thorax.  Since  the  proportion  of  the  head 
length  to  the  body  length  varies  greatly  with 
the  genera  and  cannot  be  determined  from 
the  illustrations,  measurements  are  given  for 
the  genera  (unless  it  is  impossible  to  get  an 
accurate  measurement  because  of  the  vari- 
ability of  the  curvature  of  the  head  process  as 
in  Enchophora).  The  ratio  of  head/pronotum 
is  given  for  most  species.  Male  genitalia  are 
difficult  to  inflate  consistently,  but  they  are 
figured  for  some  specimens;  I  state  when  I 
believe  they  must  be  used  for  a  valid  determi- 
nation, such  as  in  Scolopsella.  These  descrip- 
tions are  brief  and  diagnostic;  illustrations  are 
more  helpful  than  words.  A  character  matrix  is 
being  prepared  and  will  be  published  when  all 
of  the  genera  have  been  studied. 

Part  I  covers  those  genera  with  head  pro- 
cesses. These  keys  are  artificial.  When  the 
definition  of  genera  and  placement  of  species 
is  completed,  an  attempt  will  be  made  to  pro- 
duce cladograms  and  to  identify  monophyletic 
taxa.  Two  species  are  consigned  to  incertae 
sedis.  I  do  not  recognize  the  genus  of  En- 
chophora ensifera  (Germar)  among  any  New 
World  Fulgoridae  or  Dictyopharidae,  nor 
Fulgora  caerulescens  Olivier  as  figured  by 
Stoll  (1788,  plate  13,  Fig.  65)  among  any  New 
World  Fulgoridae  or  Flatidae. 

Illustrations  of  the  characters  used  are 
found  in  Figures  1-10  and  33.  Three  words 
that  may  not  be  commonly  used  elsewhere  are 
defined  as  ioWows:  fossette — a  small,  deep  pit; 
porrect — extending  forward  horizontally; 
terete — round  in  cross  section.  The  head  pro- 
cess may  have  10  carinae  or  fewer,  which  are 
named  in  Figure  4.  The  median  vein  is  pecti- 
nate in  Figure  5.  The  convention  in  Ful- 
goroidea  is  to  consider  the  "base"  of  the  insect 
the  dorsal  junction  between  head  and  thorax, 
with  the  base  of  the  head  and  pronotum  both 
here.  The  ventral  part  of  the  head  has  its  base 
anterad,  at  the  junction  with  the  vertex,  and 
its  apex  at  the  frontoclypeal  suture,  which  is 
the  base  of  the  clypeus. 

To  save  duplication  in  each  section,  1  have 
included  within  the  final  part  of  this  paper  the 
literature  cited  for  all  parts  (although  papers 
published  subsequent  to  Metcalf 's  catalog  are 
cited  here);  a  fist  of  localities  with  provinces  or 
longitude  and  latitude  (only  the  city  or  dis- 
tance for  a  city  plus  elevation  where  given  will 


be  included  earlier),  and  leclotypes  or  neo- 
types  will  be  designated.  There  is  a  list  of 
genera  and  species  and  their  distribution  by 
country  at  the  end  of  this  part.  Type  reposi- 
tory abbreviations  are  listed  in  the  acknowl- 
edgments. 

To  complete  this  monograph  successfully,  I 
will  need  help  in  locating  specimens,  since 
many  species  are  known  from  uni(}ues  or  one 
sex  only,  and  distribution  records  are  incredi- 
bly incomplete.  For  at  least  a  year  after  the 
publication  of  this  article,  I  will  accept  speci- 
mens for  identification  for  the  traditional  rec- 
ompense, a  sample  of  the  specimens  in  return 
for  identification.  Perhaps  your  specimens 
will  become  types.  The  genus  Chilohia  is  a 
case  in  point:  there  are  four  described  species, 
1  have  10  new  ones  on  hand,  and  only  one 
species  is  known  from  more  than  one  speci- 
men. Please  help  if  you  have  unidentified 
specimens. 

Key  to  Genera  of  New  World  Fulgoridae 
with  Head  Processes 

1 .  Tegmen  usually  transparent,  rarely  opaque  in 

basal  half 2 

—  Tegmen  opaque,  rarely  transparent  at  apex  .  .    4 

2(1).  Tegmen  transparent  brown  with  small  bris- 
tles on  veins;  prothorax  with  small,  white  nod- 
ules      Enhydria 

—  Tegmen  without  bristles  on  veins;  prothorax 
without  small,  white  nodules 3 

3(2).       Vertex  longer  than  broad  (Fig.  1) Stalubra 

—  Vertex  3  times  as  broad  as  long  in  midline, 
emarginate  medially  (Figs.  51,  54,  60)  or  apex 
of  vertex  hidden  by  recurved  head  process 

(Fig.  57)   Chilohia 

4(1).  Each  hind  wing  with  an  eyespot  visible  from 
below  with  wings  unspread;  head  process  gib- 
bous or  laterally  serrate  with  narrow  apex; 
large  species,  over  50  mm  in  length    5 

—  Hind  wings  without  large,  round  eyespots; 
head  process  porrect  or  curved,  but  not  as 
above    6 

5(4).  Head  process  peanut-shaped,  in  lateral  view 
with  markings  mimicking  an  alligator  head; 
eyespot  on  hind  wing  with  iris  or  pupil     Fulgora 

—  Head  process  subtriangular,  narrowing  at 
apex,  strongly  serrate  (Fig.  26);  eyespot  on 
hind  wing  without  iris  or  pupil Cathedra 

6(4).        Head  process  porrect  (Figs.   14-32,  46-50, 

102-117) 9 

—  Head  process  curved  up  and  back  (Figs. 
33-35,  40-42,  63-76) 7 

7(6).  Preocular  horizontal  flange  between  eye  and 
junction  of  frons  and  vertex;  head  process  ad- 
pressed  to  vertex,  flattened,  rugosely  tuber- 
culate  (Figs.  33-35) Artacie 


138 


Great  Basin  Naturalist  Memoirs 


No.  12 


Figs.  1-13.  See  facing  page  for  identification. 


1988 


O'BRIEN;  New  World  Fulgoridae 


139 


Figs.  1-13.  1-3,  5,  6:  Staluhra  hrevis  O'Brien:  1,  dorsal  view,  head  and  thorax;  2,  frons  and  clypeus;  3,  lateral  view, 
head;  5,  tegmen;  6,  hind  wing.  Hypothetical  insect:  4,  diagramatic  cross  section  of  head  process  showing  position  of 
carinae;  7,  stylized  hind  tibia.  Male  genitalia,  lateral,  dorsal,  and  ventral  views:  8-10,  Scolopsella  mexicana  O'Brien; 
11-13,  Scolopsella  reticulata  Ball. 


Morphological  characters: 

a-antenna 

A,-lst  anal  vein 

aed— aedeagiis 

af-anal  fold  of  hind  wing 

all- anal  flap  (10th  and  1 1th  abdominal  segments) 

an— anal  area 

ant— anterior  margin  of  wing 

ap— apical  area  of  hind  wing 

ax-apex 

b-base 

C— costa  or  costal  vein 

c— clypeus 

cl-clavus 

clas-clasper 

cm— commissural  margin 

cor-corium 

cs— claval  suture 

CU— cubitus  or  cubital  vein 

cx-coxa 

e-eye 

f-frons 

fe— femur 

fo-fossette 

g-gena 

hp-head  process 

Ic^lateral  carina 

Ih-length  of  head 


Ihp-length  of  head  process 

Ip-length  of  pronotum 

M -media,  medial  vein 

m-mesonotum 

mac-marginal  carina 

mc-median  carina 

mem-membrane  of  tegmen 

nl-nodal  line 

o-ocellus 

p-pronotum 

pc-pleural  carina 

Pcu-postcuhital  vein 

pf-postocular  flange 

post-posterior  area  of  wing 

pr-pronotum 

prf-preocular  flange  (see  Fig.  33) 

py-pygofer 

R-radius,  radial  vein 

sa— sutural  angle 

Sc-subcostal  vein 

sf-supraocular  flange 

t-tegula 

ta-tarsomeres 

ti-tibia 

tr-trochanters 

vertex  of  head 


(Continued  from  page  137) 

—  Preocular  horizontal  flange  absent;  head  process 
usually  not  adpressed  to  vertex  (see  E.  tuber- 
culata,  subviridis,  pyrrhocrifpta.   Figs.   67, 
73,  75) ' 8 

8(7).  Pronotal  median  carina  raised,  with  deep  fos- 
settes  at  base;  mesonotal  apex  depressed,  stri- 
ate (Figs.  63-76)   Enchophora 

—  Pro-  and  mesonota  smooth  and  gently 
rounded  (Figs.  40-42) Copidocephala 

9(6).        Medium-sized  species,  26-50  mm  in  length 

10 

—  Small  species,  under  25  mm  in  length 13 

10(9).  Head  process  with  apex  twice  as  wide  as  nar- 
rowest part,  dorsolateral  edges  of  process 
with  spines Phrictus 

—  Head  process  with  apex  not  so  widened,  sides 

of  process  without  spines   11 

11(10).  Sutural  angle  of  tegmen  triangularly  pro- 
duced; head  with  small,  horizontal,  preocular 
flange  (Figs.  116-117) Odontoptera 

—  Sutural  angle  of  tegmen  rounded;  head  with 
slight  vertical  flange  on  vertical  carina  (Figs. 
27-32) 12 

12(11).  Head  process  elongate,  head  1/3  length  of 
insect,  subtriangular,  slightly  bent  dorsad  at 
apex;  hind  wing  clear  with  veins  tinged  with 
green  at  base,  brown  elsewhere  ....  Ainerzanna 


—  Head  process  proportionally  shorter,  less 
than  1/5  length  of  insect,  with  sides  subparal- 
lel,  apex  slightly  enlarged,  not  bent  dorsad; 
hind  wing  brown  with  either  red  or  white 
spots Diareusa 

13(9).  Head  process  semicircular  in  cross  section; 
with  dorsal  surface  slightly  compressed,  ven- 
tral rounded Amycle 

—  Head  process  more  nearly  rectangular  or  cir- 
cular in  cross  section 14 

14(13).   Head,  including  process,  longer  than  thorax; 

tegmen  with  major  veins  straight 15 

—  Head,  including  process,  shorter  than  thorax; 
tegmen  with  major  veins  irregular,  slightly 
angulate 16 

15(14).  Head  process  subrectangular  in  cross  section, 
carinae  foliaceous,  angulate  (Figs.  24,  25)  .  .  . 
Scolopsella 

—  Head  process  smoothly  rounded,  almost  cir- 
cular in  cross  section,  carinae  normal  (Fig.  23) 
Rhabdocephala 

16(14).  Costal  margin  of  tegmen  markedly  sinuate 
(Fig.  43);  head  process  not  transversely  striate 
Sinuala 

—  Costal  margin  of  tegmen  slightly  convex;  head 

process  transversely  striate  (Fig.  49) 

Aphrodisias 


140 


Great  Basin  Naturalist  Memoirs 


No.  12 


Figs.  14-28.  Head  and  thorax,  dorsal  view:  14,  Amycle  brevis  O'Brien,  15,  A.  pinyonae  KnuW  &  Knull;  16,  A.  saxatilis 
Van  Duzee;  17,  A.  tumacacoriae  Knull  &  Knull;  18,  A.  vernalis  Manee;  19,  A.  g,randis  O'Brien;  20,  A.  sodalis  Stal;  21, 
A.  manhinsi  O'Brien;  22,  A.  amabilis  (Westwood);  2.3,  RhahdoccpJuila  hninnea  Van  Dnzee;  24,  Scolopsella  mexicana 
O'Brien;  25,  S.  reticulata  Ball;  26,  Cathedra  scrrata  (Fahricius);  27,  28,  Amcrztinna  prrnana  O'Brien  (27,  lateral  view). 


Amerzanna,  n.  gen. 

Figs.  27,  28 

Type-species:  Amerzanna  peruana  O  Brien. 

Medium-sized,     narrow,    brown    insect.s, 
31-41  mm  long,  head  1/3  length  of  insect. 


Head  process  porrect,  terete,  gradually  nar- 
rowed to  apex;  apex  bent  upward;  dorsal  and 
ventral  median  and  lateral  carinae  present 
on  anterior  third;  weak  hut  traceable  pleu- 
ral carina  cvuving  ventrally  to  end  in  slight 


1988 


O'BRIEN:  New  World  Fulgoridae 


141 


ventral  tubercle.  Preocular  flange  absent. 
Pronotum  smoothly  rounded,  traces  of  fos- 
settes  and  pleural  carinae  faintly  visible.  Teg- 
men  with  M  pectinate;  cross-veins  irregular, 
forming  diamond-shaped  cells.  Profemur  Ion- 
longer  than  metafemur,  posterior  tibia  with  6 
lateral  spines.  Ninth  abdominal  tergite  in  fe- 
males twice  as  long  as  eighth. 

I  name  this  genus  Amerzanna  because  su- 
perficially it  resembles  the  genus  Zanna  from 
Africa  and  Asia.  The  type-species  is  A.  peru- 
ana, new  species.  Atiierzanna  differs  from 
Odontoptera,  which  has  the  head  similarly 
shaped,  by  the  differences  in  the  tegmina,  for 
it  lacks  the  sharp  sutural  angle  and  the  spot  on 
the  nodal  line  of  Odontoptera. 

Amerzanna  peruana,  n.  sp. 

Figs.  27,  28 

Length. — Male  31-33  mm,  female  35-41 
mm;  head  15.6  mm;  ratio,  head/pronotum 
8.3. 

Ground  color  reddish  brown.  Tegmina  well 
pigmented,  transparent,  but  not  obviously  so 
(the  hind  wings  may  be  seen  through  the  teg- 
mina). Hind  wings  transparent,  greenish 
cloud  at  base,  brown  cloud  at  apex,  brown 
veins  between. 

HOLOTYPE  (male)  and  ALLOTYPE  (female). — 
Peru:  Madre  de  Dios:  Rio  Tambopata  Res., 
30  air  km  SW  Pto.  Maldonada,  290  m,  16-20- 
XI-1979,  J.  B.  Heppner,  subtropical,  moist 
forest  (USNM).  Paratypes  (10  female,  1  male): 
3,  same  data  (USNM);  2,  11-15-XI-1979 
(USNM);  2  females,  1  male  6-10-XI-1979 
(USNM,  LOB);  2,  Loreto,  Ucayali  R.  Yarina 
Cocha,  30-XI-1954,  leg.  Peter  Hocking,  1 
head  process  broken  (LOB,  FMNH);  1, 
Yahuarmayo,  Ape.  Group  (MCZ). 

Amycle  Stal 
Figs.  14-22,  93-101 

Amtjcle  Stal  1861:148.  Type  species:  A.  amahilis  (West- 
wood),  subsequent  designation  by  Van  Duzee 
1916:78 

Cyrpoptus  (Amyele)  [sic]  Stal,  1862:305. 

Small,  brown  insects,  12-23  mm  long,  head 
1/8  to  1/4  length  of  insect.  Head  process  por- 
rect,  dorsoventrally  flattened,  dorsal  surface 
flat,  ventral  convex;  usually  elongate  triangu- 
lar, but  sometimes  parallel-sided,  sometimes 
slightly  enlarged  at  apex;  dorsal  lateral  carinae 
sinuate,  it  and  other  carinae  variable  between 
species.  Preocular  flange  horizontal,  about  as 


produced  as  marginal  carinae.  Tegmen  with 
M  not  pectinate  (2  main  branches),  cross- 
veins  reticulate,  apical  margin  more  angulate 
than  in  most  genera.  Metatibia  with  4-5  lat- 
eral spines  (sometimes  3  or  7  on  a  single  tibia). 
Female  ninth  tergite  1.5-2  times  as  long  as 
eighth. 

This  genus  may  be  separated  from  other 
small,  brown  fulgorids  by  the  porrect  head 
being  flattened  dorsoventrally.  It  resembles 
Cyrpoptus  in  wing  coloration.  Diagnoses  are 
not  provided;  these  species  may  be  identified 
by  the  shape  of  the  head  (Figs.  14-22)  and  the 
color  and  color  pattern  of  the  hind  wings 
(Figs.  93-101). 

Key  to  the  Species  oi  Amycle 

1.        Hind  wing  with  2  colors,  brown  posterior  and 

base  white,  yellow,  pink,  or  red  (Fig.  93) 

saxatilis  Van  Duzee 

—  Hind  wing  with  3  colors,  brown  posterior, 
translucent  area  (Figs.  94-101),  and  base  white, 
yellow,  orange,  or  red 2 

2(1).    Base  of  hind  wing  yellow,  white,  or  pink    3 

—  Base  of  hind  wing  red  or  orange 4 

3(2).  Base  of  hind  wing  yellow  or  yellow-orange,  con- 
tiguous with  brown  suffusion,  apical  area  lightly 
suffused  with  brown,  intermediate  area  translu- 
cent (Fig.  101)  amahilis  (WesUvood) 

—  Base  of  hind  wing  pale  pink,  contiguous  with 
white  band,  posterior  margin  brown  (Fig.  97); 

sides  of  head  process  parallel  (Fig.  18)    

vernalis  Manee 

4(2).  Vertex  at  least  twice  as  long  as  broad  (Figs. 

17-21) 5 

—  Vertex  less  than  twice  as  long  as  broad  (Figs.  14, 
15) 8 

5(4).   Hind  wing  with  brown  apical  and  anal  areas 

contiguous  (Figs.  93-97) 6 

—  Hind  wing  with  transparent  area  between 
brown  apical  and  anal  areas  (Figs.  98-100) 7 

6(5).  Hind  wing  with  transparent  area  restricted  to 
narrow  anterior  band  (Fig.  94);  head  process 
parallel-sided  (Fig.  20)   sodalis  Stal 

—  Hind  wing  with  transparent  area  about  1/4  area 
of  wing  (Fig.  9.5);  head  process  subtriangular 
(Fig.  21)   mankinsi,  n.  sp. 

7(5).  Head  process  expanded  at  tip  (Fig.  19);  hind 
wing  with  red  area  separated  from  brown  anal 
and  apical  areas  by  transparent  band  (Fig.  98) 
grandis,  n.  sp. 

—  Head  process  subtriangular  (Fig.  17);  red  area 
and  brown  anal  area  contiguous  (Fig.  100)  .... 
tumacacoriae  Knull  &  Knull 

8(4).   Head  broader  than  long  (Fig.  14);  apex  of  hind 

wing  narrowly  brown  (Fig.  99) brevis,  n.  sp. 

—  Head  longer  than  broad  (Fig.  15);  apex  of  hind 


142 


Great  Basin  Naturalist  Memoirs 


No.  12 


wing  broadly  brown  (Fig.  96) 

pinyonae  Knull  &  Knull 

Amycle  amabilis  (Westwood) 

Figs.  22,  101 

Fttlgora  amabilis  Westwood  1842:119.  Type  repository: 

UMO. 
Amycle  amabilis  (Westwood),  Stal  1861:  148. 

Length. — Male  14  mm;  head  3.4  mm,  ra- 
tio, head  pronotum  3. 1. 

Head  process  elongate,  parallel-sided. 
Hind  wing  yellow  to  yellow-orange  at  base, 
followed  by  brown  suffusion,  apical  area  less 
darkly  suffused,  area  between  latter  two  hya- 
line. 

Distribution. — Holotype  and  paratype: 
Mexico.  No  other  specimens  seen. 

Amycle  brevis,  n.  sp. 

Figs.  14,  99 

Length. — Male  13  mm,  female  14  mm; 
head  1.8  mm;  ratio,  head/pronotum  1.0. 

Head  triangular,  not  produced  into  distinct 
process  (Fig.  14).  Hind  wing  pale  orange  at 
base,  anal  area  pale  brown,  dark  brown  area 
on  each  side  of  anal  fold,  apical  row  of  cells 
brown  (Fig.  79).  Tegmina  in  male  pale  orange 
with  brown  maculation  apically;  in  female 
mottled  brown  throughout,  forming  a  pattern 
with  a  clean,  apical,  diagonal  line. 

The  two  specimens  upon  which  this  species 
is  based  may  represent  different  species,  but 
the  head  and  hind  wings  are  similar;  examina- 
tion of  male  genitalia  will  be  necessary  to  de- 
termine whether  these  are  two  different  spe- 
cies. A.  brevis  may  be  separated  from  other 
species  by  its  short  head. 

Holotype  (male). — Mexico:  Nayarit,  San 
Bias,  4-IX-1971,  W.  J.  Hansen  (LOB).  Fe- 
male (not  allotype  or  paratype):  Guerrero: 
Ciudad  Altamirano  20-X-1983,  H.  Brailov- 
sky,  E.  Barrera  (LOB). 

Amycle  grandis,  n.  sp. 
Figs.  19,  45,  98 

Length. — Female  20-22  mm;  head  5  mm; 
ratio,  head/pronotum  4.5. 

Head  elongate,  slightly  expanded  at  apex 
(Fig.  19).  Hind  wing  orange  in  basal  1/3, 
brown  in  anal  1/3  and  apical  1/4,  translucent 
areas  between  (Fig.  98).  Tegmina  brown  with 
most  veins  orange-red;  two  diagonal,  brown 
lines  delineating  translucent  half  oval  on 
costa,  translucent  triangle  at  apical  angle,  and 


mottled  brown  and  white  pattern  in  anal  angle 
(Fig.  45). 

Holotype  (female). — Mexico:  Sinaloa,  Ma- 
zatlan,  5-VIII-1971,  D.  W.  Davis  (LOB). 
Paratype  (female):  Sonora,  Canyon  Sapopa, 
15-X-1934,  Rio  Mayo,  H.  S.  Gentry  (CIS). 
Possible  male  (not  paratype):  Jalisco,  Chapala, 
6  mi  W  Jahsco,  30-VM963,  J.  Doyen  (LOB). 

Amycle  rnankinsi,  n.  sp. 

Figs.  21,  95 

Length. — Female  17  mm;  head  3.5  mm; 
ratio,  head/pronotum  2.9. 

Head  slightly  elongate  triangular,  turned 
up  at  apex  (Fig.  21).  Hind  wing  red  in  basal 
third,  apical  and  anal  thirds  brown,  all  colored 
areas  separated  from  each  other  by  pale  areas 
(Fig.  95).  Tegmen  brown  with  most  veins  red. 

I  dedicate  this  species  to  Dr.  J.  V.  Mankins, 
with  whom  we  have  spent  many  happy  hours 
collecting. 

Holotype  (female). — Honduras:  Lago 
Yojoa,  2-IX-1977,  J.  V.  Mankins  (LOB).  Fe- 
male paratype:  same  data  except  6-X-1977 
(LOB). 

Amycle  pinyonae  Knull  &  Knull 

Figs.  15,  96 

Amycle  pinyonae  Knull  &  Knull  1947:397.  Type  reposi- 
tory: OSU. 

Length. — Male  8  mm,  female  14-14.5 
mm;  head  2.6  mm,  ratio  of  head/pronotum 
1.9. 

Head  process  triangular,  lateral  margins 
sinuate.  Hind  wing  reddish  orange  in  basal 
2/5,  anal  and  apical  angles  brown,  translucent 
area  between  with  brown  veins.  Tegmen  dark 
brown  with  many  clear  areas,  these  not  form- 
ing definite  pattern. 

Distribution. — Holotype  (male):  Califor- 
nia: Pinyon  Flat,  Santa  Rosa  Mts.  on  Pinus 
cembroides  var.  monophylla  Voss.  I  have  seen 
three  other  specimens  collected  in  August 
from  California:  Pinyon  Flats  and  Valvermo 
and  New  Mexico:  Juan  Tabo,  7,000  ft. 

Amycle  saxatilis  Van  Duzee 

Figs.  16,  93 

Amycle  saxatilis  Van  Duzee  1914:33.  Type  repository: 
CAS. 

Length. — Male  lectotype  12.5  mm;  head 
2.8  mm,  ratio,  head/pronotum  3.5. 

Head  process  elongate  triangular.  Hind 
wing  pale  orange  at  base,  followed  by  white 


1988 


O'BRIEN:  New  World  Fulgoridae 


143 


Figs  29-45  Head  lateral  view:  29,  Diareusa  conspersa  Schmidt;  30,  D.  kemneri  Ossiannilsson;  31,  D.  annularis 
(Olivier)  32  D  imitatrix  Ossiannilsson.  Dorsal  view:  33,  Artacie  haemoptera  (Perty);  34,  A.  dufouri  (Signoret).  Lateral 
view  35  A  dufouri  (Signoret)  (pronotal  nodules  shown  only  in  lateral  view);  36,  Enhydria  longicornuta  Lallemand;  37, 
£  tesseilata  (Walker);  38,  E.  cicadina  Gerstaecker.  Frontal  view:  39,  E.  cicadina  Gerstaecker  Lateral  view:  40, 
Copidocephala  merula  Distant  (nodules  not  shown);  41,  C.  viridiguttata  Stal;  42,  C.  guttata  white.  Tegmen:  43, 
Sinuala  tuberculata  O'Brien;  44,  Odontoptera  carrenoi  Signoret;  45,  Amycle  grandis  O  Bnen. 


144 


Great  Basin  Naturalist  Memoirs 


No.  12 


band,  posterior  half  light  brown.  Tegmen 
pale,  only  slightly  darker  at  apex. 

Van  Duzee  said  the  color  at  the  base  of  the 
hind  wing  varied  from  red  to  luteus  in  his 
specimens.  This  has  been  verified  (N.  Penny, 
personal  communication).  The  color  pattern, 
however,  in  both  specimens  has  no  translu- 
cent area. 

Distribution. — Lectotype,  right  wing  re- 
moved and  pinned  beneath  specimen:  Cali- 
fornia: San  Diego  County.  I  have  seen  no 
other  specimens. 

Amijcle  sodalis  Stal 

Figs.  20,  94 

Amycle  sodalis  Stal  1861:148.  Type  repository:  NRS. 

Length. — Male  13-14  mm,  female  16.5- 
18  mm;  head  4.5  mm,  ratio,  head/pronotum 
4.5. 

Head  process  elongate,  sides  parallel  to 
apex.  Hind  wing  orange  at  base,  dark  brown 
posteriorly,  anterior  1  1/2  rows  of  cells  behind 
orange  base  translucent  (Fig.  94).  Tegmina 
red  basally,  apically  translucent  with  2  dark 
apical  lines  (similar  to  Cyrpoptiis);  some  dark 
patches  caudally. 

Distribution. — Holotype:  Mexico.  I  have 
seen  six  other  specimens  collected  from  July 
through  November  from  El  Salvador:  Santa 
Tecla,  900  m  and  Mexico:  Atlixco,  Canada  de 
Negros,  Cordoba,  Huachinango. 

Amycle  tumacacorae  Knull  &  Knull 

Figs.  17,  100 

Amycle   tumacacorae   Knull   &    Knull    1947:398.    Type 
repository:  OSU. 

Length. — Male  14  mm,  female  14-15.5 
mm;  head  3.7  mm;  ratio,  head/pronotum  3.7. 

Head  process  elongate  triangular,  slightly 
widened  just  before  apex,  slightly  bent  dorsad 
in  lateral  view.  Hind  wing  red-orange  at  basal 
third,  dark  anal  and  apical  areas,  gray  mem- 
branous areas  between  with  dark  veins.  Teg- 
men dark  brown  with  few  translucent  apical 
cells.  This  species  is  very  close  to  A.  vernalis 
in  shape  and  length  of  head  process;  it  differs 
in  the  color  and  pattern  in  the  hind  wing. 

Distribution. — Holotype  (male):  Arizona: 
Tumacacori  Mts.  I  have  seen  12  other  speci- 
mens from  Arizona:  Catalina  Mts.,  Chiric- 
ahua  National  Monument,  Molino  Basin,  and 
Pepper  Sauce  Canyon;  Texas:  Eastland 
County,  Forestburg,  31  mi  W  Ozone,  Tyler. 


Amycle  vernalis  Manee 

Figs.  18,  97 

Amyele[sic]  vernalis  Manee  1910:117.  Type  repository; 

USNM. 

Length. — Female  14.5  mm;  head  4  mm; 
ratio,  head/pronotum  4.0. 

Head  elongate,  sides  gradually  narrowing 
to  apex,  apex  slightly  turned  up.  Hind  wing 
pale  pink  in  basal  fourth,  followed  by  band  of 
white,  posteriorly  black. 

Distribution. — Lectotype:  North  Caro- 
lina: Southern  Pines  (USNM).  I  have  seen 
one  other  specimen  from  Louisiana:  Gal- 
braith. 

Aphrodisias  Kirkaldy 

Figs.  49,  50 

Compsoptera  Stal  1869:236.  Type  species  C.  cacica  Stal, 
original  designation. 

Aphrodisias  Kirkaldy  1906:248,  new  name  for  Compsop- 
tera Stal. 

Small,  reddish  brown  insects,  18-23  mm 
long,  head  about  1/8  length  of  insect.  Head 
process  porrect,  terete,  transversely  striate 
on  all  sides;  dorsolateral  carinae  present; 
short,  nonstriate,  apical  portion  with  dorsal 
median  carina  and  longitudinal  ridge  on  verti- 
cal face.  Preocular  flange  horizontal.  Tegmina 
with  longitudinal  veins  irregularly  angled,  not 
straight;  few  cross-veins.  Female  ninth  tergite 
slightly  longer  than  eighth. 

This  genus  may  be  separated  from  all  other 
genera  by  the  wrinkled,  porrect  head  process. 
The  head  process  of  Artacie  also  is  textured, 
but  it  is  recurved  and  recumbent  on  the  ver- 
tex. 

Key  to  the  Species  of  Aphrodisias 

1.  Head  with  wrinkled  or  striate  portion  of  process 
shorter  than  pronotum;  hind  wings  red  at  base, 
brown  apically cacica  (Stal) 

—  Head  with  wrinkled  portion  of  process  longer 
than  pronotum;  hind  wing  red,  white,  and  brown 
shaman,  n.  sp. 

Aphrodisias  cacica  (Stal) 
Fig.  49 

Compsoptera   cacica    Stal    1869:237.    Type    repository: 

NRS. 
Aphrodisias  cacica  (Stal),  Kirkaldy  1906:248. 

Length. — Male  18-20  mm;  head  2  mm; 
ratio,  head/pronotum  1.5. 

Head  process  siibe(|ual  in  length  to  vertex. 
Hind  wing  red  at  base,  brown  posteriorly. 
Tegmen  brown. 


1988 


OBrien;  New  World  Fulc;ohidae 


145 


Figs.  46-62.  Head  and  pronotum,  dorsal  view:  46,  Sinuala  stali  O'Brien;  47,  S.  schmidti  O'Brien;  48,  S.  tuberculata 
O'Brien;  49,  Aphrodisias  cacica  (Stal);  50,  A.  shaman  O'Brien.  Head  and  thorax,  dorsal  view  and  lateral  view,  and  head, 
ventral  view:  51-53,  Chilobia  silena  Stal;  54-56,  C.  smaragadina  (Walker);  57-59,  C.  cinxia  Stal;  60-62,  C. 
dichopteroides  (Distant). 


146 


Great  Basin  Naturalist  Memoirs 


No.  12 


Distribution. — Holotype  (female):  Mex- 
ico. I  have  seen  six  specimens  collected  from 
Mexico:  Alamos,  15  mi  S  Cuernavaca,  Cuer- 
nonica,  from  September  through  November. 

Aphrodisias  shaman,  n.  sp. 
Fig.  50 

Length. — Female  23  mm;  head  3.5  mm; 
ratio,  head/pronotum  2.3. 

Head  process  1.7  x  longer  than  vertex. 
Hind  wing  pale  pink  at  base,  triangular,  white 
area  intruding  into  pink,  apex  and  anal  area 
missing,  but  edges  of  anal  area  brown,  apical 
area  brown  with  white  spots. 

The  wings  of  this  specimen  are  badly  dam- 
aged, but  it  may  be  separated  from  cacica  on 
the  basis  of  head  shape.  This  specimen  has  lost 
its  left  tegmen  and  hind  wing  and  part  of  the 
right  wing,  but  it  is  a  very  distinctive  species 
based  on  the  head  alone. 

Holotype  (female). — Mexico:  Sinaloa,  5 
mi  N  Mazatlan,  l-VII-1965,  J.  A.  and  M.  A. 
Chemsakand  E.  G.  and  J.  M.  Linsley  (UCB). 

Artacie  Stal 
Figs.  33-35 

Artacie  Stal  1866:132.  Type  species:  Flata  haemoptera 
Perty,  subsequent  designation  by  da  Costa  Lima 
1935:488. 

Brown-winged,  medium-sized  insects,  20- 
27  mm  long.  Head  process  terete,  recurved  to 
lie  against  vertex  (Fig.  35),  texture  of  apex 
unevenly,  rugosely  tuberculate,  ventral  me- 
dian and  marginal  carina  on  process,  lateral 
carinae  on  frons.  Pre-  and  supraocular  flanges 
large  (Figs.  33,  34).  Pronotum  with  white 
nodules.  Tegmina  with  main  veins  slightly 
raised,  surface  concave  between  them,  M  3- 
branched,  cross- veins  reticulate  in  corium, 
forming  rectangular  cells  in  membrane.  Red 
or  orange-red  color  at  base  of  hind  wing  al- 
most reaching  apical  margin  medially,  divid- 
ing it  into  apical  and  anal  areas. 

I  synonymized  the  two  species  in  this  genus 
in  1985.  After  studying  additional  specimens, 
I  conclude  that  I  was  in  error  and  that  the 
artist's  illustrations  of  the  species  were  correct 
in  each  case  and  there  is  a  difference  in  the 
shape  of  the  vertex  of  the  two  species.  The 
white  spots  in  the  apical  portion  of  the  wings 
do  vary  in  size,  placement,  and  number  be- 
tween specimens,  but  a  general  pattern  for 
haemoptera  is  described  below. 


Key  to  the  Species  oi  Artacie 

1.  Median  length  of  head  process  before  preocular 
flange  subequal  to  median  length  of  vertex  .... 
dufourii  (Signoret) 

—     Median  length  of  head  process  before  preocular 

flange  half  of  median  length  of  vertex 

haemoptera  (Perty) 

Artacie  dufourii  (Signoret),  new  status 
Figs.  ,34,  35 

Encophora  [sic]  dufourii  Signoret  1858:497,  pi.  12,  no.  2, 

Fig.  1.  Type  repository:  MZF? 
=  Artacie  hemoptera  (Perty),  O'Brien  1985:661.  Error. 

Length. — Male  25  mm,  female  27  mm; 
head  length  2.6  mm;  ratio,  head/pronotum 
1.4.  Length  of  head  process  in  front  of  preocu- 
lar flange  subequal  to  length  of  vertex. 

Wings  orangish  red  at  base,  white  spots 
scattered  through  brown  apical  and  anal  ar- 
eas, not  forming  a  figure  with  an  empty  center 
(a  square,  circle,  or  diamond). 

The  two  known  species  may  be  separated 
by  the  proportion  of  the  head  process  in  front 
of  the  preocular  flange  to  the  vertex,  subequal 
in  dufourii  and  about  half  in  haemoptera 
(Figs.  33,  34). 

Distribution. — Type  (not  seen):  Cayenne. 
I  have  seen  two  other  specimens  collected  in 
March  from  French  Guiana:  Maron  River; 
Guyana:  Wineperu.  Part  of  Signoret's  collec- 
tion went  to  the  Naturhistorisches  Museum, 
Vienna,  and  part  to  Museo  Zoologico 
dell'Universita  degli  Studi  di  Firenze,  Flo- 
rence, according  to  Horn  and  Kahle  (1935).  I 
did  not  see  the  type  in  Vienna  and  have  not 
heard  from  Florence. 

Artacie  haemoptera  (Perty) 
Fig.  33 

Flata  haemoptera  Perty  1833:176,  pi.  35,  Fig.  3.  Type 

repository:  ZSBS,  not  found. 
Artacie  haemoptera  (Perty),  Stal  1866:389. 
Encophora  [sic]  dufourii  Signoret,  synonymized  O'Brien 

1985:661.  Error. 

Length.— Male  20-22  mm,  female  23-25 
mm;  head  length  2.1  mm;  ratio,  head/prono- 
tum 1.2. 

Length  of  head  process  in  front  of  preocular 
flange  half  length  of  vertex.  Wings  red  at  base, 
brown  in  apical  and  anal  areas,  4-6  white 
spots  in  circle,  diamond,  or  rectangle  on  api- 
cal area,  few  smaller  dots  also. 

These  two  species  may  be  separated  by  the 
length  of  the  head  process  and  the  pattern  of 
white  dots  on  the  hind  wing. 


1988 


O'BRIEN:  New  World  Fulgoridae 


147 


Distribution. — Holotype  (not  seen) :  Ama- 
zon basin  [Brazil?].  I  have  seen  nine  other 
specimens  collected  in  March,  August,  and 
October  from  Brazil:  Canum,  Manaus, 
Obidos,  Rio  Negro,  and  Teffe.  The  type  could 
not  be  found  in  the  Zoologische  Sammlungen 
des  Bayerischen  Staates,  Munich,  in  1975. 

Cathedra  Kirkaldy 

Fig.  26 

Cathedra  Kirkaldy  1903:179.  Type-species:  Phrictus  ser- 
rata  (F.),  original  designation. 

Pristiopsis  Schmidt  1905:332.  Type-species:  Fulgora  ser- 
rata  F. ,  original  description;  synonymized  by  Dis- 
tant 1906:20. 

Large,  yellowish  brown  insects,  52-71 
mm,  head  1/3  length  of  insect.  Head  process 
porrect,  terete,  gradually  narrowed  to  apex; 
marginal  and  lateral  carinae  of  frons  produced 
into  11  and  8  spines,  respectively;  head  about 
1/3  length  of  insect.  Preocular  flange  horizon- 
tal, spinelike.  Tegmen  with  M  irregularly 
branched  (2  main  branches);  clavus  open; 
cross-veins  forming  almost  round  cells  anteri- 
orly, arched  and  parallel  behind  nodal  line. 
Hind  wings  brown  with  ochre  eye  spot  at 
apex,  pale,  elongate  spots  at  base,  some  waxy 
spots  and  dashes  throughout;  emarginate  on 
posterior  border  before  spot. 

This  monotypic  genus  has  a  head  process 
like  a  cathedral  spire,  with  many  pairs  of  lat- 
eral spines,  which  distinguishes  it  immedi- 
ately from  any  other  fulgorid  genera. 

Cathedra  serrata  (Fabricius) 

Fig.  26 

Fulgora  serrata  Fabricius  1781:313. 

Phrictus  serratus  (Fabricius),  Schaum  1850:65. 

Cathedra  serrata  (Fabricius),  Distant  1906:20. 

Length. — Male  52-53  mm,  female  61-71 
mm;  head  19  mm;  ratio  head/pronotum  6.3. 

Ground  color  ochre  mottled  with  brown. 

Distribution. — Holotype:  Suriname.  I 
have  studied  eight  other  specimens  collected 
from  January  to  November  from  Bolivia: 
Santa  Cruz;  Brazil:  Manaus-Itacoatiana  road, 
km  30;  Panama:  Barro  Colorado  Island.  Zim- 
sen  (1964)  did  not  list  the  repository  of  this 
Fabrician  type. 

Chilobia  Stal 
Figs.  51-62 

Chilobia  Stal  1863:237.  Type-species:  C.  cinxia  Stal,  sub- 
sequent designation  by  da  Costa  Lima  1935:497. 


Ecuadoria  Distant  1906:21.  Type-species:  E.  dichop- 
teroides  Distant,  original  designation.  New  syn- 
onymy. 

Tegmina  membranous  throughout  or  mem- 
branous in  apical  half,  basal  half  opaque, 
brown  or  brownish  red,  medium-sized  in- 
sects, 19-28  mm  long.  Head  process  usually 
recurved,  usually  dorsoventrally  compressed, 
usually  gradually  expanding  in  ventral  view 
towards  apex.  Lateral  carinae  of  frons  parallel 
or  diverging  to  near  base,  carinate  throughout 
or  becoming  broad,  fused  with  median  carina 
into  broad  hump.  Frons  elevated  above  level 
of  frontoclypeal  suture,  expanded  into  lobes 
near  apex,  with  transverse  carina  extending 
across  frons  from  lobe  to  lobe.  Vertex  with 
lateral  carinae  sinuate  or  angulate  in  lateral 
view,  supraocular  flanges  present,  apical  mar- 
gin V-shaped,  sometimes  angulately  emar- 
ginate medially  in  addition,  sometimes  hid- 
den under  recurved  process.  Preocular  flange 
absent.  Pronotum  with  few  white  nodules. 
Tegmen  with  M  pectinate  (5-8  branches); 
clavus  closed;  cross-veins  irregular  and  dense 
in  basal  half,  few  and  as  thick  as  longitudinal 
veins  in  apical  half;  longitudinal  veins  with 
small  spines  alternately  angled  to  left  or  right; 
transition  zone  between  membrane  and 
opaque  area,  when  present,  is  arched.  Hind 
wings  variable,  from  only  slightly  enfumed  at 
base  to  bright  red  with  sinuate,  brown,  con- 
tiguous area,  only  two  marginal  rows  of  cells 
enfumed  at  apex.  Female  ninth  abdominal 
tergite  shorter  than  eighth. 

Because  I  have  intermediates  and  because 
the  male  genitalia  are  very  similar,  I  have 
synonymized  t\i^  genera  Ecuadoria  and  En- 
hydria  in  spite  of  several  character  differ- 
ences. The  species  of  Ecuadoria  have  a  long 
head  process  and  brightly  colored  tegmen  and 
hind  wing,  whereas  the  Enhydria  species 
have  a  short  head  process,  transparent  teg- 
mina, and  clear  hind  wings. 

The  four  described  species  are  illustrat- 
ed, but  1  have  not  provided  a  key  because  I 
am  unable  to  determine  the  range  of  varia- 
tion within  a  species  due  to  lack  of  available 
material. 

Chilobia  dichopteroides  (Distant),  n.  comb. 
Figs.  60-62 

Ecuadoria  dichopteroides  Distant  1906:22.  Type  reposi- 
tory: BMNH. 


148 


Great  Basin  Naturalist  Memoirs 


No.  12 


Distribution, — Holotype:  Ecuador  (nw): 
Rio  Durango,  350  ft.  I  have  seen  no  other 
specimens. 

Chilobia  cinxia  Stal 

Figs.  57-59 

Chilobia  cinxia  Stal  1863:238.  Type  repository:  NHMV. 

Distribution. — Holotype  (female):  Vene- 
zuela. I  have  seen  no  other  specimens. 

Chilobia  silena  Stal 
Figs.  51-53 

Chilobia  silena  Stal  1863:238.  Type  repository:  NRS. 

Distribution. — Holotype  (male):  Ecua- 
dor: Quito.  I  have  seen  no  other  specimens. 

Chilobia  smaragdina  (Walker) 
Figs.  54-56 

Dichoptera  smaragdina  Walker  1851:304.  Type  reposi- 
tory: BMNH. 
Chilobia  smaragdina  (Walker),  Gerstaecker  1895:39. 

Distribution. — Holotype:  Venezuela.  I 
have  seen  no  other  specimens. 

Copidocephala  Stal 
Figs.  40-42 

Copidocephala  Stal  1869:235.  Type-species:  Enchophora 
guttata  White,  original  designation. 

Coanaco  Distant  1887:28.  Type-species:  Enchophora 
guttata  White,  original  designation;  synonymized 
by  Distant  1906:23. 

Medium-sized,  greenish  or  brown  or  red- 
dish brown  insects,  22-32  mm  long.  Head 
process  recurved  at  junction  with  head,  later- 
ally compressed,  gradually  narrowing  ante- 
rad;  lateral  carinae  of  frons  fused  at  junction 
with  process  with  medium  and  lateral  carinae 
of  process  into  one  large  mound;  dorsolateral 
carina  also  present  on  process.  Preocular 
flange  absent.  Pronotum  with  small,  white, 
hemispherical  tubercles,  median  carina  not 
strongly  humped.  Tegmen  with  M  usually 
pectinate  (5-6  branches);  corium  with  cross- 
veins  regularly,  evenly  reticulate,  with 
smooth,  sunken,  red  spots  without  cross- 
veins;  membrane  with  cells  subsquare.  Fe- 
male eighth  and  ninth  abdominal  tergites 
subequal  in  length. 

This  genus  has  the  head  process  curved 
back  at  the  base,  never  extending  foi-ward  as 
in  some  Enchophora;  it  is  scimitar-shaped  like 
Enchophora  and  Enhydria.  Copidocephala 
differs  from  Enhydria,  which  has  membran- 


ous tegmina,  and  from  Enchophora  in  the 
characters  of  the  thorax  given  in  that  generic 
discussion.  The  close  proximity  of  the  prono- 
tum and  eyes  suggests  that  the  postocular 
flange  serves  to  protect  the  eye  from  contact 
with  the  pronotum. 

Key  to  the  Species  of  Copidocephala 
1.        Hind  wing  black,  luispotted merula  Distant 

—  Hind  wing  brown,   spotted  with  red  or  bine, 
green  or  white 2 

2(1).    Large  species,   28  mm  or  longer;  hind  wing 

brown  with  red  spots guttata  (White) 

—  Small  species,  25  mm  or  less;  hind  wing  brown 

with  blue,  green,  or  white  spots    

viridiguttata  Stal 

Copidocephala  guttata  (White) 
Fig.  42. 

Enchophora  guttata  White  1846:331.  Type  repository: 

BMNH. 
Copidocephala  guttata  (White),  Stal  1869:236. 

Length. — Female  29-32  mm,  head  length 
1.4  mm;  ratio,  head/pronotum  .63.  Pronotum 
and  costal  margin  of  tegmen  green,  pronotum 
with  red,  transverse,  sometimes  broken, 
band;  ground  color  of  rest  of  insect  ranging 
from  yellow  or  green  through  red  to  olive 
green.  Corium  with  approximately  21  round, 
smooth,  sunken,  small  (1  mm  or  smaller),  red 
spots;  spots  differing  slightly  in  number,  size, 
and  placement  among  individuals.  Hind  wing 
brown  with  red  spots  in  basal  two-thirds. 

This  species  may  be  separated  from  inerida, 
which  is  similar  in  size  and  coloration,  by  the 
absence  of  red  spots  in  the  hind  wing  in 
merula.  It  differs  from  viridiguttata  in  size 
and  in  the  color  of  spots  in  the  hind  wing,  blue 
or  green  to  white  spots  in  viridiguttata. 

Distribution. — Type:  South  America.  I 
have  seen  20  other  female  specimens  col- 
lected from  March  through  September, 
November  and  January.  Panama:  Barro  Colo- 
rado Island,  Chiriqui  Provence,  Fort  Clayton 
Costa  Rica:  6  mi  W  Las  Canas,  La  Selva 
Honduras:  Catacamas,  15  km  W  La  Ceiba 
Mexico:  Tamazunchale,  Temiscal,  Volcan 
Tacana. 

Copidocephala  merula  Distant 

Fig.  40 

Copidoccpluda  iiicrula  Distant  1906:23.  Type  repository: 
BMNH. 

Coanaco  melanoptera  Schmidt  1907:361.  T\pe  reposi- 
tory: IZW. 


1988 


O'BRIEN;  New  World  Fulgoridae 


149 


Copidocephala  melanoptera  (Schmiclt),  Metcall  1947: 
176,  New  synonymy. 

Length. — Female,  34-38  mm. 

Ground  color  as  in  guttata  except  hind  wing 
black,  without  red  spots. 

The  differences  in  coloration  of  the  tegmen 
found  in  the  two  specimens  of  mcrula  and 
melanoptera  are  within  the  variation  found  in 
the  species  guttata;  so  on  the  basis  of  size  and 
the  color  of  the  hind  wing,  I  synonymize  the 
two  species. 

Distribution. — Holotypes:  Colombia  (both 
types).  I  have  seen  the  type  oimerula  but  not 
o{ melanoptera.  No  other  specimens  seen. 

Copidocephala  viridiguttata  Stal 
Fig.  41 

Copidocephala  viridiguttata  Stal  1869:236.  Type  reposi- 
tory: NHMV. 

Coanaco  ornanda  Di.stant  1887:29,  pi.  4,  Fig.  13.  Type 
repository:  BMNH. 

Copidocephala  ornanda  (Distant),  da  Costa  Lima  1935: 
491.  New  synonymy. 

Length. — Male  22-24  mm;  head  1.1  mm; 
ratio  of  head/pronotum  0.5. 

Ground  color  olive  green  with  black  or  red- 
black,  sunken,  smooth,  round  spots  in  same 
pattern  as  guttata.  Hind  wing  brown  with 
blue  or  green  to  white  spots. 

This  species  differs  from  the  other  two  in 
having  blue,  green,  or  white  spots  in  the  hind 
wing  rather  than  red  spots  or  none.  It  is  inter- 
esting that  I  have  13  males  of  this  species  for 
study  and  20  females  of  guttata.  I  will  examine 
other  collections  to  see  if  this  is  coincidence  or 
unexpected  sexual  dimorphism.  I  synonymize 
ornanda  on  the  basis  of  the  shape  of  the  head 
process  and  wing  coloration. 

Distribution. — Type:  Golombia  (viridi- 
guttata); syntypes:  Panama:  Bugaba,  Tole  (or- 
nanda). I  have  studied  13  other  specimens 
collected  from  May  through  September  and 
January  from  Panama:  Achiote  Road,  Barro 
Colorado  Island,  Fort  Clayton;  Costa  Riga: 
Golfito,  La  Selva;  Mexico:  Los  Tuxtlas,  3  mi  S 
Palenque. 

Diareusa  Walker 

Figs.  29-32 

Diareusa  Walker  1858:43.  Type-species:  Phrictus  annu- 
laris (Olivier),  by  monotypy. 

Large,  reddish  brown  insects,  32-49  mm, 
head  relatively  short,  from  .1  to  .2  length 
of  the  insect.  Head  process  porrect,  terete, 


gradually  slightly  enlarging  at  apex;  median 
ventral  carina  in  about  anterior  third  of  pro- 
cess, ventral  and  dorsal  lateral  carinae  com- 
plete; short  dorsal  median  carina  in  anterior 
portion.  Preocular  flange  vertical,  well  pro- 
duced. Tegmen  with  M  sometimes  pectinate 
(3-4  main  branches),  sometimes  evenly 
branched;  cross-veins  irregular  in  corium, 
forming  small  squares  beyond  stigmal  line. 
Tegmen  brown  throughout  with  some  ante- 
rior cells  and  many  spots  either  white  or  red; 
often  some  apical  spots  translucent.  Female 
eighth  and  ninth  abdominal  tergites  subequal 
in  length. 

Diareusa  are  large,  reddish  brown  insects 
with  a  comparatively  short  (.1  to  .2  length  of 
insect),  terete,  but  not  inflated  head  without 
transverse  striae  or  spines;  these  characters 
will  separate  them  from  other  New  World 
Fulgoridae.  The  species  differ  in  the  color  of 
spots  in  the  hind  wing  and  in  the  shape  of  the 
head  process. 

Key  to  the  Species  of  Diareusa 

1.        Spots  on  hind  wing  red  and  white  

conspersa  Schmidt 

—  Spots  on  hind  wing  white  only 2 

2(1).   Head  process  short;  vertex  and  head  process  as 

long  as  width  of  head,  including  eye  

kemneri  Ossiannilsson 

—  Head  process  longer  than  width  of  head  includ- 
ing eye  (Fig.  30) 3 

3(2).   Dorsal  surface  of  head  process  in  lateral  view 

convex  (Fig.  31) annularis  (Olivier) 

—  Dorsal  surface  of  head  process  in  lateral  view 

straight  or  concave,  not  convex  (Fig.  32) 

imitatrix  Ossiannilsson 

Diareusa  annularis  (Olivier) 

Fig.  31 

Fulgora  annularis  Olivier  1791:568.  Type  repository:  un- 
known. 
Phrictus  annularis  (Olivier),  Walker  1851:264. 
Diareusa  annidaris  (Olivier),  Walker  1858:44. 

Length. — Male  32-38  mm,  female  45  mm; 
head  6.2  mm;  ratio  of  head/pronotum  2.2. 

Head  process  in  lateral  view  convex  dor- 
sally  (Fig.  31.)  Hind  wings  with  white  spots. 
These  two  characters  separate  it  from  the 
other  species. 

Distribution. — Type  (not  seen):  Suri- 
NAME.  I  have  seen  10  other  specimens  col- 
lected June  to  August  and  January  from  PERU: 
Brazilian  frontier;  BRAZIL:  Serra  do  Navio, 
Sinop;  French  Guiana:  67  km  S  Cayenne; 
SURINAME:  Mapane;  GUYANA:  Shudihat  River. 


150 


Great  Basin  Naturalist  Memoirs 


No.  12 


Diareusa  conspersa  Schmidt 

Diareusa  conspersa  Schmidt  1906:375.  Type  repository: 

IZW. 
Diareusa   dahli   Ossiannilsson    1940:44,    Fig.    2.    Type 

repository:  UZIL.  Tslew  synonymy. 

Length. — Male  38  mm,  female  46-49  mm; 
head  5.3  mm,  ratio  of  head/pronotum  1.8. 

Hind  wings  with  red  spots.  Head  process 
similar  to  that  odinitatrix. 

This  is  the  most  easily  identified  species  in 
the  genus  with  the  spots  on  the  hind  wing  red 
and  white;  in  all  other  species  they  are  white. 
It  is  interesting  that  although  this  is  the  easiest 
character  for  identification,  both  Schmidt  and 
Ossiannilsson  described  this  character  among 
the  last,  causing  it  to  be  overlooked.  Thus, 
many  museums  have  imitatrix,  the  common 
Central  American  species,  identified  as  con- 
spersa. I  synonymize  conspersa  and  dahli  on 
the  basis  of  head  shape  and  wing  color. 

Distribution. — Holotypes:  Ecuador:  Pal- 
mar (conspersa);  Colombia:  Rio  San  Agustin 
(dahli).  I  have  seen  three  other  specimens 
collected  from  June  through  August  from 
Ecuador:  Amhuagu,  Rio  Palenque. 

Diareusa  imitatrix  Ossiannilsson 

Fig.  32 

Diareusa  itnitatrix  Ossiannilsson  1940:45,  Fig.  3.  Type 
repository:  UZIL. 

Length. — Male  35-39  mm,  female  43-48; 
head  5.8  mm;  ratio  head/pronotum  2.2. 

Hind  wing  with  white  spots.  Head  process 
similar  to  conspersa,  but  longer  and  narrower 
than  other  species  with  white-spotted  hind 
wings. 

This  is  the  only  Central  American  species 
and  often  has  been  misidentified  as  con- 
spersa. Panamanians  call  this  insect  "totoran. 

Distribution. — Holotype  (female):  Pan- 
ama: Chiriqui.  I  have  seen  50  other  specimens 
collected  all  year  around  from  Colombia: 
Turbo;  Venezuela:  Rancho  Grande;  Panama: 
Barro  Colorado  Island,  Fortuna,  Hartman's 
Finca,  Margarita;  Costa  Rica:  La  Selva,  Tur- 
rialba;  Belize:  Middlesex;  Mexico:  Bonampa 
R.,  1  mi  W  Fortin  de  las  Flores,  Los  Tuxtlas. 

Diareusa  kemneri  Ossiannilsson 

Fig.  30 

Diareusa  kemneri  Ossiannilsson  1940:43,  Fig.   1.  Type 
repository:  UZIL. 

Length. — Male  35  mm;  head  4  mm;  ratio. 


head/pronotum  1.8. 

Hind  wings  with  white  spots.  Head  process 
short  (Fig.  30).  D.  kemneri  may  be  separated 
from  other  white-spotted  species  by  the  short 
head  process. 

Distribution. — Holotype  (female):  Peru: 
Perene.  I  have  studied  one  other  specimen 
collected  in  September  from  PERU:  Aguaitia. 

Enchophora  Spinola 

Figs.  63-76 

Enchophora  Spinola  1839:221.  Type-species:  recurva, 
subsequent  designation  by  Duponchel  1840:200. 

Medium-sized,  reddish  brown,  green,  yel- 
low, or  mottled  green,  white,  and  red  insects, 
22-32  mm  long.  Head  process  recurved,  lat- 
erally compressed;  nine  carinae  present  on 
process  (all  except  dorsal  median  carina).  Pre- 
ocular  flange  present  as  vertical  carina  linking 
frons  and  vertex  laterally  (Fig.  3).  Pronotum 
with  pair  of  deep  fossettes  laterad  of  strongly 
raised  median  carina.  Apex  of  mesonotum 
with  striate  depression  between  bifurcate  me- 
dian carina.  Tegmen  with  few  strong,  longitu- 
dinal veins,  cross-veins  variable,  dense  to  av- 
erage. Female  abdominal  segments  8  and  9 
subequal  in  length. 

Enchophora  may  be  separated  from  the 
very  similar  Copidocephala  by  the  raised  me- 
dian carina  of  the  pronotum  with  a  fossette  or 
depression  on  each  side,  the  depression  at  the 
apex  of  the  mesonotum  surrounded  by  a  high 
ridge,  and  the  ventrolateral  fields  of  the 
pronotum  being  separated  from  the  eye  by  a 
space  about  the  width  of  the  last  antennal 
segment.  In  Copidocephala  the  callus  behind 
the  eye  almost  touches  the  pronotum  when 
viewed  from  above,  and  the  pronotum  and 
mesonotum  are  smoothly  rounded.  Both  gen- 
era lack  the  preocular  flange.  Enhijdria, 
which  also  has  a  scimitar-shaped  head  pro- 
cess, may  be  differentiated  by  its  pale  tegmen 
and  small  spines  on  the  tegminal  veins. 

The  species  may  be  separated  by  the  shape 
of  the  head  process  (although  this  structure 
varies  slightly  in  curvature  and  compression), 
the  color  of  the  hind  wings,  and  sometimes  by 
distinctive  marks  or  colors  on  the  body  or 
wings.  Waxy  spots  on  the  wings  may  be  help- 
ful if  they  have  not  been  rubbed  off;  often  they 
may  be  identified  by  a  smooth,  pale  brown 
area  if  the  wax  is  missing.  The  hind  wings  are 
usually  red,  yellow,  orange,  or  white  in  the 
basal  1/4  to  2/3,  measured  along  the  anterior 


1988 


OBrien:  New  World  Fulgoridae 


151 


margin,  or  5/6  if  measured  from  anterior  to 
posterior.  The  color  pattern  of  the  tegmen 
may  be  quite  variable,  as  in  sanguinea  Dis- 
tant, nigromaculata  Distant,  and  reciirva 
(Olivier)  where,  in  females,  the  tegmen  may 
be  concolorous,  marked  with  discrete  red  or 
black  spots,  or  marked  with  coalescent  red 
spots.  Although  I  have  not  seen  both  sexes  of 
every  species,  if  there  is  a  sexual  difference  in 
color  pattern,  the  male  tends  to  be  more 
evenly  mottled  and  less  variable. 

Three  species  listed  as  Enchophora  in  Met- 
calf's  (1947)  catalog  have  been  transferred 
since  to  other  genera  (see  list  of  species). 

Key  to  the  Species  of  Enchophora 

1.  Hind  wing  with  white,  waxy  points  on  red 

area stillifera  (Stal) 

—  Hind  wing  without  white,  waxy  points  on  red 
area 2 

2(1).  Head  process  about  twice  length  of  frons  be- 
fore process,  enlarged  and  narrowed  near  tip 
(Figs.  63,  64)  .3 

—  Head  process  less  than  twice  length  of  frons 
before  process,  scimitar-shaped,  curved  back 
over  vertex,  or  curved  and  then  bent  upward 
4 

3(2).        Apex  of  head  process  black,  broadly  trilobed 

(Fig.  63) reciirva  (Olivier) 

—  Head  process  brown  throughout,  narrowly 
expanded  at  apex  (Fig.  64) tuba  (Germar) 

4(2).  Hind  wing  red  in  basal  half;  posterior  half 
medium  to  dark  brown;  tegmen  brown  or 
reddish  brown,  often  mottled   5 

—  Hind  wing  white,  pale  orange,  or  brown  at 
base;  either  only  apical  1/4  pale  brown  or 
posterior  half  medium  brown  (subviridis,  tu- 
berculata),  tegmen  unmottled  green  or  yel- 
low or  green  mottled  with  red 9 

Apex  of  pronotum  with  transverse,  black 
band  (Fig.  66)    nigromaculata  Distant 

Apex  of  pronotum  without  transverse,  black 
band 6 

Mesonotum  with  black  spots  (Figs.  67,  76) ...    7 

Mesonotum  without  black  spots   8 

Mesonotum  with  pair  of  black  spots,  also  pair 
on  ventrolateral  fields  of  pronotum;  head  pro- 
cess recurved  to  near  vertex,  then  bent  up- 
ward (Fig.  67)   pyrrhocrypta  Walker 

—  Mesonotum  with  2  or  3  pairs  of  black  spots; 
head  process  short,  only  slightly  recurved 
(Fig.  76) maculata,  n.  sp. 

8(6).  Apical  fourth  of  pronotum  pale;  base  of  mem- 
brane of  tegmen  with  a  brown  spot  about  size 
of  eye  on  each  side,  3-5  other  such  spots 
usually  present  on  membrane;  ca  basal  half  of 
hind  wing  red;  abdominal  tergites  brown 
pallidiptinctata  Lallemand 


5(4). 

6(5). 

7(6). 


—  Pronotum  concolorous;  tegmen  not  as  above; 
ca  basal  2/3  of  hind  wing  red;  abdominal  ter- 
gites bright  red sanguinea  Distant 

9(4).        Hind  wing  white  at  base 10 

—  Hind  wing  yellow,  orange,  or  brown  at  base 
12 

10(9).      Tegmen  green  or  yellowish  11 

—  Tegmen  greenish  white  mottled  with  red  .  . 
rosacea  Distant 

11(10).    Tegmen  green  or  yellowish  with  white  costal 

margin;  head  process  large  (Fig.  71) 

prasina  Gerstaecker 

—  Tegmen  greenish  yellow  throughout;  head 
process  small  (F'ig.  72)    uniformis,  n.  sp. 

12(9).  Head  process  scimitar-shaped  (Fig.  74);  hind 
wing  yellow  at  base,  then  white,  then  brown 
at  apex   viridipennis  Spinola 

—  Head  process  recurved  and  wrinkled  (Figs. 
73,  75);  hind  wing  orange  at  base,  brown  at 
apex   13 

13(12).    Head  process  recurved,  then  bent  upward  at 

apex  (Fig.  75) subviridis  Distant 

—  Head  process  recurved  almost  to  vertex  (Fig. 
73) tuberculata  (Olivier) 

Enchophora  maculata,  n.  sp. 

Fig.  76 

Length. — Male  24  mm,  female  28  mm. 

Ground  color  brown  with  minute  dots  of 
red  and  green.  Head  process  short,  extending 
forward  then  dorsad  (Fig.  76).  Wings  brown 
on  apical  third,  red  basally,  red  extending 
from  anterior  to  posterior  margin.  Distinctive 
markings:  two  or  three  pairs  of  brown  spots  on 
pronotum  (Fig.  76);  anterior  spot  black,  poste- 
rior medium  brown,  sometimes  pale  brown 
spot  laterad  of  latter.  Abdominal  tergites 
brownish  red. 

This  species  may  be  identified  easily  by  its 
unusual,  short,  partially  forward-directed 
head  process  (Fig.  76)  and  the  spots  on  the 
pronotum. 

HoLOTYPE  (male)  and  allotype  (female). — 
Peru:  Madre  de  Dios:  Rio  Tambopata  Res., 
30  air  km  SW  Pto.  Maldonada,  290  m,  16-20- 
Xl-1979,  J.  B.  Heppner,  subtropical,  moist 
forest  (USNM). 

Enchophora  nigromaculata  Distant 

Fig.  66 

Enchophora  nigromaculata  Distant  1906:23.  Type  repos- 
itory: BMNH. 

£.  nigrolimbata  Lallemand  1938:350.  Type  repository: 
DEI.  New  synonymy. 

Length. — Female  24-26mm. 


152 


Great  Basin  Naturalist  Memoirs 


No.  12 


Figs.  63-76.  Head  and  pronotum,  lateral  view,  ofEnchophora:  63,  E.  recurva  (Olivier);  64,  E.  tuba  (Germar);  65,  £. 
stillifera  (Stal);  66,  E.  nigromaculata  Distant;  67,  E.  pyrrhocrypta  Walker;  68,  £.  piillidipunctata  Lallemand;  69,  E. 
sanguinea  Distant;  70,  E.  rosacea  Distant;  71,  E.  prasina  Gerstaecker;  72,  £.  uiiiformis  O  Brien;  73,  E.  tuhercidata 
(Olivier);  74,  E.  viridipennis  Spinola;  75,  E.  subviridis  Distant;  76,  E.  maculata  O'Brien. 


Ground  color  reddish  brown,  variable,  teg- 
men  may  be  clear,  mottled  with  yellow  veins, 
or  with  10  dark  spots;  membrane  brown  or 
with  dark  spots,  5  to  7  waxy  points  along  nodal 
line.  Head  process  scimitar-shaped  (Fig.  66). 
Wings  red  at  base,  brown  posteriorly. 


This  species  may  be  separated  from  all 
other  Enchophora  by  the  black  line  along  the 
hind  margin  of  the  pronotum.  I  have  not  seen 
the  type  o(nifi,rolimhata. 

Distribution. — Holotype:  Bolivia  (nigro- 
maculata), Peru  (nigrolimbata).  I  have  seen 


1988 


O'BRIEN:  New  World  Fulgoridae 


153 


11  other  specimens  collected  from  October  to 
January  from  Bolivia:  Chapare;  Brazil:  Rio 
de  Janeiro;  Peru:  Chanchamayo;  30  km  SW 
Puerto  Maldonado,  290  m;  Satipo;  Tingo 
Maria;  Explorama  Lodge;  Ecuador:  Parque 
Nacional  St.  Cecelia. 

Enchophora  pallidipunctato  Lallemand 

Fig.  68 

Enchophora  pallidipunctata  Lallemand  1966:.52.  Type 
repository;  FSAG. 

Length. — Male  23  mm,  female  24-26  mm. 

Ground  color  reddish  brown,  mottled. 
Head  process  scimitar-shaped  (Fig.  68). 
Wings  red  on  basal  half,  brown  posterior  half 
(not  the  usual  2/3  red,  1/3  brown).  Seventeen 
waxy  tufts  present  in  apical  area.  Distinctive 
markings:  pale  band  along  apical  fourth  of 
pronotum  and  five  round,  dark  spots  in  tegmi- 
nal  membrane,  one  on  each  side  just  behind 
nodal  line,  three  to  five  distad  of  these.  Ab- 
dominal tergites  brown. 

This  species  may  be  separated  from  all 
other  Enchophora  by  the  pale  apical  fourth  of 
the  pronotum,  the  dark  spots  in  the  membra- 
nous area  of  each  tegmen,  and  the  smaller, 
red  portion  of  the  hind  wings.  It  most  closely 
resembles  sanguinea,  which  has  the  abdomi- 
nal tergites  bright  red,  while  they  are  brown 
in  pallidipunctata. 

Distribution. — Holotype:  Bolivia:  Buena 
Vista.  I  have  seen  24  other  specimens  col- 
lected from  February  through  April,  August 
and  September  from  Bolivia:  Buena  Vista, 
Santa  Cruz;  Peru:  Chanchamayo,  Chontilla, 
Iquitos,  Lima,  Pileopota,  600  m,  Rio  San- 
tiago, Satipo,  Tarapoto.  Several  specimens  la- 
beled on  Copaifera  Dourajeanni  (Legumi- 
nosae)  at  Tingo  Maria,  but  I  am  unable  to  find 
this  species  listed  in  any  botanical  reference. 

Enchophora  prasina  Gerstaecker 

Fig.  71 

Enchophora  prasina  Gerstaecker  1895:37.  Type  reposi- 
tory: emau. 

Length. — Male  24-27  mm,  female,  28 
mm. 

Ground  color  green,  with  costal  margin  of 
tegmen  white  and  head  and  genitalia  reddish 
brown.  Head  process  scimitar-shaped,  long, 
almost  as  long  as  mesonotum  and  pronotum 
combined.  (Fig.  71).  Wings  white.  No  waxy 
tufts  evident. 


This  species  may  be  separated  from  all 
other  Enchophora  by  the  green  tegmen  with 
white  costal  margin. 

Distribution. — Holotype:  Colombia:  Nova 
Granada.  I  have  seen  12  other  specimens  col- 
lected from  April  through  June,  and  October 
from  Bolivia:  Buena  Vista;  Peru:  NE; 
Panama:  Barro  Colorado  Island,  Cerro  Azul, 
Santa  Rita  Ridge;  Costa  Riga:  La  Selva. 

Enchophora  pyrrhocrypta  Walker 

Fig.  67 

Enchophora  pyrrhocrijpta  Walker  1851:272.  Type  repos- 
itory: BMNH. 

Length. — Male  19-20  mm,  female  23-25 
mm. 

Ground  color  reddish  brown,  mottled. 
Head  process  recurved  over  pronotum,  then 
bent  upward  (Fig.  67).  Wings  red  at  base, 
brown  posteriorly.  Two  pairs  of  distinctive, 
black  spots,  one  on  each  side  of  mesonotum, 
and  one  on  each  ventrolateral  margin  of 
pronotum  (Fig.  67).  No  waxy  tufts  evident. 

This  species  may  be  separated  from  all 
other  Enchophora  by  the  black  marks  on  the 
lateral  fields  of  the  pronotum  and  on  the  sides 
of  the  mesonotum  (Fig.  67).  E.  subviridis  hsis 
a  similarly  recurved  head  process. 

Distribution. — Holotype:  Brazil:  Para.  I 
have  seen  seven  other  specimens  collected 
from  October  through  February  from  Brazil: 
Benjamin  Constant,  Rio  Negro,  Belem,  Man- 
gabeira;  Guyana:  Kartabo,  St.  Laurent; 
Venezuela:  Cerro  de  Neblina. 

Enchophora  recurva  (Olivier) 
Fig.  63 

Fulgora  recurva  Olivier  1791:34.  Type  repository:  un- 
known. 

Enchophora  recurva  (Olivier),  Spinola  1839:  222,  pi.  10, 
Figs.  1,  2 

Enchophora  bohemani  Stal  1854:244.  Type  repository: 
NRS.  New  synonymy. 

Length. — Male  24-26  mm. 

Ground  color  reddish  brown,  tegmen  mot- 
tled, often  with  spots  composed  of  yellow 
clumped  veinlets;  membrane  with  long  yel- 
low and  brown  markings.  Head  process  elon- 
gate, produced  in  club  that  at  apex  is  twice 
average  width  of  process,  apex  trilobed,  black 
(Fig.  63).  Wings  red  at  base,  brown  posteri- 
orly. No  waxy  tufts  evident. 

This  species  may  be  separated  from  all 
other  Enchophora  by  the  enlarged,  blackapex 
of  the  head  process. 


154 


Great  Basin  Naturalist  Memoirs 


No.  12 


Distribution. — Holotypes:  Brazil  {bohe- 
manni)  and  Suriname  (recurva).  I  have  seen 
nine  other  specimens  from  BRAZIL:  Linhares, 
Araxa,  and  Painaivas,  taken  from  October 
through  December.  I  am  unable  to  confirm 
Berg's  report  from  Argentina  (Berg  1879). 

Enchophora  rosacea  Distant 

Fig.  70 

Enchophora  rosacea  Distant  1887:27,  pi.  4,  Fig.  IL  Type 
repository:  BMNH. 

Length. — Male  22-23  mm,  female  25-26 
mm. 

Ground  color  green,  tegmen  at  base  green- 
ish white  with  distinctive  clumps  of  red  dots 
between  the  green  veins;  membrane  green 
with  20  white,  waxy  spots.  Head  process  scim- 
itar-shaped (Fig.  70).  Wings  mostly  white, 
dusted  with  red  spots  at  base,  brown  posteri- 
orly. 

This  species  may  be  separated  from  all 
other  Enchophora  by  the  red  and  green  com- 
bination on  the  tegmen. 

Distribution. — Holotype:  Nicaragua: 
Chontales.  I  have  seen  10  other  specimens 
collected  from  March  through  August  from 
Panama:  Barro  Colorado  Island;  Costa  Riga: 
La  Selva. 

Enchophora  sangtiinea  Distant 
Fig.  69 

Enchophora  sanguinea  Distant  1887:27,  pi.  4,  Fig.  16. 
Type  repository;  BMNH. 

Enchophora  florens  Distant  1887:28,  pi  4,  Fig.  12.  Type 
repository:  BMNH.  New  synonymy. 

Enchophora  longirostris  Distant  1887:28.  Type  reposi- 
tory: BMNH.  New  synonymy. 

Length.— Male  22-23,  female  24-25  mm. 

Ground  color  greenish  to  reddish,  tegmen 
greenish  or  reddish,  or  greenish  black  with 
orange  or  red  spots  encircled  with  yellow 
veins,  or  the  red  spots  may  be  fused  into  large 
areas  or  confluences,  usually  very  mottled 
with  clumps  of  yellow  veins  and  dark  and/or 
light  spots.  Head  process  scimitar-shaped 
(Fig.  69).  Wings  red  at  base,  brown  posteri- 
orly. Twenty  waxy  tufts  present  in  membra- 
nous area  of  tegmen.  Abdominal  tergites 
bright  red. 

I  synonymize  this  species  with  two  other 
species;  E.  florens  and  longirostris  have  the 
same  male  genitalia;  E.  sanguinea  differs  from 
them  only  in  the  pattern  of  the  tegmen.  When 
I  found  comparable  difference  in  coloration  in 


other  species,  I  felt  confident  in  synonymizing 
the  three.  This  species  may  be  separated  from 
all  other  Enchophora  with  red  hind  wings  by 
the  shape  of  the  head  and  the  lack  of  distinc- 
tive identifying  marks  such  as  bands  and  spots 
on  the  pronotum. 

Distribution. — Type:  Guatemala, 
Panama  (sanguinea).  Nicaragua:  Chontales 
(florens);  Costarica:  Cache.  Colombia  (lon- 
girostris). I  have  seen  39  other  specimens  col- 
lected from  July  to  September,  with  one  in 
February.  Ecuador:  Palmar;  Panama:  Barro 
Colorado  Island,  Cana  1,200  ft;  Costa  Riga:  1 
mi  N  Tucurrique,  Rincon,  Turrialba,  Cairo. 

Enchophora  stillifera  (Stal) 
Fig.  65 

Phrictus  stillifera   Stal   1862:303.   Type  repository: 

nhmv. 

Enchophora  stillifera  (Stal),  Stal  1864:49. 

Length. — Male  25-28  mm,  female  32  mm. 

Ground  color  greenish  brown,  tegmen  of- 
ten with  spots  of  clumped  yellow  veins,  also 
often  with  broad  patches  of  waxy  exudate. 
Head  process  elongate,  scimitar-shaped, 
about  twice  as  long  as  frons  below  process. 
Wings  red  at  base,  mottled  with  white,  waxy 
spots;  brown  posteriorly. 

This  distinctive  species  is  the  only  En- 
chophora with  many  white,  waxy  spots  on  the 
red  portion  of  the  hind  wing. 

Distribution. — Holotype  (male):  Mexico; 
Guatemala:  Sabo;  Panama:  Bugaba,  2,000- 
3,000  ft.  I  have  seen  24  other  specimens  col- 
lected from  June  to  October  from  Panama 
Chiriqui,  Lino;  CoSTA  RiCA:  Osa  Peninsula 
Honduras:  Orina  del  Rio  Camgre,  Juticalpa 
Belize:  Northern  Road,  mi  28-45;  Mexico 
San  Quintin. 

Enchophora  subviridis  Distant 

Fig.  75 

Enchophora  subviridis  Distant  1887:28,  pi.  4,  Fig.  17. 

Type  repository:  BMNH. 
Enchophora  subviridis  vdT.  a.  Distant  1887:497. 
Enchophora  subviridis  distanti  Metcalf  1938:358. 

Length. — Male  22  mm,  female  23-25  mm. 

Ground  color  green,  brown,  or  yellow. 
Head  process  reflexed,  then  turned  up  at  tip. 
Wings  orange  in  basal  half,  brown  posteriorly; 
8  waxy  tufts  in  costal  margin  of  tegmen,  20 
plus  in  membrane. 

This  species  may  be  separated  from  all 
other  species  but  pijrrhocnjpta  by  the  re- 
curved and  bent  dorsad  head  process,  and 


1988 


O'BRIEN;  New  World  Fulcoridae 


155 


pyrrhocrypta  has  distinctive,  black  spots  on 
the  pro-  and  mesonota  and  red  on  the  hind 
wings. 

Distribution. — Syntypes:  Panama:  Chiri- 
qui,  Bugaba.  I  have  seen  six  other  specimens 
collected  from  August  to  October  fiom 
Panama:  Chiriqui,  Bugaba,  800-1,500  ft; 
Costa  Rica:  Puntarenas  Prov.,  Osa  Penin- 
sula, Rincon;  San  Vito,  Golfito. 

Enchophora  tuba  (Germar) 

Fig.  64 

Fulgora  tuba  Germar  1830:46.  Type  repository:  Lvov. 

(All  types  there  are  lost.) 
Enchophora  tuba  (Germar),  Burmeister  1845:4. 

Length. — Female  30  mm. 

Ground  color  reddish  brown,  mottled. 
Head  process  elongate,  produced  in  club  that 
at  apex  is  twice  average  width  of  process,  apex 
trilobed,  brown  like  rest  of  process  (Fig.  64). 
Wings  red  at  base,  brown  posteriorly.  No 
waxy  tufts  evident. 

This  species  may  be  separated  from  all 
other  Enchophora  by  the  long,  thin  head  pro- 
cess with  a  trilobed,  concolorous  tip. 

Distribution. — Type:  Brazil.  I  have  seen 
two  other  specimens  collected  from  Brazil: 
Agua  Preta,  in  December. 

Enchophora  tuherculata  (Olivier) 

Fig.  73 

Fulgora  tuherculata  Olivier  1791:  569.  Type  repository: 
unknown. 

Enchophora  tuberculata  (Olivier),  Burmeister  1845:  [4]. 
Enchophora  tuberculata  sub.  sp.  fuscomaculata 
Lallemand  1956:6.  Type  repository:  FSAG. 

Enchophora  parvipennis  Walker  1858:30.  Type  reposi- 
tory: BMNH.  New  stjnonyinij. 

Length. — Male  22  mm. 

Ground  color  greenish  brown.  Head  pro- 
cess recurved  toward  vertex  (Fig.  73).  Wings 
brown,  orange  in  anterior  four  cells.  No  waxy 
tufts  evident. 

This  species  may  be  separated  from  all 
other  Enchophora  by  the  recurved  head  pro- 
cess, which  differs  from  pyrrhocrypta  and 
subviridis  by  not  being  turned  up  at  the  tip. 
The  basal  colored  area  of  the  hind  wing  is  also 
more  reduced  than  in  any  other  species;  in 
one  specimen  it  is  brown  with  the  four  cells 
slightly  paler  brown. 

Distribution. — Type:  Suriname  {tuhercu- 
lata), Brazil:  Para  {parvipennis  male).  I  have 
seen    two    other    specimens    from    French 


Guiana:   St.   Jean  du  Moroni;  Venezuela: 
Salto  de  las  Acadencias. 

Enchophora  uniformis,  n.  sp. 

Fig.  72 

Length. — Male  24-26  mm,  female  25-26 
mm. 

Ground  color  yellow  to  greenish  yellow. 
Head  process  scimitar-shaped,  shorter  than 
frons  below  process.  Hind  wing  white  with 
apical  third  yellowish.  Tegmen  with  margin 
darker  behind  stigmal  line;  this  margin  with 
eight  spots  that  may  be  indicators  of  waxy 
tufts.  Distinctive  markings,  brown  or  black: 
apex  of  supraocular  flange,  three  or  more 
dashes  along  clavus,  spots  along  stigmal  line 
(these  are  variable;  one  specimen  lacks  them 
all). 

This  species  has  a  smaller  process  than 
viridipennis  and  prasina,  the  only  two  other 
species  with  green  or  yellow  tegmen.  It  lacks 
the  white  costal  margin  of  prasina  and  the 
yellow-orange  base  of  the  hind  wing  of  viri- 
dipennis. 

Distribution. — Holotype  (male),  allotype 
(female),  and  1  paratype  (female):  Peru:  Iqui- 
tos,  F6062,  H.  Bassler  Golln.  Ace.  33591 
(AMNH).  Four  other  paratypes:  PERU:  Chau- 
chamayo  [sic],  from  W.  F.  H.  Rosenberg 
(NCU,  male);  Jauja  Prov.  Satipo,  Sept.  1946, 
A.  Mailer  Coll.,  Frank  Johnson  Donor 
(AMNH,  female).;  Dept.  Huanuco,  Vic. 
Tingo  Maria,  Jungle,  670  m,  XII- 1939,  Felix 
Woytkowski  (AMNH,  male);  Iquitos,  Rio 
Itaya,  Herbert  Osborn  Colin  (OSU,  female). 

Enchophora  viridipennis  Spinola 

Fig.  74 

Enchophora  viridipennis  Spinola  1839:225,  pi.  11,  Fig.  2. 
Type  repository:  NHMV. 

Enchophora  erninenta  Schmidt  1909:187.  Type  reposi- 
tory: IZW.  New  synonymy. 

Length. — Male  22  mm. 

Ground  color  yellow  brown  (given  as  green 
in  original  description).  Head  process  scimi- 
tar-shaped, shorter  than  most  other  species 
(Fig.  74).  Hind  wings  yellow-orange  at  base, 
posteriorly  white  with  ochre  veins.  More  than 
10  waxy  tufts  present  along  slightly  infuscate 
margin  of  tegmen. 

This  species  may  be  separated  from  all 
other  Enchophora  but  uniformis  by  the  col- 
oration of  the  hind  wings.  In  the  specimen 
examined  the  distinctive  color  markings  are 


156 


Great  Basin  Naturalist  Memoirs 


No.  12 


the  shiny,  black  apex  of  the  head  process,  a 
black  spot  on  each  side  of  the  frons  before  the 
eye,  five  very  small,  red  marks  on  the  tegmen, 
red  longitudinal  lines  on  the  clypeus,  a  diago- 
nal, red,  marginal  band  on  each  side  of  the 
pronotum,  and  the  browner  wing  margin. 
Schmidt  (1909)  described  the  clypeus  as  hav- 
ing blood-red  longitudinal  stripes,  the  apex  of 
the  clypeus,  the  pronotum  and  scutellum,  the 
legs,  and  the  five  spots  on  the  tegmen  blood 
red.  The  specimen  at  hand  has  fainter  col- 
oration. Spinola  (1839)  said  that  the  spots  on 
the  tegmen  are  black.  I  did  not  have  a  speci- 
men to  compare  with  either  of  these  types 
when  I  visited  their  respective  museums;  so,  I 
am  placing  them  in  synonymy  by  utilizing 
their  descriptions  and  illustrations  and  my 
notes. 

Distribution. — Holotype:  Brazil:  Pebas 
(eminenta) ,  locality  unknown  {viridipennis).  I 
have  seen  one  specimen  from  BRAZIL:  Jacarea- 
canga,  collected  in  February. 

Enhydria  Walker 

Figs.  36-39 

Enhydria    Walker    1858:44.    Type-species:    Dichoptera 

tesellata  Walker,  by  monotypy. 
Ulubra  Stal  1866:133.  Type-species:  U.  hrachialis  Stal, 

by  monotypy.    Synonymized  by  Stal   1870:286. 

Ulubra  resurrected  by  O  Brien  1985:662.  [Error] 

Netc  synompny. 

Medium-sized,  brown  insects  with  trans- 
parent tegmina,  19-22  mm  long.  Head  pro- 
cess recurved,  laterally  compressed,  gradu- 
ally narrowing  to  apex.  All  five  ventral  carina 
and  two  dorsolateral  carina  present,  com- 
plete; dorsal  median  carina  present  at  base  of 
process,  extending  as  far  as  dorsal  notch.  Pre- 
ocular  flange  horizontal,  small.  Pronotum 
with  white  nodules.  Tegmen  with  short  spines 
along  veins  on  upper  surface,  M  not  pectinate; 
cross-veins  along  costal  margin  dense,  diago- 
nal; other  cross-veins  not  dense,  about  same 
diameter  as  longitudinal  veins;  most  cells 
rectangular.  Female  with  eighth  and  ninth 
abdominal  tergites  subequal. 

This  genus  may  be  separated  by  the  short 
spines  on  the  veins  on  the  upper  surface  of  the 
tegmen  and  the  transparent,  brown  hind 
wing.  The  dorsal  head  measurement  and 
head/pronotal  ratio  were  not  made  in  this 
genus  because  the  length  varies  greatly  with 
the  position  of  the  head  process. 

I  resurrected  Ulubra  as  a  valid  genus  on  the 


basis  of  illustrations  and  notes  taken  from  a 
specimen  so  identified  at  the  British  Mu- 
seum. Recently,  I  discovered  a  photograph  of 
the  type  of  Ulubra  brachialis  taken  in  Stock- 
holm; and  it  is  a  synonym  of  tessellata.  The 
genus  I  resurrected  as  Ulubra  is  renamed  he- 
lowAS  Stalubra(q.  v.). 

All  species  o( Enhydria  have  a  similar  color 
pattern:  brown  insects  with  red  spots  on  the 
pronotum,  black  markings  on  the  head  pro- 
cess, etc.  They  may  be  identified  to  species  by 
the  length  and  shape  of  the  head  process. 

Key  to  the  Species  oi Enhydria 

1.  Head  in  lateral  view  with  distance  from  fronto- 
clypeal  suture  to  dorsal  notch  greater  than 
length  from  dorsal  notch  to  apex,  thus  process 
short  (Figs.  37,  38) 2 

—  Head  in  lateral  view  with  distance  from  fronto- 
clypeal  suture  to  dorsal  notch  on  process  less 
than  length  from  dorsal  notch  to  apex,  thus  pro- 
cess long  (Fig.  36)   longicornuta  Lallemand 

2(1).  Venter  of  process  black;  thickness  of  process  in 
lateral  view  about  1/2  width  of  eye  (Fig.  38)  .  .  . 
cicadina  Gerstaecker 

—  Venter  of  process  pale  with  brown  spots,  only  tip 
black;  thickness  of  process  in  lateral  view  sube- 
qual to  width  of  eye  (Fig.  37)   

tessellata  (Walker) 

Enhydria  cicadina  Gerstaecker 

Figs.  38,  .39 

Enhydria  cicadina  Gerstaecker  1895:38.  Type  repository: 
EMAU. 

Length. — Male  19.5-21.5  mm. 

This  species  has  the  shortest  head  process 
(Fig.  38). 

Male  genitalia. — The  two  pairs  of  inflat- 
able lobes  have  darkened,  tuberculate  tips, 
which  the  other  species  do  not  have. 

Distribution. — Syntypes:  Brazil:  Bahia. 
I  have  seen  six  other  specimens  collected 
in  December  from  Brazil:  Caviuna,  Iguazu 
Falls,  and  Tijuco  Preto. 

Enhydria  longicornuta  Lallemand 

Fig.  36 

Enhydria  brachialis  i.  longicorntita  Lallemand  1960:106. 

Type  repository:  NRS. 
Enhydria  longicornuta  Lallemand,  O  Brien  1985:661. 

Length— Male  20-22  mm,  female  20-22 
mm. 

This  species  has  the  longest  head  process 
(Fig.  36). 

Male  genitalia. — ^The  ventral  lobe  in  ven- 
tral view  has  a  slight  median  projection  and  is 


1988 


O'BRIEN:  New  World  Fulgoridae 


157 


about  half  as  wide  as  the  claspers. 

Distribution. — Holotype:  Brazil.  I  have 
seen  five  other  specimens  collected  from 
Brazil:  Manaus,  Serro  do  Navio,  and  Trin- 
idad. 

Enhydria  tessellata  (Walker) 

Fig.  37 

Dichoptera  tessellata  Walker  185L305.  Tvpe  repositorv: 

BMNH. 
Enhydria  tessellata  (Walker),  Walker  1858:44. 
Enchophora   brachialis   Stal    1862:  L    Type   repository: 

NHS.  New  sijnonytmj. 
Enhydria  brachialis  (Stal),  da  Costa  Lima  1935:497. 

Length. — Male  19-20  mm,  female  21-22 
mm. 

This  species  has  a  medium-length  head  pro- 
cess (Fig.  37). 

Male  genitalia. — Ventral  lobe  of  the 
aedeagus  with  median  emargination,  ventral 
lobe  about  as  wide  as  claspers. 

Distribution. — Holotype:  Brazil:  Rio  de 
Janeiro  (brachialis  male);  Para  {tessellata  fe- 
male). I  have  seen  11  other  specimens  col- 
lected in  January,  March,  June,  and  October 
from  Brazil:  Corcovado,  Faz  Morelandia, 
Corupa,  and  Para;  PERU:  Tingo  Maria. 

Fulgora  Linnaeus 

Figs.  77-92 

Fulgora  Linnaeus  1767:703.  Type-species:  F.  laternaria 
(Linnaeus),  subsequent  designation  by  Interna- 
tional Commission  on  Zoological  Nomenclature, 
Opinion  322  (1954:187). 

Large  insects,  65-105  mm  in  length,  yel- 
lowish brown  mottled  with  black  markings 
and  white,  waxy  areas,  head  .25  to  .28  length 
of  insect.  Head  process  porrect,  terete,  in- 
flated, resembling  a  peanut  from  above  and  an 
alligator  head,  including  false  eye  spots,  in 
lateral  view.  Preocular  flange  large,  horizon- 
tal. Tegmen  with  M  not  pectinate,  most  cross- 
veins  forming  many-sided,  almost  round 
cells.  Female  with  abdominal  tergites  eight 
and  nine  subequal  in  length. 

At  the  museum  in  Sao  Paulo,  1  examined 
the  specimens  used  by  da  Fonseca  (1926)  in 
his  revision;  these  did  not  fit  his  key.  B.  V. 
Ridout  (personal  communication)  examined 
previously  used  characters  and  several  new 
features,  including  male  genitalia  and  the  re- 
sults of  morphometric  analyses,  and  found  the 
best  way  to  identify  species  in  this  genus  was 
to  match  specimens  to  the  accompanying  il- 
lustrations of  head  shapes.   His  fllustrations 


are  labeled  lectotype  except  for  la7npetis 
(holotype),  crocodilia,  and  lucifera.  I  give  him 
full  credit  for  this  section  and  am  convinced 
that  this  is  the  best  that  can  be  done  at  this 
time.  Any  errors  are  mine. 

Ridout  suggested  that  species  with  narrow 
heads  might  live  in  dry  forests,  the  others  in 
rain  forests.  Janzen  (1983)  described  the  be- 
havior of  F.  laternaria,  including  at  least  100 
specimens  seen  resting  on  Hymenaea  cour- 
haril  or  guapinol,  but  questioned  whether 
this  plant  is  the  host  or  only  host,  as  it  does  not 
extend  throughout  the  range  of  the  insect  spe- 
cies. 

Fulgora  castresii  Guerin-Meneville 

Figs.  81,  82 

Fulgora  castresii  Guerin-MeneviWe  1837:3,  pis.  173,  174, 
Figs.  3,  4. 

Distribution. — Type:  Mexico.  I  have  seen 
10  specimens  collected  from  July  through 
September  from  MEXICO:  Hustusco,  Nuevo 
X-Can  [sic],  Tamazunchale;  Peru:  Tingo 
Maria:  Panama:  Barro  Colorado  Island. 

Fulgora  cearensis  Forseca 

Figs.  87,  88 

Laternaria  cearensis  Fonseca  1932:3,  Figs.  1,  2,  3h. 
Fulgora  cearensis  Fonseca,  Metcalf  1947:219. 

Distribution. — ^Type:  Brazil.  I  have  seen 
two  specimens  from  Trinidad :  Curepe  and 
Maraval,  collected  in  September. 

Fulgora  crocodilia 
Brailovsky  &  Beutelspacher 

Figs.  85,  86 

Fulgora  crocodilia  Brailovsky  &  Beutelspacher  1978:176, 
pi.  3. 

Distribution. — Type:  Mexico.  I  have 
seen  three  specimens  collected  from  MEXICO: 
Chamela,  in  September  and  November. 

Fulgora  graciliceps  Blanchard 

Figs.  83,  84 

Fulgora  graciliceps  Blanchard  1849:pl.  2,  Fig.  1. 
Laternaria  orthocephala  Fonseca  1926:493,  pi.  6,  Figs.  1, 

2.  New  synonymy,  Ridout. 
Fulgora  orthocephala  Fonseca,  da  Costa  Lima  1942:42. 

Distribution. — Type:  Brazil  (orthoceph- 
ala, graciliceps  unknown).  I  have  seen  three 
specimens  from  Bolivia:  Requena  and 
Warnes,  and  PERU:  Tingo  Maria,  collected  in 
November. 


158 


Great  Basin  Naturalist  Memoirs 


No.  12 


14.2mm 


I       \  nn   19.   Vuloin-ii  Itiuwctis  Burnieister; 

Figs.  77-101.  Head,  dorsal  view  (odd  numbers)^  ^t^J^;;;:  ^8^4  T^^S  Bbn'wl;  SS^SG,  F.  crocodiUa 
79-80,  F.iaternam(L.);  81-82,  F.casfrm.Guenn-Menevlle^  8.3   M  j^  ^^_y^    ^    riofirandensts 

BraiWsky  &  Beutelspacher;  87-88,  F.  -^Xvan  Xzt   94  1  voi^^  95,  A.  u^ankinsi  OBnen;  96^A. 

Fonseca.  Hind  wings  oi  Arnycle:93,  A.  '^""f^ll"^^^^^^^^^  99  a   brevis  O'Brien;  100,  A.  f.i»Ki«iror««e  K  &  K, 

pinyonae  K  &  K;  97,  A.  vernalis  Manee;  98,  A.  grandts  O  Bnen,  99,  A. 
101,  A.  amabilis  (Westwood). 


1988 


OBrien:  New  World  Fulgoridae 


159 


Fulgora  lampetis  Burmeister 

Figs.  77,  78 

Fulgora  lampetis  Burmeister  1845:3. 

Distribution. — Type:  Brazil.  I  have  seen 
11  specimens  collected  throughout  the  year 
from  Bolivia:  La  Cordillera  Santa  Cruz; 
Peru:  Tingo  Maria;  Brazil:  Obidos,  Sinop; 
Panama:  Barro  Colorado  Island;  Costa  Rica: 
Sarapiqui;  Nicaragua:  Leon;  Honduras:  La 
Ceiba. 

Fulgora  laternaria  (Linnaeus) 
Figs.  79,  80 

Cicada  laternaria  Linnaeus  1758:434. 
Fulgora  laternaria  Linnaeus,  Linnaeus  1767:703. 
Fulgora  servillei  Spinola  1839:214.  New  synonymy.  Rid- 
out. 

Distribution. — Type:  Tropical  America 
{laternaria);  Brazil  {servillei).  I  have  seen 
live  specimens  collected  in  January,  and  April 
through  July  in  Panama:  Barro  Colorado  Is- 
land; Honduras:  Armenia,  La  Ceiba;  Mex- 
ico: LosTuxtlas. 

Fulgora  lucifera  Germar 

Figs.  89,  90 

Fulgora  lucifera  Germar  1821:100. 

Fulgora  mitrii  Burmeister  1867;xxiii,  Berg  1879:178. 

Distribution. — ^Type:  Brazil.  I  have  seen 
16  specimens  collected  from  November 
through  February  from  Argentina:  Abra 
Grande  Iran;  Bolivia:  Coipa,  El  Torno,  Mora 
Abadolz.,  Montero,  Santa  Cruz,  andWarnes. 

Fulgora  riograndensis  Fonseca 
Figs.  91,  92 

Laternaria  servillei  riograndensis  Fonseca  1926:486,  pi. 

IV,  Fig.  1. 

Laternaria  riograndensis  Monte  1932:22. 

Fulgora  riograndensis  Fonseca,  Metcalf  1947:232. 

Distribution. — Type:  Brazil.  I  have  seen 
one  specimen  from  Panama:  Barro  Colorado 
Island,  collected  in  June. 

Odontoptera  Carreno 
Figs.  44,  116,  117 

Odontoptera    Carreno    1841:275.     Type    species:    O. 
spectabilis  Carreno,  by  monotypy. 

Medium-sized,  green  or  brown  insects, 
27-37  mm  long,  head  1/6  to  1/3  length  of 
insect.  Head  process  porrect,  terete,  elongate 
triangular;  apex  bent  dorsad;  lateral  carinae  of 
dorsum  and  venter  and  pleural  carinae  all  visi- 


ble. Preocular  flange  small,  diagonal  or  hori- 
zontal. Ventropleural  carina  of  pronotum 
present,  dorsopleural  carina  absent,  repre- 
sented by  low  ridge.  Tegmen  with  apex  trun- 
cate; anal  angle  angulate;  dark  spot  along  stig- 
mal  line;  M  greatly  expanded,  16-19 
branches  at  level  of  stigmal  line;  cross-veins 
incomplete  in  corium,  forming  rectangular 
cells  in  membrane.  Female  with  eighth  and 
ninth  abdominal  tergites  subequal  in  length. 

Cathedra  serrata  and  the  two  species  of  this 
genus  are  the  only  Fulgoridae  with  head  pro- 
cesses which  I  have  never  seen  misidentified. 

Key  to  the  Species  oi  Odontoptera 

1.  Head  about  1.5  times  as  long  as  pro-  and 
mesonota  combined;  tegmen  green  with  nar- 
row, apical,  brown  band;  hind  wing  green  with 
brown  and  orange  markings  ....  carrenoi  Signoret 

—  Head  almost  as  long  as  pro-  and  mesonota  com- 
bined; tegmen  orange-brown  basally,  brown  be- 
hind pale,  thin,  nodal  band;  hind  wing  white  at 
base,  brown  apicalUy spectabilis  Carreno 

Odontoptera  carrenoi  Signoret 

Figs.  44,  117 

Odontoptera  carrenoi  Signoret  1849:178,  pi.  6,  Fig.  4. 
Type  repository:  NHMV. 

Length. — Male  27-28  mm,  female  30-33 
mm;  head  length  9.7  mm;  ratio,  head/prono- 
tum5.4. 

Ground  color  green  with  brown  apical  band 
on  tegmen,  round,  black  spot  surrounded  by 
white  line  at  apex  of  clavus  on  stigmal  line 
(Fig.  44).  Hind  wing  green  with  two  black 
spots  near  anal  angle,  spots  surrounded  by 
orange  area. 

This  species  is  green  with  a  more  elongate 
head  than  the  brown  spectabilis. 

Distribution. — Holotype:  locality  un- 
known. 1  have  seen  26  other  specimens,  col- 
lected May  to  October  and  February,  from: 
Brazil:  Sinop;  French  Guiana:  Montague 
des  Singes  near  Kourou;  67  km  S  Cayenne; 
Panama:  Barro  Colorado  Island,  Santa  Rita 
Arriba,  Parque  Nacional  Soberania  Frijolito; 
Costarica:  La  Selva;  Mexico:  Los  Tuxtlas. 

Odontoptera  spectabilis  Carreno 

Fig.  116 

Odontoptera  spectabilis  Carreno  1841:277,  pi.  5,  Fig.  2. 
Type  repository:  ZMHB. 

Length. — Female  36-37  mm;  head  length 
5.8  mm;  ratio,  head/pronotum  2.3. 


160 


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No.  12 


Ground  color  yellow-orange  on  head,  tho- 
rax, and  apical  part  of  tegmina,  tegmen  be- 
hind stigmal  line  brown.  Head,  thorax,  and 
tegmen  with  lateral,  longitudinal,  brown 
band  extending  half  length  of  costa.  Hind 
wing  white,  black  along  apical  margins,  tinged 
with  orange  at  base. 

This  species  is  brown  and  has  a  shorter  head 
than  the  green  carrenoi. 

Distribution. — Type:  America  (?).  I  have 
seen  four  other  specimens,  collected  from 
January  to  March,  from  Brazil:  Boraceia 
Field  Station  and  Corupa. 

Phrictus  Spinola 

Figs.  102-115 

Phrictus  Spinola   1839:216.   Type-specie.s:   Fulgora   di- 
adema  Linnaeus  by  nionotypy. 

Large,  reddish  brown  or  yellowish  brown 
insects,  rarely  yellow  and  green,  29-53  mm; 
head  1/8  to  1/4  length  of  insect.  Head  process 
porrect,  terete,  usually  enlarged  into  trifur- 
cate  apex  (Figs.  102-115);  dorsolateral  carinae 
each  with  two  pairs  of  spines,  ventrolateral 
carinae  each  produced  into  flange  near  fronto- 
clypeal  suture  and  into  small,  dull  spine  at 
junction  of  frons  and  clypeus.  Preocular  flange 
horizontal,  small;  supra-  and  postocular 
flanges  produced  into  strong  spines.  Tegmen 
with  M  not  pectinate,  cross-veins  reticulate  in 
corium  forming  square  cells  behind  stigmal 
line.  Female  with  ninth  tergite  medially 
shorter  than  eighth. 

These  beautiful  insects  with  a  name  that 
means  "horrible"  or  "terrible"  may  be  identi- 
fied by  the  shape  of  the  apex  of  the  head 
process  and  the  waxy  tufts  on  it,  the  color 
pattern  of  the  tegmina,  and  the  color  of  the 
hind  wings.  Galdwell  (1945)  also  used  the  me- 
dian notch  in  the  caudal  margin  of  the  prono- 
tum  and  variation  in  the  furcation  of  the  me- 
dian carina  around  the  notch,  which  I  found 
difficult  to  quantify.  The  female  first  valvulae, 
as  he  illustrated,  also  serve  to  identify  difficult 
specimens.  The  apex  and  venter  of  the  head 
process  are  usually  reddish  orange  or  reddish 
brown  with  a  white  stripe  down  the  dorsum 
reaching  the  apex  of  the  pronotum.  There  are 
several  color  patterns  on  the  tegmina  which 
will  be  discussed  under  species  descriptions.  I 
have  illustrated  the  head  process  with  the 
insect  at  approximately  a  45-degree  angle  in 
order  to  provide  a  good  view  of  the  apex. 


Because  this  makes  the  length  of  the  head  and 
process  difficult  to  determine,  I  have  included 
a  head  length  and  head/pronotum  ratio. 

Key  to  the  Species  of  Phrictus 

1.  Head  process  flattened  apically,  transversely 

arcuate,   lacking  definite  apical  teeth  (Fig. 
1 12);  base  of  hind  wing  yellow  with  transverse 

bands  of  brown  maculae 

auromaculatus  Distant 

—  Head  process  with  3  to  5  apical  teeth  (Figs. 
106-111,  113,  114);  hind  wing  lacking  trans- 
verse bands  of  brown  maculae 2 

2(1).        Head  process  with  5  apical  teeth  (Fig.  104) 

quinquepartitus  Distant 

—  Head  process  with  3  apical  teeth 3 

3(2).  Hind  wing  with  large  (ca  2  mm),  hyaline, 
apical  spots,  visible  from  below  with  wing 
unspread   4 

—  Hind  wing  without  hyaline,  apical  spots,  but 
small  pruinose  areas  sometimes  present,  visi- 
ble when  wings  are  spread 5 

4(3).  Hind  wings  yellow  or  orange  at  base;  head 
process  with  3  black  spots  dorsally  just  behind 
apical  area  (Fig.  113)  ...  .  xanthopterus  Schmidt 

—  Hind  wing  red  at  base;  head  process  with  no 
black  markings  near  apex,  but  diamond- 
shaped,  red  area  in  same  position  (Fig.  107) 
oceUatus  Signoret 

5(3).  Basal  area  of  hind  wing  yellow  or  yellow- 
orange   moebiusi  Schmidt 

—  Basal  area  of  hind  wing  red  or  reddish  orange 

6 

6(5).        Tegmen    with    continuous,    broad,    yellow, 

transverse  fasciae  present    7 

—  Tegmen  lacking  transverse  fascia,  or,  if 
present,  interrupted  medially 8 

7(6).        Tegmen  brown,  transverse  fascia  yellow  only 

diligens,  n.  sp. 

—  Tegmen  scarlet  with  green  veins,  anterior 
edge  of  fascia  partially  bordered  with  black 
tripartitus  Metcalf 

8(6).  Transverse  fascia  absent,  apical  trident  of 
head  process  2/3  as  wide  as  distance  between 

supraocular  spines  (Fig.  108) 

delicatus,  n.  sp. 

—  Transverse  fascia  interrupted  medially;  apical 
trident  of  head  process  wider  (Figs.  105,  106, 
109,  110)    9 

9(8).  Head  process  with  apical  teeth  broader  than 
head  (Fig.  105);  hind  wing  with  black  or  fus- 
cous area  covering  apical  three-lourths 

recalls  Caldwell 

—  Head  process  with  apical  teeth  not  wider  than 
head  (Figs.  106,  109,  110);  hind  wing  with 
black  or  fuscous  areas  covering  apical  third   .    10 

10(9).  Tegmen  green  in  basal  two-thirds,  with  a  few 
small,  roinid,  red  or  orange  maculae;  apical 

third  with  large,  brown  spots 

hoffmannsi  Schmidt 


1988 


OBrien:  New  World  Fulgoridae 


161 


Figs.  102-117.  Head  and  pronotum,  dorsal  view,  drawn  with  head  process  at  45-degree  angle  so  that  the  apex  is 
horizontal;  102,  Phrictus  diadema  L.;  103,  P.  tripartitus  Metcalf;  104,  P.  quinquepartitus  Distant;  105,  P.  regalis 
Caldwell;  106,  P.  punctatus  Caldwell;  107,  P.  ocellatus  Signoret;  108,  P.  delicatus  O'Brien;  109,  P.  minutacanthis 
Caldwell;  110,  P.  hoffmansi  Schmidt;  111,  P.  moebiusi  Schmidt;  112,  P.  auromactdatus  Distant;  113,  P.  xanthopterus 
Schmidt;  114,  P.  diligens  O'Brien;  115a,  P.  diadema  (L.)  and  115b,  P.  triportitus  Metcalf  (head,  lateral  view);  116, 
Cathedra  spectabilis  Carreno;  117,  O.  carrenoi  Signoret. 


162 


Great  Basin  Naturalist  Memoirs 


No.  12 


—  Tegmen  brown  or  reddish  brown,  sometimes 
with  yellow  spots 11 

11(10).  Tegmen  some  shade  of  reddish  brown,  with 
pink,  calloused  areas  present,  especially 
basally;  head  process  toothed  on  ventral  sur- 
face (Fig.  115a) diadeina  (Linnaeus) 

—  Tegmen  brown,  maculate  with  yellow;  head 
process  not  toothed  ventrally 12 

12(1 1).  Combined  length  of  head  process  and  head  as 
long  as  pronotum;  expanse  of  trifurcate  apex 
equal  to  distance  between  ocular  spines  (Fig. 
109) minutacanthis  Caldwell 

—  Combined  length  of  head  process  and  head 
longer  than  pronotum;  expanse  of  apex  much 
greater  than  distance  between  ocular  spines 
(Fig.  106) punctatus  Caldwell 

Phrictus  auromaculatus  Distant 

Fig.  112 

Phrictus  auromaculatus  Distant  1905:672.  Type  reposi- 
tory: BMNH. 

Phrictus  notatus  Lallemand  1931:188.  Type  repository: 
FSAG,  NRS.  New  synonymy. 

Length. — Male  29-32  mm,  female  34-35 
mm;  head  length  3.7  mm;  ratio,  head/prono- 
tum  1.0. 

Head  process  semicircular  at  apex,  not  di- 
vided into  teeth,  orange-red;  three  round, 
waxy  plates  on  each  side  (Fig.  112).  Tegmen 
evenly  pale  brown  mottled  with  yellow,  veins 
dark  brown  in  corium,  no  transverse  fascia. 
Hind  wings  yellow  at  base,  brown  posteriorly; 
two  maculate,  brown  bands  from  anal  area 
running  into  yellow  area. 

This  species  is  easily  separated  by  the  apex 
of  the  head  not  being  divided  into  spinelike 
segments.  It  may  be  verified  by  the  yellow 
base  of  the  hind  wings  and  the  evenly  colored 
tegmina. 

Distribution. — Syntypes:  Bolivia  (auro- 
maculatus), Ecuador  (notatus).  I  have  seen 
five  other  specimens  collected  from  Bolivia: 
Santa  Cruz;  Prov.  Sara  [sic],  450  m;  Brazil: 
Sinop;  Peru:  Jauja  Prov.,  Satipo. 

Phrictus  delicatus,  n.  sp. 

Fig.  108 

Length. — Female  36  mm;  head  length  3.7 
mm;  ratio,  head/pronotum  1.3. 

Head  process  tripartite,  lateral  portions  di- 
rected anterad,  not  laterad,  smaller  than  that 
oi minutacanthis;  head  pale  brown  (Fig.  108). 
Mesonotum  with  three  pairs  of  dark  spots  (pin 
is  placed  through  apex;  there  may  or  may  not 
be  one  there).  Tegmen  evenly  colored,  no 


fascia.  Hind  wings  red  basally,  posteriorly 
pale  brown. 

This  species  may  be  identified  by  the  pale 
brown,  narrow  head  process,  without  a  white 
dorsal  stripe  and  red  venter  and  apex.  The 
pale  apex  of  the  hind  wings  separates  this 
species  from  all  but  moebiusi.  The  evenly  col- 
ored tegmen,  broader  and  less  rooflike  than 
most  of  the  species,  is  similar  to  that  of  auro- 
maculatus and  ocellatus. 

HoLOTYPE  (female). — Brazil:  Amazonas, 
S.  Paulo  de  Olivenca,  Vn-32,  A.  Mailer 
Colin.,  Frank  Johnson  Donor  (AMNH).  The 
printed  label  says  S.  P.  Olivenca/  Amazonas/ 
Brasil  Vn-32.  The  handwritten  label  under- 
neath gives  the  more  complete  data. 

Phrictus  diadema  (Linnaeus) 
Figs.  102,  115a. 

Fulgora  diadema  Linnaeus  1767:703.  Type  repository: 

UZIU. 
Phrictus  diadema  (Linnaeus),  Spinola  1839:  219. 

Length. — Male  38  mm,  female  45  mm; 
head  length  8  mm;  ratio,  head/pronotum  2.0. 

Apex  of  head  process  trifurcate,  three  or 
four  pairs  of  lateral,  waxy  plates  and  one  me- 
dian oval  (Fig.  102).  Ventral  margin  of  head 
process  in  lateral  view  sinuate.  Hind  wings 
red  at  base.  Tegmina  mottled  with  red  and 
brown  in  large  splotches,  transverse,  yellow- 
ish fascia  narrow,  not  bordered  anteriorly  in 
black. 

This  species  most  closely  resembles  regalis 
in  the  mottled  color  pattern  of  the  tegmen  and 
pale,  transverse  fascia  and  the  sinuate  ventral 
margin  of  the  head  process  in  lateral  view,  but 
regalis  has  a  much  wider,  trifurcate  apex  on 
the  head  process,  exceeding  the  distance  be- 
tween the  tips  of  the  supraocular  spines  (Fig. 
105).  In  diadema  the  spines  and  apex  are 
about  the  same  width. 

Distribution. — Type:  India  ?  [error].  I 
have  seen  two  other  specimens  collected  in 
November  from  BRAZIL:  Linhares. 

Phrictus  diligens,  n.  sp. 
Fig.  114 

Length. — Female  40-43  mm;  head  length 
6  mm;  ratio,  head/pronotum  1.3. 

Apex  of  head  process  trifurcate,  narrow 
band  of  waxy  plates  each  side  (Fig.  114).  Hind 
wings  red  at  base.  Tegmen  evenly  brown  with 
unbroken  clear  transverse  fascia. 

This  species  may  be  identified  easily  by  the 


1988 


O'BRIEN:  New  World  Fulgoridae 


163 


browii  tegmen  with  the  unbroken  transverse 
band.  I  name  it  diligens,  which  means  prim, 
because  it  is  more  conservative  in  color  and 
shape  of  head  process  than  the  other  species. 
It  is  most  closely  related  to  diadema  and  re- 
galis  in  the  color  pattern  of  the  tegmen. 

HoLOTYPE  (female). — Colombia:  1,500  ft, 
Anchicaya,  26-VII-1970,  J.  M.  Campbell 
(LOB).  Paratype  (female):  Colombia:  Dept. 
El  Valle,  Anchuaya,  200-300  mi,  27-IV- 
1969,  Gonin-H  Balazar-S.  Draachsler  Rec. 
(MNHP). 

Phrictus  hoffmannsi  Schmidt 

Fig.  110 

Phrictus  hoffmannsi  Schmidt  1905:338.  Type  repository: 
IZW. 

Length. — Male  37  mm,  female  48  mm; 
head  length  7  mm;  ratio,  head/pronotum  1.0. 

Apex  of  head  process  trifurcate,  large, 
squarish,  waxy  plate  area  on  each  side 
(Fig.  110).  Hind  wings  red  at  base.  Tegmina 
green  mottled  with  yellowish  orange  in 
large  splotches,  clavus  predominately  orange, 
broad,  transverse,  yellow  fascia  interrupted 
by  brown  spots  medially  and  toward  commis- 
sure, membrane  yellowish  brown  with  brown 
oval  spots. 

This  species  may  be  distinguished  easily  by 
its  bright  green  and  yellowish  orange  tegmen. 

Distribution. — Holotype  (female):  Peru: 
Chanchamayo.  I  have  seen  four  other  speci- 
mens collected  in  January  from  Peru:  Tingo 
Maria;  Ecuador:  Guayaquil,  Loja. 

Phrictus  minutacanthis  Caldwell 
Fig.  109 

Phrictus  jninutacanthis  CaldweW  1945:180,  pi.  7,  Figs.  2, 
16,  pi.  8,  Fig.  2,  pi.  9.  Type  repository:  USNM. 

Length. — Female  36  mm;  head  length  4.8 
mm;  ratio,  head/pronotum  1.5. 

Apex  of  head  process  trifurcate,  small;  waxy 
plate  on  each  side;  pale  red,  midline  near  apex 
with  reddish  triangle  (Fig.  109).  Hind  wings 
red  at  base.  Tegmina  light  brown  evenly  mot- 
tled with  yellow,  transverse  yellowish  fascia 
indistinct,  broken. 

This  species  may  be  identified  by  its  small 
head.  It  resembles  xanthopterus  and  punc- 
tatus  most  closely  in  the  light  brown  tegmen 
with  yellow  markings,  and  ocellatus  and  punc- 
tatus  in  having  the  red  subapical  triangle  on 
the  pale  dorsum  of  the  head  process. 


Distribution. — Holotype:  Peru:  Chaqui- 
mayo.  I  have  seen  only  the  type. 

Phrictus  moebiusi  Schmidt 
Fig.  Ill 

Phrictus  moebiusi  Sclimiclt  1905:335.  Type  repository: 

ZMHB. 
Phrictus  sordidus  Caldwell  1945:180,  pi.  7,  Figs.  3,  20, 

pi.  8,  Figs.  1,  13,  pi.  9.  Type  repository:  USNM. 

New  synonymy. 

Length — Male  40  mm,  female  43-48  mm; 
head  length  6  mm;  head/pronotal  ratio  1.2. 

Apex  of  head  process  trifurcate,  large,  waxy 
plate  on  apex  (Fig.  111).  Hind  wings  yellow  at 
base,  posterior  pale  brown.  Tegmina  brown 
evenly  mottled  with  yellow,  transverse  fascia 
pale,  broad,  interrupted  by  two  brown  spots 
in  middle  and  near  commissural  margin. 

This  is  one  of  three  species  with  yellow 
coloration  at  the  base  of  the  hind  wing.  P. 
auromaculatus  has  a  head  process  without 
apical  spines,  and  xanthopterus  has  white 
spots  in  the  posterior  brown  area  of  the  hind 
wing.  The  type  of  sordidus  is  a  pale  specimen 
that  agrees  in  all  external  characters  with  moe- 
biusi. 

Distribution. — Syntype  (female):  Colom- 
bia: Antioquia  (moebiusi);  Ecuador  (male 
holotype  sordidus).  I  have  seen  four  other 
specimens  collected  in  January  from  "New 
Granada"  and  Ecuador:  Route  Mono-Los 
Banos,  km  37,  2,385  m. 

Phrictus  ocellatus  Signoret 
Fig.  107 

Phrictus   ocellatus   Signoret    1855:v.    Type   repository: 

nhmv. 

Length. — Male  35  mm,  female  42  mm; 
head  length  6.0;  ratio,  head/pronotum  1.9. 

Apex  of  head  process  trifurcate;  large,  waxy 
plates  on  each  side;  red  diamond  and  median 
line  along  pale  dorsum  of  process  (Fig.  107). 
Hind  wings  red  at  base,  white  spots  in  brown 
posterior  area.  Tegmina  evenly  mottled  with 
pale  yellow  and  red  brown,  no  fascia. 

Two  species  have  hind  wings  with  white 
spots  in  the  brown  posterior  area,  xan- 
thopterus with  a  yellow  base  and  ocellatus 
with  a  red  base. 

Distribution — Holotype  (female):  Co- 
lombia: Fusagasuga.  I  have  seen  two  other 
specimens  collected  in  June  from  VENEZUELA: 
Rancho  Grande. 


164 


Great  Basin  Naturalist  Memoirs 


No.  12 


Phrictus  punctatus  Caldwell 

Fig.  106 

Phrictus  punctatus  Ca\dwe\\  1940:181,  pi. 7,  Figs.  12,  23, 
pi.  8,  Fig.  6,  pi.  9.  Type  repository:  USNM. 

Length. — Female  40  mm,  head  length  6.5 
mm;  ratio,  head/pronotum  1.4. 

Apex  of  head  process  trifurcate,  with  trans- 
verse, waxy  plates;  triangular  red  area  on  mid- 
line behind  apex  (Fig.  106).  Hind  wings  red  at 
base,  pale  brown  posteriorly.  Tegmina  brown 
mottled  with  yellow  in  large  splotches,  clavus 
predominantly  yellow;  transverse,  yellowish 
fascia  narrow,  interrupted  medially;  diagonal 
yellow  line  from  clavus  to  costal  margin. 

In  coloration,  the  tegmen  o( punctatus  most 
closely  resembles  xanthoptertis,  while  the 
color  of  the  head  process  resembles  oceUatus 
and  minutacanthis. 

Distribution. — Holotype  (female):  Pana- 
ma: Bugaba.  Paratype:  El  Volcan.  I  have  seen 
two  other  specimens,  one  without  data,  the 
other  collected  from  CosTA  RiCA:  Palo  Verde. 

Phrictus  quinquepartitus  Distant 

Fig.  104 

Phrictus  quinquepartitus  Distant  1883:24,  pi. 4,  Fig.  8. 
Type  repository:  BMNH. 

Length. — Male  40-42  mm,  female  45-48 
mm;  head  length  9.3  mm;  ratio,  head/prono- 
tum 2.06. 

Apex  of  head  process  trifurcate,  one  median 
and  three  lateral  pairs  of  waxy  tufts  in  apex 
(Fig.  104).  Hind  wings  red  at  base.  Tegmina 
red  with  green  veins;  transverse,  yellowish 
fascia  bordered  anteriorly  in  black. 

This  species  is  closely  related  to  tripartitus 
in  the  color  of  the  tegmen  but  differs  in  the 
apex  of  the  head  process. 

Distribution. — Syntypes:  Panama:  Buga- 
ba, 3,000-4,000  ft;  Colombia:  Bogota.  I  have 
seen  nine  other  specimens  collected  in  Feb- 
ruary, June,  August,  October  and  December 
from  Panama:  Barro  Colorado  Island,  Code, 
Chiriqui;  CoSTA  RiCA:  La  Selva. 

Phrictus  regalis  Caldwell 
Fig.  105 

Phrictus  regalis  Ca\dwe\\  1945:183,  pi.  7,  Figs.  7,  19,  pi. 
8,  Fig.  8,  pi.  10.  Type  repository;  USNM. 

Length. — Male  42  mm,  female  49-53  mm; 
head  length  10;  ratio,  head/pronotum  2.3. 

Apex  of  head  process  trifurcate,  three  waxy 
tufts  on  each  side,  two  larger  ones  medially. 


semicircular  carinate  area  astride  median 
carina  (Fig.  105).  Hind  wings  red  at  base. 
Tegmina  mottled  with  red  and  brown  in  large 
splotches;  transverse,  yellowish  fascia  nar- 
row, not  bordered  anteriorly  in  black. 

This  species  is  very  similar  to  diadema  in 
the  color  of  the  tegmen,  but  the  apex  of  the 
head  process  is  much  wider  in  regalis  (Figs. 
105,  102). 

Distribution. — Holotype  (female):  French 
GuiANA:  Maroni  River,  near  Duserre.  I  have 
seen  three  other  specimens  collected  in 
March  from  French  Guiana:  St.  Jean  du  Ma- 
roni; Suriname:  Nassaugebergte. 

Phrictus  tripartitus  Metcalf 

Figs.  103,  115b 

Phrictus  tripartitus  Metcalf  1938:365,  pis.  20,  21.  Type 
repository:  AMNH. 

Length. — Male  35-40  mm,  female  43-45 
mm;  head  length  9.4  mm;  ratio,  head/prono- 
tum 1.8. 

Apex  of  head  process  trifurcate,  three 
round,  waxy  plates  and  one  oval  on  each  side 
(Fig.  103).  Head  process  sinuate  ventrally  in 
lateral  view  (Fig.  115b).  Hind  wings  red  at 
base.  Tegmina  red  with  green  veins;  trans- 
verse, yellowish  fascia  bordered  anteriorly  in 
black. 

This  species  most  closely  resembles  quin- 
quepartitus in  the  color  of  the  tegmen,  but  the 
species  may  be  separated  by  the  shape  of  the 
apex  of  the  head  process.  Some  specimens  of 
tripartitus  do  have  five  spines,  but  the  inter- 
mediate spines  are  smaller  than  those  of  quin- 
quepartitus (Fig.  104)  and  the  shape  of  the 
other  teeth  is  like  that  illustrated  for  triparti- 
tus (Fig.  103). 

Distribution. — Type:  Belize.  I  have  seen 
21  other  specimens  collected  from  August 
through  October  from  Belize:  24  mi  SE  Bel- 
mopan, 10-11  mi  S  Georgeville;  Honduras: 
La  Ceiba;  MEXICO:  Nuevo  X-Can  [sic],  San 
Quintin,  Yucatan. 

Phrictus  xanthupterus  Schmidt 

Fig.  113 

Phrictus  xanthopterus  Schmidt  1910:144.  Type  reposi- 
tory: IZN. 

Length. — Female  39-41  nnn;  head  length 
5  mm;  ratio,  head/pronotum  1.0. 

Head  process  with  apex  trifurcate,  without 
waxy  plates,  orange-red,  with  three  triangu- 
lar, black  spots  behind  apex  (Fig.  113).  Hind 


1988 


O'BRIEN;  New  World  Fulgoridae 


165 


wings  yellow  at  base  with  white  spots  in 
brown  posterior  area.  Tegmen  brown  with 
yellow  markings,  half  of  clavus  predominantly 
yellow;  transverse,  yellow  fascia  interrupted 
medially. 

This  species  is  one  of  the  most  distinctive 
with  the  hind  wings  yellow  at  base  and  with 
white  spots  in  the  apical  brown  area  and  the 
three  black,  subapical  spots  on  the  head  pro- 
cess. The  tegmen  most  closely  resembles 
punctatus. 

Distribution. — Holotype  (female):  Ecua- 
dor: Canelos.  I  have  seen  six  other  specimens 
collected  in  June  and  July  and  December 
from  Peru:  Cusco;  Sta.  Isabel;  Route  Olmoc- 
Tarapoto,  km  385,  1,800  m;  Tingo  Maria; 
Ecuador:  Lumbagui. 

Rhabdocephala  Van  Duzee 

Fig.  23 

Rhabdocephala  Van  Duzee  1929;  190.  Type-species;  R. 
brunnea  Van  Duzee,  original  designation. 

Small,  brown,  narrow  insects,  12-16  mm 
long;  head  about  1/3  length  of  insect.  Head 
process  porrect,  terete,  sides  parallel;  apex 
slightly  widened;  dorsal  lateral  carinae  com- 
plete, ventral  incomplete,  present  only  in 
middle  of  process.  Preocular  and  supraocular 
flanges  absent.  Tegmen  with  M  not  pec- 
tinate, cross-veins  reticulate,  apical  margin 
angled  at  apex  of  clavus,  forming  angulate,  not 
smoothly  rounded,  apical  margin.  Ninth  ab- 
dominal tergite  1.5  times  length  of  eighth. 

These  small  insects  may  be  separated  from 
other  small  Fulgoridae  by  the  simple  cephalic 
process  with  no  dorsolateral  undulate  carinae 
as  in  Scolopsella  and  the  head  process  not 
dorsoventrally  flattened  as  in  Amycle. 

Rhabdocephala  brunnea  Van  Duzee 

Fig.  23 

Rhapdocephala   brunnea   Van    Duzee    1929;  191.    Type 
repository;  CAS. 

Length. — Male  12-14  mm,  female  14.5- 
16  mm; "head  5.2  mm;  ratio,  head/pronotum 
2.1. 

Tegmen  brown  mottled  with  white.  Hind 
wing  red  in  basal  third,  brown  posteriorly. 

Distribution. — Holotype  (female):  USA: 
Arizona.  I  have  seen  15  other  specimens  col- 
lected from  May  to  October  from  USA:  ARI- 
ZONA: Pima,  Cochise,  and  Santa  Rita  counties; 
Mexico:  Sonora,   15  mi  S  Hermosfllo.  Two 


specimens    were    collected    on    Baccharis 
sarothroides  Gray. 

Scolopsella  Ball 

Figs.  8-13,  24-25 

Scohypsella  Ball   190.5;  118.   Type-species;   S.    reticulata 
Hall  by  original  designation. 

Small,  brown  insects,  14-21  mm  long,  head 
.23-. 28  length  of  insect.  Head  process  por- 
rect, terete,  very  slightly  enlarged  at  apex; 
dorsolateral  carinae  undulate;  median  and  lat- 
eral ventral  and  marginal  carinae  complete  on 
process  and  frons.  Preocular  flange  indistinct. 
Tegmen  with  M  pectinate  (3  branches),  com- 
missural margin  emarginate  between  claval 
apex  and  wing  tip. 

This  genus  of  small,  brown  insects  may  be 
separated  from  other  similar  genera  by  the 
straight  head  process  with  dorsolateral  mar- 
gins crenulate  or  undulate  and  without  the 
apex  greatly  expanded.  The  head  process  is 
not  as  unadorned  as  Rhabdocephala,  lacks  the 
transverse  wrinkles  o(  Aphrodisias ,  and  is  not 
flattened  as  in  Amycle.  Diareusa  specimens 
are  much  larger  insects  with  much  shorter 
heads. 

Key  to  the  Species  oi  Scolopsella 

1.      Length  of  anal  flap  of  male  greater  than  width  at 
apical    expansion    (Fig.    23);    apical    margin    of 

pygofer  emarginate  ventrally  (Fig.  25)    

reticulata  Ball 

—    Length  of  anal  flap  of  male  less  than  width  at 
apical    expansion    (Fig.    20);    apical    margin    of 

pygofer  smoothly  convex  (Fig.  22) 

mexicana  O'Brien 

Scolopsella  niexicana  O'Brien 

Figs.  8-10,  24 

Scolopsella  niexicana  O'Brien  1985:73.  Type  repository; 
LOB. 

Length. — Male  14-15  mm,  female  17-21 
mm;  head  length  5.6  mm;  ratio,  head/prono- 
tum 3.7. 

Tegmen  various  shades  of  brown,  cross- 
veins  and  reticulations  pale,  some  apical  cells 
white.  Structural  characteristics  common  to 
both  species  given  in  generic  description. 

Scolopsella  reticulata  and  S.  mexicana  may 
be  separated  by  the  shape  of  the  anal  flap, 
which  can  be  seen  without  dissection,  the 
applicable  characters  being  given  in  the  key. 
Once  males  are  identified  in  this  way,  females 
may  often  be  associated  with  them  because 
the  golden  brown  reticulata  is  slightly  larger 


166 


Great  Basin  Naturalist  Memoirs 


No.  12 


and  the  head  process  is  wider,  and  mexicana  is 
darker. 

Distribution. — Type:  Mexico.  I  have 
seen  seven  other  specimens  collected  from 
June  through  September  from  Baja  Califor- 
nia. 

Scolopsella  reticulata  Ball 
Figs.  11-13,  25 

Scolopsella  reticulata  Ball   1905:118.   Type  repository: 
CAS. 

Length. — Male  14-16  mm,  female  17-19 
mm;  head  length  5  mm;  ratio,  head/pronotum 
4.1. 

The  structural  characters  and  coloration  do 
not  differ  sufficiently  from  S.  mexicana  to  de- 
termine the  species  without  either  using  the 
shape  of  the  male  anal  flap  or  comparing  the 
two  species  as  discussed  above. 

Distribution. — Holotype:  Arizona.  I  have 
seen  other  specimens  collected  from  April  to 
November  in  Arizona:  Maricopa,  Pima, 
Pinal,  and  Santa  Cruz  counties;  California: 
Joshua  Tree  National  Momument,  Riverside 
or  San  Bernardino  County. 

Sinuala,  n.  gen. 
Figs.  43,  46-48 

Type-species:  Sintiala  stall  O  Brien. 

Reddish  brown,  medium-sized  insects, 
19-21  mm  long,  head  .12  to  .15  length  of 
insect.  Head  process  porrect,  terete,  en- 
larged at  apex;  vertex  truncated  triangle,  basal 
angle  rounded,  supraocular  lobes  variable. 
Frons  with  lateral  carinae  diverging  anteriorly 
to  form  process;  lateral  margins  expanded 
near  frontoclypeal  suture,  ridge  from  lateral 
carina  to  margin  at  this  expanded  area.  Frons 
and  clypeus  about  in  same  plane,  frons  not 
raised  above  clypeus;  clypeus  medially  and 
marginally  carinate.  Preocular  flange  length- 
ened into  pleural  carina.  Pronotum  without 
raised  median  carina,  lateral  fossettes  and  two 
pleural  carinae  present;  apex  of  mesonotum 
with  striate  triangle,  delimited  by  carinae,  not 
high  ridge.  Tegmen  with  M  not  pectinate, 
clavus  open,  longitudinal  veins  angulate, 
cross-veins  variable.  Fore  and  mid  femora  ex- 
panded. Posterior  tibiae  with  eight  lateral 
spines.  Female  abdominal  tergite  nine  twice 
length  of  eight. 

This  genus  is  similar  to  Aphrodisias ,  from 
which  it  differs  by  the  costal  margin  of  the 


tegmen  sinuate  and  the  head  process  not 
transversely  ridged. 

Key  to  the  Species  oi  Sinuala 

1.  Supraocular  flange  expanded,  height  equal  to 
width  of  eye;  tegmen  with  10  strong  tubercles 
tuberculata,  n.  sp. 

—  Supraocular  flange  smaller  than  above;  tegmen 
with  tubercles  evanescent    2 

2(1).  Vertex  as  long  as  broad  (Fig.  46);  hind  wing  red 
at  base stali,  n.  sp. 

—  Vertex  1.7  times  longer  than  broad  (Fig.  48); 
hind  wing  pink  at  base    sclimidti,  n.  sp. 

Sinuala  stali,  n.  sp. 
Fig.  46 

Length. — Male  19-20  mm;  head  2.3  mm; 
ratio,  head/pronotum  1.4. 

Head  process  short,  vertex  as  long  as  broad. 
Hind  wing  red  at  base,  narrow  anterior  mar- 
gin and  broad  posterior  area  brown,  apical 
angle  brown  with  some  white  spots  and 
crossveins. 

This  species  may  be  easily  separated  from 
tuberculata,  which  it  resembles  in  wing  color 
and  head  length,  by  the  small,  supraocular 
flange  oi  stali.  I  name  this  species  in  honor  of 
Carl  Stal,  who  provided  a  fine  basis  for  generic 
work  in  this  family. 

Holotype  (male). — Honduras:  Lago 
Yojoa.,  6-VI-1976,  J.  V.  Mankins,  coll.  (LOB). 
Paratype  (male):  El  Salvador:  Alfredo  Mar- 
tinez Cuestas,  no  further  data  (NCSR). 

Sinuala  schmidti,  n.  sp. 

Fig.  47 

Length. — Female  21.5  mm;  head  3.3  mm; 
ratio,  head/pronotum  2.2. 

Head  process  elongate,  vertex  1.7  times  as 
long  as  broad.  Hind  wing  pink  at  base,  then 
white;  anal  margin  pale  brown,  posterior  mar- 
gin with  cells  infuscate,  veins  white;  apical 
third  brown  with  white  lines  following  each 
cross- vein. 

This  species  may  be  separated  from  the 
others  by  the  long  head  and  the  pale  base  of 
the  hind  wings.  I  name  this  species  in  honor  of 
Edmund  Schmidt,  who  had  an  excellent  eye 
for  species  and  a  gift  for  describing  the  most 
significant  characters  precisely. 

HoLOT\'PE  (female):  BELIZE:  Corazol,  1  mi 
NW  Corozal,  12-VIII-1977.  C.  W.  and  L.  B. 
O'Brien  and  Marshall  (LOB). 

Sinuala  tuberculata,  n.  sp. 

Figs.  43,  48 


1988 


O'BRIEN:  New  Would  Fulgoridae 


167 


Length. — Female  21  mm;  head  2.6  mm; 
ratio,  head/pronotum  1.6. 

Head  process  short,  vertex  with  postocular 
flange  raised  above  vertex  equal  to  width  of 
eye,  vertex  as  long  as  broad.  Hind  wing  red  in 
basal  half,  anal  area  brown,  apical  2/5  brown 
with  some  white  spots  and  cross-veins.  Five 
strong  tuberculations  on  veins  in  anterior  half 
of  tegmen,  five  smaller  in  apical  half. 

This  species  may  be  separated  easily  from 
the  others  by  the  strong  tubercles  on  the  teg- 
men and  the  strong  supraocular  flange. 

Holotype  (female):  Costa  Rica:  P.  R.  Uh- 
ler  Colin.  (NCSR). 

Stalubra,  n.  gen. 
Figs.  1-3,  .5,  6 

Type-species:  Stalubra  brunnea  O'Brien. 

Medium-sized,  brown  or  reddish,  transpar- 
ent-winged insects,  19-28  mm  long,  head  .  14 
to  .19  length  of  insect.  Head  process  terete, 
gradually  narrowing  to  apex,  upturned  at 
apex.  Vertex  twice  as  long  as  broad,  elongate 
medially  (Fig.  1).  Lateral  carinae  of  frons  par- 
allel or  diverging  toward  head  process,  me- 
dian carina  on  process  and  basal  2/3  of  frons; 
frons  only  very  slightly  raised  above  fronto- 
clypeal  suture;  lateral  margins  of  frons  ex- 
panded near  apex;  weak  ridge  from  lateral 
carina  to  lateral  margins.  Clypeus  laterally 
carinate.  Preocular  flange  reduced  to  slight 
tubercle  on  vertical  carina  joining  marginal 
carinae  of  frons  and  vertex  (Fig.  3).  Pronotum 
without  median  carina  raised,  fossettes  and 
two  pleural  carinae  present;  mesonotal  apex 
not  depressed  or  striate.  Tegmen  with  M 
pectinate  (5-7  branches);  clavus  open;  cross- 
veins  undulate,  parallel;  no  small  spines  on 
veins,  only  on  wing  margins.  Posterior  tibiae 
with  six  lateral  spines.  Female  abdominal  ter- 
gite  nine  shorter  than  eight. 

This  genus  is  similar  to  Enhydria  in  the 
compressed,  recurved  head  process  and 
translucent  tegmen;  however,  it  lacks  the 
hairs  on  the  veins  of  the  tegmen  and  the 
dense,  diagonal  cross-veins  along  the  costal 
margin  found  in  Enhydria,  and  the  head  pro- 
cess is  less  strongly  curved.  It  differs  from 
Chilobia  in  the  vertex  being  elongate  medi- 
ally, whereas  the  vertex  in  Chilobia  is  broader 
than  long  and  emarginate  medially  (Figs.  51, 
54,  60)  or  hidden  by  the  recurved  head  pro- 
cess (Fig.  57). 


Lallemand's  species  Enhydria  rufula  (1966) 
belongs  in  this  genus,  but  I  cannot  tell  from 
his  description  which  it  is  of  two  of  the  three 
species  I  have  before  me.  The  requested  type 
has  not  yet  arrived,  and  so  these  two  species 
will  be  discussed  in  Part  II. 

Stalubra  brunnea,  n.  sp. 

Figs.  l-,3,  5,  6 

Length. — Male  19-22  mm;  head  2.6  mm; 
ratio,  head/pronotum  1.9. 

Head  short,  vertex  1.5  times  as  long  as 
wide.  Hind  wing  clear  with  brown  veins.  Teg- 
men translucent  with  brown  markings  and 
minute  red  dots  along  veins,  apex  pale  brown 
with  white  circles  plus  some  darker  brown 
spots. 

Holotype  (male). — Brasil  [sic]:  Mato 
Grosso,  Sinop  (12°31'S,  55°37'W),  X-1975, 
M.  Alvarenga  (LOB).  Paratypes  (25  male): 
same  data  (LOB,  MZSP);  Guyana:  1  male. 
Upper  Mazaruni  River,  IX-X-1938,  A.  S. 
Pinkus(AMNH). 

ACKNOWLE  DGM  ENTS 

It  is  with  great  pleasure  that  I  dedicate  this 
paper  to  our  Latin  American  colleagues  and  to 
Paul  Oman,  among  whose  many  accomplish- 
ments is  a  paper  on  a  U.S.  fulgorid  (Oman 
1936). 

I  thank  the  American  Philosophical  Society 
for  a  grant  to  study  at  European  museums 
(Grant  No.  5771,  Penrose  Fund),  without 
which  this  work  would  have  been  impossible. 
I  am  deeply  indebted  to  B.  V.  Ridout  for  his 
work  on  Fulgora.  I  greatly  appreciate  the  crit- 
ical reading  of  the  manuscript  by  F.  W.  Mead, 
G.  J.  Wibmer,  and  S.  W.  Wilson  and  the 
sending  and  examining  of  types  and  paratypes 
of  R.  C.  Froeschner  and  N.  D.  Penny.  I  am 
also  grateful  to  the  following  museums  for 
allowing  me  to  examine  types  and  borrow 
specimens  in  their  collections:  AMNH — 
American  Museum  of  Natural  History,  New 
York;  BMNH — British  Museum  (Natural  His- 
tory), London,  England;  CAS — California 
Academy  of  Sciences,  San  Francisco;  DEI — 
Institute  fiir  Pflanzenschutzforschung  (for- 
merly Deutsches  Entomologisches  Institut), 
Kleinmachnow,  Eberswalde,  East  Germany; 
EMAU — Zoologisches  Institut  und  Museum, 
Ernst  Morits  Arndt  Universitat,  Greifswald, 
East  Germanv;  FMNH — Field  Museum  of 


168 


Great  Basin  Naturalist  Memoirs 


No.  12 


Natural  History,  Chicago,  Illinois;  FSAG — 
Faculte  des  Sciences  Agronomiques,  Gem- 
bloux,  Belgium;  IZW — Instytut  Zoologii 
PAN,  Warsaw,  Poland;  LOB— Lois  O'Brien 
collection,  Tallahassee,  Florida;  MCZ — 
Museum  of  Comparative  Zoology,  Harvard 
University,  Cambridge,  Massachusetts; 
MHNP — Museum  National  d'Histoire  Na- 
turelle,  Paris,  France;  MRSN — Museo  Re- 
gionale  di  Scienze  Naturali,  Turin,  Italy; 
MZF — Museo  Zoologico  dell'  Universita  de- 
gli  Studie  di  Firenze,  Florence,  Italy; 
MZSP — Muzeo  de  Zoologia,  Universidade  de 
Sao  Paulo,  Brazil;  NCSU— North  Carolina 
State  University,  Raleigh;  NHMV — Natur- 
historisches  Museum,  Vienna,  Austria; 
NRS — Naturhistoriska  Riksmuseet,  Stock- 
holm, Sweden;  OSU — Ohio  State  University, 
Columbus;  UCB — University  of  California, 
Berkeley;  UMO — University  Museum,  Ox- 
ford University,  Oxford,  England;  UNAM — 
Instituto  de  Biologia  and  Museo  de  Zoologia, 
Universidad  Nacional  Autonoma  de  Mexico, 
Mexico  City,  D.F.;  USNM— United  States 
National  Museum,  Washington,  D.C.; 
UZIL — Universitets  Zoologiska  Institut, 
Lund,  Sweden;  UZIU — Universitets  Zoolo- 
giska Institut,  Uppsala,  Sweden;  ZMHB — 
Zoologisches  Museum,  Humboldt  Universi- 
tat.  East  Berlin,  East  Germany;  ZMUC- 
Zoologisk  Museum,  Universitets  Copen- 
hagen, Denmark;  ZSBS — Zoologische  Samm- 
lungen  des  Bayerischen  Staates,  Munich, 
East  Germany. 

Literature  on  New  World  Fulgoridae 
Since  Metcalfs  Catalog 

(a  complete  bibliography  will  be  included  with  the 
final  part.) 

Brailovsky,  a.,  H.  and  C.  R.  Beutelspacher  B  1978. 
Una  nueva  especie  de  Fulgora  Linneo  (Ho- 
moptera:  Fulgoridae)  de  Mexico.  An.  Inst.  Biol. 
Univ.  Nal.  Auton.  Mexico,  Ser.  Zool.  49: 
175-182. 

Dlabola,  J  ,  AND  F  Zayas  1980.  Eine  neue  Fulgoride  der 
Gattung  javiaicastes  von  Cuba  (Homoptera, 
Auchenorrhyncha).  Acta  Faun.  Entomol.  Mus. 
Nat.  Pragae  16:  83-84. 

Emelyanov,  A.  F  1979.  The  problem  of  the  family  dis- 
tinction between  the  Fulgoridae  and  the  Dic- 
tyopharidae  (Homoptera,  Auchenorrhyncha).  In: 
Phylogeny  and  systematics  of  insects.  Acad.  Sci. 
USSR,  Proc.  Zool.  Inst.  82:  3-22. 

Fennah,  R  C  ,  and  J.  C  M.  Carvalho.  1963.  On  a  new 
genus  and  species  of  fulgorid  from  Brazil  (Ho- 


moptera, Lystrini).  An.  Acad.  Brasileira  Cienc. 
35:  459-461. 

Hemming,  F  1955.  Validation,  under  the  plenary  pow- 
ers, of  the  generic  name:  "Fulgora"  Linnaeus, 
1767  (Class  Insecta,  Order  Hemiptera)  and  desig- 
nation for  the  genus  so  named  of  a  type  species  in 
harmony  with  current  nomenclatorial  practice. 
Opinion  322.  International  Commission  on  Zo- 
ological Nomenclature.  Opin.  &  Declar.  ICZN. 
9(13):  185-208. 

Horn.  W.,  and  I  Kahle  1935.  Uber  entomologische 
Sammlungen  (Ein  Beitrag  zur  Geschichte  der  En- 
tomo-Museologie).  Ent.  Beih.  Berlin-Dahlem  2: 
1-540. 

Janzen.  D  H  ,  andC  L  Hogue  1983.  Fulgora  late rnaria 
(machacha,  peanut-head  bug,  lantern  fly.)  In: 
Costa  Rican  natural  history.  University  of  Chicago 
Press,  Chicago,  Illinois,  xi  +  816  pp. 

Johnson,  L.  K  ,  and  R  B  Foster  1986.  Associations  of 
large  Homoptera  (Fulgoridae  and  Cicadidae)  and 
trees  in  a  tropical  forest.  J.  Kansas  Entomol.  Soc. 
59:  41.5-422. 

Knull,  D  J  .  and  J.  N  Knull  1947.  Two  new  Amycle 
(Homoptera:  Fulgoridae).  Ann.  Entomol.  Soc. 
Amer.  60:  397-400. 

Kramer,  J  P  1978.  Taxonomic  study  of  the  American 
planthopper  genus  Cyrpoptus  (Homoptera:  Ful- 
goroidea:  Fulgoridae).  Proc.  Biol.  Soc.  Washing- 
ton 91:  303-335. 

Lallemand,  V  1956.  Contribution  a  I'etude  des  Fulgori- 
dae (Hemiptera)  (Ire.  note).  Bull.  Inst.  R.  Sci. 
Nat.  Belgique.  32:  1-7. 

1959.  Description  de  nouvelles  especes  de  Ful- 

gorides  d'Asie  et  d'Afrique.  Zool.  Meded.  36: 
267-272  [no  African  species,  five  New  World]. 

1960.    De  quibusdam   Fulgoris.    Ent.    Mitt.    24: 

101-107. 

1963.    Nouvelles    especes   d'Homopteres.    Bull. 

Inst.  R.  Sci.  Nat.  Belgique.  39:  1-5. 

1966.  Fulgorides  nouveaux  du  Musee  Zoologique 

de  Berlin  et  de  ma  collection.  Bull.  Recher. 
Agron.  Gembloux.  N.  S.  1:  51-54. 

Metcalf,  Z  p.  1947  General  catalogue  of  the  Hemiptera. 
Ease.  IV.  Fulgoroidea,  Part  9.  Fulgoridae.  Smith 
College,  Northampton,  Massachusetts.  280  pp. 

Nast,  J  1950.  New  genera  and  species  of  Neotropical 
Fulgoridae  in  the  collection  of  the  British  Museum 
(Homoptera).  Ann.  Mus.  Zool.  Polonici  14:  167- 
175. 

1951.  Some  remarks  on  Neotropical  Fulgoridae 

with  descriptions  of  new  genera  and  species  (Ho- 
moptera). Ann.  Mus.  Zool.  Polonici  14:  267-279, 
3  plates. 

O  Brien,  L.  B  1985.  New  synonymies  and  combinations 
in  New  World  Fulgoroidea  (Achilidae,  Del- 
phacidae,  Flatidae,  Fulgoridae:  Homoptera). 
Ann.  Entomol.  Soc.  Amer.  78:  657-662. 

1986.  Five  new  species  of  Fulgoroidea  (Ho- 
moptera) from  the  western  United  States  and 
Mexico.  Southwestern  Entomol.  11:67-74. 

O  Brien,  L  B  ,  and  S  W  Wilson  1985.  Planthopper 
systematics  and  external  morphology.  Pages 
61-102  in  L.  R.  Nault  and  J.  G.  Rodriguez,  eds., 
The  leafhoppers  and  planthoppers.  Wiley  and 
Sons,  Inc. 


1988 


O'BRIEN:  New  World  Ful(;oridae 


169 


RiDOUT.  B  V  1983.  Structure,  form  and  function  of  the 
lantern  fly  head  process  {Fulgora  laternaria  Linn.) 
Unpublished  dissertation,  Birheck  C'ollege. 

Wilson,  S  W  ,  and  L  B  O  Brikn  1986.  Descriptions  of 
nymphs  of  Itzalana  suhinaculata  Schmidt  (Ho- 
moptera:  Fulgoridae),  a  species  new  to  the  United 
States.  Great  Lakes  Entomol.  19:  101-105. 

1987.  A  survey  of  planthopper  pests  of  economi- 
cally important  plants  (Homoptera:  Fulgoroidea). 
Pages  343-360  in  Proc.  Second  Annual  Workshop 
on  Leafhoppers  and  Planthoppers  of  Economic 
Importance,  held  Provo,  Utah,  USA,  28  July-1 
August  1986.  Commonwealth  Inst.  Entomol., 
London. 

ZiMSEN,  E  1964.  The  type  material  of  1.  C.  Fabricius. 
Munksgaard,  Copenhagen.  280  pp. 


List  OF  Species 

(Countries  are  listed  from  south  to  north,  west  before 
east.) 


imitatrix  Ossiannilsson 


Venezuela,  Panama, 
Costa  Rica,  Belize,  Mexico 
Peru 


Amerzanna  O  Brien,  new  genus 


peruana  O'Brien 

Peru 

Amycle  Stal 

aniahilis  (Westwood) 

Mexico 

brevis  O'Brien 

Mexico 

grandis  O  Brien 

Mexico 

mankinsi  O  Brien 

Honduras 

pinyonae  Knull  &  Knull 

USA:CA,  AZ,  NM 

saxatilis  Van  Duzee 

USA:CA 

sodalis  Stal 

Mexico,  El  Salvador 

tumacacoriae  Knull  &  Knull 

USA:  TX,  AZ 

vernalis  Manaee 

USA:  NC,  LA 

Aphrodisias  Kirkaldy 

cacica  (Stal) 

Mexico 

shaman  O'Brien 

Mexico 

Artacie  Stal 

dufourii  (Signoret), 

resurrected  name 

Fr.  Guiana,  Guyana 

haemoptera  (Perty) 

Brazil 

Cathedra  Kirkaldy 

serrata  (Fabricius) 

Bolivia,  Peru,  Brazil, 

Fr.  Guiana,  Suriname, 
Colombia,  Panama,  Costa  Rica 
Chilobia  Stal 
=  Ecuadoria  Distant,  new  synonymy 

cinxia  Stal  Venezuela 

dichopteroides  (Distant),  (Ecuadoria), 

new  combination  Ecuador 

silena  Stal  Ecuador 

sinaragdina  (Walker)  Venezuela 

Copidocephala  Stal 

guttata  (White)  Panama,  Costa  Rica, 

Guatemala,  Mexico 
merula  Distant  Colombia 

=  melanoptera  (Schmidt),  new  synonymy 
viridiguttata  Stal  Columbia,  Panama,  Costa  Rica, 

Honduras,  Mexico 
=  ornanda  (Distant),  new  synonymy 
Diareusa  Walker 

annularis  (Olivier)  Peru,  Brazil,  Fr.  Guiana, 

Suriname,  Guyana 
conspersa  Schmidt  Ecuador,  Colombia 

=  dahli  Ossiannilsson,  new  synonymy 


keinneri  Ossiannilsson 

Ecuadoria,  sec  Chilobia 

Encho})hora  Spinola 

atonuiria  Brancsik,  see  Cornelia, 

teste  Lallcmand  1959:88 
dufourii  Signoret,  see  Artacie, 

teste  O'Brien  1985:661 
ensifera  (Germar),  to  inccrtae  sedis 
maculata  O'Brien  Peru 

nigroinaculata  Distant  Bolivia,  Peru,  Ecuador, 

Brazil 
=  nigrolimbata  Lallcmand 

pallidipunctata  Lallcmand  Bolivia,  Peru 

prasina  Gerstaecker  Bolivia,  Peru,  Panama, 

Costa  Rica 
pyrrhocrypta  Walker  Brazil,  Guyana,  Venezuela 

recurva  (Olivier)  Suriname,  Brazil 

=  bohe7nani  Stal,  new  synonymy 

rosacea  Distant  Panama,  Costa  Rica,  Nicaragua 

sanguinea  Distant  Ecuador,  Colombia,  Panama, 

Costa  Rica,  Nicaragua,  Guatemala 
=florens  Distant,  new  synonymy 
=  longirostris  Distant,  new  synonymy 
stillifera  (Stal)  Panama,  Costa  Rica,  Honduras, 

Guyana,  Belize,  Mexico 
subviridis  Distant  Panama,  Costa  Rica 

=  subviridisv ar.  distanti  Metcalf 

tuba  (Germar)  Brazil 

tuberctdata  (Olivier)       Brazil,  Fr.  Guiana,  Suriname, 

Venezuela 
=  tuberculata  vm.  fuscoinaculata  Lallcmand 
=  parvipennis  Walker,  new  synonymy 
unifonnis  O  Brien  Peru 

viridipennis  Spinola  Brazil 

=  eminenta  Schmidt,  new  synonymy 

Enhydria  Walker 

=  Ulubra  Stal,  new  synonymy 

cicadina  Gerstaecker  Brazil 

ferruginea  (Walker)  to  Hydriena  (Dictyophariadae) 
longicornuta  Lallcmand  Brazil,  Trinidad 

tessellata  (Walker)  Peru,  Brazil 

=  brachialis  (Stal),  new  synonymy 

Fulgora  L.  (New  synonymies  in  this  genus  are  attributed 

to  B.  V.  Ridout) 

caerulescens  Olivier,  to  incertae  sedis 
castresii  Guerin-Meneville         Peru,  Panama,  Mexico 
cearensis  (Fonseca)  Brazil,  Trinidad 

crocodilia  Brailovsky  &  Beutelspacher  Mexico 

graciliceps  Blanchard  Peru,  Bolivia,  Brazil 

=  orthocephala  (Fonseca),  new  synonymy,  Ridout 
lampetis  Burmeister  Bolivia,  Peru,  Brazil, 

Panama,  Costa  Rica,  Nicaragua,  Honduras 
laternaria  (Linne)  Brazil,  Panama,  Honduras, 

Mexico 
=  servillei  Spinola,  new  synonymy,  Ridout 
lucifera  Germar  Argentina,  Bolivia,  Brazil 

riograndensis  (Fonseca)  Brazil 

Odontoptera  Carreno 

carrenoi  Signoret  Brazil,  Fr.  Guiana,  Guyana, 

Panama,  Costa  Rica,  Guatemala,  Mexico 

spectabilis  Carreno  Brazil 

Phrictus  Spinola 

auromaculatus  Distant  Bolivia,  Peru,  Ecuador, 

Brazil 


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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 


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No.  12 


Material 

The  data  on  which  this  study  is  based  were 
taken  from  dried  male  specimens  (OTUs)  of  32 
Cicada  orni  and  32  C.  harhara  lusitanica 
(Table  1).  These  samples  were  mostly  taken 
by  the  author  in  Portugal:  all  32  males  of  orni 
were  collected  in  central  Portugal;  an  equal 
number  of  males  of  harhara  hisitanica  were 
taken  in  several  areas  of  Algarve  (the  southern 
province  of  Portugal),  where  the  species  ap- 
pears to  be  particularly  common,  with  the 
exception  of  two  specimens  only  that  were 
collected  in  Sesimbra  (south  of  Lisbon).  The 
localities  and  sample  sizes  are  C.  orni:  Albu- 
ritel,  Vila  Nova  de  Ourem  (n  =  32);  C.  har- 
hara hisitanica:  Carvoeiro  (n  =  25),  Praia  da 
Rocha  (n  =  4),  Serra  de  Monchique  (n  =  1), 
and  Sesimbra  (n  =  2). 

Methods 

Measurements  and  Counts 

Thirty-seven  of  the  40  characters  were  mea- 
surements; the  remaining  3  were  counts. 
Measurements  were  made  using  a  Wild  M3 
microscope  with  a  graduated  eyepiece  and 
were  taken  as  described  in  Table  2  or  as  illus- 
trated in  Figures  1-9.  Of  these  40  characters, 
33  refer  to  external  morphology  and  the  re- 
maining 7  to  male  genitalia. 

Data  Analysis 

Data  processing  was  carried  out  on  the 
CDC  6500  computer  at  the  Imperial  College 
Computer  Center  (University  of  London)  us- 
ing two  multivariate  statistical  programs  de- 
veloped by  Prof.  R.  G.  Davies  (Department  of 
Pure  and  Applied  Biology,  Imperial  College) 
for  cluster  analysis  and  ordination  (Quartan 
and  Davies  1983,  1985). 

In  most  analyses,  characters  were  standard- 
ized by  expressing  each  state  as  a  deviation 
from  the  mean  in  standard  deviation  units. 

For  Q-mode  analysis,  taxonomic  distances 
as  well  as  product-moment  correlations  were 
found  and  structured  by  the  WPGMA  method 
of  cluster  analysis  (Weighted  Pair  Group 
Method  with  Arithmetic  Averaging).  Pheno- 
grams,  expressing  the  phenetic  relationships 
among  the  OTUs  in  a  hierarchy  of  increasingly 
larger  clusters,  were  thus  obtained  (Figs. 
10-16). 

For  R-mode  analysis,  character  correlations 


Table  L    List  of  specimens  (males)  oi  Cicada  orni  Lin- 
naeus and  of  C.  harhara  lusitanica  Boulard  investigated 

(OTUs). 

OTUs      Locality,  date  of  capture,  and  collector 

Cicada  orni 

1-15      Alburitel,  10.viii.l979,  J.  A.  Quartau 
16-32      Alburitel,  vii.  1971,  J.  A.  Quartau 

C.  harhara  hisitanica 

33-34  Carvoeiro,  14.viii.l966,  P.  D.  Rodrigues 

35-38  Praia  da  Rocha,  IT.viii.  1973,  J.  A.  Quartau 

39-40  Carvoeiro,  14. vii.  1978,  J.  A.  Quartau 

41-47  Carvoeiro,  30. vii.  1978,  J.  A,  Quartau 

48-55  Carvoeiro,  28. vii.  1978,  J.  A.  Quartau 

56-57  Carvoeiro,  9.viii-10.i.x.l980,  L.  Mendes 

58-59  Sesimbra,  2.viii.  1980,  J.  A.  Quartau 

60  Carvoeiro,  31. vii.  1978,  J.  A.  Quartau 

61-62  Carvoeiro,  18. vii.  1978,  J.  A.  Quartau 

63  Monchique,  2.i.x.l971,  F.  Carvalho 

64  Carvoeiro,  24.viii.1981,  J.  A.  Quartau 


based  on  data  standardized  by  OTUs  were 
subjected  to  principal  component  analysis 
(PGA).  This  ordination  method  transforms  the 
original  characters,  generally  continuous,  cor- 
related characters,  into  a  suite  of  uncorre- 
cted, composite  variables — the  principal 
components  (principal  axes).  In  addition  to 
being  mutually  independent,  these  compo- 
nents account  for  maximum  variance  as  fol- 
lows: the  variance  along  the  first  axis  (i.e.,  the 
corresponding  eigenvalue)  is  the  maximum 
possible.  The  second  axis  describes  the  next 
largest  variance  orthogonal  to  (uncorrected 
with)  the  first.  The  third  axis  follows  similarly 
but  is  independent  of  both  first  two  axes,  and 
so  on,  for  as  many  axes  as  one  wishes  to  extract 
(e.g.,  Gibson  et  al.  1984).  A  transposed  matrix 
of  the  character  loadings  was  post-multiplied 
by  the  standardized  data  matrix  to  yield  a 
matrix  of  OTU  projections  in  the  principal 
component  space.  Two-dimensional  ordina- 
tion diagrams  of  the  representations  of  the  two 
species,  together  with  the  character  loadings 
(scaled  eigenvectors),  were  thus  obtained 
(Figs.  17-18,  Table  3). 

Results 

Phenograms 

The  seven  phenograms  resulting  from  vari- 
ous analyses  based  on  all  characters,  on  the 
genitalia  only,  or  on  the  external  characters 
alone  are  shown  in  Figures  10-16.  The 
WPGMA  clustering  technique  was  followed 


1988 


QuARTAU:  Cicada  Analysis 


173 


Table   2.    Description   of  characters:    measurements 
and  counts  (terminology  mostly  follows  Myers  [1928]). 


Character  No. 


Description 


1.  Overall  length  measured  from  tip  of  crown  to  apical 
margin  of  the  right  forewing  with  the  latter  in  posi- 
tion of  rest  alongside  the  body  (Fig.  1). 

2.  Length  of  crown  measured  along  a  medial  line  pass- 
ing through  the  median  ocellus  (Fig.  I). 

3.  Minimum  distance  between  the  ocular  sutures  mea- 
sured along  the  paired  ocelli  (Fig.  2). 

4.  Medial  length  of  frons  measured  dorsally  as  indicated 
(Fig.  2). 

5.  Medial  length  of  pronotum  measured  dorsally  as  in- 
dicated (Fig.  1). 

6.  Medial  length  of  mesonotum  measured  dorsally  from 
anterior  margin  to  posterior  margin  of  cruciform  ele- 
vation or  scutellum  (Fig.  1). 

7.  Width  of  pronotum  measured  at  the  level  of  anterior 
lateral  margins  (Fig.  1). 

8.  Width  of  pronotum  measured  at  the  level  of  postero- 
lateral margins  (Fig.  1). 

9.  Width  of  crown  measured  at  the  level  of  median 
ocellus  and  as  indicated  (Fig.  2). 

10.  Inner  distance  between  the  paired  ocelli  (Fig.  2). 

11.  Distance  between  the  right  paired  ocelli  and  the 
right  ocular  suture  as  indicated  (Fig.  2). 

12.  Distance  between  the  base  of  the  left  antenna  and  the 
left  ocular  suture  as  indicated  (Fig.  3). 

13.  Inner  distance  between  the  base  of  antennae  (Fig.  3). 

14.  Length  of  frons  as  illustrated  (Fig.  3). 

15.  Length  of  clypeus  as  illustrated  (Fig.  3). 

16.  Length  of  the  exposed  part  of  beak. 

17.  Length  of  dorsal  margin  of  the  left  fore  femur  as 
illustrated  (Fig.  5). 

18.  Length  of  ventral  margin  of  the  left  fore  femur  as 
illustrated  (Fig.  5). 


19.  Length  of  basal  spine  in  ventral  margin  of  left  fore 
femur  as  illustrated  (Fig.  5). 

20.  Length  of  apical  spine  in  ventral  margin  of  left  fore 
femur  as  illustrated  (Fig.  5). 

21.  Distance  between  tips  of  the  apical  and  basal  spines 
in  the  ventral  margin  of  the  left  fore  femur  as  illus- 
trated (Fig.  5). 

22.  Distance  from  anterior  right  corner  to  posterior  left 
corner  of  left  operculum  as  illustrated  (Fig.  6). 

23.  Distance  from  anterior  left  corner  to  posterior  right 
corner  of  left  operculum  as  illustrated  (Fig.  6). 

24.  Length  of  right  forewing  as  illustrated  (Fig.  1). 

25.  Greatest  width  of  right  forewing  as  illustrated  (Fig. 
4). 

26.  Length  of  subcostal  cell  ("gancho"  cell  of  G6mez- 
Menor  1957)  in  right  forewing  (Fig.  4). 

27.  Length  of  anterior  margin  of  basal  cell  in  right 
forewing  (Fig.  4). 

28.  Length  of  posterior  margin  of  basal  cell  in  right 
forewing  (Fig.  4). 

29.  Maximum  width  of  basal  cell  in  right  forewing. 

30.  Minimum  width  of  basal  cell  in  right  forewing. 

31.  Number  of  apical  cells  in  right  forewing. 

32.  Number  of  cells  other  than  apicals  of  right  forewing. 

33.  Number  of  spots  in  cross-veins  of  right  forewing. 

34.  Length  of  pygofer  in  lateral  view  as  indicated  (Fig.  7). 

35.  Overall  length  of  tenth  abdominal  segment  as  indi- 
cated (Fig.  7). 

36.  Overall  length  of  appendages  of  tenth  abdominal 
segment  as  indicated  (Fig.  7). 

37.  Distance  in  basal  curvature  of  shaft  of  aedeagus  as 
indicated  (Fig.  9). 

38.  Width  of  shaft  of  aedeagus  as  indicated  (Fig.  9). 

39.  Width  of  shaft  of  aedeagus  in  area  of  curvature  as 
illustrated  (Fig.  9). 

40.  Medial  length  of  eighth  sternite  or  hypandrium  (Fig. 


in  all;  and  Pearson's  product-moment  coeffi- 
cient and  the  taxonomic  distance  coefficient 
were  used  as  measures  of  taxonomic  proxim- 
ity. 

(a)  Genital  analyses 

Figures  10  and  11  illustrate  a  correlation 
and  a  distance  phenogram,  respectively,  both 
based  on  the  seven  standardized  variables  of 
the  male  genitalia.  Both  analyses,  notwith- 
standing their  being  based  on  a  small  number 
of  variables,  resulted  in  two  main  clusters, 
one  with  C.  orni  and  the  other  with  C.  bar- 
bara  lusitanica.  However,  in  the  former  phe- 
nogram, the  cluster  oi  barbara  lusitanica  in- 
cludes one  specimen  of  orni  (No.  13). 

(b)  External  characters 

These  analyses  resulted  in  the  production 
of  the  phenograms  depicted  in  Figures  12  and 
13.  Both  were  based  on  standardized  data,  but 
only  the  correlation  coefficient  succeeded  in 


giving  an  almost  complete  separation  of  the 
two  species  of  cicadas.  In  fact,  OTUs  were 
grouped  into  two  main  clusters  as  in  the  geni- 
tal analyses,  but  specimen  No.  19  belonging 
to  C.  orni  appeared  misplaced  within  C.  bar- 
bara lusitanica  (Fig.  12).  On  the  contrary,  the 
distance  phenogram  provided  much  less  satis- 
factory results  than  the  previous  analysis, 
since  each  of  the  two  major  clusters  incorpo- 
rates elements  of  both  species  of  cicadas  (Fig. 
13). 

(c)  Combined  characters 

The  phenograms  of  this  group  of  analyses, 
involving  all  40  characters  combined,  are  il- 
lustrated in  Figures  14-16. 

Considering  the  phenograms  based  on 
standardized  data  (Figs.  14,  16),  it  is  clear  that 
the  correlation  phenogram  gave  a  much  bet- 
ter distinction  between  the  two  species  than 
the  distance  analysis.  In  fact,  the  latter  (Fig. 
16)  clustered  six  specimens  of  C.  orni  with  C. 


174 


Great  Basin  Naturalist  Memoirs 


No.  12 


1                          10                           'i 
i           ' 

1                       1      9      '                         1 

1    '         1     :     n     1  : 

Figs  1-6.  Diagrams  of  a  male  Cicada  orni  Linnaeus  illustrating  most  of  the  measurements  taken;  1,  body,  dorsal 
view;  2,  head,  dorsal  view;  3,  head,  anterior  view;  4,  right  forewing,  dorsal  view;  .5,  left  foreleg,  mner  view;  6,  lett 
operculum,  ventral  view. 


1988 


QuARTAU:  Cicada  Analysis 


175 


Figs.  7-9.  As  in  Figures  1-6:  7,  pygofer  and  tenth  abdominal  segment,  ventral  view;  8,  eighth  stemite  or 
hypandrium,  ventral  view;  9,  aedeagus,  lateral  view. 


barbara  lusitanica.  Even  when  the  data  were 
unstandardized,  correlations  gave  a  good  pic- 
ture of  the  relationships  between  these  two 
species  (Fig.  15). 

Principal  Component  Analysis 
This  analysis  involved  all  40  characters  and 
was  computed  from  a  between-character  cor- 
relation matrix  based  on  data  standardized  by 
OTUs. 

As  in  similar  analyses  carried  out  with 
leafhoppers  of  the  genus  Batracomorphus 
(Quartan  1983),  slightly  more  than  half  (54%) 
of  the  total  variation  in  the  study  was  ex- 
plained by  the  first  three  axes. 

The  first  component  accounted  for  38.90% 
of  the  variation  in  the  data  and  is  interpreted 
as  a  contrast  between  the  lengths  of  the 
pygofer,  tenth  abdominal  segment,  or  ap- 
pendages of  the  latter  and  the  width  of  the 
shaft  of  the  aedeagus.  It  does  not  represent 
overall  size  as  commonly  is  the  case,  since 
many  of  the  characters  (Table  3)  are  not  posi- 
tively correlated  with  it  (e.g.,  Jolicoeur  and 
Mosimann  1960,  Blackith  and  Reyment  1971, 
Baker  1980,  Gibson  et  al.  1984,  Shea  1985). 
In  fact,  it  must  represent  both  size  and  shape 
as  has  been  pointed  out  by  several  authors 


(Mosimann  1970,  Oxnard  1978,  Humphries  et 
al.  1981).  A  complete  separation  of  C.  orm  and 
C.  barbara  lusitanica  was  given  by  the  dis- 
crimination afforded  by  this  axis,  which  is 
probably  close  to  the  orientation  of  the  opti- 
mum discriminant  function.  The  characters 
loading  most  heavily  on  this  component 
(Table  3)  are  therefore  of  considerable  taxo- 
nomic  interest,  since  they  are  diagnostic  for 
this  pair  of  species.  The  highest  negative 
scores,  in  decreasing  order,  were  for  charac- 
ters numbered  35  (length  of  tenth  abdominal 
segment),  34  (length  of  pygofer),  and  36 
(length  of  appendages  of  tenth  abdominal  seg- 
ment). The  highest  positive  score  was  for 
character  numbered  38  (width  of  shaft  of 
aedeagus). 

The  second  principal  component  accounted 
for  8.51%  of  the  total  variation  and  was  inter- 
preted as  a  contrast  between  the  number  of 
spots  in  cross-veins  of  the  wings  and  the  width 
of  the  crown.  It  was  most  heavily  loaded,  neg- 
atively and  positively,  on  characters  num- 
bered 33  and  9,  respectively. 

The  third  principal  component  accounted 
for  6.61%  of  the  total  variation  and  was  inter- 
preted as  a  factor  resulting  from  the  lengths  of 


176 


Great  Basin  Naturalist  Memoirs 


No.  12 


Table  3.  Eigenvector  matrix  (character  loadings)  in  a 
principal  component  analysis  of  the  matrix  of  correlations 
among  the  40  variables  (data  standardized  by  OTUs.) 


Variables 

Scaled 

eigenvectors 

I 

II 

III 

1 

0.296 

-0.1.58 

0.441 

2 

0.404 

0.123 

0. 155 

3 

0.531 

0.557 

-0.174 

4 

0.843 

-0.160 

0.126 

5 

0.492 

0.300 

-0.119 

6 

0.037 

-0.096 

0.280 

7 

0.567 

0.280 

-0.069 

8 

0.314 

0.365 

-0.131 

9 

0.336 

0.633 

-0.1,33 

10 

0.738 

-0.237 

-0.000 

11 

0.687 

0.155 

-0.200 

12 

0.859 

-0.127 

-0.142 

13 

0.313 

0.423 

0.414 

14 

0.117 

0.422 

0.306 

15 

-0.604 

-0.196 

-0.120 

16 

-0.769 

0.104 

-0.192 

17 

-0.316 

0.338 

0,524 

18 

-0.667 

0.174 

0.,375 

19 

0.565 

0.2.52 

-0.091 

20 

0.531 

-0.066 

0.111 

21 

-0.132 

0.019 

0.257 

22 

0.784 

0.203 

0.160 

23 

0.847 

0.112 

0.000 

24 

0.524 

-0,340 

0.060 

25 

0.683 

-0.104 

0.154 

26 

0.187 

-0.207 

-0.412 

27 

-0.334 

0.126 

-0,561 

28 

0. 133 

0.117 

-0.,549 

29 

-0.011 

-0.207 

0.471 

30 

0.129 

-0.287 

0.511 

31 

0.818 

-0.490 

-0.049 

32 

0.818 

-0.479 

-0.059 

33 

0.266 

-0.745 

-0.163 

34 

-0.952 

-0.011 

-0.040 

35 

-0.958 

-0.153 

-O.OOI 

36 

-0.944 

-0.180 

0.043 

37 

-0.869 

-0.250 

0.028 

38 

0.942 

-0.065 

-0.023 

39 

0.892 

-0.249 

0.024 

40 

-0.858 

-0.125 

-0.090 

Latent  roots 

15.559 

3.406 

2.644 

Percentage  of 

component    38.898 

8.514 

6.609 

variation 

cumulative    38.898 

47.412 

,54.021 

the  anterior  and  posterior  margins  of  the  basal 
cell  of  the  wings.  It  was  most  heavily  loaded 
(negative  scores)  on  characters  numbered  27 
and  28. 

Neither  the  second  nor  the  third  axis  leads 
to  a  separation  of  the  two  species  of  cicadas. 
Figures  17  and  18  are  two-dimensional  views 
of  the  relationships  among  specimens  of  both 
species  in  the  space  determined  by  compo- 
nent I  combined  with  component  II  and  by 


component  I  with  component  III,  respec- 
tively. It  is  clear  that  these  two  plots  gave  a 
good  separation  between  C.  orni  and  C.  bar- 
bara  lusitanica.  The  plot  combining  axes  II 
and  III  did  not,  however,  succeed  in  giving  a 
correct  assignment  of  the  two  species  and 
therefore  was  not  illustrated  here. 

Conclusions 

Cluster  analysis  and  principal  component 
analysis  are  two  of  the  methods  most  com- 
monly used  for  recognition  of  group  structure 
in  numerical  ta.xonomy  (e.g.,  Quartau  1987). 
This  study  suggests  that  the  two  general 
methods  are  also  very  useful  for  discriminat- 
ing between  pairs  of  closely  related  species. 
In  fact,  because  of  the  uniformity  of  the  gen- 
eral clustering  pattern  that  resulted  from  the 
application  of  both  methods,  it  is  clear  that 
two  distinct  species  exist,  a  fact  also  indicated 
by  behavioral  data. 

However,  concerning  cluster  analysis,  it  is 
worthwhile  noting  that  the  hierarchical  struc- 
ture within  each  of  the  two  major  groups  of 
OTUs  differs  a  good  deal  from  one  particular 
method  to  another.  Moreover,  it  appeared 
that  correlations  were  more  effective  than  tax- 
onomic  distances  in  describing  relationships 
between  the  two  cicadas,  a  result  in  keeping 
with  Boyce  (1964)  or  Cheetam  (1968),  for  in- 
stance. It  is  interesting  to  note,  in  this  regard, 
that  such  finding  is  in  disagreement  with  a 
similar  study  carried  out  with  leallioppers 
(Quartau  and  Davies  1983)  or  with  results 
based  on  other  groups  (e.g..  Smith  1972).  Fi- 
nally, the  failure  of  the  cluster  analysis  using 
distances  with  nongenital  characters  shows 
that  the  use  of  such  techniques  needs  to  be 
undertaken  with  care  and  that  it  might  be  best 
to  use  a  consensus  of  several  clustering  tech- 
niques when  applying  numerical  methods  to  a 
novel  taxonomic  situation. 

Principal  component  analysis  succeeded  in 
giving  a  good  distinction  of  the  two  species 
along  axis  I,  in  spite  of  its  accounting  for  only 
.38.90%  of  the  total  variation.  This  analysis  also 
showed  that  the  main  distinguishing  charac- 
ters between  C.  orni  and  C.  barbara  lusitan- 
ica are  the  following:  lengths  of  the  pygofer, 
the  tenth  abdominal  segment,  and  the  ap- 
pendages of  the  latter  (which  are  smaller  in  C. 
barbara  lusitanica.  Figs.  19,  22),  as  well  as 
the  width  of  the  shaft  of  the  aedeagus  (which  is 


1988 


QuARTAU;  Cicada  Analysis 


177 


-0.8 


Correlations 
Data  standardized 
Genital  characters 

D  C.  orni 

•  C.    barbara    lusitanica 


10 


it 


Distances 

Data  standardized 

Genital  characters 

DC.  orni 

•  C.    barbara   lusitanica 


11 


Figs.  10-11.  10,  correlation  phenogram  based  on  the  seven  genital  characters  with  standardized  data;  11,  distance 
phenogram  based  on  the  seven  genital  characters  with  standardized  data. 


thinner  in  orni.  Figs.  20,  23).  Moreover,  de- 
tailed examination  of  the  male  genitalia  also 
showed  that  the  dorsal  spine  of  the  pygofer  is 
smaller  in  barbara  lusitanica  than  in  orni 
(Figs.  19,"  22). 

Acknowledgments 

1  thank  Prof.  R.  G.  Davies,  Department  of 
Pure  and  Applied  Biology,  Imperial  College, 
for  having  made  his  computer  programs  avail- 
able for  my  use  as  well  as  for  his  critical  com- 
ments on  the  manuscript.  Prof.  R.  M.  Ander- 
son is  also  acknowledged  for  having  granted 


working  facilities  at  the  Imperial  College 
(University  of  London).  1  am  also  grateful  to 
my  colleague  Dr.  M.  L.  Mathias  for  her  help 
with  the  preparation  of  most  phenograms,  and 
to  Miss  A.  R.  Cruz  for  help  with  other  illustra- 
tions and  the  measurements.  For  enabling  me 
to  publish  this  study  in  honor  of  Prof.  P.  Oman 
(Oregon  State  University,  USA),  who  has 
been  a  source  of  encouragement  since  the 
beginning  of  my  taxonomic  studies  on 
leafhoppers,  I  also  acknowledge  Dr.  M.  W. 
Nielson  (Brigham  Young  University,  USA) 
and  Dr.  H.  D.  Blocker  (Kansas  State  Univer- 
sity, USA). 


178 


Great  Basin  Naturalist  Memoirs 


No.  12 


Correlations 

Data  standardized 

Cxt.  Morphology  characters 
D  C.  orni 
•  F.  barbara  lusitanica 


Figs.  12-13.  12,  correlation  phenogram  based  on  the  33  external  morphological  characters  with  standardized  data; 
13,  distance  phenogram  based  on  the  33  external  morphological  characters  with  standardized  data. 


This  paper  was  completed  at  Imperial  Col- 
lege (University  of  London)  under  an  INIC 
(Ministerio  da  Educagao,  Lisbon)  sabbatical 
fellowship. 

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Gibson,  A  R  ,  A  J  Baker,  and  A  Moeed  1984.  Morpho- 
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common  myna  (Acridotheres  tristis):  an  applica- 
tion of  the  jackknife  to  principal  component  analy- 
sis. Systematic  Zoology  33(4):  408-421. 

Gomez-Menor,  O  J  1957.  MonografiadeCicadidos  (Ho- 
moptera) de  Esparia.  Academia  de  Madrid, 
Madrid. 


1988 


QUARTAU:  CiCADA  ANALYSIS 


179 


0.0 


0.993 


0.998 


Correlat ions 
Data   standardized 
All    characters 

DC.    orni 

o  C.    barbara    lusitanica 


14 


T 31 


Correlat ions 

Data   unstandardized 

All    ctiaracters 
n  £.    orni 
o  C.    barbara    lusitanica 


Figs.  14-15.  14,  correlation  phenogram  based  on  all  40  combined  characters  with  standardized  data;  15,  correlation 
phenogram  based  on  all  40  combined  characters  with  unstandardized  data. 


Humphries,  J  M  ,  F  L  Bookstein,  B  Chernoff,  G  R 
Smith,  R  L  Elder,  and  S  G  Poss  1981.  Multi- 
variate discrimination  by  shape  in  relation  to  size. 
Systematic  Zoology  30:  291-308. 

JOLICOEUR,  P  ,  AND  J  E  MosiMANN  1960.  Size  and  shape 
variation  in  the  painted  turtle.  A  principal  compo- 
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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 
(Homoptera).  Proceedings  of  the  Zoological  Soci- 
ety of  London  1928:  265-472. 

OXNARD,  C  E  1978.  One  biologist's  view  of  morphomet- 
ries. Annual  Review  of  Ecology  and  Systematics  9: 
219-241. 


QuARTAU,  J  A  1983.  A  comparison  of  different  numerical 
methods  applied  to  the  taxonomy  of  Batracomor- 
phus  Lewis  (Insecta,  Homoptera,  Cicadellidae, 
lassinae):  ordination  techniques.  Journal  of  the 
Entomological  Society  of  Southern  Africa  46(1): 
9-35. 

1987.  Ordination  vs.  cluster  analysis  in  the  numer- 
ical taxonomy  of  some  leafhoppers  (Homoptera, 
Cicadellidae).  Pages  207-215  in  M.  R.  Wilson  and 
L.  R.  Nault,  eds..  Proceedings  of  the  2nd  Interna- 
tional Workshop  on  Leafhoppers  and  Planthop- 
pers  of  Economic  Importance,  CLE.,  London. 

QuARTAU,  J  A  ,  AND  R  G  Davies  1983.  A  comparison  of 
different  numerical  methods  applied  to  the  taxon- 
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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 
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Smith,  J  D  1972.  Systematics  of  the  chiropteran  family 
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Temple,  J  T  1968.  The  Lower  Llandovery  (Silurian)  bra- 
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tographical  Society  521:  1-58. 


1988 


QuARTAU:  Cicada  Analysis 


181 


Figs.  19-24.  Diagrams  of  the  male  genitalia  oiCicada  orni  (Figs.  19-21)  and  of  C.  barbara  lusitanica  (Figs.  22-24): 
19,  22,  pygofer  and  tenth  abdominal  segment,  lateral  view;  20,  23,  aedeagus,  lateral  view;  21,  24,  eighth  sternite  or 
hypandrium,  ventral  view  (scale  =  0.5  mm). 


REVISION  OF  THE  NIRVANINAE  (HOMOPTERA:  CICADELLIDAE) 
OF  THE  INDIAN  SUBCONTINENT 

C.  A.  Viraktamath'  andC.  S.  Wesley^ 

Abstract — ^Three  tribes  of  Nirvaninae,  namely,  Balbillini  (two  genera  and  three  species),  Nirvanini  (five  genera  and 
26  species),  and  Occinirvanini  (one  genus  and  one  species),  occur  on  the  Indian  subcontinent.  The  tribes,  genera,  and 
species  are  redescribed,  illustrated,  and  keyed.  New  ta.xa  recognized  are  BalbiUus  indicus  n.  sp.  (India;  Kerala), 
Stenotortor  stibhimalaya  n.  sp.  (India:  West  Bengal),  Kana  bispinosa  n.  sp.  (India:  Tamil  Nadu),  K.  nigropicta  n.  sp. 
(India:  Kerala),  Sophonia  bakeri  n.  sp.  (India:  Karnataka,  Kerala,  Tamil  Nadu,  Uttar  Pradesh;  Nepal),  S.  bifida  n.  sp. 
(India:  West  Bengal),  S.  complexa  n.  sp.  (India;  Meghalaya),  S.  compUcata  n.  sp.  (India:  Mizoram,  Meghalaya),  S. 
keralica  n.  sp.  (India;  Kerala),  Nirvana  peculiarisn.  sp.  (India:  Mizoram,  Meghalaya),  and N.  striata  n.  sp.  (India;  West 
Bengal,  Uttar  Pradesh,  Himachal  Pradesh).  The  genus  Qitercinirvana  Ahmed  &  Mahmood  (type  species;  Q.  longi- 
cephala  Ahmed  &  Mahmood)  is  treated  as  a  junior  synonym  oi  Sophonia  Walker.  Kana  signata  Distant  and  Nirvana 
greeni  Distant  are  suppressed  as  junior  synonyms  of  K.  illuminata  and  N.  Hnealis ,  respectively,  and  are  transferred  to 
Sophonia .  Three  other  species  transferred  to  Sophonia  are  Kana  modesta  Distant,  Quercinirvana  longicephala  Ahmed 
&  Mahood,  and  Nirvana  insignis  Distant.  The  relationships  among  various  genera  are  discussed  briefly.  The  genera 
Crispina  Distant  and  Mohiinia  Distant  are  excluded  from  Nirvaninae. 


The  Nirvaninae,  one  of  the  smaller  sub- 
families of  the  leafhopper  family  Cicadellidae, 
include  a  predominantly  tropical  group  of 
leafhoppers.  They  are  fragile,  often  brightly 
colored,  depressed  leaflioppers  and  common 
on  lush  green  vegetation.  They  are  often  mis- 
taken for  Typhlocybinae,  but  the  structure  of 
the  head  and  the  more  depressed  form  and 
truncate  basitarsus  of  the  hind  leg  distinguish 
Nirvaninae.  The  subfamily  includes  111  spe- 
cies (excluding  Mukariinae  and  other  non- 
Nirvaninae  genera,  which  were  traditionally 
included  in  the  Nirvaninae)  distributed  in  the 
Oriental  (56),  Afrotropical  (24),  Neotropical 
(7),  Palearctic  (7),  Australian  (11),  and  Pacific 
island  (6)  regions.  The  fauna  of  Afrotropical 
(Linnavuori  1979),  Australian  (Evans  1966), 
Papua  New  Guinea  (Evans  1973),  Neotropical 
(Kramer  1964),  and  Palearctic  (Kuoh  and 
Kuoh  1983)  regions  have  recently  been  re- 
vised. The  tribe  Mukariini  earlier  included  in 
the  subfamily  by  Distant  (1908g)  and  Metcalf 
(1963)  is  now  considered  a  subfamily  distinct 
from  Nirvaninae  (Linnavuori  1979). 

Information  on  host  plants  of  Nirvaninae  is 
very  meagre.  Nirvana  pallida  Melichar  and 
N.  suturalis  Melichar  are  destructive  to  sugar- 
cane and  grasses  in  Taiwan  (Schumacher 
1915a).  They  also  feed  on  rice,  mulberry,  and 
camphor  tree  (Esaki  and  Ito  1954a).   Baker 


(1923a)  found  Stenotortor  inocarpi  Baker  on 
Otaheite  chestnut,  Inocarpus  edulis,  in  Sin- 
gapore. In  Australia,  Nirvaninae  feed  on  /. 
edulis,  casuarina,  and  croton  (Evans  19411). 
Teco7naria  capensis  is  recorded  as  the  host  of 
Narecho  tecomariae  Theron  in  South  Africa 
(Theron  1970).  Quercinirvana  longicephala 
Ahmed  &  Mahmood  feeds  on  Aesculus  indica, 
Quercus  dilatata,  and  Vihurmmi  nervosum 
(Ahmed  and  Mahmood  1970).  Chudania 
delect  a  Distant  breeds  on  Ficus  carica 
(Ahmed  and  Mallik  1972).  In  South  America, 
Tahura  fowleri  Kramer  is  found  on  Passiflora 
(Kramer  1964).  Kuoh  and  Kuoh  (1983) 
recorded  Camellia  sinensis.  Acacia  confusa. 
Citrus  reticulata,  Pterocarpus  indicus,  Orijza 
sativa,  Psidium  guajava,  and  Chimonanthus 
praecox  as  host  plants  of  species  oi  Pseudonir- 
vana  Baker  {  =  Sophonia  Walker)  in  China. 
Nirvana  pallida  and  N.  greeni  Distant  breed 
on  black  gram,  green  gram,  cowpea,  field 
bean,  pigeon  pea,  and  soybean  in  India  (Ra- 
makrishna  1980). 

The  Nirvaninae  of  the  Indian  subcontinent 
consist  of  eight  genera  and  30  species  grouped 
in  three  tribes,  namely,  Balbillini,  Nirvanini, 
and  Occinirvanini.  The  genera  Mohunia  Dis- 
tant and  Crispina  Distant  are  excluded  from 
the  study,  as  they  probably  belong  to  the  sub- 
family Deltocephalinae. 


Department  of  Entomology,  University  of  Agricultnral  Sciences.  CKVK  Campus,  Bangalore  560  065,  India. 
^Department  of  Ecology  and  Evolution,  S.U.N.Y.,  Stony  Brook,  New  York  11794,  US.\. 


182 


1988 


VlRAKTAMATII,  WESLEY:  REVISION  OF  INDIAN  NiKVANINAE 


183 


Material  and  Abbreviations 

This  study  was  based  mainly  on  specimens 
collected  over  the  years  and  deposited  in  the 
insect  collection  of  the  Department  of  Ento- 
mology, University  of  Agricultural  Sciences 
by  the  senior  author  and  his  students.  Addi- 
tional specimens  were  borrowed  from  various 
institutions  for  the  study.  Abbreviations  used 
for  these  institutions  and  for  the  depositories 
of  the  types  of  new  taxa  are  as  follows: 

BMNH  -  British  Museum  (Natural  History),  London. 

lARI  -Indian  Agricultural  Re.search  Institute,  New 
Delhi. 

IRSNB  -  Institute  Royal  des  Sciences,  Naturelles  de  Bel- 
gique,  Bru.xelles. 

JU  -  Department  of  Biosciences,   Jammu   Univer- 

sity, Jammu. 

PAU      -  Punjab  Agricultural  University,  Ludhiana. 

UAS       -  University  of  Agricultural  Sciences,  Bangalore. 

USNM  -U.S.  National  Museum  (Natural  History), 
Washington,  D.C. 

ZMHU  -Museum  fiir  Naturkunde  der  Humboldt,  Urii- 
versitat  zu  Berlin,  Berlin. 

ZSI        -Zoological  Survey  of  India,  Calcutta. 

Distant  (1908g,  1918b)  did  not  mention  the 
number  of  specimens  (syntypes)  of  each  new 
species  he  described,  except  when  a  single 
specimen  was  involved,  viz.,  Chundania  de- 
lecta  Distant  (1908g;  268)  In  this  study  lecto- 
type  designations  were  made  even  though  a 
single  specimen  of  the  type  series  (syntypes?) 
was  present  in  the  BMNH  collection.  Unless 
otherwise  stated  they  were  considered  as  part 
of  the  syntype  series. 

The  terminology  used  for  describing  hind 
leg  spinulation  follows  Davis  (1975). 

Description  of  Taxa 

Nirvaninae 

Delicate,  fragile,  depressed,  small  to  mod- 
erately large  (4.0-9.0  mm)  leafhoppers.  Usu- 
ally yellow  or  white,  often  marked  with  red, 
black,  orange,  or  brown  fascia.  Head  as  wide 
as  or  wider  than  pronotum.  Vertex  usually 
flattened,  strongly  produced,  and  with  rugae 
or  sculpturing  on  lateral  and  anterior  regions. 
Ocelli  on  crown  along  lateral  margin  in  front  of 
eyes.  Frontoclypeus  and  clypellus  either  flat 
or  tumid,  in  the  latter  case  often  provided 
with  a  median  keel.  Lora  often  small,  extreme 
anterior  margin  of  genae  bluntly  prominent, 
extending  beyond  apex  of  clypellus.  Antennal 
pits  deep,  antennal  ledges  more  or  less 
strong,  antennae  long.  Lower  margin  of  eyes 


sinuate  ventrally.  Anterior  tentorial  branches 
L-shaped.  Lateral  margins  of  pronotum  cari- 
nate  (except  in  Omaranus  Distant),  moder- 
ately long.  Forewing  venation  reduced,  with- 
out cross-veins  at  base  (except  Balbillini), 
longitudinal  veins  represented  by  a  series  of 
paired  pits  basally,  appendix  either  narrow 
(Nirvanini)  or  broad  (BcdhiUus  Distant).  Hind- 
wing  with  three  or  lour  apical  cells,  veins  lA 
and  2A  fused  basally.  Fore  and  middle  tibiae 
cylindrical  or  flattened  (Balbillini),  hind 
femoral  spinulation  2+1  +  1  (Nirvanini), 
2+1  +  0  (Balbillini  and  Occinirvanini),  or 
2+0+0  (Balbilhni). 

Male  pygofer  either  cylindrical  or  de- 
pressed, with  or  without  anal  collar  process, 
but  may  be  armed  with  ventral  process,  heav- 
ily macrosetose.  Valve  small,  fused  with 
pygofer  laterally.  Plates  usually  parallel- 
sided.  Style  variable.  Connective  Y-shaped 
(Nirvanini)  or  platelike  (Balbillini).  Aedeagus 
with  single  shaft  and  often  provided  with  pro- 
cesses. 

Linnavuori  (1979)  suggested  that  the  sub- 
family Nirvaninae  is  a  derivative  of  the  Aphro- 
dinae-Cicadellinae  stock  and  that  Nirvanini  is 
the  most  advanced  tribe. 

Key  to  Tribes 

1.  Fore  and  middle  tibiae  flattened;  head  notched 
in  front  of  eyes  so  that  the  scape  is  visible  in 
dorsal  aspect    Balbillini 

—  Fore  and  middle  tibiae  rounded;  scape  not  visi- 
ble from  above 2 

2(1).  Ocelli  nearer  to  apex  of  head  than  to  eyes; 
forewing  appendix  wide  and  extending  around 
apical  wing  margin;  lora  large;  vertex  about  half 
as  long  as  wide,  obliquely  produced  in  front  of 
eyes,  with  hind  femoral  spinulation  2+1  +  0; 
hind  basitarsus  with  six  platellae  .  .  .  Occinirvanini 

—  Ocelli  nearer  to  eyes  than  to  apex  of  head; 
forewing  appendix  narrow  and  does  not  extend 
around  apex  of  wing;  lora  small;  vertex  as  long  as 
wide  or  longer,  not  obliquely  produced  in  front 
of  eyes;  hind  femoral  spinulation  2+1  +  1;  hind 
basitarsus  with  two  or  three  platellae    .  .    Nirvanini 

Tribe  Balbillini 

This  tribe  has  been  well  characterized  by 
Linnavuori  (1979).  The  following  additional 
characters  are  noted.  Forewing  with  supernu- 
merary cross-veins  along  costal  margin  and 
with  two  m-cu  cross-veins,  four  to  five  apical 
cells,  two  anteapical  cells,  and  appendix  (ab- 
sent in  Stenotortor  Baker).  Hindwing  with 
four  apical  cells.    Hind  femoral  spinulation 


184 


Great  Basin  Naturalist  Memoirs 


No.  12 


2  +  2  +  0  or  2  +  0+0.  Male  connective  long  and 
incrassate  or  lamellate  and  fusiform.  Aedea- 
gus  simple  with  large  gonopore. 

Key  to  Genera  of  Balbillini 

1.    Forewing  with  a  well-developed  appendix;  hind 
femoral  spinulation  2+1+0 Balhillus  Distant 

—  Forewing  without  an  appendix;  hind  femoral  spin- 
ulation 2  +  0  +  0    Stenotortor  Baker 

Genus  Balhillus  Distant 

Balhillus  Distant  1908g:  287.  Type  species;  Balhillus 
granulosus  Distant,  by  original  designation  and 
monotypy. 

Head  narrower  than  pronotum.  Crown 
bluntly,  angularly  produced,  convex,  de- 
clivous anteriorly,  with  median  and  submar- 
ginal,  anterior  carinae  between  ocelli.  Face 
horizontal,  flat.  Clypellus  twice  as  wide  at 
base  as  at  apex;  lora  small.  Lateral  margins  of 
head  in  front  of  eyes  notched,  exposing  scape 
of  antenna.  Pronotum  declivous  laterally, 
with  two  lateral,  carinate  margins;  mese- 
pisternum  attaining  lateral  position  in  front  of 
base  of  wing  and  strongly  ridged.  Fore  and 
middle  tibiae  angular;  hind  femoral  spinula- 
tion 2+1  +  0,  inner  spine  very  slender  and 
minute.  Apex  of  hind  basitarsus  with  three 
platellae.  Hind  tibial  spinulation  Rj  12  ±  1,  R, 
19±1,  R3  25±l,  R4  25±l. 

Pygofer  much  shorter  than  its  height,  with- 
out processes.  Anal  tube  short.  Genital  plate 
parallel-sided,  3.4  times  as  long  as  wide  with 
uniseriate,  submarginal  setae  and  few  hairlike 
setae.  Apophysis  of  style  simple,  without 
preapical  lobe.  Connective  lamellate  and  fusi- 
form. Aedeagus  articulate  with  connective, 
simple  and  with  large  gonopore. 

Remarks. — As  pointed  out  bv  Linnavuori 
(1979)  and  Baker  (1923),  Balhillus  and  Steno- 
tortor are  closely  related.  They  can  be  sepa- 
rated by  the  characters  used  in  the  key.  The 
bodies  of  both  genera  remain  appressed  to  the 
leaf  surface,  which  explains  the  flat  nature  of 
the  tibiae  and  the  undersurface  of  the  body. 

The  Indian  species,  B.  granulosa  and  B. 
indica,  have  hind  femoral  spinulation  2+1  +  0 
and  three  platellae  at  the  apex  of  the  hind 
basitarsus.  The  male  plate  of  B.  indica  is  paral- 
lel-sided. The  connective  and  aedeagus  are 
articulated  rather  than  fused.  The  connective 
is  spindle-shaped,  lamellate  rather  than  long 
and  incrassate,  and  the  preapical  lobe  of  the 
style  is  wanting.  These  characters  strongly 


suggest  that  the  Afrotropical  species  of  Balhil- 
lus, namely,  B.  ahas  Linnavuori  and  B.  tri- 
maculatus  Linnavuori,  may  not  belong  to  Bal- 
hillus. 

Key  to  Species  of  Balhillus 

1.    Vertex  and  scutellum  with  dark  brown  spots  .... 
granulosus  Distant 

—  Vertex  and  scutellum  immaculate  .  .  .    indicus,  n.  sp. 

Balhillus  granulosus  Distant 
Figs.  1-5 

Balhillus  granulosus  Distant  1908g:  288.  Holotype  9  ,  Sri 
Lanka  (BMNH,  examined). 

Yellow.  Vertex  with  a  median  spot,  a  spot  in 
apical  half  on  either  side  divided  by  carina, 
and  a  spot  near  inner  margin  of  eye  dark 
brown.  Pronotum  with  a  median,  short  stripe 
on  anterior  margin  fuscous.  Basal  triangles  on 
scutum  and  apex  of  scutellum  dark  brown; 
blackish,  median  line  on  scutellum  not  reach- 
ing apex  of  scutellum.  Longitudinal  veins  and 
claval  veins  lemon  yellow,  entire  surface  of 
forewing  punctate,  punctations  along  radial 
vein  and  on  outer  apical  cell  and  a  spot  at  base 
of  appendix  fuscous. 

Vertex  bluntly  conical,  1.24  times  as  wide 
between  eyes  as  its  median  length,  median 
carina  prominent  in  its  apical  half,  apical  0.75 
of  disc  sculptured,  basal  0.25  polished.  Face 
about  as  long  as  wide.  Pronotum  widened 
posteriorly.  Scutellum  longer  than  pronotum, 
transversely,  finely  rugulose  beyond  sulcus. 
Forewing  3.4  times  as  long  as  wide,  with  one 
m-cu  cross-vein  abut  0.33  distance  from  base 
and  another  near  apex  limiting  the  short,  in- 
ner anteapical  cell,  outer  anteapical  cell  open 
behind,  numerous  veins  arising  from  apical 
half  of  outer  anteapical  cell  and  reaching 
costa,  few  of  them  forked  at  costal  margin, 
third  apical  cell  divided  by  cross-vein.  Hind- 
wing  with  four  apical  cells. 

Female  genitalia. — Seventh  sternum 
about  twice  as  long  as  sixth,  its  hind  margin 
slightly  concave  with  a  strong,  U-shaped,  me- 
dian excavation,  very  finely,  transversely 
rugulose.  Ovipositor  exceeding  pygofers. 

Measurements. — Female  7.10  mm  long, 
head  1.85  mm  wide,  pronotum  1.87  mm 
wide. 

Spegimen  e.xamined. — Holotype  9  la- 
beled /Type/  /Balhillus  granulosus  Dist., 
tvpe/  /Ceylon  (Green)/  /Distant  Coll., 
1911-383/ (BMNH). 


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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. 


1988 


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187 


Figs.  18-27.  Stenotortor  stibhimalaya,  n.  sp.:  18,  habitus;  19,  face;  20,  head  and  thorax,  profile;  21,  forewing;  22, 
pygofer;  23,  male  plate;  24,  connective;  25,26,  aedeagus,  lateral  and  caudal  views;  27,  style. 


Tribe  Nirvanini 

Pale  yellow  with  dark  or  red  markings. 
Body  long,  narrow,  more  or  less  depressed. 
Head  as  long  as  or  longer  than  its  width,  disc 
of  vertex  longitudinally  rugose  in  anterior  re- 
gion. Ocelli  near  lateral  margins  in  front  of 
eyes.  Face  with  anterior,  median  carina  and 
usually  with  lateral,  oblique  ridges  on  fronto- 
clypeus.  Gena  narrow,  lora  small.  Antennal 
pits  rather  deep,  near  anterior  margin  of  eyes 
not  visible  from  above;  antennae  long,  reach- 
ing at  least  hind  margin  of  pronotum. 
Forewings  narrow,  with  reduced  appendix 
and  venation  indistinct  basally,  second  api- 
cal cell  not  widened  apically.  Hindwing 
with  three  or  four  apical  cells.  Fore  and  mid- 


dle tibiae  rounded.  Hind  femoral  spinulation 
2+1  +  1.  Hind  basitarsus  terminated  by  two 
platellae. 

Pygofer  with  well-developed,  lateral  lobes. 
Anal  tube  elongate,  often  with  apical  or  cau- 
doventral  process.  Male  plate  elongate,  Ion- 
longer  than  pygofer  and  apically  rounded. 
Style  variable,  usually  with  slender  apophysis 
that  is  curved,  hooked,  or  avicephaliform. 
Connective  free,  Y-shaped.  Aedeagus  weakly 
sclerotized,  usually  with  appendages. 

Remarks. — This  group  is  most  abundant 
among  Nirvaninae  and  appears  highly 
evolved  compared  to  other  tribes,  consider- 
ing the  reduced  wing  venation,  the  more 
streamlined  body  form,  and  the  specialized 
male  genitalia. 


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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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189 


Figs.  28-36.  Kana  ordinate  Distant;  28,  head  and  thorax;  29,  same,  profile;  30,  ventral  pygofer  process;  31,  male 
plate;  32,  connective;  33,  style;  34,  connective  and  aedeagus,  lateral  view;  35,  aedeagus,  caudal  view;  36,  forewing. 


6(5).  Vertex  with  pair  of  anteriorly  converging,  or- 
ange-yellow stripes  continued  on  pronotum  .  .  . 
fasciata  Pruthi 

—  Vertex  without  stripes;  pronotum  with  four  or- 
ange stripes;  male  genitalia  as  in  Figs.  30-35  .  . 
ordinata  Distant 

7(3).  Pronotum  with  two  median,  red  stripes  and  two 
lateral,  anterior,  lemon  yellow  spots  (Figs.  44); 
forewing  with  two  red,  transverse  bands  on 
corium  (Fig.  47) thoracica  Distant 

—  Pronotum  with  anterior  margin  and  two  spots  on 
posterior  margin  orange-yellow  (Fig.  42); 
corium  of  forewing  with  three  transverse,  red 
bands  (Fig.  43);  male  genitalia  as  in  Figs.  37-41 
ramificata  Distant 

Kana  ordinata  Distant 

Figs.  28-36 

Kana  ordinata  Distant   1908g:  287.    Lectotype   6,   Sri 
Lanka  (BMNH,  examined). 


Yellow.  Pronotum  with  two  inner  and  two 
outer  orange  stripes  joined  posteriorly  in 
male.  Scutellum  with  brownish  basal  trian- 
gles. Forewing  with  reddish  orange  band 
across  basal  area,  fuscous  speckled  band  about 
midlength  and  series  of  fuscous  spots  at  apices 
of  anteapical  cell. 

Head  slightly  narrower  than  pronotum. 
Vertex  of  head  slightly  longer  than  its  width 
between  eyes  (112:123),  apex  bluntly  conical. 
Face  with  four  prominent,  lateral  ridges. 

Male  genitalia. — Pygofer  rather  squarish 
with  caudodorsal  area  angulate,  and  with  two 
rows  of  stout,  long  setae;  ventral  process  with 
triangular,  dorsal  lobe  and  median,  dorsal 
hump.  Plate  with  rounded,  caudal  apex,  with 
row  of  stout  setae  in  caudal  0.66  and  with 


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No.  12 


marginal  row  of  long,  hairlike  setae.  Style 
with  well-developed,  preapical  lobe,  apophy- 
sis short,  stout,  its  apical  extension  slender, 
avicephaliform.  Connective  Y-shaped  with 
stem  as  long  as  the  length  of  arms.  Aedeagus 
tubular  with  unsclerotized,  poorly  developed 
dorsal  apodeme.  Shaft  strongly  bent  and  di- 
rected ventrad  and  cephalad  in  apical  0.33, 
appearing  rather  like  a  compressed,  inverted 
J,  cephalic  part  of  shaft  laterally  expanded  and 
constricted  in  middle  in  cephalic  view;  caudal 
part  with  two  long,  curved  processes  that 
cross  over  and  are  longer  than  those  in  K. 
ramificata. 

Measurements. — Male  body  3.00  mm 
long,  forewing  4.10  mm  long,  head  1.17  mm 
wide,  pronotum  1.25  mm  wide.  Female  6.40 
mm  long,  head  1.30  mm  wide,  pronotum  1.32 
mm  wide. 

Specimens  examined. — Lectotype  S  la- 
beled /Bogawantalawa,  Ceylon,  April/  /Dis- 
tant Coll.,  1911-383/  /1625/  /Nirvana  ordi- 
nata  Dist.,  type/  here  designated  (BMNH). 
Paralectotype  9  labeled  /Maskeliya,  Ceylon, 
8-05/  /Distant  Coll.,  1911-383/  here  desig- 
nated (BMNH). 

Remarks. — Kana  ordinata  is  closely  re- 
lated to  K.  ramificata  and  differs  in  coloration 
and  structure  of  the  pygofer  process.  Kana 
ordinata  has  a  shorter  caudal  part  of  the 
aedeagus  and  longer  processes  that  cross 
over. 

Kana  ramificata  Distant 

Figs.  37-43 

Kana  ramificata  Distant  1908g:  286.  Lectotype  6,  Sri 
Lanka  (BMNH,  examined). 

Pale  yellow,  with  submarginal  band  on  ver- 
tex, anterior  margin  of  pronotum  and  two 
spots  on  posterior  margin  orange.  Scutellum 
golden  yellow.  Forewing  with  basal  band  con- 
nected by  similar  but  more  oblique  band  from 
costa  to  clavus  and  then  continued  on  clavus 
but  not  reaching  claval  tip,  orange;  another 
oblique  band  with  well-defined,  orange,  ante- 
rior margin  and  cross-veins  limiting  apical 
cells,  reddish;  spot  at  apex  of  clavus  and  an- 
other on  third  apical  cell  fuscous. 

Head  slightly  narrower  than  pronotum 
(46:49).  Vertex  slightly  longer  than  width  be- 
tween eyes,  bluntly  conical.  Lateral  ridges  on 
face  four  in  number,  not  prominent.  Scutel- 
lum longer  than  pronotum. 


Male  genitalia. — Pygofer  rather  squar- 
ish, with  caudal  tufts  of  long,  stout  setae;  ven- 
tral process  stout  with  dorsal,  triangular  pro- 
cess near  caudal  apex.  Anal  tube  stout.  Male 
plate  with  oblique  row  of  stout  setae  in  caudal 
0.66,  with  long,  hairlike  setae,  apex  bluntly 
rounded.  Style  and  connective  as  in  K.  ordi- 
nata. Aedeagus  as  in  K.  ordinata,  but  caudal 
bent  part  0.66  as  long  as  tubular  cephalic  part, 
apical  processes  shorter. 

Measurements. — Male  5.20  mm  long, 
head  1.15  mm  wide,  pronotum  1.22  mm 
wide. 

Specimen  examined. — Lectotype  S  la- 
beled/Green, Ceylon,  95-221/ /Ntroana  ram- 
ificata Dist.,  type/  here  designated  (BMNH). 

Remarks. — Kana  ordinata  and  K.  ramifi- 
cata share  a  similar  basic  pattern  of  male  geni- 
talia; however,  they  differ  in  the  relative  size 
of  the  aedeagal  processes  and  the  shape  of  the 
pygofer  process.  Both  hindwings  and  the  right 
side  of  the  forewing  of  the  lectotype  are  dam- 
aged and  were  placed  in  a  microvial. 

Kana  thoracica  Distant 
Figs.  44-48 

Kana  thoracica  Distant  1908g;  285.   Lectotype   9 ,   Sri 
Lanka  (BMNH,  examined). 

Pale  yellow.  Two  anteriorly  converging, 
submarginal  stripes  on  vertex,  two  stripes  one 
on  either  side  of  median  line  red,  two  lateral 
spots  on  anterior  margin  of  pronotum  lemon 
yellow.  Forewing  with  red  bands  (Fig.  47) 
along  anal  margin,  obliquely  crossing  over  to 
clavus;  short  band  on  corium  near  base,  an- 
other longer,  oblique  band  about  midlength 
from  costal  margin  to  clavus.  Series  of  spots  on 
either  side  of  cross-veins  separating  apical 
cells  fuscous. 

Head  narrower  than  pronotum  (46:50). 
Vertex  longer  than  its  width  between  eyes 
(37:26).  Face  with  three  lateral,  oblique 
ridges;  longer  than  width  including  eyes. 
Pronotum  shorter  than  both  vertex  (24:37) 
and  scutellum  (24:28). 

Female  genitalia. — Hind  margin  of  sev- 
enth sternum  slightly  concave  with  a  median 
protuberance. 

Measurements. — Female  5.90  mm  long, 
head  1.15  mm  wide,  pronotum  1.25  mm 
wide. 

Specimen  examined. — Lectotype  9  la- 
beled /Pundaloya,   Ceylon/  /Ceylon,   Green 


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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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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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193 


Figs.  49-68.  Species  ofKana:  K.  bispinosa,  n.  sp.:  49,  head  and  thorax,  profile;  50,  same,  dorsal  view;  51,  face;  52, 
male  pygofer;  53,  style;  54,  connective;  55,  56,  aedeagus,  lateral  and  ventral  views;  57,  male  plate;  58,  forewing;  59, 
hindwing.  K.  nigropicta,  n.  sp.:  60,  head  and  thorax;  61,  forewing;  62,  male  pygofer;  63,  apex  of  ventral  process  of 
pygofer  in  two  views;  64,  style;  65,  connective;  66,  male  plate;  67,  68,  aedeagus,  lateral  and  caudal  views. 


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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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Great  Basin  Naturalist  Memoirs 


No.  12 


appendix  wanting,  apical  cells  four  and  an- 
teapical  cells  two.  Hindwing  with  four  apical 
cells.  Hind  tibial  spinulation  Rj  18  ±2,  Ro 
12±1,  R3l2±l. 

Male  pygofer  with  rounded,  caudal  margin, 
length  shorter  than  height,  strongly  sclero- 
tized  with  single,  submarginal  row  of  long 
setae  and  ventral  process.  Anal  tube  stout  and 
large,  about  0.8  times  as  high  as  height  of 
pygofer.  Male  plate  with  uniseriate  macrose- 
tae  in  apical  0.66  and  with  long,  hairlike  setae. 
Style  with  prominent  preapical  lobe,  apophy- 
sis slender,  apex  avicephaliform.  Style  rather 
T-shaped.  Aedeagus  membranous  for  most 
part,  bent  in  midlength,  laterally  expanded, 
with  ventrally  directed,  subapical,  pronglike, 
sclerotized  processes  surrounding  gonopore. 

Remarks. — This  genus  is  related  to  Kana 
and  Afrokana  Heller  with  which  it  shares 
hindwing  venation;  with  Afrokana  it  shares 
the  structure  of  the  male  pygofer.  However, 
they  differ  in  the  shape  of  the  head.  Lin- 
navuori  (1979)  stated  that  A/roniruana  Evans, 
Chudania,  and  Afrokana  are  very  close, 
sharing,  for  instance,  the  same  unique  genital 
structure.  He  believes  the  recorded  differ- 
ences are  too  slight  for  separating  them  as 
valid  genera.  Afrokana  is  closer  to  Kana  than 
to  Chudania  considering  the  peculiar  shape  of 
the  aedeagus  they  share.  It  is  likely  that  the 
study  of  more  material  of  these  genera,  espe- 
cially from  Malaysia,  Indonesia,  and  the 
Philippines,  may  prove  that  these  taxa  are 
congeneric  with  Chudania. 

Chudania  delecta  Distant 

Figs.  82-96 

Chudania  delecta  Distant  1908g:  268.  Holotype  9 ,  India 
(BMNH,  examined). 

Male.  Slender,  darker  than  female.  Vertex, 
pronotum,  scutellum,  upper  part  of  face  at 
level  with  lower  margins  of  eyes,  chocolate 
brown.  Ocelli  white,  eyes  dark  brown. 
Forewing  yellow,  with  posteriorly  widening, 
black  stripe  along  inner  margin,  occupying 
entire  apical  0.33  of  forewing  (Fig.  82);  two 
white,  hyaline,  triangular  areas  on  costal  area 
near  apex  and  some  cross-veins  enclosing  api- 
cal cells  paler.  Lower  half  of  face,  sterna,  and 
legs  yellow.  Pygofer  and  fifth  to  eighth  terga 
blackish. 

Female.  Similar  to  male  but  paler.  Black 
markings  of  male,  chocolate  brown.   Entire 


face,  lateral  margins  of  vertex  in  front  of  eyes, 
and  pronotum  yellow. 

Male  genitalia. — Pygofer  rounded  cau- 
dally,  with  submarginal  row  of  macrosetae, 
ventral  process  broad  caudodorsally  with 
finely  serrated,  dorsal  margin.  Male  plate 
more  or  less  of  uniform  width,  caudally 
rounded,  with  outer,  marginal,  long,  hairlike 
setae.  Anal  tube  large,  about  as  long  as  length 
of  pygofer.  Style  with  anterior  part  larger  than 
posterior  part  to  the  point  of  articulation  with 
connective.  Stem  of  connective  thrice  as  long 
as  arms,  apex  expanded.  Aedeagus  membra- 
nous for  most  part,  processes  sclerotized, 
bent  about  its  midlength,  apical  half  robust, 
preatrium  with  short  ventral  and  longer  lat- 
eral processes;  two  pairs  of  processes  arising 
where  shaft  bends,  dorsal  pair  elongate, 
curved  at  midlength,  the  ventral  pair  shorter, 
three  to  five  branched. 

Female  genitalia. — Hind  margin  of  sev- 
enth sternum  straight.  Ovipositor  exceeding 
length  of  pygofer.  Distal  half  of  pygofer  black. 

Measurements. — Male  4.62  (4.60-4.70) 
mm  long,  head  1.07  mm  wide,  pronotum  1.11 
(1.07-1.12)  mm  wide.  Female  5.38  (5.20- 
5.40)  mm  long,  head  1.20  mm  wide,  prono- 
tum 1.22  (1.20-1.25)  mm  wide. 

Specimens  examined. — Holotype  9  la- 
beled /Type,  H.T./  /Chudania  delecta  Dist., 
type/  /Kurseong/  Distant  Coll.  1911-383/ 
(BMNH).  India:  West  Bengal:  8  d,  7  9, 
Kurseong,  1,483  m,  22.x.  1981,  C.  A.  &  S. 
Viraktamath;  1  9,  15km  E  Kahmpong,  1,780 
m,  27.x.  1981,  C. A. v.;  4  c?,  11  9,  Kalimpong, 
1,768  m,  29.x.  1981,  S.  V. ;  9  c? ,  19  9  ,  8  km  E  of 
Kalimpong,  1,968  m,  29.x.  1981,  C.  A.  V.  Mi- 
zoram:  11  d,  11  9,  Aizawl,  18.xi.l981,  C.  S. 
Wesley;  11  c?,  14  9  ,  Lungleh,  20-25. xi.  1981, 
C.S.W.  (UAS). 

Remarks. — Chudania  delecta  and  C.  afri- 
cana  Heller  are  closely  related  but  differ  in 
coloration,  shape  of  the  ventral  pygofer  pro- 
cess, and  the  processes  on  the  aedeagus.  Also, 
the  dorsal  half  of  the  aedeagus  is  more  robust 
in  C.  delecta  than  in  C.  africana.  The  male 
genitalia  of  the  third  species,  C.  exposita  Ja- 
cobi,  is  not  known. 

Genus  Sophonia  Walker 

Sophonia  Walker  1870b:  .327.  Type  species:  Sophonia 

rufitehim  Walker  by  monotypy. 
Pseudonirvana     Baker     1923a:     386.     Type     species: 

Pseudonirvana  sandakanensis  Baker,  by  original 

designation. 


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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. 


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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. 


200 


Great  Basin  Naturalist  Memoirs 


No.  12 


Figs.  103-121.  Sophonialinealis  {Distant):  103,  head  and  thora.x  of  holotype  female  Niruaria  greeni  Dist.;  104,  head 
and  thorax  of  holotvpc  of  Nirvana  linealis  Dist.;  105,  profile;  106,  ovipositor;  107,  male  pygofer;  108,  forewing;  109 
110,  aedeagus,  lateral  and  dorsal  views;  HI,  connective  and  style.  S.  km^icephala  {Ahmed  in  Mdhmood):  112,  head  and 
thorax;  113,  face;  114,  male  plate;  115.  male  pygofer;  116,  ventral  process  of  pygofer;  117,  style;  118,  119,  aedeagus, 
lateral  and  caudal  views;  120,  connective;  121,  forewing. 


1988 


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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 


202 


Great  Basin  Naturalist  Memoirs 


No.  12 


Figs.  122-141.  Sophonialongitudinalis  (Vistsint):  122,  head  and  thorax;  123,  face;  124,  ovipositor;  125,  male  pygofer; 
126,  127,  male  plate;  128,  connective  and  stvle;  129,  130,  aedeagus,  lateral  and  caudal  views;  131,  forewing.  S.  I^^ken, 
n.  sp.:  132,  head  and  thorax;  133,  face;  134,  forewing;  135,  male  pygofer;  136,  pygofer  process;  137,  male  plate;  138, 
connective;  139-141,  aedeagus,  lateral,  caudal,  and  dorsal  views. 


1988 


ViRAKTAMATH,  WeSLEY:  REVISION  OF  INDIAN  NiRVANINAE 


203 


half  with  long,  slender,  arcuate,  apically 
pointed,  ventrally  curved  process  that  bears  a 
short  tooth  on  its  outer  margin  at  base. 

Female  genitalia. — Hind  margin  of  sev- 
enth sternum  slightly  concave.  Ovipositor 
slightly  exceeding  pygofer. 

Measurements— Male  4.43  (3.80-5.40) 
mm  long,  head  0.96  (0.80-1.43)  mm  wide, 
pronotum  0.96  (0.85-1. 13)  mm  wide.  Female 
4.90  (4.60-5.30)  mm  long,  head  1.02 
(0.98-1.10)  mm  wide,  pronotum  1.05 
(1.00-1. 10)  mm  wide. 

Specimens  examined. — Lectotype  9  la- 
beled /Myitta  (Doherty)/  /Distant  Coll. 
1911-383/  /Nirvana  longitudinalis  Dist., 
type/ here  designated  (BMNH).  India:  Karna- 
taka:  6  (5,  Mudigere,  4-6. iv.  1980,  C.  A.  Vi- 
raktamath;  1  9,  7.iv.l975,  C.A.V.;  1  9, 
Kodlipet,  6.iv.l975;  1  S,  Hunsur,  10.  i.  1978; 
1  9,  Bannerghatta,  12. ix.  1976,  B.  Mallik; 
Kerala:  1  9,  Thekkadi,  27.iii.1977,  C.A.V. 
Nepal:  Nagarkot,  1  (5,  1  9,  15.ix.l979,-  I. 
Dworakowska  (UAS).  Burma:  1  9,  Amberst 
Dist.,  Kawkareik,  19-20.xi.l911,  F.H.G. 
(ZSI). 

Remarks. — This  species  can  be  distin- 
guished from  all  other  species  of  Sophonia  by 
its  lamellate  aedeagal  shaft  and  tooth  on  the 
apex  of  the  male  plate. 

Sophonia  bakeri,  n.  sp. 
Figs.  132-141 

Pale  yellow  to  whitish  yellow.  Vertex  with 
black  spot  near  apex,  joined  by  two  black, 
longitudinal  lines  that  fuse  basally  and  extend 
to  scutellum.  Pronotum  and  scutellum  some- 
times with  reddish  spots,  scutellum  in  male 
entirely  reddish.  Forewing  with  brown  stripe 
close  to  or  along  anal  margin,  marked  with 
yellow  along  inner  margin,  apex  of  clavus  and 
apical  margin  brown,  costal  margin  with  ante- 
rior, oblique  and  posterior,  transverse,  brown 
fasciae,  second  apical  cell  with  large,  black 
spot. 

Vertex  about  1.5  times  as  long  as  wide,  apex 
subtriangular,  disc  depressed.  Posterior  half 
of  pronotum  transversely  rugulose.  Second 
apical  cell  narrowed  at  apex. 

Male  genitalia. — Pygofer  with  ventrocau- 
dal  angle  produced  to  process  with  attenuated 
apex.  Plate  3.5  times  as  long  as  wide.  Apophy- 
sis of  style  with  long,  beaklike  extension. 
Stem  of  connective  1.5  times  as  long  as  each 
arm.  Aedeagus  with  each  arm  of  dorsal  apo- 


deme  robust,  bearing  triangular  process  on 
caudal  margin;  preatrium  short;  shaft  tubular, 
caudodorsally  curved,  apex  strongly  incurved 
with  ventrally  directed,  short,  pronglike  pro- 
cess on  caudal  margin  subapically. 

Female  genitalia. — Hind  margin  of  sev- 
enth sternum  concavely  emarginate.  Oviposi- 
tor exceeding  length  of  pygofer. 

Measurements,— Male  4.48  (4.30-5.00) 
mm  long,  head  0.90  (0.85-1.00)  mm  wide, 
pronotum  0.88  (0.78-1.03)  mm  wide.  Female 
5.08  (4.70-5.30)  mm  long,  head  1.04  (0.93- 
1.23)  mm  wide,  pronotum  1.05  (1.00-1.10) 
mm  wide. 

Specimens  examined. — Holotype  6,  In- 
dia: Karnataka:  Mudigere,  at  light, 
4-6.iv.l980,  C.  A.  Viraktamath  (UAS). 
Paratypes:  India:  Karnataka:  2  6,  Dharwar, 
— .i.l970,  C.  A.  Viraktamath;  1  9,  Kodlipet, 
6.iv.l975,  C.A.V. ;  1  9,  Bangalore,  9.iii.l975, 
C.A.V.;  1  9,  Kalhalla,  18.i.l978,  C.A.V.,  1 
9,  Mudigere,  23. ix.  1973,  C.A.V.;  1  (?,  1  9, 
Kemmanagundi,  8.iv.l975,  K.  D.  Ghorpade; 
Kerala:  1  c?,  Thekkadi,  27.iii.1977,  S.  Virakta- 
math; Maharashtra:  1  9 ,  Matheran,  915  m, 
23.xi.1977,  C.A.V.;  Tamil  Nadu:  1  6,  Ko- 
daikanal,  2,250  m,  10. vi.  1980,  C.  S.  Wesley; 
Uttar  Pradesh:  1  6,  1  9,  Dehra  Dun, 
28. iv.  1975,  C.A.V.;  Nepal:  2  9,  Gokarna, 
14.  ix.  1979,  I.  Dworakowska;  1  9,  Sarankot, 
17. ix.  1979,  exQiiercus  sp.,  I.D.  (1  6  and  1  9 
in  BMNH,  lARI,  USNM,  and  rest  in  UAS). 

Remarks. — Sophonia  bakeri  is  related  to  S. 
longitudinalis  and  S.  longicephala  as  they 
share  similar  external  coloration  and  male 
genitalia.  They  form  a  closely  knit  group.  S. 
bakeri  differs  from  the  other  two  by  its  shorter 
process  of  the  dorsal  apodeme  and  aedeagal 
process. 

Sophonia  keralica,  n.  sp. 
Figs.  142-153 

Uniformly  yellow.  Vertex  with  round,  black 
spot  near  apex  connected  by  single,  median, 
longitudinal,  black  line,  discontinuous  for 
short  distance  near  black  spot  and  then  con- 
tinued to  apex  of  scutellum.  Lateral  margin  of 
vertex  orange  basally.  Forewing  with  claval 
suture  and  punctures  along  cubital  vein 
brown,  second  and  part  of  third  apical  cells 
occupied  by  large,  black  spot;  costal  margin 
with  anterior,  oblique  and  posterior,  trans- 
verse, black  fasciae,  apex  of  clavus  and  apical 
margin  brownish. 


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Great  Basin  Naturalist  Memoirs 


No.  12 


Figs.  142-153.  Sophonia  keralica,  n.  sp.:  142,  head  and  thorax;  143,  face;  144,  head  and  thorax,  profile;  145,  male 
pygofer;  146,  pygofer  process;  147,  connective;  148,  style;  149,  male  plate;  150,  151,  aedeagus,  lateral  and  cephalic 
views;  152,  forewing;  153,  hindwing. 


Head  nearly  as  wide  as  pronotum.  Vertex 
slightly  longer  than  wide,  disc  flat,  depressed 
near  apex.  Scutellum  slightly  longer  than 
pronotum.  Second  apical  cell  of  forewing  nar- 
rowed slightly  near  apex. 

Male  genitalia. — Pygofer  caudally 
rounded,  its  ventrocaudal  angle  produced  to 
sharp,  thin  process,  five  macrosetae  along 
caudal  margin  in  straight  line,  black,  rest  hya- 
line. Anal  tube  as  long  as  pygofer.  Plate  four 
times  as  long  as  wide.  Style  with  strongly 
hooked  apophysis.  Stem  of  connective  1.5 
times  as  long  as  each  arm.  Dorsal  apodeme  of 
aedeagus  slender,  each  arm  with  ventral,  dor- 
sal, and  short,  median  process;  shaft  tubular 
with  pair  of  laterally  directed  processes, 
gonopore  subapical. 

Measurements. — Male  3.90  mm  long, 
head  0.90  mm  wide,  pronotum  0.93  mm 
wide. 

Specimen  examined. — Holotvpe  6,  India: 
Kerala:  Thekkadi,  27.iii.1977,  s!  Viraktamath 
Coll.  (UAS). 


Remarks. — Sophonio  keralica  is  related  to 
S.  iUuminata.  Both  have  similar  male  geni- 
talia. They  differ,  however,  in  coloration  and 
detailed  structure  of  the  aedeagus. 

Sophonia  iUuminata  (Distant),  n.  comb. 
Figs.  154-163 

Kana  iUuminata  Distant  1918b:  35.  Lectotype  d,  India 

(BMNH,  examined). 
Kana   sigruiffl   Distant   1918b:  34.    Lectotype    9 ,   India 

(BMNH,  examined).  New  synonymy. 

Female.  Vertex  and  pronotum  creamy 
white  with  yellowish  lateral  areas,  median 
stripe  narrowing  to  apex  of  vertex  san- 
guineous on  vertex  and  anterior  half  of  prono- 
tum, widened  in  posterior  half  where  it  is 
black.  Scutelhun  black  except  for  two  lateral, 
basal  spots.  Clavus  along  claval  suture  and 
apical  half  piceous  except  for  subapical,  elon- 
gate, orange  spot.  Corium  along  its  inner  mar- 
gin piceous  which  widens  near  apex  of  cla- 
vus and  changes  to  orange;  appendix  fuscous, 
two  large,  round  spots  in  second  apical  cell. 


1988 


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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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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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ViRAKTAMATH,  WeSLEY:  REVISION  OF  INDIAN  NiRVANINAE 


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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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ViRAKTAMATH,  WESLEY;  REVISION  OF  INDIAN  NiRVANINAE 


215 


Figs.  233-243.  Nirvana  suturalis  Melichar:  233,  head  and  thorax  of  holotype  female;  234,  female  from  India;  235, 
face;  236,  ovipositor;  237,  male  pygofer;  238,  connective;  239,  240,  aedeagus,  lateral  and  cephalic  views;  241,  style;  242, 
male  plate;  243,  forewing. 


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Great  Basin  Naturalist  Memoirs 


No.  12 


and  Tamil  Nadu  from  January  to  December, 
at  an  altitudinal  range  of  534  m  (Jog  Falls  in 
Karnataka)  to  2,350  m  (Ootacamund  in  Tamil 
Nadu)  above  mean  sea  level. 

Remarks. — This  is  the  second  most  com- 
monly collected  Nirvaninae  in  southern  In- 
dia. Nirvana  suturalis  and  N.  striata  are  very 
closely  related.  Both  differ  from  other  species 
of  Nirvana  and  Sophonia  in  having  a  broad, 
black,  median  stripe  running  from  the  apex  of 
the  vertex  to  the  corium,  but  the  stripe  is  not 
as  broad  as  in  species  of  Chudania.  Nirvana 
suturalis  has  a  serrated  lateral  margin  to  the 
median  stripe  compared  to  the  smoother  lat- 
eral margin  in  N.  striata. 

Nirvana  striata,  n.  sp. 

Figs,  244-255 

Coloration  as  in  N.  suturalis,  median  stripe 
more  or  less  of  uniform  width  and  even  mar- 
gined, black  spot  on  second  apical  cell  of 
forewing,  larger  than  in  N.  suturalis,  extend- 
ing to  first  apical  cell. 

Head  narrower  than  pronotum.  Vertex 
slightly  longer  than  width  between  eyes, 
anterior  half  sparsely  rugulose.  Pronotum 
slightly  more  than  1.5  times  as  wide  as  long, 
longer  than  scutellum. 

Male  genitalia. — Pygofer  bluntly  conical 
with  macrosetae  confined  to  caudal  0.33  area, 
ventral  margin  with  short,  spinelike  process. 
Plate  caudally  rounded,  about  seven  times  as 
long  as  wide.  Style  with  rounded  preapical 
lobe,  apical  extension  of  apophysis  beaklike 
and  long.  Stem  of  connective  twice  as  long  as 
each  arm.  Aedeagus  with  shaft  strongly 
curved  (more  than  in  N.  suturalis),  apex 
abruptly  directed  cephalad  and  with  membra- 
nous area;  each  arm  of  dorsal  apodeme  with 
basal,  short  process. 

Female  genitalia. — Hind  margin  of  sev- 
enth sternum  straight.  Ovipositor  exceeding 
pygofer. 

Measurements.— Male  4.90  (4.80-5.00) 
mm  long,  head  0.98  (0.95-1.00)  mm  wide, 
pronotum  1.06  (1.05-1.08)  mm  wide.  Female 
5.46  (5.30-5.60)  mm  long,  head  1.08  (1.05- 
1.10)  mm  wide,  pronotum  1.12  (1.08-1.13) 
mm  wide. 

Spegimens  examined. — Holotype  9,  In- 
dia: West  Bengal:  Kurseong,  1,483  m, 
22.x.  1981,  S.  Viraktamath  (UAS).  Paratypes: 
India:  Uttar  Pradesh:  1  9,  Mussoorie,  2,005 
m,  27. iv.  1975,  C.  A.  Viraktamath;  Himachal 


Pradesh:  3  9,  Simla,  2,133  m,  14.x.  1979,  C. 
A.  Viraktamath;  West  Bengal:  3  d,  3  9, 
Kurseong,  1,483  m,  22.x.  1981,  S.  Virakta- 
math (1  (5 ,  1  9 )  and  C.  A.  Viraktamath  (2  c? ,  2 
9);  1  9,  15  km  E  of  Kalimpong,  1,780  m, 
27.x.  1981,  S.  Viraktamath;  1  <5 ,  2  9  ,  8  km  E  of 
Kalimpong,  1,768  m,  29.x.  1981,  C.  A.  Virak- 
tamath (1  6  and  1  9  paratypes  in  BMNH, 
USNM,  and  rest  in  UAS). 
Remarks. — See  N.  suturalis. 

Nirvana  peculiaris,  n.  sp. 

Figs.  256-267 

Yellow.  Small,  black  spot  at  apex  of  vertex 
visible  both  dorsally  and  ventrally  (absent  in 
female  from  Shillong).  Fuscous,  median 
stripe  extending  posteriorly  to  0.25  length 
(absent  in  some  females).  Lateral,  orange 
stripe  on  vertex  mesad  of  ocelli,  continued  on 
pronotum,  scutellum  and  then  on  to  inner 
claval  margin  where  they  converge  and  con- 
tinue to  apex  of  clavus;  outer  margin  of  stripe 
on  clavus  often  fuscous.  Transverse  spot  at 
apex  of  clavus,  small  spot  at  apex  of  inner, 
anteapical  cell,  and  anterior,  oblique  fascia 
from  costa  fuscous.  Black  spot  on  each  cau- 
dodorsal  angle  of  male  pygofer. 

Head  as  wide  as  pronotum  or  slightly  nar- 
rower. Disc  of  vertex  depressed  with  rolled- 
up,  sharp  margin,  basal  half  with  median  sul- 
cus, apical  half  with  a  carina,  apical  half  of  disc 
very  finely  pitted,  posterior  half  with  minute 
rugulae  on  polished  surface. 

Male  genitalia. — Pygofer  elongate,  nar- 
rowed caudally  to  acute,  dorsally  curved  pro- 
cess. Anal  collar  elongate,  anterior  margin 
armed  with  two  pronglike  processes  on  each 
side.  Setae  on  pygofer  confined  to  caudodor- 
sal  area  where  it  articulates  with  anal  tube. 
Male  plate  elongate,  5.4  times  as  long  as  me- 
dian width,  macrosetae  and  hairlike  setae  con- 
fined to  apical  half  Connective  robust,  stem 
broad,  and  arms  as  long  as  stem.  Apophysis  of 
style  fairly  long.  Aedeagus  with  dorsal  apo- 
deme poorly  developed,  shaft  tubular,  nar- 
rowed caudally,  strongly  curved  dorsally  near 
apex  and  with  elongate,  caudoventrally  di- 
rected process  and  two  lateral,  anteriorly 
directed  processes  surrounding  small  gono- 
pore. 

Measurements.— Male  5.80  (5.70-5.90) 
mm  long,  head  1.23  (1.22-1.25)  mm  wide, 
pronotum  1.24  (1.22-1.25)  mm  wide.  Female 
7.40  (7.00-7.80)  mm  long,  head  1.50  (1.45- 


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ViRAKTAMATH,  WeSLEY:  REVISION  OF  INDIAN  NiRVANINAE 


217 


Figs.  244-255.  Nirvana  striata,  n.  sp.:  244,  head  and  thorax;  245,  face;  246,  profile;  247,  ovipositor;  248,  forewing; 
249,  hindwing;  250,  male  pygofer;  251,  male  plate;  252,  style;  253,  254,  aedeagus,  lateral  and  cephalic  views;  255, 
connective. 


1.55)  mm  wide,  pronotum  1.54  (1.50-1.60) 
mm  wide. 

Specimens  examined. — Holotype  6,  In- 
dia: Mizoram:  Aizawl,  18. xi.  1981,  C.  S. 
Wesley  (UAS).  Paratypes:  India:  Mizoram:  2 
S,4  9,  Aizawl,  18.xi.l981,  C.  S.  Wesley;  1 
d,  2  $,  Lungleh,  23.xi.1981,  C.  S.  Wesley; 
Meghalaya:  1  9,  Shillong,  1,961  m,  3.xi.l981, 
S.  Viraktamath  (1  9  and  1  9  paratypes  in 
BMNH;  1  9  paratype  in  lARI,  USNM,  ZSI, 
and  rest  in  UAS). 

Remarks. — This  species  is  unique  among 
Nirvanini  in  having  an  armed  tenth  segment. 
In  coloration  it  resembles  species  of  Kana, 
especially  K.  fasciata,  but  it  differs  from  those 
species  in  having  three  apical  cells  in  the  hind- 
wing  and  unique  male  genitalia. 


Ophiuchus  Distant 

Ophiuchus  Distant  1918b:  33.  Type  species:  Ophiuchus 
princeps  Distant,  by  original  designation  and 
monotypy. 

Pale  yellow  with  bright,  contrasting  col- 
oration of  black  and  red  or  pink.  Head  about  as 
wide  as  pronotum,  spatulate,  disc  of  vertex 
depressed  in  middle,  margined  by  carina,  lat- 
eral margin  of  vertex  straight  in  front  of  eyes  to 
ocelli  where  it  is  widened  and  then  slightly 
narrowed  to  apex.  Face  with  five  lateral 
ridges,  concave  medially,  with  short,  median 
carina  near  apex.  Pronotum  twice  as  wide  as 
median  length,  disc  minutely,  transversely 
rugose.  Hindwing  with  three  closed  apical 
cells.  R2  of  hind  tibia  with  12  spines. 


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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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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 


1988 


ViRAKTAMATH,  WESLEY:  REVISION  OF  INDIAN  NiRVANINAE 


221 


Genus  Omaranus  Distant 

Omaranus  Distant  1918b:  5.  Type  species:  Omaranus 
typictis  Distant,  by  original  designation  and  inoiio- 
typy. 

Didius  Distant  1918b:  36.  Type  species:  Didius  sexualis 
Distant,  by  original  designation  and  nionotypy. 

Sexually  dimorphic.  Head  narrower  than 
pronotum  in  male,  wider  than  pronotum  in 
female.  Vertex  with  median  carina  in  apical 
half,  sulcus  in  basal  half.  Face  longer  than 
broad,  frontoclypeus  tumid.  Clypellus  longer 
than  broad;  frontoclypeus,  lora,  genae,  and 
clypellus  with  scattered,  short  setae.  Prono- 
tum gradually  widened  posteriorly  in  male, 
parallel-sided  and  with  faint,  median  ridge 
near  posterior  half  in  female;  posterior  margin 
strongly  inwardly  concave;  disc  llattish  in  fe- 
male, slightly  raised  posteromedially  in  male, 
lateral  margins  not  carinate.  Male  forewing 
exceeds  abdomen,  appendix  runs  around  api- 
cal margin,  with  four  apical  and  three  anteapi- 
cal  cells;  hindwing  well  developed  with  three 
apical  cells.  Female  forewings  cover  only 
three  basal,  visible,  abdominal  terga,  while 
hindwings  cover  two  basal,  visible  terga;  ap- 
pendix wanting.  Hind  tibial  spinulation  Rj 
16  ±  2,  R,  10  ±  1,  R3  3.  Hind  basitarsus  shorter 
than  combined  length  of  second  and  third 
tarsi  and  terminated  by  six  platellae. 

Male  pygofer  without  processes.  Tenth  seg- 
ment stout,  rather  short.  Ovipositor  not  ex- 
ceeding pygofer. 

Remarks. — This  genus  and  the  Austrahan 
Occinirvana  are  closely  related.  The  shape  of 
the  head  differentiates  the  two  genera. 

Omaranus  typicus  Distant 

Figs.  277-289 

Omaranus  typicus  Distant  1918b;  6.  Lectotype  S ,  India 

(BMNH,  examined). 
Didius  sexualis  Distant  1918b:  36.  Lectotype  9,  India 

(BMNH,  examined). 

Male. — Ochraceous  (probably  green  in 
life).  Anterior  margin  of  vertex  narrowly  black 
margined,  with  median,  black  spot,  two 
smaller,  somewhat  elongate  spots  near  poste- 
rior margin,  ocelli  black.  Dorsal-most  area  of 
frontoclypeus  and  face  ventrad  of  antennal 
bases,  blackish  brown,  marginal  band  dorsad 
of  antennal  base  and  longitudinal  stripe  on 
proepimeron  blackish  brown.  Apical  0.25  of 
forewing  and  stripe  along  inner  margin  fus- 
cous. Abdomen  with  four  black  stripes;  sterna 
marked  with  brown  stripes.  Apical  half  of  hind 


femora  ventrally  streaked  with  brown.  A  ven- 
tral spot  near  apex  of  fore  femora  black. 

Genitalia. — Pygofer  lobe  caudoventrally 
produced,  caudodorsal  area  beset  with  stout, 
short  setae.  Plate  caudally  rounded  with  scat- 
tered, stout  setae.  Valve  straplike  with  caudal 
margin  roundly  and  medially  produced.  Apo- 
physis of  style  curved  laterally  with  series  of 
teeth  on  lateral  margin.  Stem  of  connective 
0. 75  as  long  as  each  arm .  Aedeagus  with  short, 
dorsal  apodeme,  shaft  stout  at  base,  narrowed 
caudally  in  lateral  aspect  with  a  large  gono- 
pore  on  dorsal  aspect. 

Female. — Ochraceous.  Head  with  median 
spot  on  anterior  margin,  ocelli,  and  two  spots 
near  base  of  vertex  black;  median  stripe,  a 
transverse  band  across  ocelli,  hind  margin, 
except  medially,  and  two  spots  in  front  of  eyes 
reddish.  Dorsal  margin  of  face  black  from 
which  two  apically  divergent  stripes  traverse 
face.  Proepisternum  traversed  by  longitudi- 
nal, piceous  stripe,  much  longer  on  pro- 
episternum; mesopleura  and  mesocoxae  with 
piceous  spot.  Pronotum  with  scattered,  dark 
brown  spots,  with  obscure  stripes,  two  lateral 
and  two  median  reddish.  Scutellum  except  for 
two  lateral,  basal  angles  in  basal  half  black, 
apical  half  blackish  brown.  Forewing  veins 
marked  with  red.  Fore  tibiae  and  tarsi  red- 
dish; fore-  and  mesothoracic  femora  black, 
mesocoxae  with  piceous  spot.  Abdominal  col- 
oration as  in  male. 

Genitalia. —  Hind  margin  of  seventh  ster- 
num with  median,  U-shaped  excavation  and 
translucent  in  middle.  Anal  segment  con- 
cealed by  pygofer. 

Measurements. — Male  4.30  mm  long, 
head  1.00  mm  wide,  pronotum  1.10  mm 
wide.  Female  7.90  mm  long,  head  1.00  mm 
wide,  pronotum  1. 10  mm  wide. 

Specimens  examined. — Lectotype  6  la- 
beled /  Calcutta,  28.viii.07,  Mus.  Coll./  / 
Omaranus  typicus  Dist.,  type/  here  desig- 
nated. Paralectotype  S  labeled  /  Calcutta, 
23.viii.07/  /Omaranus  typicus  Dist.,  type/ 
here  designated.  Paralectotype  has  the  abdo- 
men and  left  fore  and  hindwings  missing.  Lec- 
totype 9  labeled  /  Calcutta/  /Distant  Coll. 
1911-384/  /Didius  sexualis  Dist.  type/  here 
designated  (BMNH).  India:  Punjab:  1  c?,  1  9 
nymph,  Ludhiana,  13. ix.  1966,  A.  S.  Sohi 
(PAU).  India:  Madhya  Pradesh:  3  6 ,  18  9 , 
Nebudda  Survey,  Sta.  No.  31,  37,  58, 
21.ii-9.iii.l927,  H.S.  Pruthi  (ZSI). 


222 


Great  Basin  Naturalist  Memoirs 


No.  12 


Figs.  277-289.  Omamnus  ti/picus  Distant:  277,  head  and  thorax  of  lectotype  male;  278,  same  of  lectotype  female, 
Didius  sexiialis  Distant;  279,  ovipositor;  280,  281,  fore  and  hindwings  of  male;  282,  male  pygofer;  283,  male  plate;  284, 
male  valve;  285,  connective;  286,  style;  287,  288,  aedeagus,  lateral  and  cephalic  views. 


1988 


ViRAKTAMATH,  WeSLEY:  REVISION  OF  INDIAN  NiRVANINAE 


223 


Acknowledgments 

The  authors  express  thanks  to  Dr.  W.  J. 
Knight,  Mr.  M.  D.  Webb  (BMNH),  Dr.  U. 
Gollner-Scheiding  (ZMHU),  Dr.  H.  Synave 
(IRSNB),  Dr.  A.  S.  Sohi  (PAU),  and  Dr. 
Baldev  Sharma  (JU)  for  lending  specimens  for 
study,  including  primary  types  under  their 
care.  Also,  the  director  of  the  Zoological  Sur- 
vey of  India  allowed  the  senior  author  to  ex- 
amine Dr.  H.  S.  Pruthi's  type  material  during 
his  visit  to  ZSI.  The  authors  are  also  grateful  to 
Dr.  P.  H.  Freytag,  University  of  Kentucky, 
Lexington,  for  critically  reviewing  the  manu- 
script. 

Literature  Cited 

The  references  cited  in  the  text  prior  to  1956  are  found 
in  Metcalf  (1964),  wherein  Pruthi  s  works  have  been  cited 
under  the  name  Singh-Pruthi. 

Ahmed,  M.,  and  S.  H  Mahmood  1970.  A  new  genus  and 
two  new  species  of  Nirvaninae  (CicadeUidae:  Ho- 
moptera)  from  Pakistan.  Pakistan  Journal  of  Scien- 
tific and  Industrial  Research  12:  260-263. 

Ahmed,  M  ,  and  K  F  Mallik  1972.  On  the  true  identifi- 
cation of  genus  Chudania  Distant  collected  on  fig 
{Ficiis  carica )  in  forests  of  Azad  Kashmir  and  Kap- 
tai  (Pakistan).  Pakistan  Journal  of  Forestry  22; 
103-108. 

Davis,  R.  B.  1975.  Classification  of  selected  higher  cate- 
gories of  Auchenorrhynchous  homoptera  (Cica- 


deUidae and  Actalionidae).  Technical  Bulletin 
No.  1494,  Agricultural  Research  Service,  United 
States  Department  of  Agriculture.  .52  pp. 

EvAN.s.  J  W  1966.  The  lealhoppers  and  froghoppers  of 
Australia  and  New  Zealand  (Homoptera:  Cicadel- 
loidea  and  Cercopoidea).  The  Australian  Museum 
Memoir  12:  1-347. 

1973.  Some  new  genera  and  species  of  Cicadel- 

loidea  from  Australia  and  New  Guinea  (Ho- 
moptera). Pacific  Insects  15:  185-197. 

KUOH.  C  L ,  AND  J  L  KUOH  1983.  New  species  of 
Pseudonirvana  (Homoptera:  Nirvanidae).  Acta 
Entomologica  Sinica  26:  316-.325  [In  Chinese]. 

Kramer,  J  P  1964.  A  review  of  the  Neotropical  Nirvan- 
inae (Homoptera:  CicadeUidae).  Entomological 
News  75:  113-127. 

LiNNAVUORl,  R  1979.  Revision  ofthe  African  CicadeUidae 
(Homoptera  Auchenorrhyncha),  Part  II.  Revue  de 
Zoologie  Africaine  93:  929-1010. 

Metcalf,  Z  P  1963.  General  Catalogue  of  the  Ho- 
moptera, Fascicle  VI.  Cicadelloidea,  Part  7  Nir- 
vanidae. Agricultural  Research  Service,  United 
States  Department  of  Agriculture.  35  pp. 

1964.  General  Catalogue  of  Homoptera,  Fa.scicle 

VI.  Cicadelloidea,  Bibliography  of  the  Cicadel- 
loidea (Homoptera:  Auchenorrhyncha).  Agricul- 
tural Research  Service,  United  States  Depart- 
ment of  Agriculture., 349  pp. 

Ramakrishna,  B  V.  1980.  Leaihopper  fauna  of  pulse 
crops  and  biology  of  Empoasca  (Distantasca)  ter- 
minalis  Distant  (Homoptera:  CicadeUidae).  Un- 
published thesis.  University  of  Agricultural  Sci- 
ences, Bangalore.  197  pp. 

Theron,  J  G  1970.  A  new  species  of  Narecho 
(Hemiptera,  Cicadelloidea:  Nirvanidae)  from 
Africa.  Novos  Taxa  Entomologicos  No.  83,  1-7. 


GENUS  FLEXAMIA:  NEW  SPECIES,  PHYLOGENY,  AND  ECOLOGY 

Robert  F.  Whitcomb'  and  Andrew  L.  Hicks' 

Abstract — Eight  new  species  of  Flexamia  DeLong  are  described,  and  two  Mexican  species  previously  syn- 
onymized  are  reinstated.  Species  are  assigned  to  13  species  groups;  keys  are  presented  for  the  groups  and  for  the  44 
recognized  species.  Host  data,  in  many  cases  with  ohgophagy  coefficients,  are  presented  for  37  of  the  species.  Many 
species  speciaUze  on  native,  dominant,  perennial,  choridoid  or  panicoid  grasses;  some  are  monophagous.  Seven  sister 
species  pairs  specialize  on  the  same  (or  a  closely  related)  grass  species;  in  addition,  four  closely  related  species  appear  to 
be  restricted  to  Bouteloua  curtipendtila.  Few  if  any  specialists  colonize  their  host  throughout  its  entire  range. 
Ecological  factors  such  as  phenology  and/or  host  patchiness  strongly  influence  geographic  distribution.  An  intuitive 
phylogeny  is  proposed  and  is  used  as  a  basis  for  constructing  18  character  transformations.  The  genus  Flexamia 
probably  originated  in  Mexico  by  division  of  an  ancestral  lineage  from  which  the  modern  genus  Spartopyge  also 
diverged. 


The  first  specimen  of  the  genus  Flexa7nia 
DeLong  was  apparently  collected  at  Jack- 
sonville, Florida,  in  the  height  of  the  Lin- 
naean  explosion;  it  was  deposited  in  the 
British  Museum  (Natural  History)  and  named 
Jassus  productus  by  Walker  (1851).  In  their 
studies  of  Iowa  prairies  in  the  latter  part  of  the 
nineteenth  century,  Osborn  and  Ball  encoun- 
tered other  species,  which  they  referred  to 
the  genus  Deltocephalus.  DeLong  (1926) 
erected  a  subgenus  Flexamia  DeLong  within 
Deltocephalus  Burmeister.  DeLong  and 
Sleesman  (1929)  recognized  that  Delto- 
cephalus was  an  unworkably  large  assemblage 
of  distantly  related  species  and  divided  the 
genus  into  several  genera  (e.g.,  Laevi- 
cephalus  DeLong  &  Sleesman,  Pohjamia  De- 
Long  &  Sleesman,  and  Flexamia).  The  genus 
concept  for  Flexamia  was  refined  by  Oman 
(1949)  in  his  revision  of  North  American  ci- 
cadellid  genera,  but  the  modern  generic  con- 
cept was  established  by  Young  and  Beirne 
(1958).  These  authors  segregated  Spartopyge 
Young  &  Beirne,  synonymized  Secopennis 
Oman,  described  genitalic  characters  of  males 
and  females  (the  first  use  of  genitalic  charac- 
ters for  Flexamia  females),  and  presented  a 
species  key.  The  30  species  they  described 
included  all  of  the  widely  distributed  and 
commonly  collected  representatives  of  the 
genus,  and  their  key  proved  to  be  accurate 
and  easy  to  use.  Young  and  Beirne  also  de- 
scribed possible  relationships  among  species 


and,  without  specifying  an  explicit  phylogeny, 
proposed  broad  outlines  of  hypothetical  evo- 
lutionary events  in  the  genus.  The  genus  was 
incorrectly  synonymized  with  Acurhinus  Os- 
born by  Linnavuori  (1959).  Comparison  of 
the  specimens  (all  females)  of  Acurhinus 
(  =Dorydium)  maculatum  Osborn  in  the  Ohio 
State  University  collection  led  Linnavuori 
and  DeLong  (1978)  to  synonymize  Acurhinus 
with  the  African  stirelline  genus  Hododoecus 
Jacoli.  The  genus  Flexamia  as  defined  herein 
is  an  entirely  North  American  taxon  related  to 
Spartopyge,  to  Alapus  DeLong  &  Sleesman, 
and,  perhaps  distantly,  to  Aflexia  Oman.  The 
closest  Old  World  relative  may  be  Enantio- 
cephalus  Haupt. 

Although  the  taxonomic  status  of  Flexamia 
has  been  generally  adequate  since  1958,  host 
data  have  been  difficult  to  obtain.  Earlier  col- 
lectors were  not  knowledgeable  agrostologists 
and  often  reported  only  that  deltocephaline 
species,  including  Flexamia,  resided  on 
"grasses. "  Astute  collectors  such  as  Osborn 
and  Ball  recognized  the  importance  of  biologi- 
cal data,  and  their  notes  contain  some  host 
records.  In  1967,  H.  H.  Ross  encouraged  the 
senior  author  to  ascertain  the  hosts  of  Flex- 
amia species,  in  large  part  to  test  his  intuitive 
belief  that  host  relationships,  especially  host 
transfers,  could  be  invoked  to  explain  specia- 
tion  of  phytophagous  insects.  Accordingly,  we 
sought  to  determine  hosts  of  the  species  de- 
scribed by  Young  and  Beirne.  Much  to  our 


'insect  Pathology'  Laboratory,  AEriciiltural  Research  Service,  Beltsville,  Maryland  20705. 


224 


1988 


Whitcomb,  Hicks:  Flexamia  Revision 


225 


Fig.  1.  Flexamia  serrata. 

surprise,  in  the  course  of  this  search,  unde- 
scribed  species  emerged,  particularly  when 
we  examined  dominant,  perennial  Muhlen- 
bergia  or  Bouteloua  species  (Whitcomb  et  al. 
1986,  1987).  Under  these  circumstances,  it 
seems  appropriate  to  review  the  species  con- 
cept and  to  emend  the  generic  concept, 
which,  although  basically  intact  in  its  essential 
aspects,  must  accommodate  the  new  species. 

Genus  Flexamia  DeLong 

Deltocephalus  subgenus  Flexamia  DeLong  1926:20,  22. 

Type  species:  Deltocephalus  reflextis  Osborn  & 

Ball  1897,  by  original  designation. 
Flexatnius,  DeLong  and  Sleesman  1929:82. 
Flexamius  subgenus  Secopennis  DeLong  &  Sleesman 

1929:85.    Type    species:    Deltocephalus   slossoni 

Ball. 
Acurhinus  Oshorn,  Linnavuori  1959:89. 
Flexamia  DeLong,  Young  and  Beirne  1958;  Linnavuori 

and  DeLong  1978:208. 

Small  deltocephaline  leafhoppers.  Length 
of  male  2.4-5.0  mm,  of  female  2.6-5.0  mm. 
Head  produced,  median  length  of  crown  usu- 
ally greater  than  interocular  width;  clypellus 
broader  at  base  than  at  apex;  forewing  with 
reflexed  veinlets  to  costal  margin  in  vicinity  of 
small,  outer  anteapical  cell. 

Dried  specimens  yellowish  brown  or  gray, 
with  ferruginous  to  black  markings;  live  in- 
sects in  some  cases  with  green  pigmentation 
changing  to  orange  or  stramineous  when 
dried.  Apical  area  of  crown  partly  or  com- 
pletely surrounded  by  darker  markings,  in 
latter  case  with  clearly  defined,  apical  areolet. 
Crown  often  with  pair  of  oblique,  dark  mark- 
ings at  base;  pair  of  transverse,  dark  markings 
at  about  midlength;  pair  of  darker  markings 
parallel  to  lateral  margins  near  apex.  Face 
color  varying  from  pale  to  black,  often  useful 


in  species  discrimination.  Pronotum  of  some 
species  with  conspicuous,  longitudinal 
stripes,  but  many  species  with  faint,  wider 
stripes.  Veins  of  forewings  usually  dark-mar- 
gined; distinct,  dark-margined  areole  fre- 
quently near  center  of  corium. 

Male, — Plates  usually  shorter  than  pygo- 
fer,  frequently  fused  along  mesal  margins; 
pygofer  usually  with  pronounced  posterior 
lobe  in  lateral  aspect;  style  with  longitudinally 
grooved,  preapical  lobe,  which  is  usually  not 
pronounced;  connective  linear,  arms  contigu- 
ous basally,  sometimes  fused  to  aedeagus. 
Connective  with  upper  portion  less  heavily 
sclerotized  than  lower  portion,  often  thinner 
and  extended  dorsad  in  thin  keel  that  varies  in 
width  interspecifically;  dorsal  aedeagal  apo- 
deme  with  pair  of  conspicuous  appendages 
directed  caudoventrad  or  caudad.  Aedeagus 
interspecifically  variable,  of  great  value  in 
species  discrimination. 

Female. — Sternum  VII  nearly  always 
longer  than  VI,  often  2X  or  more;  hind  margin 
with  2-4  teeth  and  usually  small,  median 
notch.  Pygofer  with  well-dispersed  setae; 
ovipositor  barely  reaching  pygofer  apex. 

Flexamia  Species  Groups 

In  the  following  discussion,  we  describe  13 
species  groups  and  provide  keys  to  species. 
We  describe  8  new  species  and  reinstate  2 
species  previously  synonymized.  We  also  re- 
view important  characteristics  of  previously 
described  species,  emphasizing  diagnostic 
morphology,  ecology,  and  biogeography.  Our 
brief  descriptions  of  previously  described  spe- 
cies complement  the  descriptions  of  Young 
and  Beirne  (1958). 

Key  to  Flexamia  Species  Groups 

1 .  Dorsum  with  pair  of  stripes  extending  from  hind 
portion  of  disc  of  crown  across  pronotum  and 
scutellum  (Figs.  2A-D) 2 

—  Dorsum  without  such  stripes  (Figs.  2E,F) 4 

2(1).   Individuals  large,  at  least  4.5  mm   

IV.  grammica  group 

—  Individuals  smaller 3 

3(2).   Forewing  broadly  truncate  (Fig.  2D);  Arizona 

VIII.  ritana  group 

—  Forewing  obliquely  truncate  (Fig.  2B)  or  not 

truncate  (Fig.  2A)  (prairie);  Florida 

VII.  alhida  group 

4(1).  Face  entirely  black,  genae  at  most  with  oblique, 
pale  streak;  crown  without  midlength  transverse 
lines    5 


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Great  Basin  Naturalist  Memoirs 


No.  12 


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


1988 


Whitcomb,  Hicks:  Flexamia  Revision 


227 


Fig.  3.  Faces  of  Flexamia  species:  A,  serrata-  B,  stylata;  C,  prairiana-  D,  prairiana;  E,  picta;  F,  decora;  G,flexulosa; 
H,  sandersi. 


—  Face  black  basally,  pale  apically,  or  brownish 
in  part;  at  least  genae  pale,  crown  usually  with 
midlength  transverse  lines 6 

5(4).  Forewing  with  conspicuous  black  spot  (Fig. 
2F)  in  corium;  crown  with  median  length 
more  than  one-half  greater  than  interocular 
width    XII.  areolata  group 

—  Forewing  without  conspicuous  black  spot  in 
corium;  crown  with  median  length  less  than 

one-half  greater  than  interocular  width 

XI.  impiitans  group 

6(4).  Face  pale  with  several  narrow,  parallel,  fus- 
cous, interocular  lines  interrupted  at  middle 

(Fig.  3A);  northwestern  (Fig.  30) 

XI.  serrata  group 

—  Face  with  (Figs.  3B,D-G)  or  without  (Figs. 
3C,H)  interocular  lines  but  not  as  above 7 

7(6).  Either  with  crown  more  than  twice  interocu- 
lar width  (Fig.  2E)  {pyrops  Crumb),  or  with 
conspicuous,  dark,  apical  wing  blotch  (Fig. 
4C)  and  sordid  yellow  face  with  black  interoc- 
ular band  {picta  [Osborn]);  viewed  laterally, 
male  pygofer  with  conspicuous,  acutely  angu- 
late,  ventral  lobe  (Figs.  5A,B);  aedeagus  (Fig. 
6D)  with  pair  of  anteapical  processes  and  api- 
cal, flangelike  processes VI.  picta  group 

Habitus  not  as  above;  male  pygofer  with  pos- 
terior lobe  rounded  ventrally;  aedeagus  not  as 
above    8 

8(7).        Males   9 

—  Females 14 

9(8).       Aedeagus  and  connective  fused  (Figs.  6F,G) 

XIII.  prairiana  group 


—  Aedeagus  and  connective  distinctlv  articu- 
lated (Figs.  6A-C,  E)    ' 10 

10(9).     Aedeagus  without  apical  processes  (Fig.  6C)     11 

—  Aedeagus  with  (at  least  minute)  apical 

processes 12 

11(10).    Male  plates  broad,  not  tapered  (Figs.  IIA-I) 

I.  pectinata  group 

—  Male  plates  more  elongate  and  tapered  (Figs. 
9C,F)   III.  zacate  group 

12(10).  Aedeagus  (Figs.  7F,G)  with  2  pairs  of  pro- 
cesses   V.  curvata  group 

—  Aedeagus  not  so 13 

13(12).   Aedeagus  with  2  processes  on  shaft  (Fig.  7H) 

II.  abbreviata  group 

—  Aedeagus  with  3  processes  on  shaft  (Figs.  6E, 
31A,D-M)   X.  flextdosa  group 

14(8).  Face  white,  ivory,  or  very  pale  yellow, 
sharply  contrasting  with  black  interocular  line 
(Figs.  3B,F,G) X.  flexulosa  group  (in  part) 

—  Face  with  lower  part  not  white  or  pale  (Fig. 
3y),  or  if  pale,  not  contrasting  sharply  with 
interocular  band,  or  interocular  band  brown, 
not  black  (Fig.  3D)    15 

15(14).    Body  color  stramineous    16 

—  Body  color  not  stramineous   17 

16(15).  Hind  margin  of  sternum  VII  with  medial  pro- 
jection (Fig.  lOO) 

X.  flexulosa  group  (inflata  subgroup) 

—  Hind  margin  of  sternum  VII  without  projec- 
tion     20 


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Great  Basin  Naturalist  Memoirs 


No.  12 


Fig.  4.  Wings  of  Flexamia  species:  A,  inflata;  B,  cur- 
vata;  C,  picta. 


17(15).  Sternum  VII  (Fig.  lOE)  with  median,  trape- 
zoidal projection,  with  a  medial  notch,  usu- 
ally with  smaller,  shallower  notches  on  either 
side II.  abbreviata  group 

—  Sternum  VII  not  as  above 18 

18(17).    Sternum  VII  produced  medially  in  a  distinct 

lobe  or  projection  (Figs.  10S,T) 

XIII.  prairiana  group 

—  Sternum  VII  not  produced  medially  in  dis- 
tinct lobe,  or  if  produced,  with  deep,  median 
incision 19 

19(18).  Hind  margin  of  sternum  VII  incised  to  form 
pattern  of  four  teeth;  hind  margin  usually 
heavily  infuscated  on  hind  margin  in  vicinity 
of  teeth  (Figs.  14A-H)    I.  pectinata  group 

—  Hind  margin  of  sternum  VII  not  so 20 

20(19).  Hind  margin  of  sternum  VII  slightly  concave 
on  each  side  of  median  excision;  infuscated 
notches  usually  contrasting  sharply  with 
stramineous  color  of  sternum  (Figs.  10J,K) 
V.  curvata  group 

—  Hind  margin  of  sternum  VII  not  so  (Figs. 
101, L) III.  zacate  group 

I.  The  pectinata  Group 
The  greatest  proposed  changes  in  Flexamia 


species  concepts  are  in  the  pectinata  species 
group  to  which  we  add  five  new  species  and 
reinstate  two  others.  All  species  of  the  group 
have  aedeagal  shafts  without  apical  processes. 
Although  aedeagal  characters  are  not  as  useful 
for  species  discrimination  in  this  group  as  in 
other  groups,  the  male  plates  and  female  ster- 
num VII  provide  useful  apomorphies  that  de- 
fine the  group.  The  plates  (except  in  collo- 
rurn)  are  broad  and  basally  parallel-sided.  The 
hind  margins  of  the  female  sternum  VII  of  all 
species  possess  four  medial  teeth  with  sur- 
rounding infuscation  (but  note  the  extensive 
modification  in  mescalero  [Fig.  21H]).  The 
size  and  shape  of  the  teeth  vary  interspecifi- 
cally.  The  species  are  largely  southwestern 
and/or  Mexican,  and  many  appear  to  have 
small  ranges.  The  nominate  species  F.  pecti- 
nata, however,  has  a  wide  geographic  range. 
As  defined  by  Young  and  Beirne  (1958),  this 
species  ranges  from  western  Mexico  {zamora 
DeLong  &  Hershberger)  through  the  eastern 
highlands  (Valles  and  Monterrey  [7ninima 
DeLong  &  Hershberger]),  the  Davis  Moun- 
tains of  west  Texas,  Otero  and  Guadalupe 
counties  of  New  Mexico  north  to  North  Da- 
kota, and  as  far  east  as  Akron,  Ohio,  in  the 
prairie  peninsula.  However,  our  studies  indi- 
cate that  zamora  and  minima  should  be  rein- 
stated. Recognition  herein  of  the  new  species 
batidarita,  gila,  coUorum,  jacala,  and  mesca- 
lero makes  it  clear  that  this  group  is  diverse 
and  has  speciated  frequently.  The  host  of 
pectinata  is  side-oats  grama  (Bonteloua  cur- 
tipendula),  a  widespread  prairie  species.  In 
addition  to  pectinata,  there  are  three  south- 
western side-oats  grama  specialists  of  the 
pectinata  group.  Flexamia  doeringae  and  F. 
gila  reside  in  the  southeastern  Arizona  moun- 
tains and  Mogollon  Rim  region  and  extended 
conifer-oak  savanna  of  Mexico,  respectively. 
Flexamia  bandarita,  as  far  as  known,  is  con- 
fined to  the  Chisos  Mountains  and  Marathon 
Basin  of  Trans-Pecos  Texas.  We  suspect  that 
side-oats  grama,  which  also  has  a  wide  range 
in  Mexico,  will  prove  to  be  the  host  of  some 
Mexican  species  of  the  pectinata  group.  The 
most  distinctive  species  of  the  group  {mesca- 
lero), however,  appears  to  specialize  on  Muh- 
lenhergia  pauciflora. 

Several  factors  may  account  for  speciation 
in  the  pectinata  group.  Side-oats  grama  is  it- 
self a  complex  of  ecotypes  that  vary  greatly  in 
structure  and  seasonality  (Gould  1979).  This 


1988 


Whitcomb,  Hicks:  Flexamia  Revision 


229 


Fig.  5.  Pygofers  oi Flexamia  species:  A,  picta;  B,  pyrops;  C,  stylata;  D,  producta.  Redrawn  from  Young  and  Beirne 

(1958). 


Fig.  6.  Aedeagus  and  connective  of  Flexamia  species:  A,  curvata;  B,  abbreviata;  C,  canyonensis;  D,  picta;  E, 
flexulosa;  F,  prairiana;  G,  graminea.  Redrawn  from  Young  and  Beirne  (1958).  Aspects;  A-F,  caudoventral;  G,  lateral. 


chloridoid  grass  crosses  many  biome  lines  and 
is  a  dominant  in  mesic  Sonoran  and  Chi- 
huahuan  grasslands  and  in  montane  systems 
within  these  semiarid  grasslands.  It  is  com- 
mon through  much  of  the  prairie  and  occurs 
eastward  on  nonforested  slopes  in  the  Ap- 
palachian mountains.  Like  other  major  domi- 
nants of  North  American  grasslands,  there- 
fore, side-oats  grama  has  a  much  wider 
geographic  range  than  any  of  the  cicadellid 
species  that  exploit  it. 

The  Chihuahuan  desert  system  presents  a 
major  barrier  to  geographical  continuity  of 
prairie  grasses,  and  side-oats  grama  is  no  ex- 
ception. Many  prairie  cicadellid  taxa  occur 
west  of  this  desert  system,  but,  depending 
upon  the  case,  the  taxonomic  results  vary.  For 
example,  populations  related  to  the  buffalo- 
grass  specialist  Athysanella  (Gladionura) 
emarginata  occur  in  disjunct  stands  of 
Buchloe  in  New  Mexico's  Hidalgo  County. 
This  population,  until  very  recently,  had  been 
regarded  as  a  subspecies  (sobrina)  but  will  be 
elevated  to  a  species  in  a  forthcoming  revision 


oi  Gladionura  (H.  D.  Blocker,  personal  com- 
munication). On  the  other  hand,  forms  of  A. 
(Gladionura)  argenteola  (a  Botiteloua  gracilis 
specialist)  in  southeastern  Arizona  are  not  dis- 
tinguishable from  short-grass  prairie  forms. 
The  connecting  link  between  the  relatively 
mesic,  temperate  grasslands  of  the  northern 
Mexican  highlands  and  the  prairie  is  tenuous 
at  best,  consisting  of  a  small  set  of  "stepping 
stone"  montane  islands  within  the  Chi- 
huahuan desert  of  Coahuila  and  Chihuahua, 
or  of  the  relatively  narrow  grassland  periph- 
ery surrounding  the  desert  on  the  east  and 
west  and  meeting  at  the  south  in  the  state  of 
San  Luis  Potosi.  Also,  the  mesquite-acacia 
savanna  and  Trans-Pecos  shrub  savanna  of 
Texas  are  further  barriers  to  geographic  conti- 
nuity of  B.  curtipendula.  During  full  glacia- 
tions,  these  arid  regions  had  climates  that 
were  much  more  moist  than  they  are  today 
(Van  Devender  and  Burgess  1985). 

One  of  the  new  species,  collorum,  from  the 
Edwards  Plateau  of  central  Texas,  is  appar- 
ently a  specialist  on  Nealley  grama,  Bouteloua 


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Great  Basin  Naturalist  Memoirs 


No.  12 


LT     CA 


CV      LT 


Fig.  7.  Aedeagal  apices  oi Flexamia  species;  A,  scrrata-  B,  alhida;  C,  slossonac-  D,  litana-  E,  ^rammica;  F,  curvata; 
G,  sitrcuhi;  H,  abbreviatii;  I,  zflcafc,  J,  canijonensis;  K,  /;icf«,  L,  ptjrop.s.  Aspects:  AD,  anterodorsal;  CA,  caudal;  CV, 
caudoventral;  DS,  dorsal;  LT,  lateral.  Redrawn  from  Young  and  Beirne  (1958). 


uniflora.  Flexamia  coUonim  may  be  related  to 
another  new  species,  jacala,  whose  type  lo- 
cahty  in  Mexico  is  close  to  the  intersection  of 
eastern  and  western  montane  Mexican  grass- 
lands. The  latter  species  is  represented  by  a 
single  male  specimen  from  the  USNM,  so 
nothing  is  known  about  its  biology. 

Finally,  another  new  species,  niescalero, 
discovered  in  a  single  location  in  southeastern 
New  Mexico,  presumably  arose  by  host  trans- 
fer to  New  Mexican  muhly,  Muhlenbergia 
paucijlora. 

Description  of  the  pectinata  Group 

Medium-sized  to  small  deltocephaline  leaf- 
hoppers.    Length  of  6    2.7-3.8  mm,   of  9 
2.8-4.2  mm.  Species  with  general  facies  of 
genus;  brown  or  in  dark  specimens,  almost 
black.  Crown  variably  produced.  \6]  Plates 


(Figs.  IIA-I)  elongate,  broad,  parallel-sided 
basally  (except  coUorum).  Pygofer  with  dis- 
tinct, posterior  lobe,  strongly  constricted  in 
lateral  aspect,  both  dorsally  and  ventrally. 
Aedeagus  and  connective  distinctly  articu- 
lated. Aedeagus  (Figs.  12A-I)  with  no  apical 
processes,  or  at  most  slightly  capitate 
(mescalero),  often  appearing  flared  in  dorsal 
aspect.  [Note:  Caution  must  be  exercised 
when  using  the  aedeagal  apex  as  a  diagnostic 
character,  since  this  structure  is  membranous 
and  is  subject  to  tearing  and  breaking  during 
copulation  or  preparation  of  the  specimen.] 
Styles  variable  interspecifically  (Figs.  13A-I). 
Gonopore  subapical  on  caudoventral  surface. 
[  9  ]  Sternum  VII  (Figs.  14A-H)  with  four  me- 
dial teeth  on  hind  margin;  teeth  varying  inter- 
specifically  in  length  and  shape,  surrounded 
by  infuscated  region. 


1988 


Whitcomb,  Hicks;  Flexamia  Revision 


231 


Fig.  8.  Male  plates  and  pygolers  oi  Flexamia  species:  A,  alhida:  B,  slossonae:  C,  serrata,  D,ritana;  E,  abbreviata;  F, 
grammica. 


Fig.  9.  Male  plates  and  pygofers  oi  Flexamia  species:  A,  picta;  B,  pyrops;  C,  canyunensis;  D,  curvata; 
E,  surcula;  F,  zacate. 


232 


Great  Basin  Naturalist  Memoirs 


No.  12 


Fig.  10.  Female  sternum  VII  of  Flexarnia  species:  A,  albida^  B.  slossonac,  C,  ^errata^D,  t"7;,L"  o'llXa-  P 
grammica;  G,  picta;  H,  pyrops;  I,  canyonensis;],  surcula;  K,  curvata;  L,  zacate;  M,flexulosa;  N,  decora,  O,  mjlata,  F, 
imputans;  Q,  areolata;  R,  sfyiata;  S,  prainana; T,  sandersi;  U,  fitammea. 


1988 


Whitcomb,  Hicks:  Flexamia  Revision 


233 


Fig.  11.  Male  plates  and  pygofers  oiFlexamia  pectinata  group,  ventral  aspect:  A,  mminui;  B 
D,  handarita-  E,  gila;  F,  doeringae;  G,  collorurn;  HJacala;  I,  mescalero. 


zamora;  C,  pectinata; 


234 


Great  Basin  Naturalist  Memoirs 


No.  12 


A  B  u  u  t  .  -  ^ 

Fig.  12.  Aedeagal  apices  of  Flexamia  pectinata  group,  caudoventral  aspect:  A,  minmui;  B,  zainora;  C,  pectinata;  D, 
bandarita;  E,  gila;  F,  doeringae;  G,  coUorum;  H,jacala;  I,  mescalero. 


Key  to  Males  of  the  pectinata  Group 
1.       Plates  long  and  parallel-sided  at  bases 2 

—  Plates  short,  not  parallel-sided 

collorum  Whitcomb  &  Hicks 

2(1).   Plates  subrectangular  (Figs.  IIC.F)    3 

—  Plates  with  apices  rounded 4 

3(2).  Aedeagus  in  ventral  aspect  (Fig.  12F)  with  pro- 
tuberance close  to  tip;  face  pale;  Arizona   

doeringae  Beamer  &  Tuthill 

—  Aedeagus  in  ventral  aspect  with  protuberance 
considerably  removed  from  tip  (Fig.  12C);  face 
brown;  prairie   pectinata  (Osborn  &  Ball) 

4(2).   Plates  (Fig.  11  A)  with  apices  acute;  Mexico   .  .  . 

minima  DeLong  &  Hershberger 

—  Plates  with  apices  not  acute  5 

5(4).   Plates  almost  as  long  as  pygofer  (Fig.  HE)  .... 

gila,  n.  sp. 

—  Plates  no  more  than  3/4  length  of  pygofer 6 

6(5).  Aedeagus  expanded  apically  in  ventral  aspect, 
appearing  bluntly  saggitate  (Figs.  121,  22)  .... 
mescalero,  n.  sp. 

—  Aedeagus  not  expanded  apically   7 

7(6).   Style  apex  (Fig.  13H)  avicephaliform  in  lateral 

aspect jacala,  n.  sp. 

—  Style  apex  not  avicephaliform    8 

8(7).   Style  apex  (Fig.  13D)  ventrally  directed,  chelate 

bandarita,  n.  sp. 

—  Style  apex  (Fig.  13B)  not  chelate 

zamora  DeLong  &  Hershberger 

Key  to  Females  of  the  pectinata  Group 

1.        Sternum  VII  with  4  teeth  of  approximately  equal 

length  and  3  relatively  shallow  incisions 2 

—  Sternum  VII  with  deep  incisions  and/or  teeth  of 
different  length  4 

2(1).  Sternum  VII  with  infuscations  on  either  side  of 
middle  incision  (Fig.  14G);  Texas,  host  Butite- 
loua  uniflora    coUorum,  n.  sp. 

—  Sternum  VII  not  as  above 3 

3(2).   Sternum   VII  with   heavy   infuscation   around 


teeth  on  hind  margin  (Fig.  14C);  USA,  prairie 
pectinata  (Osborn  &  Ball) 

—  Sternum  VII  with  light  infuscation  on  hind  mar- 
gin (Fig.  14A);  Mexico 

minima  DeLong  &  Hershberger 

4(1).   Sternum  VII  with  outer  teeth  longer  than  inner 

teeth 5 

—  Sternum  VII  with  inner  teeth  longer  than  outer 
teeth 6 

5(4).  Sternum  VII  with  outer  teeth  sharply  pointed, 
median  incision  shallow  (Fig.  14B);  Mexico  .  .  . 
zamora  DeLong  &  Hershberger 

—  Sternum  VII  with  outer  teeth  blunt,  median 
incision  deep  (Fig.  14H) mescalero,  n.  sp. 

6(4).   Inner  teeth  of  sternum  VII  together  forming 

blunt,  median  projection  (Fig.  14F)    

doeringae  Beamer  &"Tuthill 

—  Inner  teeth  of  sternum  VII  separated,  acute  at 
tip,  separated  by  tapered  incision   7 

7(6).   Sternum  VII  with  three  incisions  of  equal  depth 

(Fig.  14E) gila,  n.  sp. 

—  Sternum  VII  with  median  incision  deeper  than 
lateral  incisions  (Fig.  14D) bandarita,  n.  sp. 

1.  Flexamia  minima  DeLong  & 
Hershberger,  n.  stat. 

Flexamia  minima  DeLong  &  Hershberger  1947:  138. 
Flexamia  pectinata.  Young  &  Beirne  1958:  46.  (In  part.) 

Length  of  c?  2.9  (2.8-2.9)  mm,  9  3.0 
(2.7-3.2)  mm;  head  width  of  6  0.84  mm,  9 
0.92  mm.  Crown  produced;  median  length  of 
crown  0.74  x  head  width  and  L45  x  interocu- 
lar  width  (c?  n  =  6;  9  n  =  3).  Crown  hght 
brown  to  pale  yellow  with  dark,  circular  spot 
around  apex,  pair  of  broken,  transverse  lines 
at  midlength,  two  obli(|ue  markings  at  rear, 
appearing  continuous  with  the  medial  pair  of 
six  pale  brown,  pronotal  stripes.  Face  with 
broad,  dark  brown,  interocular  band,  fading 
to  pale  yellowish.    Brown  basally,  disc  and 


1988 


Whitcomb,  Hicks:  Flexamia  Rf.vision 


235 


blX 


Fig.  13.  Styles  and  style  tips  oiFlexamia  pectinata  group:  A,  minima;  B,  zamora;  C,  pectinate;  D,  bandarita;  E,  gila; 
F,  doeringae;  G,  collorum;  H,jacala;  I,  mescalero. 


apex  of  forewings  usually  with  irregular,  dark 
spot. 

Male. — Pygofer  much  like  that  o( pectinata 
but  shorter;  plates  (Fig.  11  A)  barely  exceed- 
ing ventral  lobe  in  lateral  view  or  2/3  length  of 
pygofer,  unevenly  tapering  to  blunt  apices. 
Genitalia  similar  to  those  of  pectinata  and 
zamora,  but  little  more  than  half  their  size.  In 
ventral  aspect,  apex  of  style  not  exceeding 
plane  of  well-developed,  preapical  lobe,  angle 
formed  between  preapical  lobe  and  style  apex 
more  acute  than  in  pectinata.  Connective 
similar  to  that  of  pectinata,  but  with  a  nar- 
rower, deeper,  U-shaped  incision  at  the  joint 
with  the  aedeagus.  Other  characters  as  in 
pectinata. 

Female. — Sternum  VII  with  posterior 
margin  broadly,  shallowly  emarginate,  hind 
margin  with  four  teeth  of  approximately  equal 
length  (Fig.  14A),  lightly  infuscated.  Oviposi- 
tor with  bases  of  first  valvulae  as  in  zamora. 

Types. — Holotype  i:  Valles,  San  Luis  Po- 
tosi,  Mexico,  1  December  1938,  J.  S.  Cald- 
well. 

Remarks. — An  additional  series  o{ minima, 
collected  by  E.  D.  Ball  at  Monterrey,  Nuevo 
Leon  (5d,  39,  11  August  1936,  USNM),  has 


been  examined.  On  the  basis  of  characters  of 
the  male  plates,  the  styles,  and  the  distinc- 
tiveness of  the  female  sternum  VII  from  that 
of  zamora,  we  reinstate  minima.  Species 
problems  in  this  group  are  discussed  under 
"Species  Concept."  This  species,  like  zamora, 
may  be  entirely  Mexican  in  distribution  and  is 
represented  by  the  two  collections  discussed 
above  (Fig.  15).  The  host  or  hosts  are  un- 
known, but  the  closely  related  pectinata  spe- 
cializes on  Bouteloua  curtipendula.  The 
grasslands  of  San  Luis  Potosi  are  extensive, 
and  many  other  warm  season  grasses,  includ- 
ing other  Bouteloua  spp.,  are  present  (Rze- 
dowski  1966);  further  fieldwork  is  required  to 
clarify  the  biology  oiininima. 

2.  Flexamia  zamora  DeLong  & 
Hershberger,  n.  stat. 

Flexamin  zamora  DeLong  and  Hershberger  1947:  137. 
Flexamia  pectinata.  Young  and  Beirne  1958:  46.  (In  part.) 

Length  of  c?  3.6  (3.5-3.7)  mm,  9  3.9 
(3.7-4.2)  mm;  head  width  of  S  1.08  mm,  9 
1.15  mm.  Crown  not  strongly  produced;  me- 
dian length  of  crown  0.61  x  head  width  and 
1.27  X  interocular  width  ((5  n  =  7;  9  n  =  3). 


236 


Great  Basin  Naturalist  Memoirs 


No.  12 


Fig.  14.  Female  sternum  VII  ofFlexamia  pectinata  group:  A,  minima:  B,  zamora:  C,  pectinata:  D,  bandarita;  E,  gila; 
",  doeringae;  G,  collorum;  H,  mescalero. 


Crown  light  brown  with  dark,  circular  spot 
around  apex,  pale  brown,  transverse  line  on 
either  side  at  middle.  Pronotum  light  brown, 
usually  with  traces  of  four  wide  longitudinal 
stripes.  Elytra  subhyaline,  dark  brown  spot 
on  disc;  costal  veinlets  heavily  margined  with 
dark  brown.  Face  black  above,  pale  brown  on 
lower  portion. 

Male. — Pygofer  as  in  pectinata.  Plates 
(Fig.  IIB)  extending  to  approximately  3/4 
length  of  pygofer,  with  apices  less  produced 
than  in  pectinata,  bluntly  rounded.  Style 
apices  exceeding  the  plane  of  the  moderately 
developed,  preapical  lobe,  forming  an  angle 
with  it  smaller  than  that  in  pectinata.  Other 
characters  as  in  pectinata. 

Female. — Sternum  VII  (Fig.  14B)  with 
posterior  margin  shallowly  emarginate;  four 
produced  teeth  at  middle;  outer  teeth  longer, 
acute,  curved  inward.  Ovipositor  with  basal 
processes  of  first  valvulae  similar  to  those  of 
pectinata,  but  recurved  portion  not  separated 
along  the  middle. 

Types. — Holotype  6:  Zamora,  Michoacan, 
Mexico,  2  October  1941,  Plummer,  Good, 
Caldwell  and  DeLong  (OSU).  Paratvpes:  5  6 , 
3  $ ,  same  collection  data  (OSU  and  USNM). 

Remarks. — Although  the  male  genitalia  of 


zamora  are  similar  to  those  o{  pectinata,  the 
styles  oi  zamora  differ  in  having  a  more  acute 
angle  between  the  apex  and  the  less  well- 
developed  preapical  lobe.  Also,  the  plates  are 
more  evenly  rounded  than  those  of  pectinata 
and  longer  than  those  of  minima.  The  female 
sternum  VII  of  zamora  differs  from  that  of 
pectinata  in  that  the  outer  teeth  of  the  hind 
margin  are  much  longer  than  the  inner  teeth; 
in  pectinata  and  minima  the  teeth  are  of  equal 
length.  This  species,  almost  certainly  entirely 
Mexican  in  distribution,  is  known  only  from 
its  type  locality  (Fig.  15).  The  host  or  hosts  are 
unknown,  but  other  members  of  the  complex 
feed  on  Bouteloua  curtipendula. 

3.  Flexamia  pectinata  (Osborn  &  Ball) 

Dehocephahis  pectinatus  Osborn  &  Ball  1897:  205. 
Deltucephalus (Flexamia) pectinatus  DeLong  1926:  32. 
Flexamius  pectinatus.  DeLong  and  Sleesman  1929:  83. 
Flexamius  zatnora.  Young  and  Beirne  1958:  46.  Incorrect 

synonymy. 
Flexamia  minima,  Yoimg  and  Beirne  19.58:  46.  Incorrect 

synonymy. 

Important  CHARACTERS. — Length  of  S  3.4 
(2.9-3.8),  of  9  3.6  (3.1-4.0).  Head  slightly 
produced  (median  crown  length  1.40  x  inter- 
ocular  width;  0.62  X  head  width)  (c?  n  =  37; 


1988 


WHITCOMB,  HiCKS:  FLEXAMIA  REVISION 


237 


Fig.  15.  Geographic  distribution  of  Flexaniia  pectinata  {i 
miniirui  (O),  And  jacala  (ir)  with  Botiteloua  curfipendtila. 


),  doeringae  (■),  bandarita  (A),  gila  (D),  zatnora  (A), 


9  n  =  29).  [6]  Pygofer  (Fig.  62F)  strongly 
constricted  both  ventrally  and  dorsally;  plates 
(Fig.  IIC)  wide  and  parallel-sided  for  much  of 
their  length,  almost  subrectangular,  but  with 
rounded  tips.  Aedeagus  (Fig.  12C)  symmetri- 
cal, without  lateral  processes;  gonopore  sub- 
apical  on  caudoventral  surface.  [9]  Sternum 
VII  (Fig.  14C)  with  four  teeth  of  approxi- 
mately equal  length.  Ovipositor  with  bases  of 
first  valvulae  recurved  and  calipterate  (Fig. 
63G). 

Geographic  distribution. — This  species 
is  a  specialist  of  side-oats  grama  {Bouteloiia 
curtipendula)  in  prairie  and  mesic  grasslands; 
it  occurs  (Fig.  15)  at  higher  elevations  of 
the  desert  plains  of  eastern  New  Mexico  east 
to  Akron,  Ohio,  the  eastern  extension  of  the 


prairie  peninsula  (Transeau  1935). 

Biology. — Like  other  Flexamia  species, 
pectinata  is  probably  multivoltine  and  is 
present  throughout  the  growing  season.  In 
New  Mexico  it  is  often  absent  from  its  host 
(Botiteloua  curtipendula,  side-oats  grama)  at 
lower  elevations  where  host  growth  may  be 
unpredictable  because  of  sporadic  drought.  In 
Chihuahuan  and  Sonoran  highlands  it  is  re- 
placed by  bandarita  and  doeringae,  respec- 
tively, and  in  the  intervening  mountains,  by 
gila. 

Oligophagy  coefficients. — Gramineae 
1.000;  Chloridoideae  0.973;  Bouteloua  0.926; 
Bouteloua  curtipendula  0.825  (n  =  57). 

Remarks  and  diagnosis. — Flexamia  pecti- 
nata can  be  recognized  by  a  combination  of  its 


238 


Great  Basin  Naturalist  Memoirs 


No.  12 


Fig.  16.  Flexamia  gila,  n.  sp. :  A,  aedeagus  and  connective,  lateral  aspect;  B,  aedeagus  and  connective,  dorsal  aspect; 
C,  apex  of  aedeagus,  caudoventral  aspect;  D,  male  plates  and  pygofers,  ventral  aspect;  E,  male  pygofer,  lateral  aspect; 
F,  right  style,  dorsal  aspect;  G,  ape.x  of  right  style,  lateral  aspect;  H,  female  sternum  VII;  I,  bases  of  first  valvulae  of 
female,  dorsal  aspect. 


external  genitalic  characters  and  its  geo- 
graphic range.  Other  species  (doeringae,  ja- 
cala)  of  the  pectinata  group  with  somewhat 
similar  male  plates  do  not  occur  with  pecti- 
nata in  its  range.  Of  the  nine  members  of  the 
group,  only  mescalero  (host:  Muhlenhergia 
pauciflora)  and  colloruin  (host:  Bouteloua 
uniflora )  have  ranges  that  overlap  the  range  of 
pectinata.  The  male  plates  (Fig.  Ill)  and  fe- 
male sternum  VII  (Fig.  14H)  of  mescalero 
distinguish  it  from  pec^inaf a  (Figs.  IIC,  14C). 
The  male  plates  oi collorum  (Fig.  IIG)  differ 
from  those  of  pectinata  in  general  aspect  and 
in  the  absence  of  the  sclerotized  ridge  on  the 
inner  plate  surface  that  characterizes  coUo- 
rum.  The  female  sternum  VII  of  collorum  is 
similar  to  that  ofpectinata.  Although  the  char- 
acters discussed  above  permit  tentative 
recognition  of  the  commonly  encountered 
pectinata,  definitive  identification  requires 
examination  of  the  genitalia. 


4.  Flexamia  gila,  n.  sp. 
Length  of  6   3.4  (3.1-3.7)  mm, 


9    3.8 


(3.6-4.0)  mm;  head  width  of  d  1.03  mm, 
9  1.08  mm.  Crown  not  strongly  produced; 
median  length  of  crown  approximately 
0. 63  x  head  width  and  1 .  32  x  interocular  width 
(d  n  =  6;  9  n  =  4). 

Crown  light  brown  with  dark,  circular  spot 
around  apex,  pale  brown,  transverse  line  on 
either  side  at  middle.  Pronotum  light  brown, 
usually  with  traces  of  four  wide,  longitudinal 
stripes.  Elytra  subhyaline,  dark  brown  spot 
on  disc;  costal  veinlets  heavily  margined  with 
dark  brown.  Face  black  above,  pale  brown  on 
lower  portion. 

Male. — Pygofer  (Fig.  16E)  much  as  in  ban- 
darita;  posterior  margin  slightly  more  pro- 
duced. Plates  (Fig.  16D)  broad,  gradually 
rounded  from  base  to  apex,  extending  to  ap- 
proximately 4/5  length  of  pygofer.  Connective 
in  lateral  view  with  dorsal  keels  narrow  (or 
rarely  absent),  approximately  1/4  height  of 
dorsal  apodeme;  apodemal  processes  as  in 
bandarita;  styles  (Fig.  16F)  long,  diverging 
apically  in  ventral  aspect,  preapical  lobe 
rounded,  produced  ventrad  and  laterad;  style 


1988 


Whitcomb.  Hicks;  Flexamia  Revision 


239 


portion  distad  from  preapical  lobe  gradually 
narrowing  to  embrowned,  strongly  divergent, 
acute  apex  in  ventral  aspect,  or,  in  lateral 
aspect  (Fig.  16G),  appearing  bladelike,  pro- 
duced ventrally  and  flattened,  bearing  minute 
teeth  along  the  irregularly  curved  ventral 
margin  of  style  ending  in  a  truncate  apex  that 
is  slightly  curved  ventrad.  Aedeagus  (Figs. 
16B,C)  symmetrical,  short,  slender.  Apical 
margin  flared  and  convex,  much  as  in  pecti- 
nata.  Distance  between  gonopore  and  apex 
less  than  in  pectinata.  Gonopore  subapical  on 
caudoventral  surface. 

Female.— Sternum  VII  (Fig.  16H)  with 
posterior  margin  broadly  but  shallowly  emar- 
ginate,  with  four  produced  teeth;  middle 
teeth  broad.  Ovipositor  with  basal  processes 
of  first  valvulae  (Fig.  161)  recurved;  recurved 
portion  constricted  above  proximal  margin, 
expanded  slightly  mesad  and  distad  into  dif- 
ferentially sclerotized  lobes. 

Types. — Holotype  6 :  New  Mexico,  Sierra 
Co.,  Kingston,  17  August  1985,  R.  F.  Whit- 
comb (6,300  ft,  IPL  001875,  Bouteloua  cur- 
tipendula).  Paratypes:  3  6  and  1  9,  same 
collection  data;  2  6  and  2  9 ,  Mexico,  5  km  S 
Casas  Grandes,  Chihuahua  (5,600  ft,  Boute- 
loua curtipendula,  3  September  1987,  A.  L. 
Hicks,  IPL  001774);  2  c?  and  1  9 ,  New  Mex- 
ico, Socorro  Co.,  Magdalena  Mtns.,  Water 
Canyon,  7,000  ft.,  16  August  1987,  IPL 
001756,  A.  L.  Hicks.  Deposited  BARC,  CNC, 
KSU,  KU,  OSU,  and  USNM. 

Remarks. — The  aedeagal  characters  of  gi/a 
are  intermediate  between  those  of  doeringae 
and  pectinata.  The  styles  in  gila  are  less  pro- 
duced ventrally  than  in  doeringae.  The 
longer,  bladelike  styles  of  gila  differentiate 
males  oi  gila  from  those  of  pectinata;  females 
can  be  readily  differentiated  on  the  basis  of 
the  morphology  of  the  female  sternum  VII 
and  the  bases  of  the  first  valvulae.  The  host  of 
gila  is  B.  curtipendula.  The  name  gila,  a  noun 
in  apposition,  denotes  the  New  Mexican 
mountains  where  the  holotype  was  collected. 

5.  Flexamia  bandarita,  n.  sp. 

Length  of  6  3.4  (3.3-3.5)  mm,  9  3.7 
(3.4-4.0)  mm;  head  width  of  6  1.09  mm,  9 
1.04  mm;  head  not  strongly  produced;  median 
length  of  crown  0.64  x  head  width  and  1.33  x 
interocular  width  (n  =  8  d ,  3  9 ). 

Color  variable,  ranging  from  very  dark  gray 
to  pale,  with  irregular,  dark  markings  on  dor- 


sum and  forewings;  face  very  dark  with  pale 
arcs  on  upper  half;  variable  in  lower  half. 
Head  stramineous  in  lighter  specimens  with 
paired,  transverse  lines  at  midpoint  of  crown 
and  obli(}ue  markings  at  rear;  venter  and  legs 
with  irregular,  fuscous  markings. 

Male. — Pygofer  strongly  produced  ven- 
trally in  rounded  lobe;  plates  (Fig.  17D) 
contiguous  on  basal  half,  diverging  apically 
into  narrow  V;  extending  to  approximately 
3/4  length  of  pygofer;  style  (Fig.  17F)  heavily 
sclerotized,  sinuate  in  ventral  aspect,  extend- 
ing almost  to  the  apex  of  the  aedeagus,  gradu- 
ally narrowing  to  acute,  ventrally  directed, 
chelate  apex  (Fig.  17G)  that  is  closely  associ- 
ated with  sclerotized  ridge  on  inner  surface  of 
each  plate;  connective  in  lateral  aspect  with 
dorsal  keels  narrow;  aedeagus  (Figs.  17B,C) 
symmetrical  with  slender  shaft,  straight  in 
ventral  aspect,  without  apical  processes.  Apex 
flared  in  caudoventral  aspect,  but  varying 
from  concave  to  convex.  Dorsal  apodemal 
process  small,  not  attaining  the  plane  of  the 
shaft  of  the  aedeagus,  appearing  suboval  in 
caudodorsal  aspect,  with  lobelike,  converg- 
ing, ventral  apices.  Gonopore  minute,  sub- 
apical  on  caudoventral  surface. 

Female. — Sternum  VII  (Fig.  17H)  with  in- 
ner teeth  of  the  hind  margin  longer  than  outer 
pair.  Ovipositor  with  basal  processes  of  first 
valvulae  (Fig.  171)  recurved. 

Types. — Holotype  6:  Brewster  County, 
Texas,  Chisos  Mountains,  Big  Bend  National 
Park,  Texas,  7  August  1987,  R.  F.  Whitcomb 
(5,200  ft,  Bouteloua  curtipendula,  IPL 
003262).  Deposited  in  USNM.  Paratypes: 
2  (5 ,  1  9 ,  same  collection  data;  1  S ,  Texas, 
Marathon  Basin,  8  August  1984,  deposited 
USNM,  BARC,  KSU,  KU;  5  c?,  1  9,  Chisos 
Mountains,  D.  J.  and  J.  N.  KnuU,  17  July 
1946,  deposited  OSU. 

Remarks. — Flexainia  bandarita  resembles 
pectinata,  gila,  and  doeringae,  but  the  sinuate 
styles  of  bandarita  separate  it  from  these  spe- 
cies. The  middle  teeth  of  the  female  sternum 
VII  are  longer  than  the  outer  pair,  suggesting 
an  intermediate  stage  in  development  of  the 
median  process  that  is  present  in  doeringae. 
Like  pectinata  and  doeringae,  bandarita  is  a 
specialist  of  side-oats  grama  and,  so  far  as 
known,  is  restricted  to  that  host.  Its  known 
geographic  range  (Fig.  15)  includes  the 
higher,  more  mesic  grasslands  of  the  Chisos 
Mountains  in  Big  Bend  National  Park  and  the 


240 


Great  Basin  Naturalist  Memoirs 


No.  12 


Fig.  17.  Flexamia  bandarita,  n.  sp.;  A,  aedeagus  and  connective,  lateral  aspect;  B,  aedeagus  and  connective,  dorsal 
aspect;  C,  ape.x  of  aedeagus,  caudoventral  aspect;  D,  male  plates  and  pygofers,  ventral  aspect;  E,  male  pygofer,  lateral 
aspect;  F,  right  style,  dorsal  aspect;  G,  apex  of  right  style,  lateral  aspect;  H,  female  sternum  VII;  I,  bases  of  first  valvulae 
of  female,  dorsal  aspect. 


Marathon  Basin  north  of  the  park.  This  region 
is  rich  in  endemism  presumably  derived  by 
insularization  of  high  grasslands  by  desertifi- 
cation in  the  surrounding  true  Chihuahuan 
desert.  All  insects  in  our  recent  series,  taken 
August  1987,  are  extremely  dark;  the  season 
had  been  one  of  record  rainfall,  and  vegetative 
growth  of  all  plants  in  the  Chisos  Mountains 
was  very  lush. 

The  name  bandarita,  a  noun  in  apposition, 
is  one  of  the  common  names  for  side-oats 
grama  in  Mexico. 

6.  Flexamia  doeringae  Beamer  &  Tuthill 
Flexamia  doeringae  Beamer  &  Tuthill  1934:  3. 

Important  characters. — Length  of  6  3.5 
(3.2-3.7)  mm,  of  9  3.7  (3.3-4.1)  mm.  Head 
produced  (median  crown  length  1.40  x  inter- 
ocular  width;  0.70  X  head  width)  (d  n  =  20;  9 
n  =  20).  Face  ivory  with  broad,  black,  basal, 
interocular  line,  occasional  dark  markings  be- 


low. [6]  Plates  (Fig.  IIF)  broad  through  most 
of  their  length,  then  narrowing  sharply  to 
rounded  apices.  Style  with  preapical  lobe 
strongly  developed,  flattened  and  produced 
posteriorly;  aedeagus  (Fig.  12F)  with  protu- 
berance near  shaft  apex  close  to  tip  in  ventral 
aspect.  [  9  ]  Sternum  VII  (Fig.  14F)  with  inner 
teeth  strongly  produced  and  closely  associ- 
ated to  form  median  process;  ovipositor  with 
each  first  valvula  (Fig.  63A)  recurved  at  base; 
recurved  portion  with  transverse  lobe  cepha- 
lad  of  basal  extremity. 

Geographic  distribution. — This  species 
is  known  primarily  from  the  mountains  (Babo- 
quivari,  Catalina,  Chiricahua,  Huachuca,  and 
Santa  Rita  ranges)  of  southeastern  Arizona, 
but  it  also  occurs  in  the  Mexican  highlands  of 
Sonora  and  Chihuahua  (Fig.  15).  There  is  one 
record  from  Yavapai  Co.,  Arizona  (1  6 ,  Gran- 
ite Dell,  10  July  1933,  R.  H.  Beamer). 

Biology. — This    species    is    apparently    a 


1988 


Whitcomb,  Hicks:  Flexamia  Revision 


241 


specialist  on  side-oats  grama,  Bouteloua  ciir- 
tipendula.  Because  this  grass  is  a  dominant  in 
the  mountains  of  southeastern  Arizona  and 
because  many  collectors  have  visited  this  re- 
gion, doeringae  is  a  common  component  of 
large  Flexamia  collections  (Appendix  I).  The 
climate  of  the  southeastern  Arizona  moun- 
tains is  similar  to  that  of  the  Mexican  high- 
lands but  shows  a  definite  Sonoran  influence. 
Flexamia  doeringae  may  therefore  be  a  Sono- 
ran vicariant  that  separated  from  the  Chi- 
huahuan  bandarita  and  the  prairie  pectinata, 
which  are  also  specialists  of  B.  curtipendula. 

Oligophagy  coefficients. — Gramineae 
1.000;  Chloridoideae  1.000;  Bouteloua  1.000; 
Bouteloua  curtipendula  0.943  (n  =  35). 

Remarks  and  diagnosis. — The  range  oi do- 
eringae does  not  overlap  that  of  any  other 
member  of  the  pectinata  species  group.  This 
species  can  be  readily  distinguished  by  its 
ivory  face,  the  pair  of  transverse  lines  on  a 
pale,  well-produced  crown,  and  the  structure 
of  the  external  genitalia.  The  female  sternum 
VII  (Fig.  14F)  is  particularly  distinct  in  having 
a  medially  produced  structure  consisting  of  a 
pair  of  long,  central  lobes  and  much  shorter 
lobes  on  each  side.  The  middle  pair  of  teeth, 
in  their  size  and  degree  of  fusion,  are  maxi- 
mally developed  in  this  species. 

7.  Flexamia  collorum,  n.  sp. 

Length  of  c?  2.9  (2.7-3.0)  mm,  9  3.0 
(2.8-3.2)  mm;  head  width  of  6  0.87  mm,  9 
0.89  mm.  Crown  not  strongly  produced;  me- 
dian length  of  crown  1.53  x  head  width  and 
0.68  xinterocular  width  (c?  n  =  11;  9  n  =  20). 

Crown  stramineous  to  light  brown,  prono- 
tum  with  four  faint,  wide,  brown  stripes, 
forewings  gray  with  irregular,  dark  markings. 
Face  color  and  markings  variable,  without  in- 
terocular  line,  but  usually  black  in  interocular 
area  and  lower  area.  Usually  at  least  lorae 
pale.  Crown  with  pair  of  oblique  markings  at 
rear,  pair  of  transverse  lines  at  midlength. 
Venter  usually  predominantly  dark. 

Male. — Pygofer  produced  ventrally  into  an 
angulate  lobe;  caudoventral  margin  sloping 
dorsally  to  a  broadly  rounded,  caudal  lobe. 
Plates  (Fig.  18D)  short,  extending  to  1/2 
length  of  pygofer,  fused  basally,  contiguous 
medially  for  2/3  length,  acute  apically.  Plates 
produced  on  inner  surface  at  about  midlength 
into  well-developed,  sclerotized  ridge  that 
interfaces    with    style    apices.     Connective 


(Fig.  18A)  in  lateral  view  with  dorsal  keels 
narrow,  approximately  1/3  height  of  dorsal 
apodeme;  apodemal  processes  as  in  ban- 
darita; style  (Fig.  18F)  short,  abruptly  nar- 
rowing beyond  preapical  lobe  to  avicephali- 
form,  mesoventrally  directed  apices  (Fig. 
18G).  Aedeagus  (Figs.  18B,C)  symmetrical, 
shaft  slender,  without  processes,  slightly 
flared  apically  in  caudoventral  aspect. 
Gonopore  subapical  on  caudoventral  surface. 

Female.— Sternum  VII  (Fig.  18H)  with 
four  teeth  on  hind  margin;  outer  pair  not  well 
developed,  inner  pair  distinct,  but  short  and 
rounded;  ovipositor  with  basal  processes  of 
first  valvulae  (Fig.  181)  recurved. 

Types. — Holotype  <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, 


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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 


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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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Whitcomb,  Hicks:  Flexamia  Revision 


245 


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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Great  Basin  Naturalist  Memoirs 


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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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Whitcomb,  Hicks:  Flexamia  Revision 


247 


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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No.  12 


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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Whitcomb,  Hicks:  Flexamia  Rkvision 


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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No.  12 


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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Whitcomb,  Hicks;  Flexamia  Revision 


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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No.  12 


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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Whitcomb,  Hicks:  Flexamia  Revision 


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 


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Great  Basin  Naturalist  Memoirs 


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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Whitcomb,  Hicks;  Flexamia  Revision 


255 


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 


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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 


Great  Basin  Naturalist  Memoirs 


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 


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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 


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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. 


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Whitcomb,  Hicks:  Flexamia  Revision 


263 


^A 


{ 

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iL'^'--' 

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'^'":"""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 


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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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Whitcomb,  Hicks:  Flexamia  Revision 


265 


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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Whitcomb,  Hicks;  Flexamia Revision 


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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Great  Basin  Naturalist  Memoirs 


No.  12 


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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Whitcomb,  Hicks:  Flexamia  Revision 


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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Great  Basin  Naturalist  Memoirs 


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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Whitcomb,  Hicks:  Flexamia  Revision 


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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Great  Basin  Naturalist  Memoirs 


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  = 


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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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Whitcomb,  Hicks;  Flexamia  Revision 


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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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279 


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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283 


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. 


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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 


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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 


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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 


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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 


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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 


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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. 


292 


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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 


Great  Basin  Naturalist  Memoirs 


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 


Great  Basin  Naturalist  Memoirs 


No.  12 


=2       5 


II  III 


7       8 


10 


IV 


15 


19 


VI 


16 


VII 


14 


12 


13 


Fig.  56.  See  facing  page. 


20 


VIII 

18 


23 


XI 

XII 

30 

31 

2< 

XIII 

32 


25 


IX 


21         22 


28 


26 


24 


hypothesize  (with  Young  and  Beirne,  and 
Ross)  that  Spartopyge  is  an  outgroup  of  the 
Flexamia  hneage.  We  beheve,  with  these 
workers,  that  a  primitive  Flexamia  ancestor 
arose  from  the  Spartopyge  hneage  and  later 
gave  rise  to  a  hneage  with  paired,  aedeagal 
processes.  This  hneage  in  some  way  gave  rise 
to  grammica  but,  ultimately,  to  all  Flexamia 
groups  IV-XIII.  We  assume  that  the  dorsal 
location  of  the  gonopore  in  grammica  and 
prairiana  is  a  result  of  independent  evolution- 
ary events,  since  these  species  share  few  other 
significant  characters.  Whether  the  minute, 
paired  processes  that  occur  midlength  on  the 
Spartopyge  aedeagus  are  homologous  with 
the  Flexamia  aedeagal  processes  is  conjec- 
tural. 

Phylogeny  of  Flexamia  groups  and  spe- 
cies. The  postulated  relationships  and  polari- 


ties permit  a  cladogram  to  be  drawn  relating 
Flexamia  groups  and  key  species  to  the  out- 
groups  Alapus  and  Spartopyge  (Fig.  56).  Sev- 
eral significant  possible  variations  of  the  pro- 
posed tree  are  given  in  Figure  57.  Finally,  the 
phylogeny  of  Flexamia  species  was  inferred  on 
the  basis  of  the  characters  discussed  above. 
We  present  cladograms  for  groups  I-IX  (Fig. 
58),  groups  VII-XI  (Fig.  59),  and  groups 
XI-XIII  (Fig.  60).  Synapomorphies  used  to 
define  clades  within  each  group  are  given  in 
the  legends  to  the  figures. 

Character  transformations. — Our  intu- 
itive phylogeny  provided  general  guidelines 
concerning  character  transformation  in  Flex- 
amia. From  these  concepts  we  constructed 
branched  transformations  for  the  PHYSYS 
program  of  Mickevich  and  Farris  (1984). 
These  transformations,  which  are  described 


1988  Whitcomb,  Hicks  Flexamia  Revision  297 

Fig.  56.  Proposed  phylogeny  of  Flexamia  groiip.s  and  key  species.  Synapoinorphies  [s]  or,  for  monobasic  groups, 
autapomorphies  [an]  are  indicated.  We  hypothesize  an  ancestral  form  (1)  that  possessed  dorsal  stripes  (Fig.  54 A)  and  a 
pale  face  with  a  contrasting  black  interoeiilar  band  (Fig.  54B)  similar  to  those  of  Figs.  3A  and  3B,  relatively  wide  male 
plates  (Fig.  54C);  an  aedeagus  distinctly  jointed  with  the  connective,  without  apical  processes,  and  with  an  anteapical 
gonopore  on  the  caudoventral  surface.  Divergence  of  this  lineage  led  (3)  to  Alapus  (Figs.  55F,(v),  represented  in  this 
diagram  by  elongatus,  and  to  Spartopy^e,  represented  (4)  in  this  diagram  by  miranda  and  mexicana,  in  which  [s]  the 
dorsal  aedeagal  apodeme  is  large  and  hoodlike  (Fig.  55C).  The  transformations  between  Flexamia  and  its  outgroups  are 
tenuous  (see  te.xt).  In  Flexamia  [s]  the  dorsal  aedeagal  apodeme  (Fig.  6)  has  a  pair  of  conspicuous  appendages  directed 
caudoventrad  or  caudad  (5).  In  the  first  lineage  (6)  to  diverge,  the  plesiomorphic  aedeagal  apex,  which  lacked  processes, 
was  retained,  as  were  the  caudoventral  position  of  the  gonopore  and  broad,  ancestral  plates;  the  contrasting  face  was 
retained  in  a  minority  of  species,  but  the  striped  habitus  was  completely  lost.  Synapoinorphies  defining  this  lineage 
include  [s]  the  curved  aedeagus  lacking  processes  (Figs.  71,  J;  16A,  17A,  2()A)  and  the  male  pygofers,  which  possess  both 
dorsal  and  ventral  constrictions  (Figs.  62A-I).  This  lineage  diverged  into  the  pectinata  (7)  and  the  abhreviata-zacate 
lineages  (8).  In  the  pectinata  group  the  plates  (Fig.  11)  are  distinctively  parallel-sided  basally  (except  in  collorum).  In 
the  abhreviata-zacate  lineage  the  gonopore  was  at  first  elongate  (Fig.  24)  and  [s]  the  posterior  lobe  of  the  male  pygofer 
strongly  produced  (Figs.  62H,I).  In  the  ahbreviata  group  (II)  the  plates  (Fig.  8E)  are  reduced  (9),  and  [au]  a  distinctive 
pair  of  minute  apical  processes  (Fig.  7H)  developed.  The  small  aedeagal  processes  of  the  abbreviata  group  are  not 
clearly  homologous  with  other  aedeagal  processes  in  Flexamia.  In  the  zacate  group  (III)  the  gonopore  (Fig.  24)  was  at 
first  elongate  (10)  but  in  canyonensis  moved  to  the  aedeagal  midpoint.  At  this  point  a  significant  apomorphy  appeared. 
An  ancestral  form  (11)  is  hypothesized  in  which  two  pairs  of  aedeagal  processes  were  present.  It  is  unclear  whether  this 
form  had  fully  developed  before  divergence  of  grammica  (IV),  in  which  the  plesiomorphic  feature  of  dorsal  stripes  was 
retained  (12),  but  in  which  [au]  the  aedeagal  processes  were  distinctively  modified  (Fig.  7E)  and  the  gonopore  had 
moved  to  the  dorsal  surface  (12).  A  possible  apomorphy  linking  g,rammica  with  the  curvata  group  is  the  foot-shaped 
style  apex  (see  illustrations  of  Young  and  Beirne  1958).  In  the  cttrvata-picta  lineage  [s]  both  pairs  of  aedeagal  processes 
are  present  (14),  but  many  habitus  features,  including  stripes  and  contrasting  face,  have  been  lost.  In  the  curvata  group 
(V)  two  pairs  of  processes  are  present  (15)  and  [s]  unmodified  (Figs.  7F,G).  In  the  picta  group  (VI)  the  apical  processes 
are  [s]  modified  (Figs.  7K,L)  as  flangelike,  toothed  appendages  (16).  Remaining  species  are  defined  by  an  apomorphy 
that  developed  with  the  presumed  fusion  (17)  of  the  anteapical  processes  into  a  single,  unpaired  process  that  was 
plesiomorphically  located  ventrally  in  the  sagittal  plane  (Figs.  31,  61A).  A  second  apomorphy  characterizing  this 
divergence  is  the  bifurcation  (Figs.  7A-C)  of  the  apical  aedeagal  processes,  a  feature  that  was  subsequently  lost  (22). 
The  plesiomorphic  state  of  dorsal  stripes  and  contrasting  face  was  preserved  (18)  during  the  fusion  of  subapical 
processes  and  is  present  (Figs.  2A,B)  in  the  albida  group  (VII).  This  lineage,  which  can  be  defined  [s]  in  terms  of  the 
absence  of  dorsal  or  ventral  constrictions  in  the  posterior  lobe  of  the  male  pygofer  (Figs.  62Q,T),  is  represented  today  by 
the  distantly  related  sister  species  albida  (19)  and  slossonae  (20);  for  autapomorphies  defining  these  species,  see  text.  In 
serrata  (VIII)  [au]  additional  dorsal  aedeagal  processes  (Fig.  7A)  developed  (21),  and,  while  the  contrasting  face  was 
retained,  dorsal  stripes  were  lost  (Fig.  1).  In  ritana  (IX)  the  dorsal  stripes  were  retained  (Fig.  2D)  but  greatly  reduced. 
This  monobasic  group  (23)  is  defined  [au]  by  the  unique,  pointed  aedeagal  apex  (Fig.  7D).  In  one  lineage  of  the^^xw/osa 
group  (25),  the  unpaired  aedeagal  process  is,  plesiomorphically,  ventral  in  the  sagittal  plane,  permitting  retention  of 
aedeagal  symmetry  (Figs.  31G,I,J,K,L).  The  plesiomorphic  pale  face  with  black  interocular  band  (Figs.  3B,F,G)  was 
also  retained.  This  clade  is  defined  [s]  by  the  triangulate  plates  (Figs.  33A-F,  34D)  with  conspicuous,  apical  divergence. 
(But  in  stylata  [Fig.  34E],  the  plates  [au]  are  truncated.)  In  the  remaining  species  (26)  the  unpaired  process  underwent 
phyletic  twisting  (Figs.  61B-H)  and  the  contrasting  pale  face,  which  had  persisted  through  many  nodes,  was  finally  lost. 
In  the  inflata  subgroup  of  group  X,  [s]  the  displacement  of  the  unpaired  process  (Fig.  61 B)  was  relatively  small  (27),  but 
[s]  in  the  imputans-areolata  (XI-XII)  lineage  the  process  had  become  lateral  (Figs.  31B,C)  and  bore  the  gonopore  as  an 
elongate  slit  (29).  Before  divergence  of  groups  XI-XIII,  the  connective  and  aedeagus  had  remained  articulated  (but  see 
discussion  of  group  VI  in  text).  We  hypothesize  that  after  divergence  of  the  imputans-areolata  lineage,  a  fusion  of 
aedeagus  and  connective  occurred  (31)  in  areolata .  The  imputans-areolata  lineage  is  very  specialized  in  the  location  of 
the  unpaired  process,  in  the  black  face,  and  the  greenish  dorsal  pigmentation.  Of  the  two  species,  areolata  is  the  more 
highly  specialized  (Figs.  2F,  lOQ);  in  fact,  it  may  be  the  most  highly  specialized  of  all  Flexamia  species.  It  is  most 
unlikely  that  the  widely  distributed  and  variable  prairiana  (see  text)  and  the  entire  prairiana  group  (XIII)  were  derived 
from  the  specialist  areolata  lineage.  Instead,  we  propose  that  divergence  of  a  second  lineage  in  which  aedeagus  and 
connective  fused  (32)  led  to  the  prairiana  group.  This  fusion,  the  traces  of  which  are  evident  (Fig.  6F)  in  prairiana ,  is 
designated  [s]  as  a  synapomorphy  defining  group  XIII,  and  the  fusion  in  areolata  is  defined  as  an  autapomorphy 
defining  that  species.  Details  of  species  phylogenies  are  presented  in  detail  in  Figures  58-60. 


in  Appendix  II,  Part  III,  involved  the  follow-  ness  (lateral  aspect)  of  the  aedeagus;  (7)  length 

ing  characters:  (1)  relationship  between  ae-  ofthe  aedeagus;  (8)  form  of  the  aedeagal  shaft; 

deagus  and  connective;  (2)  morphology  ofthe  (9)  form  of  margins  of  paired,  aedeagal  pro- 

aedeagal  apex;  (3)  morphology  of  anteapical  cesses;  (10)  form  and  position  ofthe  gonopore; 

aedeagal  processes;  (4)  phyletic  twisting  ofthe  (11)  morphology  of  apices  ofthe  apodemal 

unpaired  ventral  process  (Fig.  61);  (5)  curva-  arms;  (12)  position  of  apodemal  arms;  (13) 

ture  (lateral  aspect)  ofthe  aedeagus;  (6)  thick-  width  of  dorsal  keels  of  the  connective;  (14) 


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III 

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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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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 


302 


Great  Basin  Naturalist  Memoirs 


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. 


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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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No.  12 


Fig.  61.  Postulated  character  transformation:  phyletic  twisting  and  modification  of  the  Flexamia  aedeagus.  In  decora 
(A)  a  plesiomorphic  state  exists  in  which  the  unpaired  process  is  ventral  and  in  the  sagittal  plane;  the  aedeagus  is 
therefore  symmetrical.  In  inflata  (B)  there  is  a  small  displacement  of  the  unpaired  process.  In  areolata  (C)  the  unpaired 
process  is  lateral  and  bears  the  gonopore.  In  prairiana  (D)  the  process  is  dorsal,  as  is  the  gonopore.  In  reflexa  (E)  the 
gonopore  has  become  associated  with  the  dorsal  unpaired  process.  In  sandersi  (F)  the  unpaired  process  is  reduced  and 
appressed  to  the  shaft.  In  producta  (G)  the  process  is  reduced  or  absent  and  the  gonopore  is  lateral.  In  ^raininea  (H)  the 
process  has  disappeared  entirely,  the  apical  processes  are  reduced,  and  the  gonopore  is  spiral.  Aspects;  AD, 
anterodorsal;  CV;  caudoventral;  DS,  dorsal;  LT,  lateral;  VN,  ventral.  Aedeagal  structures  redrawn  from  Young  and 
Beirne(1958). 


adapted  to  northeastern  {minima)  and  west 
central  (zamora)  Mexican  grasslands,  to  Chi- 
huahuan  (bandarita )  and  Sonoran  {doeringae) 
montane  grasslands  of  the  Southwest,  to  the 
interdesert  montane  region  of  the  Gila  Moun- 
tains igila),  and  to  the  tall-grass  and  mixed- 
grass  prairie  of  the  United  States  (pectinata). 
Also  derived  from  the  side-oats  lineage  were 
jacala  (Mexico:  host  unknown)  and  collorum 
(Edwards  plateau;  host:  Bouteloua  uniflora), 
a  close  relative  of  side-oats  grama).  Clearly, 
most  or  all  of  the  nodes  in  the  pectinata  lin- 
eage are  Mexican  or  southwestern;  none  rep- 
resent events  that  assuredly  took  place  north 
of  the  Mexican-U.S.  border. 

A  second  lineage  of  the  pectinata  subtree 
resulted  in  the  divergence  of  mescalero,  a 
distinctive  species  that  represents  a  host 
transfer  to  New  Mexican  muhly,  Midilenher- 
gia  pauciflora.  Another  branch  of  this  lineage 
transferred  to  temperate  Bouteloua  grass- 
lands of  Mexico.  One  of  the  branches  (abbre- 
viata  group)  inhabited  the  mixed  Bouteloua 
grasslands  of  Mexico  and,  by  gradual  adapta- 
tion, to  similar  grasslands  of  the  United 
States.  Another  branch  adapted  to  semiarid 
grasslands  by  colonizing  bush  muhlenbergia 
{Muhlenbergia  porteri).,  divergence  of  this 
lineage  led  to  the  contemporary  canijonensis 
(Sonoran)  and  zacate  (Chihuahuan). 

Consider  next  the  surcula-curvata  lineage. 
Both  species  possess  the  four  aedeagal  pro- 
cesses that  Young  and  Beirne  proposed  as  ple- 
siomorphic; both  species  are  Buchloe  special- 


ists. We  propose  that  an  ancient  Mexican  an- 
cestral line  adapted  to  B.  dactyloides,  which 
led  to  the  present-day  surcula  and,  later,  to 
curvata.  This  divergence  (Fig.  64)  may  have 
been  mediated  by  climate.  The  grasslands  of 
northeastern  Mexico,  where  surcula  may 
have  arisen,  have  escaped  major  disturbance 
since  the  Cretaceous  (Rosen  1978).  This  re- 
gion has  an  extremely  warm,  temperate  or 
subtropical  climate  and  a  Chihuahuan  precip- 
itation pattern,  both  of  which  differ  im- 
mensely from  the  dry,  temperate  grasslands 
where  curvata  occurs  today. 

Another  branch  of  the  curvata  subtree  di- 
verged, perhaps  in  Mexico  or  south  Texas, 
leading  to  the  picta-pyrops  sister  set.  These 
species  are  Aristida  specialists. 

One  of  the  major  events  in  Flexamia  phy- 
logeny  was  marked  by  the  fusion  of  the  an- 
teapical  pair  of  aedeagal  processes  into  a  sin- 
gle, mesal  process.  The  most  primitive 
species  in  which  this  feature  is  retained  are 
members  of  the  albida,  serrata,  and  ritana 
groups.  Young  and  Beirne  argued  (and  Ross 
concurred)  that  slossonae,  albida,  and  serrata 
were  related.  We  agree  but  consider  their 
divergence  to  be  an  ancient  one,  given  their 
entirely  different  climatic  reciuirements  (sub- 
tropical, prairie,  and  northern  grasslands,  re- 
spectively), and  their  retention  of  the  Spar- 
topyge  habitus  type.  Perhaps  the  divergence 
of  the  albida  lineage  into  four  species,  each 
with  imiciue  climatic  adaptations,  may  have 
occupied  a  distinct  stage  in  Flexamia  evolu- 
tion. 


1988 


Whitcomb,  Hicks:  Flexamia  Revision 


305 


Fig.  62.  Character  transformation  of  the  Flexamia  pygofer.  Diagrammed  Hneage  was  based  in  part  on  phylogeny  of 
groups  (see  Figure  56).  A,  collorum;  B,  mescalero;  C,  handarita;  DJacala;  E,  gila;  F,  pectinata,  G,  doeringae;  H, 
ahhreviata-  I,  canyonensis;  J,  pijrops-  K,  curvata,  L,  surcula-  M,  picta;  N.  Spartopijge  miranda;  O,  grammica;  P, 
serrata,-  Q,  a/foida;  R,  A/apus  elongatus;  S,  Spartopyge  iijexicana;  T,  .s/ossoriae;  U,  ritana;  V,  impt^fans;  W,  prainana; 
X  re/Zexa-  Y  af/anfica,  Z,  areo/ata,  AA,  producta;  BB,  ffa/coto;  CC,  sanrfersi;  DD,  c/ayi,  EE,  graminea,  ¥¥  decora; 
GG,  inflata;  HH,  texaria,  II,  foeani^ri,  JJ,  modica,  KK,  yowngj,  LL,  arenicola;  MM.  ari^onenm;  NN,/^cxu/osa;  OO, 
sfy/ato;  PP,  celata .  Some  pygofers  were  redrawn  from  Young  and  Beirne  (1958)  and  Lowry  and  Blocker  (1987). 


306 


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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 


Great  Basin  Naturalist  Memoirs 


No.  12 


Fig.  65.  Geographic  interpretation  of  phylogeny  of  the  pectmata  group.  A  focal  origin  in  grasslands  of  eastern  Mexico 
is  hypothesized,  A  western  lineage  comprised  gila,  handarita,  and  doeringae.  This  lineage  speciated  exclusively  on 
Bouteloua  curtipendtda.  Morphological  data  suggest  that  mescalcro,  which  apparently  transferred  to  Muhlenbergia 
pauciflora,  belongs  on  a  deep  branch.  The  subtree  consisting  of  za»iora,  minima,  and  pectinata  is  closely  related  to  the 
doeringae  subtree;  like  all  members  of  that  subtree,  pectinata  specializes  on  B.  ctitiipendiila  (side-oats  grama).  F. 
coUorum  may  have  been  derived  from  a  lineage  on  B.  curiipendula,  since  its  host  is  B.  uniflora,  which  is  closely  related 
to  side-oats  grama.  The  host  ofjacala  is  unknown. 


grasslands).  (Interestingly,  in  contrast  to 
Athysanella  species  [Hicks  et  al.  1988],  both 
Flexamia  blue  grama  specialists  colonize  this 
host  throughout  most  of  its  range  in  New  Mex- 
ico). A  sister  species,  arizonensis,  transferred 
to  another  dominant  southwestern  grass, 
Aristida  purpurea.  We  emphasize  that  con- 
siderable climatic  diversity  exists  today  in  the 
Southwest  and  no  doubt  has  existed  through- 
out the  Pleistocene  and  Holocene  as  the  result 
of  elevational  gradients  (Axelrod  and  Raven 
1985,  Cole  1986,  Cronquist  1978,  Thompson 
and  Mead  1982,  VanDevender  and  Spaulding 
1979).  Therefore,  evolutionary  processes  that 
might  take  place  only  over  wide  geographic 


distances  in  regions  of  relatively  constant  ele- 
vation may  be  focused  in  small  geographic 
areas  in  the  Southwest. 

Another  lineage,  possibly  more  closely  al- 
lied to  the  ritana  than  serrata  lineage,  pre- 
sumably diverged  in  grasslands  of  the  south- 
ern plains.  This  is  the  inflata  subgroup  of  the 
flexulosa  group.  One  contemporary  represen- 
tative (inflata)  exhibits  one  of  the  widest  host 
ranges  in  the  genus;  largest  populations  occur 
on  Muhlcnhcrgki  aspcrifolia  in  wetland  areas. 
If  this  apparent  preference  indicates  an  ances- 
tral Muhlenbergia  host,  it  would  be  necessary 
to  postulate  a  monumental  subsequent 
widening  of  host  range  to  pooid  grasses  or 


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Whitcomb,  Hicks:  Flexamia  Revision 


309 


Fig.  66.  Geographic  interpretation  of  phylogeny  of  the  serrata,  ritana,  flexulosa,  imptitans,  and  areolata  groups. 
Texan  grasslands  may  have  been  a  focal  area  for  initial  divergence  of  lineages.  F.  serrata  appears  to  be  a  Muhlenbergia 
specialist.  The  ritana  group  appears  to  have  diverged  in  the  Southwest.  The  contemporary  distributions  of  the  albida, 
serrata,  and  ritana  groups  suggest  that  they  may  have  speciated  by  vicariant  mechanisms  or,  in  the  case  oi serrata,  by 
a  combination  of  vicariance  and  host  transfer.  An  eastern  lineage  consists  oiinflata  and  beameri.  From  a  biogeographic 
perspective,  it  is  most  reasonable  to  propose  an  independent  origin  oitexana  and  beameri,  each  from  inflate  or  a  closely 
related  lineage.  The  present-day  distribution  oiinflata  suggests  that  it  diverged  in  cool,  temperate  grasslands;  Texas,  of 
course,  was  cool  during  glacial  maxima.  In  general,  events  postulated  as  northern  in  these  reconstructions  could  have 
taken  place  much  farther  south  during  glaciations.  The  imputans-areolata  lineage  appears  to  have  diverged  in  the 
prairie,  on  Muhlenbergia  cuspidata  ind  Eragrostis  spectabilis ,  respectively.  The  western  lineage  of  the/7ex«/osa  group 
may  be  related  (see  Fig.  59)  to  the  serrata  group;  if  this  association  can  be  confirmed,  the  derivation  of  the  decora  set  of 
sister  species  from  the  serrata  lineage  would  make  an  attractive  hypothesis  from  a  biogeographic  standpoint,  since  all  of 
the  species  (including  youngi)  specialize  on  Muhlenbergia  richardsonis.  Derivation  oi  modica  (host:  Muhlenbergia 
ripens)  and  arenicola  (host;  Muhlenbergia  pungens)  represents  further  evolution  that  occurred  by  host  transfers  within 
Muhlenbergia.  The  derivation  of  flexulosa  and  arizonensis  by  host  transfer  may  have  been  a  consequence  of  the 
regional  abundance  (in  New  Mexico  and  eastern  Arizona)  of  their  hosts,  Bouteloua  gracilis  and  Aristida  purpurea, 
respectively;  presumably,  the  region  was  colonized  from  the  north.  Finally,  derivation  of  celata  (host:  Redfieldia 
flexuosa  and  perhaps  other  sandhill  grasses)  and  stylata  (hosts:  Muhlenbergia  spp.)  is  hypothesized  to  have  followed  a 
high-plains  route  around  the  eastern  edge  of  the  high  Rocky  Mountains. 


310 


Great  Basin  Naturalist  Memoirs 


No.  12 


Fig.  67.  Geographic  interpretation  of  the  phylogeny  of  the  prairiana  group.  Although  present-day  distributions  and 
host  preferences  do  not  offer  obvious  hypotheses  for  speciation  mechanisms,  an  origin  for  the  prairiana  group  in  Texan 
grasslands  is  hypothesized.  From  this  region  prairiana  presumably  moved  northward  in  postglacial  episodes,  where  it 
adapted  to  little  bluestem  (Schizachyrium  scoparium )  or  other  andropogonoid  grasses.  In  the  tall-grass  prairie,  reflexa 
diverged  from  this  lineage;  this  species  often  seems  to  be  associated  with  big  bluestem  (Andropogon  gerardii), 
particularly  in  glades  of  the  eastern  United  States.  F.  atlantica  may  have  transferred  in  postglacial  times  from 
chlorodoid  grasses,  on  which  it  predominates  in  the  Southwest,  to  switchgrass,  Panicum  virgattim.  This  host  has  been 
a  conduit  for  colonization  by  atlantica  of  the  eastern  United  States,  including  the  Atlantic  coastal  prairies,  where  its  host 
achieves  high  importance.  Derivation  oi sandersi  may  have  occurred  by  transfer  to  Andropogon  virginicus,  the  first 
colonist  in  early  fire  succession  in  the  Southeast.  From  this  lineage,  presumably  post-Wisconsinan,  delongi  may  have 
diverged  by  colonization  of  little  bluestem  in  the  glaciated  area  northwest  of  Illinois.  Specialization  on  Schizachyrium 
scopariuin  and  subsequent  divergence  led  to  dakota  in  the  dry  prairies  of  the  plains,  to  graminea  in  Texas  and 
northward  through  the  tall-grass  prairie,  and  to  a  lineage  of  eastern  species.  These  species,  also  residents  of 
andropogonoid  grasses,  are  residents  of  the  Gulf  Coast  (satilla),  or  the  Appalachian  mountains  to  Maine  and  cool, 
temperate  savannas  of  Illinois  and  Ohio  (clayi). 


Juncus.  It  may  be  more  likely  that  the  cur- 
rently observed  host  range  breadth  repre- 
sents simply  a  retention  of  plesiomorphic  gen- 
eralism.  In  any  event,  this  lineage  has 
spawned  two  additional  species:  texana  (south 
central  Texas)  and  beameri  (west  slope  of  the 
Adirondack  Mountains,  New  York).  The  cir- 
cumstances surrounding  these  divergences 
are  unclear,  since  we  have  no  data  on  their 


host  relationships. 

After  divergence  ofinflata,  another  lineage 
diverged  in  the  prairie,  leading  to  imputans 
(probable  host:  Muhlenbergia  cuspidata)  and 
areolata  (host:  Erag^rostis  spectabilis)  on  one 
hand,  and  to  the  prairiana  group  on  the  other. 

All  speciation  events  of  the  prairiana  group 
(Fig.  67)  occurred  either  in  prairie,  savanna, 
or  southeastern  grasslands.  We  hypothesize 


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Whitcomb,  HickS:  Flexamia  Revision 


311 


an  ancestor  for  this  group  that  was  adapted  to 
both  chloridoid  and  panicoid  grasses.  From 
such  an  ancestor,  prairiana  (liost:  Schiza- 
chyrium  or  Bothriochloa)  and  reflexa  (host: 
Andropogon  gerardii)  were  derived,  as  were 
atlantica  (hosts:  chloridoid  grasses  and/or 
Panicum  virgatum),  and  prodiicta  (hosts 
chloridoid  grasses,  or  [possibly]  a  habitat  spe- 
cialist). Finally,  specialization  on  Schiza- 
chyrium  scoparium  and/or  Andropogon  vir- 
ginicus  led  (Fig.  67)  to  lineages  that  adapted  to 
the  most  xeric  of  Andropogon  habitats  (da- 
kota,  a  Schizachyriiim  specialist  of  the  west- 
ern Great  Plains),  graminea  (a  Schizachyriiim 
specialist  of  the  tall-grass  prairie),  and  the 
sa7idersi-delongi  and  satilla-clayi  sister  pairs 
of  eastern  andropogonoid  grasslands. 

Stages  of  Speciation 

Mayr  (1942)  believed  that  at  any  given  time 
all  stages  of  evolution  should  be  observable. 
We  believe  that  this  is  indeed  the  case  in  the 
genus  Flexamia.  We  recognize  at  least  10  lev- 
els in  the  process  of  speciation  in  Flexamia: 

1.  Nonisolated  ephemeral  populations:  (i) 
Many  populations  of  Flexamia  inflata  appear 
to  be  ephemeral.  These  populations  often  oc- 
cur on  pooid  grasses  such  as  Poa  or  Festuca; 
presence  of  immature  insects  confirms  ovipo- 
sition;  (ii)  F.  abbreviata  colonizes  Miihlenber- 
gia  richardsonis  in  the  Sacramento  Moun- 
tains of  New  Mexico  and  M.  reverchonii  in  the 
Edwards  Plateau  of  Texas,  although  in  the 
latter  case  we  have  not  confirmed  oviposition. 

2.  Nonisolated  stable  populations:  (i)  F.  in- 
flata consistently  colonizes  path  rush  (Juncus 
tenuis)  in  Illinois  and  Maryland;  populations, 
including  immatures,  are  found  regularly  on 
that  host.  This  is  an  especially  interesting 
case,  since  it  may  represent  an  interfamily 
transfer  by  a  species  that  has  limited 
oligophagy.  (ii)  Flexamia  atlantica  regularly 
colonizes  large  plantings  of  weeping  love  grass 
(Eragrostis  spectahilis)  and  bermudagrass 
{Cynodon  dactylon)  in  Oklahoma  and  Texas. 
(iii)  Similarly,  prodiicta  colonizes  bermuda- 
grass and  Zoysia  japonica  in  Maryland.  These 
are  essentially  "natural  experiments "  that  may 
mimic  to  a  large  degree,  but  in  a  shorter  geo- 
logic time  period,  the  effect  of  radical  changes 
in  the  dominance  hierarchy  of  grasses  induced 
by  cyclic  climatic  change. 

3.  Disjunct  populations  that  have  not 
diverged  morphologically:  In  the  following 


cases  we  have  found  no  morphological  varia- 
tion associated  with  disjunct  populations,  (i)  A 
disjunct  population  oi  dakota,  a  specialist  on 
Schizachyriiim  scoparium  in  the  western 
Great  Plains,  occurs  on  that  host  in  the  Loess 
Hills  of  western  Iowa,  (ii)  Flexamia  reflexa 
occurs  (always,  it  seems,  in  association  with 
big  bluestem,  Andropogon  gerardii)  in  iso- 
lated grassland  islands  in  the  eastern  decidu- 
ous forest  as  far  east  as  the  Soldier's  Delight 
serpentine  barrens  in  Baltimore  County, 
Maryland. 

4.  Isolated  stable  populations  with  minor 
morphological  divergence:  In  each  of  the  fol- 
lowing cases  the  populations  may  or  may  not 
represent  biological  species,  (i)  Two  disjunct 
populations  (Nebraska  Sand  Hills  and  Four 
Corners  or  Anasazi  populations)  of  arenicola 
have  been  discovered;  the  unpaired,  aedeagal 
processes  of  the  Anasazi  population  always 
break,  whereas  the  processes  of  the  Sand  Hills 
population  rarely  break.  Other  characters 
have  not  been  found  to  separate  these  popula- 
tions, each  of  which  specializes  on  Muhlen- 
bergia  pungens.  The  host  population  is  also 
separated  into  disjunct  populations,  (ii)  The 
Mammoth  Lakes  population  oiyoungi,  prob- 
ably isolated  in  eastern  California  in  a  region 
rich  in  endemics,  has  diverged  somewhat 
from  the  population  in  the  eastern  Great 
Basin,  (iii)  Populations  of  prairiana  on  Both- 
riochloa in  the  Southwest  have  begun  to  di- 
verge from  the  prairie  Schizachyriiim  popula- 
tions, perhaps  as  the  result  of  phenological 
isolation.  In  this  case  divergence  of  female 
rather  than  male  structures  may  signal  the 
emergence  of  a  new  species. 

5.  Recent  (Holocene)  speciations:  (i)  The 
separation  of  delongi  from  the  sandersi  lin- 
eage may  represent  a  postglacial  transfer  from 
Andropogon  virginicus  to  Schizachyrium  sco- 
parium in  the  glaciated  region  of  the  northern 
prairie,  (ii)  The  divergence  ofclayi  and  satilla 
may  reflect  phenological  isolation  between 
Gulf  Coast  and  Appalachian  populations. 

6.  Pleistocene  speciations:  Biological  and 
biogeographic  circumstances  strongly  suggest 
that  decora  and  youngi  represent  vicariant 
populations  of  a  common  lineage  that  di- 
verged during  Pleistocene  climatic  cycles,  re- 
taining Miihlenbergia  richardsonis. 

7.  Vicariance  and  semiisolation:  In  the  case 
of  Mexican  and  Southwestern  grasslands,  spe- 
ciation was  perhaps  achieved  by  vicariance 


312 


Great  Basin  Naturalist  Memoirs 


No.  12 


during  long  periods  of  semiisolation.  It  is  pos- 
sible, in  the  case  of  certain  species,  that  there 
may  have  been  a  punctuated  infusion  of  genes 
into  semiisolated  populations  as  the  climate 
became  more  mesic  (Milstead  1960)  and  the 
distance  between  host  stands  of  side-oats 
grama  decreased.  In  any  event  the  emergence 
of  gila,  doeringae,  bandarita,  zamora,  min- 
ima, and  pectinata,  closely  related  species 
that  appear  to  partion  the  range  of  side-oats 
grama,  must  have  entailed  different  specia- 
tion  mechanisms  than  those  described  above 
and  are  clearly  enforced  today  by  phenologi- 
cal  isolation. 

8.  Distinct  species  with  obvious  sisters:  We 
consider  the  sister  pairs  of  picta-pyrops,  ciir- 
vata-surcida,  canyonensis-zaccite ,  and  prairi- 
ana-reflexa  to  be  obvious  sisters  that  occur  in 
ambiguous  circumstances  that  suggest  no  par- 
ticular age  for  their  divergence.  In  some  cases 
(surcula-curvata  [tropical/temperate]  and  za- 
cate-canyonensis  [Chihuahuan/Sonoran])  the 
biogeographic  circumstances  for  the  diver- 
gences are  clear. 

9.  Species  with  distant  sisters:  We  consider 
the  albida-slossonae  sister  pair  to  represent  a 
historically  remote  divergence.  Flexamia  slos- 
sonae  is  a  subtropical  species  that  has  many 
autapomorphies  separating  it  from  other  Flex- 
amia species.  However,  common  characters 
of  habitus  and  male  genitalia  leave  no  doubt 
that  it  is  a  sister  to  the  tall-grass  prairie  species 
albida. 

10.  Orphans:  The  characteristics  of  gram- 
mica  are  sufficiently  unique  that  it  must  be 
considered  an  orphan  within  the  genus.  In  the 
absence  of  a  contemporary  link,  the  relation- 
ships to  other  extant  species  of  the  genus  are 
obscure. 

From  the  above  examples  we  feel  that  our 
experience  with  Flexamia  elegantly  confirms 
Mayr's  hypothesis. 

Conclusions 

The  initial  choice  of  Flexamia  as  a  model  for 
the  study  of  the  role  of  host  plants  in  insect 
evolution  was  a  fortuitous  one.  Whereas  some 
speciation  events  in  Flexamia  must  have  oc- 
curred by  host  transfer,  others  surely  oc- 
curred by  vicariance.  Further,  there  is  a  ten- 
dency of  individuals  representing  advanced 
lineages  to  be  reduced  in  size  and  complexity, 
and  to  be  more  highly  specialized  in  their  host 


choice  than  their  ancestors.  There  is  therefore 
an  immense  stake  in  arriving  at  a  well-sub- 
stantiated phylogenetic  hypothesis  for  the 
genus  that  is  supported  by  independent  lines 
of  evidence.  In  particular,  if  advances  in  the 
study  of  molecular  evolution  prove  to  be 
adaptable  to  the  study  of  small  insects,  we 
believe  that  Flexamia  would  provide  an  excel- 
lent model  for  study.  Because  the  knowledge 
of  Flexamia  hosts  is  now  adequate,  obtaining 
live  material  of  most  species  for  such  a  study 
would  be  an  achievable  task.  The  rewards  of 
further  detailed  studies  would  be  many  but 
would  include,  especially,  important  insights 
into  the  evolution  of  life-history  strategies  of 
sap-sucking  insects. 

When  we  began  our  studies  of  Flexamia, 
whose  species  number  had  remained  rela- 
tively constant  for  a  number  of  years,  we  did 
not  dream  that  there  were  many  remaining 
species  awaiting  discovery.  It  is  difficult  for  us 
to  guess,  given  the  results  reported  here,  how 
far  we  have  fallen  short,  even  now,  of  a  com- 
plete inventory.  Our  discovery  of  the  role  of 
climate  in  inducing  and  reinforcing  cicadellid 
speciation  makes  it  clear  that  the  biogeo- 
graphically  rich  areas  of  the  Southwest  may 
well  have  many  interesting  stories  that  still 
await  us. 

In  particular  there  may  well  be  Mexican 
species  awaiting  discovery.  For  example,  our 
phylogenetic  hypothesis  suggests  that  there 
may  have  been  at  one  time  a  species  with  the 
plesiomorphic  dorsal  stripes  of  Spartopyge, 
grammica,  and  albida,  but  which  possessed  a 
full  complement  of  four  aedeagal  processes. 
Because  of  the  compelling  logic  of  all  phyloge- 
netic schemes  for  its  existence,  such  a  species 
seems  so  real  to  us  that  we  have  jokingly  re- 
ferred to  it  as  ancestris.  Is  it  possible  that 
this  species,  or  a  closely  related  descendant  of 
it,  may  actually  exist  in  an  unexplored  Mexi- 
can grassland?  It  is  questions  such  as  these 
that  have  driven  us  to  return  repeatedly  to  the 
grasslands  of  the  Southwest,  which  prove  to 
be  every  bit  as  full  of  enigmas  as  they  are  of 
answers. 

Acknowledgments 

There  is  a  Japanese  proverb  that  advises 
those  who  drink  water  from  a  well  today  to 
remember  the  ancestors  who  dug  the  well. 
The  Flexamia  well  has  been  dug  with  many 


1988 


WniTcoMB,  HiCKS;  Flexamia  Revision 


313 


spades.  The  early  biological  work  of  E.  D. 
Ball,  H.  Osborn,  D.  M.  DeLong,  and  R.  H. 
Beamer  is  implicit  in  our  work.  H.  H.  Ross 
introduced  us  to  the  genus,  framed  many  of 
the  questions  that  we  discuss,  and  left  much 
unpublished  material  from  his  grasslands 
(GL)  project.  D.  A.  Young,  whose  1958  mono- 
graph with  B.  P.  Beirne  continues  to  be  an 
important  tool  for  students  of  Flexamia, 
kindly  permitted  us  to  redraw  many  struc- 
tures from  drawings  in  his  monograph.  We  are 
particularly  pleased  to  acknowledge  the  ad- 
vice and  encouragement  of  P.  W.  Oman, 
given  especially  to  the  senior  author  during 
his  early  years  at  Beltsville.  Further,  a  large 
fraction  of  USNM  Flexamia  had  been  col- 
lected and  correctly  determined  by  Oman; 
this  material  was  extremely  valuable  in  our 
study.  Earlier  studies  on  Flexamia  at 
Beltsville  were  assisted  by  M.  E.  Coan,  J.  P. 
Kramer,  and,  in  later  years,  J.  E.  Lowry, 
whose  illustration  oLserrata  is  presented  here 
as  Figure  1.  All  other  drawings  were  done  by 
Debi  Roney.  Robert  Almond  and  Edward 
Clark  gave  advice  and  assistance  on  the  graph- 
ics. Host  records  were  retrieved  from  a  com- 
puterized index  developed  by  D.  E.  Lynn. 
T.  Mogel  assisted  with  the  scanning  electron 
microscopy.  Assistance  with  phylogenetic 
programs  was  provided  by  Mary  Mickevich. 
We  acknowledge  further  the  assistance  of 
G.  W.  Byers,  P.  S.  Cwikla  and  C.  P.  Triple- 
horn,  M.  W.  Nielson,  and  T.  J.  Henry  and 
R.  C.  Froeschner,  who  made  available  the 
Flexamia  collections  of  Kansas  University, 
Ohio  State  University,  Monte  Bean  Life  Sci- 
ence Museum  (Brigham  Young  University), 
and  the  United  States  National  Museum,  re- 
spectively. K.  G.  A.  Hamilton  made  available 
the  Flexamia  material  from  the  Canadian  Na- 
tional Collection  and  provided  a  wealth  of 
stimulating  comments  on  the  manuscript. 
M.  SchaufF  made  many  useful  comments  on 
the  manuscript.  So  many  friends  have  helped 
in  our  field  studies  that  all  cannot  be  acknowl- 
edged. We  are  especially  indebted,  however, 
to  Kelly  Allred  of  New  Mexico  State  Univer- 
sity for  continuing  lessons  on  New  Mexico 
agrostology.  Charles  T.  Mason,  Jr.,  assisted 
with  grasses  of  Arizona.  R.  Chambers  of  the 
U.S.  Soil  Conservation  Service  drove  us  to 
the  type  locality  of  collortwi  and  called  our 
attention  to  the  host.  Mike  Fleming  of 
Big  Bend  National  Park  and  Jim  Walters  of 


Guadalupe  National  Park  assisted  field  stud- 
ies there,  and  Barton  Warnock  identified 
many  plant  specimens  from  Big  Bend  Na- 
tional Park  and  the  Davis  Mountains.  Riley 
Nelson  of  BYU  provided  collections  from  the 
Colorado  Plateau.  P.  Cubbage  assisted  with 
fieldwork.  Judith  Leach,  wife  of  the  senior 
author,  assisted  during  many  field  trips  and 
also  made  valuable  editorial  comments  on  the 
manuscript.  The  editorial  staff  of  The  Great 
Basin  Naturalist  helped  in  many  unusual 
ways  to  assure  timely  completion  of  the  paper. 
Finally,  H.  D.  Blocker  suggested  that  the 
time  for  this  paper  had  come,  sorted  more 
than  1,000  unidentified  Flexamia  from  the  KU 
cicadellid  collection,  provided  Flexamia  ma- 
terial and  records  from  the  KSU  collection, 
read  various  drafts  of  the  manuscript,  and  en- 
couraged us  in  countless  other  ways.  To  all 
these  workers,  and  many  that  remain  un- 
named, we  offer  our  sincerest  gratitude. 

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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 

0 

0 

0 

1 

8 

zamora 

OSU(h) 

OSU(p) 

0 

0 

0 

9 

0 

pectinata 

USNM(c) 

USNM(c) 

57 

5 

150 

45 

75 

gila 

USNM(h) 

P^ 

9 

0 

0 

0 

0 

handarita 

USNM(h) 

P^ 

5 

0 

0 

6 

0 

doeringae 

KU(h) 

USNM(p) 

35 

0 

68 

24 

91 

collorum 

USNM(h) 

P^ 

81 

0 

0 

0 

0 

jacala 

USNM(h) 

none 

0 

0 

0 

0 

1 

mescalero 

USNM(h) 

P^ 

26 

0 

0 

0 

0 

abbreviata 

USNM(c) 

USNM(c) 

765 

141 

661 

74 

409 

zacate 

USNM(h) 

P^ 

96 

0 

0 

0 

0 

canyonensis 

USNM(h) 

USNM(p) 

9 

0 

0 

0 

2 

surcula 

osu 

OSU 

72 

0 

10 

57 

37 

curvata 

osu 

osu 

560 

81 

209 

77 

107 

grammica 

USNM(l) 

KU(p) 

47 

64 

164 

48 

109 

picta 

OSU(h) 

USNM(c) 

285 

107 

202 

135 

112 

pyrops 

KU(1) 

KU(c) 

56 

39 

56 

83 

162 

albida 

USNM(c) 

USNM(c) 

46 

1 

7 

17 

19 

slossonae 

USNM(h) 

none 

12 

0 

50 

0 

18 

serrata 

KU(h) 

KU(p) 

56 

98 

20 

0 

14 

ritana 

KU(h) 

USNM(p) 

0 

0 

17 

0 

5 

arizonensis 

USNM 

USNM(p) 

177 

11 

114 

20 

94 

flexulosa 

USNM(l) 

USNM(c) 

380 

211 

254 

51 

353 

arenicola 

USNM(h) 

USNM(p) 

91 

0 

0 

0 

5 

celata 

USNM(h) 

USNM(p) 

49 

0 

4 

0 

5 

stylata 

USNM(l) 

USNM(c) 

83 

67 

66 

9 

70 

decora 

KU 

USNM(p) 

95 

246 

177 

3 

28 

youngi 

USNM 

P^ 

30 

0 

5 

0 

0 

modica 

KU 

USNM(p) 

377 

0 

110 

3 

8 

inflata 

USNM(c) 

USNM(c) 

665 

103 

257 

146 

155 

beameri 

KU 

USNM(p) 

0 

0 

7 

0 

0 

texana 

USNM 

USNM(p) 

0 

0 

0 

0 

10 

imputans 

OSU(c) 

OSU(c) 

4 

0 

10 

11 

19 

areolata 

USNM(l) 

USNM(c) 

37 

51 

42 

51 

88 

prairiana 

OSU 

OSU 

612 

44 

193 

23 

172 

reflexa 

USNM 

USNM(c) 

25 

1 

1 

8 

26 

atlantica 

OSU 

OSU(p) 

378 

17 

8 

12 

44 

dakota 

USNM(h) 

USNM(p) 

209 

3 

1 

0 

2 

producta 

(BNM) 

USNM(c') 

4 

5 

136 

117 

267 

sandersi 

OSU(h) 

OSU(h^) 

45 

3 

60 

31 

126 

delongi 

(INHS) 

none 

0 

26 

1 

39 

13 

graminea 

OSU(h) 

none 

176 

28 

105 

10 

190 

clayi 

USNM(h) 

USNM(p) 

148 

9 

115 

47 

3 

satilla 

CNC(h) 

CNC(p) 

0 

2 

2 

0 

0 

Numbers  reflect  content  of  collection  before  redistribution  at  end  of  study.  Type  distribution  occurred  after  this  enumeration  (c  ~  cotype.  h      holotype; 

1  ~  lectotype;  p  -  paratype).  Parentheses  -  specimen  not  examined. 
See  text  for  distribution  of  paratypes. 
Cotype  o( vise nda('  producta). 
Hohtype  o( hidentata  (~  sandersi). 


1988 


Whitcomb,  Hicks:  F'lexamia  Revision 
Appendix  II  Part  I.  Character  matrix  for  PHYSYS. 


317 


DISTIN 

SIMPLE 

ABSENT? 

7 

SINUATC 

THICK 

1 

ELONGA 

SHORTL 

TAPER 

ABSNT 

FLAREDG 

APODNOX 

APOPRIM 

2 

ELONGA 

CONNAR 

FORCEPY 

PRIMPL 

DEEPST 

ABSTR 

BRFACE 

3 

ELONGA 

DISTIN 

SIMPLE 

ABSENTP 

7 

LEFTC 

THICK 

1 

MEXICAN 

SHORTL 

TAPER 

ABSNT 

APICALG 

APODEXP 

APOPRIM 

2 

MEXICAN 

CONNAR 

PRIMPY 

PRIMPL 

PARALST 

PRESTR 

CONBLF 

3 

MEXICAN 

DISTIN 

SIMPLE 

ABSENTP 

7 

LEFTC 

THIN 

1 

MIRAND 

SHORTL 

TAPER 

ABSNT 

APICALG 

APODEXP 

APOPRIM 

2 

MI RAND 

CONNAR 

CNSTRPY 

PRIMPL 

EXCVST 

PRESTR 

CONBLF 

3 

MIRAND 

DISTIN 

FLARED 

ABSENTP 

7 

LEFTC 

THIN 

1 

MINIMA 

SHORTL 

TAPER 

ABSNT 

FLAREDG 

APODEXP 

APOPARL 

2 

MINIMA 

CONNAR 

BULBPY 

ACUTPL 

TOOTHST 

ABSTR 

BRFACE 

3 

MINIMA 

DISTIN 

FLARED 

ABSENTP 

7 

LEFTC 

THIN 

1 

ZAMORA 

SHORTL 

TAPER 

ABSNT 

FLAREDG 

APODEXP 

APOPARL 

2 

ZAMORA 

CONNAR 

BULBPY 

LNGBRPL 

OUTRST 

ABSTR 

BRFACE 

3 

ZAMORA 

DISTIN 

FLARED 

ABSENTP 

7 

LEFTC 

THIN 

1 

PECTIN 

SHORTL 

TAPER 

ABSNT 

FLAREDG 

APODEXP 

APOPARL 

2 

PECTIN 

CONNAR 

BULBPY 

RECTPL 

TOOTHST 

ABSTR 

BRFACE 

3 

PECTIN 

DISTIN 

FLARED 

ABSENTP 

7 

LEFTC 

THIN 

1 

BANDAR 

SHORTL 

TAPER 

ABSNT 

FLAREDG 

APODEXP 

APORECU 

2 

BANDAR 

CONNAR 

BULBPY 

LNGBRPL 

INNRST 

ABSTR 

BRFACE 

3 

BANDAR 

DISTIN 

FLARED 

ABSENTP 

7 

LEFTC 

THIN 

1 

GILA 

SHORTL 

TAPER 

ABSNT 

FLAREDG 

APODEXP 

APORECU 

2 

GILA 

CONNAR 

BULBPY 

LNGBRPL 

INNRST 

ABSTR 

BRFACE 

3 

GILA 

DISTIN 

FLARED 

ABSENTP 

7 

LEFTC 

THIN 

1 

DOERIN 

SHORTL 

TAPER 

ABSNT 

FLAREDG 

APODEXP 

APOPARL 

2 

DOERIN 

CONNAR 

BULBPY 

RECTPL 

PROCST 

ABSTR 

CONBLF 

3 

DOERIN 

DISTIN 

FLARED 

ABSENTP 

7 

LEFTC 

THIN 

1 

COLLOR 

SHORTL 

TAPER 

ABSNT 

FLAREDG 

APODEXP 

APORECU 

2 

COLLOR 

CONNAR 

BUDPY 

SHORTPL 

TOOTHST 

ABSTR 

BRFACE 

3 

COLLOR 

DISTIN 

FLARED 

ABSENTP 

7 

LEFTC 

THIN 

1 

JACALA 

SHORTL 

TAPER 

ABSNT 

FLAREDG 

APODNOX 

APORECU 

2 

JACALA 

CONNAR 

BULBPY 

LNGBRPL 

7 

ABSTR 

BRFACE 

3 

JACALA 

DISTIN 

CAPITAT 

ABSENTP 

7 

LEFTC 

THIN 

1 

MESCAL 

SHORTL 

TAPER 

ABSNT 

FLAREDG 

APODNOX 

APOCLOS 

2 

MESCAL 

CONNAR 

BULBPY 

LNGROPL 

DEEPST 

ABSTR 

BRFACE 

3 

MESCAL 

DISTIN 

SHORTPR 

ABSENTP 

7 

STRAIT 

LESTHIC 

1 

ABBREV 

LONGERL 

UNTAPER 

ABSNT 

SLITG 

APODEXP 

APOPARL 

2 

ABBREV 

CONNAR 

PRODPY 

LOBATPL 

INCISST 

ABSTR 

BRFACE 

3 

ABBREV 

DISTIN 

FLARED 

ABSENTP 

7 

LEFTC 

THICK 

1 

CANYON 

SHORTL 

TAPER 

ABSNT 

MIDLENG 

APODNOX 

APORECU 

2 

CANYON 

CONNAR 

PRODPY 

LOBATPL 

LONGST 

ABSTR 

BRFACE 

3 

CANYON 

DISTIN 

FLARED 

ABSENTP 

7 

LEFTC 

THIN 

1 

ZACATE 

SHORTL 

TAPER 

ABSNT 

SLITG 

APODNOX 

APORECU 

2 

ZACATE 

CONNAR 

PRODPY 

LOBATPL 

PROJST 

ABSTR 

BRFACE 

3 

ZACATE 

DISTIN 

ASYMPR 

ABSENTP 

7 

LEFTC 

THICK 

1 

GRAMMI 

SHORTL 

TAPER 

SERRATE 

BIPROVG 

APODNOX 

APOWING 

2 

GRAMMI 

CONNAR 

NARRPY 

SQUATPL 

ROUNDST 

PRESTR 

CONBLF 

3 

GRAMMI 

DISTIN 

LONGPR 

SUBAPP 

7 

STRAIT 

LESTHIC 

1 

SURCUL 

LONGERL 

UNTAPER 

ENTIRE 

BIPROVG 

APODNOX 

APOFORK 

2 

SURCUL 

CONWIDE 

SHORTPY 

SHORTPL 

BOXST 

ABSTR 

BRFACE 

3 

SURCUL 

DISTIN 

LONGPR 

DISTALP 

7 

STRAIT 

LESTHIC 

1 

CURVAT 

LONGERL 

UNTAPER 

ENTIRE 

BIPRSLG 

APODEXP 

APOFORK 

2 

CURVAT 

CONNAR 

SHORTPY 

SHORTPL 

WIDINST 

ABSTR 

BRFACE 

3 

CURVAT 

INDIST 

FLANGED 

SUBAPP 

7 

STRAIT 

THIN 

1 

PICTA 

LONGSTL 

,  UNTAPER 

SERRATE 

SUBAPIG 

APODNOX 

APOFORK 

2 

PICT  A 

CONNAR 

ANGLPY 

TAPERPL 

SERRST 

ABSTR 

CONBLF 

3 

PICTA 

INDIST 

FLANGED 

APPOSP 

? 

SINUATC 

THIN 

1 

PYROPS 

LONGSTL 

,  UNTAPER 

SERRATE 

SUBAPIG 

APODNOX 

APOFORK 

2 

PYROPS 

CONNAR 

ANGLPY 

REDUCPL 

EXTHST 

ABSTR 

TOOTHF 

3 

PYROPS 

318 


Great  Basin  Naturalist  Memoirs 
Appendix  II.  Part  I  continued. 


No.  12 


DISTIN 

BIFURC 

FUSEDP 

SAGGITV 

LESSC 

THICK 

1 

ALB I DA 

SHORTL 

TAPER 

SERRATE 

SUBVNPG 

APODEXP  APOCLUB 

2 

ALBIDA 

CONWIDE 

WIDEPY 

TAPERPL 

TTHEXST 

PRESTR 

CONBLF 

3 

ALBIDA 

PARTLY 

BIFURC 

FUSEDP 

SAGGITV 

RIGHTC 

THICK 

1 

SLOSSO 

SHORTL 

UNTAPER 

ENTIRE 

SYMSLTG 

APODNOX 

APOWING 

2 

SLOSSO 

CONPAFU 

UNDIFPY 

ACUTPL 

TRILBST 

PRESTR 

CONBLF 

3 

SLOSSO 

DISTIN 

BIFURC 

FUSEDP 

SAGGITV 

STRAIT 

THICK 

1 

SERRAT 

SHORTL 

UNTAPER 

ENTIRE 

SUBVNPG 

APODEXP 

APOCLUB 

2 

SERRAT 

CONWIDE 

IRREGPY 

CLEAVPL 

LOBEDST 

ABSTR 

CONBLF 

3 

SERRAT 

DISTIN 

LYRIFO 

FUSEDP 

SAGGITV 

LESSC 

THICK 

1 

R I TANA 

SHORTL 

UNTAPER 

ENTIRE 

SUBVNPG 

APODNOX 

APOCLAW 

2 

RITANA 

CONNAR 

OVATEPY 

TAPERPL 

ANGLST 

REDUSTR 

CONBLF 

3 

RITANA 

DISTIN 

SYMMET 

SHORTV 

SAGGITV 

LESSC 

LESTHIC 

1 

ARENIC 

LONGERL 

UNTAPER 

ENTIRE 

SUBAPIG 

APODNOX 

APOAWAY 

2 

ARENIC 

CONNAR 

ROUNDPY 

INCISPL 

NOTCHST 

ABSTR 

CONBLF 

3 

ARENIC 

DISTIN 

SYMMET 

SHORTV 

SAGGITV 

STRAIT 

LESTHIC 

1 

CELATA 

LONGERL 

UNTAPER 

ENTIRE 

SUBAPIG 

APODNOX 

APOPARL 

2 

CELATA 

CONWIDE 

THSTYPY 

INCISPL 

NOTCHST 

ABSTR 

CONBLF 

3 

CELATA 

DISTIN 

ASYMME 

TWISTV 

DISPLCV 

RIGHTC 

LESTHIC 

1 

STYLAT 

LONGERL 

UNTAPER 

ENTIRE 

SUBAPIG 

APODNOX 

APOTHIC 

2 

STYLAT 

CONBROD 

STYLPY 

SQUARPL 

BINOTST 

ABSTR 

CONBLF 

3 

STYLAT 

DISTIN 

SYMMET 

SHORTV 

SAGGITV 

STRAIT 

LESTHIC 

1 

ARIZON 

LONGERL 

UNTAPER 

ENTIRE 

SUBAPIG 

APODNOX 

APOFORK 

2 

ARIZON 

CONNAR 

VNPRPY 

MINUTPL 

SQUARST 

ABSTR 

CONBLF 

3 

ARIZON 

DISTIN 

SYMMET 

SHORTV 

SAGGITV 

LEFTC 

LESTHIC 

1 

FLEXUL 

LONGERL 

UNTAPER 

SERRATE 

SUBAPIG 

APODNOX 

APOFORK 

2 

FLEXUL 

CONNAR 

CAUDPY 

MINUTPL 

MEDNST 

ABSTR 

CONBLF 

3 

FLEXUL 

DISTIN 

SYMMET 

FUSEDP 

SAGGITV 

LEFTC 

THICK 

1 

DECORA 

SHORTL 

TAPER 

SERRATE 

APPRG 

APODNOX 

APOAWAY 

2 

DECORA 

CONBROD 

LOBEDPY 

INCISPL 

BINOTST 

ABSTR 

CONBLF 

3 

DECORA 

DISTIN 

ASYMME 

RECURVP 

DISPLCV 

LESSC 

LESTHIC 

1 

YOUNG I 

LONGERL 

TAPER 

SERRATE 

APPRG 

APODNOX 

APOTHIC 

2 

YOUNG I 

CONBROD 

BROADPY 

INCISPL 

BINOTST 

ABSTR 

CONBLF 

3 

YOUNGI 

DISTIN 

ASYMME 

DECURVP 

DISPLCV 

LEFTC 

THICK 

1 

MODICA 

SHORTL 

TAPER 

SERRATE 

SUBAPIG 

APODNOX 

APOAWAY 

2 

MODICA 

CONBROD 

OVATEPY 

DEEPL 

NOTCHST 

ABSTR 

CONBLF 

3 

MODICA 

DISTIN 

ASYMME 

DECURVP 

DISPLCV 

RIGHTC 

THIN 

1 

INFLAT 

LONGSTL 

UNTAPER 

ENTIRE 

SUBAPIG 

APODNOX 

APOFORK 

2 

INFLAT 

CONWIDE 

BOATPY 

TAPERPL 

BINOTST 

ABSTR 

CONBRF 

3 

INFLAT 

DISTIN 

TWISTED 

RECURVP 

DISPLCV 

RIGHTC 

THIN 

1 

BEAMER 

LONGSTL 

UNTAPER 

ENTIRE 

APPRG 

APODNOX 

APOTHIC 

2 

BEAMER 

CONBROD 

SHIPPY 

SHORTPL 

BINOTST 

ABSTR 

CONBRF 

3 

BEAMER 

DISTIN 

TWISTED 

TWISTV 

DISPLCV 

STRAIT 

THIN 

1 

TEXANA 

LONGSTL 

UNTAPER 

ENTIRE 

APPRG 

APODNOX 

APOREDU 

2 

TEXANA 

CONWIDE 

SHIPPY 

SHORTPL 

BINOTST 

ABSTR 

CONBRF 

3 

TEXANA 

DISTIN 

TWISTED 

? 

LATERLV 

STRAIT 

THIN 

1 

IMPUTA 

LONGSTL 

UNTAPER 

SERRATE 

LATSLTG 

APODNOX 

APOHOOK 

2 

IMPUTA 

CONWIDE 

LGOVPY 

TAPERPL 

LOBEDST 

ABSTR 

BLFACE 

3 

IMPUTA 

FUSED 

TWISTED 

? 

LATERLV 

LEFTC 

LESTHIC 

1 

AREOLA 

LONGERL 

UNTAPER 

SERRATE 

LATSLTG 

APODNOX 

APOAWAY 

2 

AREOLA 

CONWIDE 

BIZARPY 

TAPERPL 

PROCST 

ABSTR 

BLFACE 

3 

AREOLA 

BARELY 

INVERT 

7 

DORSALV 

STRAIT 

LESTHIC 

1 

PRAIRI 

LONGERL 

UNTAPER 

SERRATE 

DORSALG 

APODNOX 

APOPEN 

2 

PRAIRI 

CONFUSE 

ELLIPY 

NARRPL 

NAROWST 

ABSTR 

VARCONF 

3 

PRAIRI 

FUSED 

INVERT 

7 

BASFUSV 

STRAIT 

LESTHIC 

1 

REFLEX 

LONGERL 

UNTAPER 

SERRATE 

PROFUSG 

APODNOX 

APOCLOS 

2 

REFLEX 

CONFUSE 

UPTNPY 

NARRPL 

NAROWST 

ABSTR 

VARCONF 

3 

REFLEX 

FUSED 

WINDSW 

7 

FUSEDV 

STRAIT 

LESTHIC 

1 

ATLANT 

LONGERL 

UNTAPER 

SERRATE 

LATG 

APODNOX 

APOWIDE 

2 

ATLANT 

CONFUSE 

THINPY 

NOTCHPL 

NAROWST 

ABSTR 

VARCONF 

3 

ATLANT 

1988 


Whitcomb,  HiCKS;  Flexamia  Revision 
Appendix  II.  Part  I  continued. 


319 


FUSED 

INVTWO 

7 

TINYV 

STRAIT 

THICK 

1 

PRODUC 

FUSEDL 

UNTAPER 

SERRATE 

FUSEDG 

APODNOX 

APOPARL 

2 

PRODUC 

CONFUSE 

VNTLBPY 

BOTTLPL 

NAROWST 

ABSTR 

VARCONF 

3 

PRODUC 

FUSED 

INVTWO 

? 

FUSEDV 

STRAIT 

THICK 

1 

DAKOTA 

FUSEDL 

UNTAPER 

SERRATE 

DORFUSG 

APODNOX 

APOCLOS 

2 

DAKOTA 

CONFUSE 

POSTPY 

BOTTLPL 

NAROWST 

ABSTR 

BRFACE 

3 

DAKOTA 

FUSED 

FUSPR 

7 

REDUCV 

STRAIT 

THICK 

1 

SANDER 

FUSEDL 

UNTAPER 

SERRATE 

APPRSDG 

APODNOX 

APOWIDE 

2 

SANDER 

CONFUSE 

HIGHPY 

NOTCHPL 

NAROWST 

ABSTR 

BRFACE 

3 

SANDER 

FUSED 

FUSPR 

? 

REDUCV 

STRAIT 

THICK 

1 

DELONG 

FUSEDL 

UNTAPER 

SERRATE 

APPRSDG 

APODNOX 

APOWIDE 

2 

DELONG 

CONFUSE 

HIGHPY 

NOTCHPL 

NAROWST 

ABSTR 

BRFACE 

3 

DELONG 

FUSED 

ASKEW 

? 

FUSEDV 

STRAIT 

THICK 

1 

GRAMIN 

FUSEDL 

SPIRALT 

SERRATE 

SPIRALG 

APODNOX 

APOPARL 

2 

GRAMIN 

CONFUSE 

THICKPY 

RECURPL 

NAROWST 

ABSTR 

BRFACE 

3 

GRAMIN 

FUSED 

ASKEW 

? 

FUSEDV 

LEFTC 

THICK 

1 

CLAY  I 

FUSEDL 

SPIRALT 

SERRATE 

WIDSPIG 

APODNOX 

APOCLOS 

2 

CLAYI 

CONFUSE 

UPLOBPY 

RECURPL 

NAROWST 

ABSTR 

BRFACE 

3 

CLAYI 

FUSED 

ASKEW 

? 

FUSEDV 

LEFTC 

THICK 

1 

SATILL 

FUSEDL 

SPIRALT 

SERRATE 

WIDSPIG 

APODNOX 

APOCLOS 

2 

SATILL 

CONFUSE 

UPLOBPY 

RECURPL 

NAROWST 

ABSTR 

BRFACE 

3 

SATILL 

Appendix  II.  Part  II.  Hypothesized  character  transformation  in  Flexamia.  Data  for  PHYSYS. 


aedconn;distin-indist-partly-barely-fused; 
aedapex;simple-flared-capitat,shortpr-[l6ngpr-flanged, 
bifurc, lyrifo, symmet-asymme-twisted- [ invert- windsw, 
[fuspr- askew] , invtwo] ] ; 

ANTEPR;ABSENTP-SUBAPP-DISTALP-APP0SP-[FUSEDP-DECURVP, 

RECURVP, [LONGV-SHORTV-TWISTV] ] ; 

VENTRALP;SAGGITV-DISPLCV-LATERLV-D0RSALV-BASFUSV-REDUCV 

TINYV-FUSEDV; 

AEDCURV;LEFTC-LESSC-STRAIT-SINUATC-RIGHTC; 

AEDTHIC ; THICK-LESTHIC-THIN ; 

AEDLENG ; SHORTL-LONGERL-LONGSTL-FUSEDL ; 

AEDTAPER; TAPER-UNTAPER- SPIRALT ; 

AEDPRMG; ENTIRE-SERRATE; 

GONOLOC;APICALG-FLARED-[SLITG-MIDLENG] , [BIPROVG- [ SUBVNPG- 

SYMSLTG] ,BIPRSLG, [ SUBAPIG-APPRG, LATSLTG-DORSALG- [PROFUSG- 

[APPRSDG-SPIRALG-WIDSPIG] , [FUSEDG-DORFUSG] , LATG] ] ] ; 

APOAPEX ; APODEXP- APODNOX ; 

APOARM;APOPRIM- [APOPARL- [APOCLAW-APOHOOK- APOPEN-APOCLOS- 

APOWIDE] , [APOAWAY-APOTHIC-APOREDU] ,APOFORK, [APOCLUB- 

APOWING ] , APORECU ] ; 

CONKEEL;CONNAR-CONWIDE-CONBROD-CONPAFU-CONFUSE; 

PYGMAL ; PRIMPY-FORCEPY- [ CNSTRPY- [ NARRPY- IRREGPY-WIDEPY- 

UNDIFPY], [BULBPY-BUDPY,PRODPY, [ SHORTPY-ANGLPY, [OVATEPY- 

[LOBEDPY-[BOATPY-SHIPPY] ,BROADPY] , [ROUNDPY- [ VNPRPY-CAUDPY] , 

[STYLPY-THSTPY] ] , [ LGOVPY-BIZARPY, [ELLIPY- VNTLBPY, [UPTNPY- 

POSTPY,THINPY, [HIGHPY-UPLOBPY-THICKPY] ]]]]]]]; 

PLATES ;  PRIMPL-  [  LNGBRPL-RECTPL ,  SHORTPI^  ]  ,  SQUATPL ,  [  LNGROPL- 

LOBATPL-REDUCPL] , [TAPERPL- [NARRPL-BOTTLPL-NOTCHPL-RECURPL] , 

ACUTPL, [CLEAVPL-[INCISPL-SQUARPL, [DEEPL-MINUTPL] ] ] ] ; 

STERNUM; PAEIALST-EXCVST-ROUNDST, [TOOTHST- [OUTRST-DEEPST] , 

[INNRST-PROCST] , [ INCISST-LONGST-PROJST] , [BOXST-WIDINST, 

[SERRST-EXTHST] ] ] , [TTHEXST-TRILBST, LOBEDST- [ ANGLST- 

[NOTCHST-BINOTST] , [ PROCST-NAROWST ] , [ SQUARST-MEDNST ] ] ] ; 

DORSVIT;  PEIESTR-REDUSTR- ABSTR; 

FACE;CONBLF-TOOTHF, [CONBRF-BRFACE] , [ VARCONF -BLF ACE] ; 


1  AEDCONN 

1  AEDAPEX 

2  AEDAPEX 

3  AEDAPEX 

1  ANTEPRO 

2  ANTEPRO 

1  VENTRLP 

2  VENTRLP 
1  AEDCURV 
1  AEDTHIC 
1  AEDLENG 

1  AEDTAPER 
1  AEDPRMG 

1  GONOLOC 

2  GONOLOC 

3  GONOLOC 
1  APOAPEX 

1  APOARM 

2  APOARM 

3  APOARM 

1  CONKEEL 

1  PYGMAL 

2  PYGMAL 

3  PYGMAL 

4  PYGMAL 

5  PYGMAL 

1  PLATES 

2  PLATES 

3  PLATES 

1  STERNUM 


2  STERNUM 

3  STERNUM 

4  STERNUM 
1  DORSVIT 
1  FACE 


320 


Great  Basin  Naturalist  Memoirs 


No.  12 


Appendix  II.  Part  III 

Character  Codes  of  Flexomia  Species: 

Code  Identification* 
P  =  plesiomorphic;  A  ^  apomorphic 

1.  AEDCONN:  Articulation  (P)  or  fusion 
(A)  of  aedeagus  and  connective:  DISTIN  (P) 
(Figs.  6A,B,C,E);  INDIST  (Fig.  6D);  BARE- 
LY (Fig.  6F);  PARTLY  (Y&B  Fig.  9B)  FUSED 
(Fig.  60). 

2.  AEDAPEX:  Morphology  of  aedeagal 
apex  which  may  be  simple,  without  processes 
or  special  structures  (P),  or  with  paired  pro- 
cesses (A):  SIMPLE  (P)  (Figs.  55C,F); 
FLARED  (Figs.  12A-H);  CAPITAT  (Figs. 
121,22);  SHORTPR  (Fig.  7H);  ASYMPR  (Fig. 
7E);  LONGPR  (Figs.  7F,G);  FLANGED 
(Figs.  7K,L);  BIFURC  (Figs.  7A-C);  LYRIFO 
(Fig.  7D);  SYMMET  (Figs.  31G,K); 
ASYMME  (Figs.  31E,H,M);  TWISTED 
(Fig.  31A);  INVERT  (Figs.  44D,G); 
WINDSW  (Fig.  44A);  INVTWO  (Fig.  441); 
FUSPR  (Fig.  44B);  ASKEW  (Figs.  44F,H). 

3.  ANTEPR:  Anteapical  processes,  which 
are  absent  (P),  present  as  paired  processes,  or 
fused  (A)  into  a  single  process:  ABSENTP 
(Figs.  55C,F);  SUBAPP  (Fig.  7G);  DISTALP 
(Fig.  7F);  APPOSP  (Fig.  7L);  FUSEDP  (Figs. 
7A-D);  LONGV  (Fig.  31G);  RECURVP 
(Fig.  38A);  DECURVP  (Fig.  31E);  SHORTY 
(Figs.  31I-L);  TWISTY  (Fig.  31A). 

4.  YENTRLP:  Phyletic  twisting  of  the  un- 
paired ventral  process  (see  3.  ANTEPR), 
which  was  (P)  symmetrically  situated  on  the 
ventral  surface  of  the  aedeagus,  but  moved  to 
the  dorsum,  and  was  (A)  ultimately  lost 
(Fig.  61),  presumably  by  fusion:  SAGGITY 
(Fig.  61A);  DISPLCY  (Fig.  61B);  LATERLY 
(Fig.  61C);  DORSALY(Fig.  61D);  BASFUSY 
(Fig.  61E);  REDUCY  (Fig.  61F);  TINYY 
(Fig.  61G);  FUSEDY(Fig.  61H). 

5.  AEDCURY:  Curvature  (lateral  aspect)  of 
the  aedeagus,  which  may  be  curved  sinistrad 
(P),  without  curvature,  sinuate,  or  curved 
dextrad  (A):  LEFTC  (Figs.  71, J);  LESSC 
(Y&B  Figs.  10B,11A);  STRAIT  (Fig.  7H); 
SINUATC  (Y&B  Fig.  2B);  RIGHTC  (Fig. 
4()A). 

6.  AEDTHIC:  Thickness  (lateral  aspect)  of 
the  aedeagus,  which  may  be  thick  (P),  less 
thick,  or  thin  (A);  THICK  (Fig.   7J);   LES- 


♦Characters  are  illustrated  herein,  in  Vonnj;  and  Beirne  (1958)  (V&B); 
Lowry  and  Blocker  (1987)  (L&B);  or  Hamilton  and  Koss  (1975)  (H&R). 


THICK  (Fig.  38A);  THIN  (Figs.  16A,  17A, 
ISA). 

7.  AEDLENG:  Length  of  the  aedeagus, 
which  may  be  short  (P),  longer,  even  longer, 
or,  finally  (A),  fused  with  the  connec- 
tive: SHORTL  (Figs.  16A,7I,J);  LONGERL 
(Fig.  6B);  LONGSTL  (Fig.  6D);  FUSEDL 
(Fig.  6G). 

8.  AEDTAPR:  Form  of  the  aedeagal  shaft 
which  may  be  tapered  (P),  untapered,  or,  ulti- 
matelv  (A),  tapered  in  association  with  a  spiral 
gonopore:  TAPER  (Figs.  7I,J);  UNTAPER 
(Fig.  7H);  SPIRALT(Fig.  6G). 

9.  AEDPRMG:  Form  of  the  margins  of  the 
paired,  apical,  aedeagal  processes  which,  if 
present,  may  have  entire  (P)  or  serrate  mar- 
gins (A):  ABSNT  (absent);  ENTIRE  (Figs. 
7F,G);  SERRATE  (Figs.  44B-D). 

10.  GONOLOC:  Form  and  position  of  the 
gonopore.  In  the  plesiomorphic  condition, 
this  structure  was  apical  and  subcircular  or 
ovate.  In  early  stages  in  transformation  the 
gonopore  was  subapical  on  the  ventral  sur- 
face. In  the  flexulosa  group  the  gonopore  may 
have  reversed  its  distad  movement  from  the 
aedeagal  apex  and  returned  to  the  apex,  or 
nearly  so;  but  in  imputans  and  areolata  the 
gonopore  became  associated  with  the  un- 
paired process,  which  had  moved  to  a  lateral 
position  on  the  aedeagal  shaft.  In  prairiana 
the  gradual  twisting  (see  4.  YENTRLP)  of  the 
processes  had  stabilized  with  the  gonopore  in 
a  dorsal,  anteapical  position.  Transformation 
from  this  condition  involved  association  with 
the  unpaired  process  and  movement  with  the 
process  as  it  became  appressed  to  the  aedea- 
gal shaft  and,  eventually,  fused  with  the  shaft. 
Ultimately,  the  fusion  led  to  an  elongate 
structure  that  winds  around  the  shaft  in  a 
helical  configuration:  APICALG  (Y&B  Fig. 
32B);  FLAREDG  (Figs.  12A-H);  SLITG  (Fig. 
7H);  MIDLENG  (Fig.  6C);  BIPROYG 
(Fig.  7G);  SUBYNPG  (Fig.  7B);  SYMSLTG 
(Fig.  7C);  BIPRSLG  (Fig.  7F);  SUBAPIC 
(Figs.  31E,J,M);  APPRO  (Fig.  31A);  LAT 
SLTO  (Fig.  610)  DORSALO  (Fig.  440) 
PROFUSO  (Fig.  44D);  APPRSDO  (Fig.  44B) 
SPIRALG  (Fig.  44F);  WIDSPIG  (Figs 
44H,J);  FUSEDO  (Fig.  44E);  DORFUSC 
(Fig.  441);  LATO  (Fig.  44A). 

11.  APOAPEX:  Apices  of  the  apodema 
arms  mav  be  expanded  (P)  or  not  expandet 
(A):  APODEXP  (Fig.  6B);  APODNOX  (Figs 
6A,D,E). 


1988 


WHITCOMB,  HICKS:  Fl.EXAMIA  RH\  ISION 


321 


12.  APOARM:  In  dorsal  aspect  the  apode- 
mal  arms  in  the  plesioniorphic  condition  were 
parallel  to  the  aedeagal  shaft.  Transformations 
from  this  state  (A)  involved  several  lines  of 
development,  some  of  which  involved  reduc- 
tion of  the  arms,  or  direction  of  the  apices 
toward  or  awav  from  the  sagittal  plane:  APO- 
PRIM  (Y&B  Fig.  32B);  APOPARL  (Fig.  6B); 
APOCLAW  (Y&B  Fig.  UB);  APOHOOK 
(Y&B  Fig.  21B);  APOPEN  (Fig.  6F);  APOC- 
LOS  (Y&B  Fig.  28C);  APOWIDE  (R&C  Fig. 
2);  APOAWAY  (Y&B  Fig.  14B);  APOTHIC 
(Fig.  40B);  APOREDU  (Y&B  Fig.  16B);  APO- 
FORK  (Figs.  6A,D,E);  APOCLUB  (Fig.  8B); 
APOWING  (Y&B  Fig.  27B);  APORECU 
(Figs.  6C,  16B,  17B,  19B,  20B). 

13.  CONKEEL:  In  lateral  aspect  the  dorsal 
keels  of  the  connective  may  be  narrow  (P), 
widened,  or  very  much  widened.  Alterna- 
tively (A),  the  connective  fused  with  the 
aedeagus,  a  process  that  resulted  in  the  loss  of 
an  independent  identity  for  the  keels:  CON- 
NAR  (Figs.  16A,  17A,  18A);  CONWIDE 
(Y&B  Fig.  12A);  CONBROD  (Fig.  40A); 
CONPAFU  (Y&B  Fig.  9B);  CONFUSE  (Fig. 
6G). 

14.  PYGMAL:    The    male    pygofer,    ple- 
siomorphically,  was  apparently  undifferenti- 
ated. From  this  state  a  defined  posterior  lobe 
differentiated,    as    in    Spartopyge   miranda. 
Several  lines  of  development  (A)  are  postu- 
lated (Fig.   62):  PRIMPY  (Fig.  62S);  FOR- 
CEPY  (Fig.  55G);  UNDIFPY  (Fig.  62T);  CN- 
STRPY  (Fig.  62N);  BULBPY  (Figs.  62C-G) 
BUDPY  (Fig.  62A);  PRODPY  (Figs.  62H,I) 
NARRPY  (Fig.  620);  IRREGPY  (Fig.  62P) 
SHORTPY   (Figs.    62K,L);    ANGLPY   (Fig 
62M);  WIDEPY  (Fig.  62Q);  OVATEPY  (Fig 
62U);  ROUNDPY  (Fig.  62LL);  STYLPY  (Fig 
6200);  THSTYPY  (Fig.  62PP);  LOBEDPY 
(Fig.      62FF);     BROADPY     (Fig.      62KK); 
BOATPY     (Fig.     62GG);     SHIPPY     (Figs. 
62HH,II);   LGOVPY  (Fig.   62V);   BIZARPY 
(Fig.  62Z);  ELLIPY  (Fig.  62W);  VNTLBPY 
(Fig.  62AA);  UPTNPY  (Fig.  62X);  POSTPY 
(Fig.  62BB)  THINPY  (Fig.  62Y);  HIGHPY 
(Fig.     62CC);     UPLOBPY     (Fig.     62DD); 
THICKPY    (Fig.     62EE)    CAUDPY    (Fig. 
62NN);  VNPRPY  (Fig.  62MM). . 

15.  PLATES:  The  male  plates,  plesiomor- 
phically,  were  apparently  long  and  relatively 
wide.  The  broad  aspect  of  the  plates  was  re- 
tained in  the  pectinata  group,  but  in  subse- 
quent lineages  tended  to  be  modified  to  be- 


come tapered.  In  the  flexulosa  group  the  ta- 
pered plates  became  notched  to  accommo- 
date dorsally  produced  pygofers;  in  arizonen- 
si.s  and  flexulosa  this  tendency  is  maximized. 
In  tlu>  praihana  group  the  tapered  design  was 
modified  by  narrowing  of  the  apical  third  of 
the  plates,  by  the  development  of  small  apical 
notches,  and  bv  the  plates  ultimately  becom- 
ing recurved:  PRIMPL  (Fig.  54C);  SQUATPL 
(Fig.  8F);  LNGROPL  (Fig.  Ill);  LNGBRPL 
(Fig.  UB);  RECTPL  (Figs.  11C,F); 
SHORTPL  (Fig.  IIG);  LOBATPL  (Fig.  8E); 
REDUCPL  (Fig.  9B);  TAPERPL  (Fig.  8A); 
NARRPL  (Fig.  45A);  BOTTLPL  (Fig.  45E); 
NOTCHPL  (Fig.  45C);  RECURPL  (Fig. 
45H);  ACUTPL  (Fig.  8B);  CLEAVPL  (Fig. 
8C);  MINUTPL  (Figs.  33A,B);  INGISPL 
(Figs.  33C,D,E);  SQUARPL  (Fig.  34E); 
DEEPL(Fig.  33F). 

16.  STERNUM:  The  sternum  VII  was  ap- 
parently plesiomorphically  excavated.  From 
this  condition  several  lines  of  development  (A) 
ensued.  Many  of  these  lines  involved  two 
pairs  of  teeth  separated  by  an  incision  on  the 
posterior  margin  of  the  sternum.  In  the  pecti- 
nata group  the  development  of  the  inner  or 
outer  sets  of  teeth  relative  to  one  another  is  an 
important  character  in  species  discrimination. 
In  the  flexulosa  and  prairiana  groups,  at  a 
time  that  aedeagal  evolution  was  proceeding 
rapidly,  the  sternum  VII  became  a  relatively 
stable  apomorphv  that  underwent  minimal 
transformation:  PARALST  (Y&B  Fig.  32E); 
EXCVST  (Y&B  Fig.  33F);  ROUNDST  (Fig. 
lOF);  TOOTHST  (Figs.  14A,C,G);  OUTRST 
(Fig.  14B);  DEEPST  (Fig.  14H);  INNRST 
(Figs.  14D,E);PROCST(Fig.  14F);  INCISST 
(Fig.  lOE);  LONGST  (Fig.  101);  PROJST 
(Fig.  10L);BOXST(Fig.  10J);WIDINST(Fig. 
lOK);  SERRST  (Fig.  lOG);  EXTHST  (Fig. 
lOH);  TTHEXST  (Fig.  lOA);  TRILBST  (Fig. 
lOB);  LOBEDST  (Fig.  IOC);  ANGLST  (Fig. 
lOD);  NOTCHST  (L&B  Figs.  IF,  2F); 
BINOTST  (Fig.  lOR);  PROCST  (Fig.  lOQ); 
NAROWST  (Fig.  lOS);  SQUARST  (Y&B  Fig. 
17F);MEDNST(Fig.  lOM). 

17.  DORSVIT:  Species  of  the  genus  Spar- 
topijge  possess  conspicuous  dorsal  stripes  (P) 
that  appear  to  be  homologous  with  the  stripes 
in  albida,  slossonae,  ritana,  and  grammica. 
The  loss  of  stripes  (A)  is  presumed  to  be  homo- 
plastic: PRESTR  (Figs.  2A-C,  54A);  RE- 
DUSTR  (Fig.  2D);  ABSTR  (Figs.  2E,F). 


322 


Great  Basin  Naturalist  Memoirs 


No.  12 


18.  FACE:  The  plesiomorphic  face  of  Flex- 
amia  (and  its  ancestor  Spartopyge)  is  pre- 
sumed to  be  pale,  with  a  conspicuously  con- 
trasting, black  interocular  band.  In  later 
lineages  this  habitus  aspect  was  modified,  as 
the  interocular  band  became  brown  rather 
than  black,  as  the  distinction  between  the  line 
and  the  remainder  of  the  face  became 
blurred,  and,  finally  (A),  as  the  face  became 
essentially  brown.  In  some  transitional  spe- 
cies, such  as  prairiana,  the  face  varies  widely 
fi-om  pale  with  a  contrasting,  brown  interocu- 
lar line  to  brown  to  (occasionally)  nearly  en- 
tirely black.  We  presume  that  loss  of  the  ple- 
siomorphic face  condition,  as  with  dorsal 
stripes,  probably  occurred  in  several  lines: 
CONBLF  (Figs.  3A,B,E,F,G);  TOOTHF 
{pyrops:  see  text);  BRFACE  (Fig.  3H);  CON- 
BRF  (contrasting  brown  face)  VARCONF 
(variable  contrast);  BLFACE  (Fig.  3C). 

Appendix  111 

Notes 

Plant  names. — Andropogon  gerardii  Vit- 
man,  big  bluestem;  Andropogon  virginicus 
L.,  broomsedge;  Aristida  dichotoina  Michx. 
var.  curtisii  Gray;  Aristida  purpurea  Nutt.; 
Bouteloua  curtipendula  (Michx.)  Torr.,  side- 
oats  grama;  Bouteloua  eriopoda  (Torr.)  Torr., 
black  grama;  Bouteloua  gracilis  (Willd.  ex 
H.  B.  K.)  Lag.  ex  Griffiths,  blue  grama; 
Bouteloua  hirsuta  Lag.,  hairy  grama;  Boute- 
loua pectinata  Featherley,  tall  grama;  Boute- 
loua uniflora  Vasey,  Nealley  grama;  Buchlo'e 
dactyloides  (Nutt.)  Engelm.,  buffalograss; 
Calamovilfa  gigantea  (Nutt).  Scribn.  & 
Merr.,  big  sandreed;  Calamovilfa  longifolia 
(Hook.)  Scribner.,  prairie  sandreed;  Cynodon 
dactylon  (L.)  Pers.,  bermudagrass;  Distichlis 
spicata  (L.)  Greene,  salt  grass;  Eragrostis 
curvula  (Schrad.)  Nees,  weeping  lovegrass; 
Eragrostis  pectinacea  (Michx.)  Nees,  tufted 
lovegrass;  Eragrostis  spectahilis  (Pursh) 
Steud.,  purple  lovegrass;  Juncus  tenuis 
Willd.,  poverty  rush;  Leptoloma  cognatum 
(Shult)  Chase,  fall  witchgrass;  Muhlenbergia 
asperifolia  (Nees  &  Mey.)  Paradi,  scratch- 
grass;  Muhlenbergia  cuspidata  (Torr.  in 
Hook.)  Rydb.,  plains  muhly;  Midilenhergia 
pauciflora  Buckley,  New  Mexican  muhly; 
Muhlenbergia  porteri  Scribn.  ex  Beal,  bush 
muhly;  Muhlenbergia  pungens  Thurb.,  sand 


muhly;  Muhlenbergia  repens  (Presl.)  A.  S. 
Hitchc,  creeping  muhly;  Muhlenbergia 
reverchonii  Vasey  &  Scribn.,  seep  muhly; 
Panicum  capillare  L.,  witchgrass;  Panicum 
virgatum  L.,  switchgrass;  Poa  pratensis  L., 
Kentucky  bluegrass;  Redfieldia  flexuosa 
(Thurb.)  Vasey,  sand  blowout  grass;  Schiz- 
achyrium  scoparium  (Michx.)  Nash,  little 
bluestem;  Sporobolus  virginicus  (L.)  Kunth., 
seashore  dropseed;  Zoysia  japonica  Steudl., 
zoysia  grass. 

Plant  ranges. — Plant  ranges  were  mapped 
using  published  range  maps  (Pohl  1968; 
Hitchcock  1971;  McGregor  1977;  Stubben- 
dieck  et  al.  1986)  and/or  various  floras  (Gould 
1975;  Harrington  1954;  Jepson  1925;  Johnston 
1943,  1963;  Kearny  and  Peebles  I960;  Kiich- 
ler  1964;  Lesueur  1945;  Lonard  and  Judd 
1980;  Muller  1947;  Rzedowski  1966,  1973, 
1975,  1983);  Shreve  1939,  1942;  Steyermark 
1963;  Pohl  1966;  and  Welsh  et  al.  1987). 

Insect  ranges. — Occurrence  of  specimens 
was  mapped  on  maps  of  the  United  States  or 
North  America  (Goode  Base  Map,  Depart- 
ment of  Geography,  University  of  Chicago, 
was  used  as  a  base).  Specimens  examined 
from  the  collections  listed  below  were 
mapped.  Canadian  and  Mexican  records  are 
mapped  by  locality;  U.S.  records  were 
mapped  by  county  to  avoid  extensive  overlap 
of  symbols.  Some  records  from  the  collections 
of  the  Illinois  State  Natural  History  Survey  (as 
reported  in  DeLong  1948),  and  the  Kansas 
State  University  Collection  were  also 
mapped. 

Other  Records. — Not  all  "records"  were 
mapped.  We  refrained  from  mapping  speci- 
mens that  in  our  judgment  represented  possi- 
ble or  probable  errors.  These  records  in- 
cluded the  following  record  for  clayi:  "Colfax, 
Gal.,  8-7-30,"  from  the  OSU  collection.  Since 
clayi  is  a  common  Ohio  species,  but  appar- 
ently does  not  occur  west  of  the  Mississippi 
River,  this  record  is  probably  in  error.  Three 
records  for  picta  seem  questionable.  Perhaps 
"Coronado  N.F.,  Pena  Blanca  Resort,  Ari- 
zona, 19  Aug.  1970,  Harris  and  Harris, 
GL2019"  (CNC)  is  valid.  However,  "Panama 
Citv,  C.  Z.,  Julv  23,  1920"  (OSU),  and  "Mina, 
Mineral  Co.  Nevada,  Aug.  20,  1955,  GL274, 
Galloway"  (CNC)  seem  less  likely  to  be  cor- 
rect. For  other  unusual  records,  see  discus- 
sion under  decora,  areolata,  and  graminea. 


1988 


Whitcomb,  Hicks:  Flexamia  Revision 


323 


In  all  studies,  some  specimens  (usually  fe- 
males) remain  unidentified  at  the  end  of  the 
study.  One  record  of  this  kind  is  intriguing. 
The  label  "Cowan,  Manitoba,  8  Aug.,  1946, 
R.  H.  Beamer"  (KU)  is  affixed  to  a  single 
female  that  we  cannot  name.  See  field 
notes  (later  in  this  appendix)  concerning  this 
record. 

Collections  utilized. — We  examined  the 
Flexamia  collections  of  the  United  States  Na- 
tional Museum  (USNM),  Ohio  State  Univer- 
sity (OSU),  University  of  Kansas  (KU),  and 
the  Canadian  National  Collection  (CNC).  Our 
own  collection  (Beltsville  Agricultural  Re- 
search Center  [BARC])  contains,  in  addition 
to  mounted  specimens,  a  large  inventory  of 
unmounted  but  accessioned  (Lynn  and  Whit- 
comb 1987)  specimens.  The  inventories  given 
in  Appendix  I  are  summations  of  cleared  and 
uncleared  specimens.  We  feel  that  the  risk  of 
misdeterminations  of  uncleared  specimens  is 
very  low,  with  some  important  caveats.  Fe- 
males of  the  prairiana  and  flexiilosa  groups 
must  be  cleared  for  determination,  and  males 
of  the  prairiana  group  should  be  cleared.  In 
many  cases  large  series  were  available,  in 
which  case  we  cleared  several  specimens  and 
tentatively  identified  others  by  inference.  For 
these  reasons  the  numbers  given  probably  are 
an  accurate  reflection  of  the  relative  abun- 
dance of  each  species.  We  attempted  to  curate 
the  collections  examined  and  to  supplement 
them,  where  possible,  with  additional  mate- 
rial. 

IPL  ACCESSIONS. — The  BARC  collection  is 
largely  computerized  (Lynn  and  Whitcomb 
1987);  collection  records  are  given  in  terms  of 


the  accession  number  in  this  system.  Collec- 
tion methods  have  been  described  (Whit- 
comb etal.  1986,  1987). 

Original  Beamer  field  notes. — Relevant 
excerpts  from  field  notes  written  by  Mrs. 
Lucy  D.  Beamer  on: 

ritana.  "08-18-35.  Santa  Rita  Mts.,  Ariz.  (E. 
side)  11:30-5  PM.  mild,  fair  to  cloudy  ...  Dr. 
Ball,  RHB  (R.  H.  Beamer)  and  Jack  climbed 
the  trail  up  the  mountain.  Just  as  they  were 
abandoning  the  search  RHB  took  a  new  spe- 
cies of  Gladioniira  or  Athysanella  which  Dr. 
Ball  is  naming.  Worked  for  a  good  while,  fi- 
nally had  30  or  40  of  them.  Before  this  only  3 
specimens  had  been  taken."  There  are  no  ex- 
plicit references  to  Flexamia,  or  to  the  exact 
collecting  location.  The  Athysanella  species 
mentioned  was  presumably  A.  (Gladionura) 
furculata. 

beameri.  "07-26-46.  Otter  Lake,  New  York 
(5  miles  west).  Collected  just  inside  Adiron- 
dack Park  on  Highway  365  at  one  of  the  first 
little  lakes.  R.  [R.  H.  Beamer]  swept  tiny 
sedges  along  the  shore — took  80  or  so  of  a  tiny 
fulgorid  he  thought  was  Bakerella  sp.  Later 
not  so  sure. "  Again,  there  was  no  explicit  men- 
tion of  Flexamia. 

Unidentifiable  female.  "08-07-37.  Cowan, 
Manitoba  (3  miles  north).  R.  swept  in  the  tall 
green  grass  growing  up  through  the  heavy  mat 
of  fallen  grass  which  surrounds  a  lake  for  fifty 
or  a  hundred  yards  in  all  directions.  Found 
interesting  cicadellid — long  wings — proba- 
bly belonging  to  the  genus  Polyamia.  Their 
wings  were  ruffed  even  when  he  took  them." 
We  have  been  unable  to  find  other  specimens 
from  this  collection  in  the  KU  collection. 


AFRONISIA,  A  NEW  AFRICAN  GENUS  OF  MEENOFLIDAE 
(HOMOPTERA:  FULGOROIDEA) 

Michael  R.  Wilson' 

Abstract — The  new  genus  Afronisia  is  founded  with  Eponisia  albovittata  Fennah  as  the  type-species.  Eponisia 
briinnescens  Synave,  £.  flavescens  Synave,  E.  pallida  Linnavuori,  and  Nisia  muiri  Metcalf  are  transferred  to  this 
genus,  and  E.  albinervosa  Muir  is  placed  in  synonymy  with  Kcnnesia  albinervosa  Muir.  Two  new  species  are 
described:  Afronisia  bredoi,  from  Zaire,  and  A.  gembtiensis,  from  Nigeria.  Keys  are  provided  for  separation  of  the 
African  genera  of  Kermesiinae  and  for  the  seven  species  oi Afronisia  that  are  described  and  figured. 


The  Old  World  planthopper  family  Meeno- 
plidae  is  one  of  the  smaller  fulgoroid  families, 
consisting  of  slightly  more  than  100  described 
species.  Until  recently  all  species  were  placed 
in  eight  genera  in  two  subfamilies,  the 
Meenoplinae  and  the  Kermesiinae  (  =  Nisii- 
nae).  However,  Emeljanov  (1984)  and  Tsaur 
et  al.  (1986)  described  new  genera  for  existing 
species  in  the  Oriental  region  and  named  sev- 
eral new  species.  Over  50  species  are  known 
from  Africa  through  the  work  of  Muir  (1927, 
1934)  and  other  early  workers  and,  more  re- 
cently, by  Fennah  ('l955,  1957,  1958),  Lin- 
navuori (1973),  and  Synave  (1957a,  1957b, 
1971).  It  is  now  apparent  that  only  a  small 
proportion  of  the  species  has  been  described 
so  far,  as  has  been  found  for  other  African 
fulgoroid  groups  (e.g.,  Derbidae,  Wilson 
1987). 

Meenoplids  are  small  fulgoroids  with  ten- 
tiform  forewings  and  one  or  both  claval  veins 
with  sensory  pits  ("granulations"  in  older  liter- 
ature) on  either  side  of  the  vein.  Females 
possess  wa.x-producing  plates  on  abdominal 
segments  6-8.  The  family  belongs  to  a  group 
that  includes  Achilixiidae,  Kinnaridae,  and 
Achilidae.  Meenoplidae  are  divided  into  two 
subfamilies,  the  Meenoplinae  and  the  Kerme- 
siinae. In  the  Meenoplinae  the  first  claval  vein 
has  a  single  row  of  sensory  pits,  and  the  sec- 
ond claval  vein  is  more  or  less  covered  with 
them.  In  the  Kermesiinae  the  claval  veins  are 
fused  near  the  apex  of  the  clavus  (e.g. ,  Fig.  4). 
The  first  claval  vein  has  a  row  of  sensory  pits 
on  either  side  of  the  vein.  Three  African  gen- 
era are  placed  in  this  subfamily:  Kermcsia, 
Nisia,  and  Afronisia. 


Kermesiinae  was  first  used  by  Kirkaldy 
(1906:427)  as  a  subfamily  of  Derbidae  to  in- 
clude Nisia,  Phaconcnra,  Suva,  and  several 
derbid  genera.  Although  the  subfamily  was 
named  Kermesiinae,  the  genus  Kermesia  was 
not  treated  at  that  time  but  was  included  by 
implication  in  comments  on  Nisia  by  Kirkaldy 
(1906:427),  who  stated  that  "[Nisia]  seems 
hardly  differentiated  from  Kermesia."  How- 
ever, in  thefollowingyear  Kirkaldy  (1907:163) 
stated  that  two  subfamilies  of  the  Derbidae 
may  be  recognized,  including  the  Nisiinae 
represented  by  Nisia.  The  name  Nisiinae  was 
accepted  by  most  subsequent  workers,  in- 
cluding Metcalf  (1945).  Recentlv,  Emeljanov 
(1984)  and  Tsaur  et  al.  (1987)  used  Kermesi- 
inae as  the  subfamily  name  in  preference  to 
Nisiinae;  this  name  has  been  followed  here. 

The  African  meenoplid  species  have  been 
placed  in  the  genera  Nisia,  Kermesia,  Epon- 
isia (Kermesiinae),  Anifinis,  and  Meenoplus 
(Meenoplinae).  Because  these  genera  are  all 
poorly  defined,  it  is  necessary  for  several  new 
genera  to  be  erected  to  accommodate  some 
existing  species  and  some  undescribed  ones. 
The  five  African  species  ciurently  placed  in 
Eponisia  have  little  in  common  with  the  re- 
maining Asiatic  species,  which  include  the 
type-species,  E.  giittula  Matsumiua,  de- 
scribed from  Taiwan  (Tsaur  et  al.  1986).  This 
has  also  been  noted  by  Emeljanov  (1984).  The 
present  paper  transfers  four  of  them  to  Afro- 
nisia and  the  fifth  to  synonymy  in  Kermesia. 

Abbreviations  of  Depositories 

BMNH    British  Museum  (Natural  History),  London,  UK 
IRSNB    Institut  Royal  des  Sciences  Naturelles  de  Bel- 
gi(liie,  Brussels,  Belgium 


'cab  International  Institute  of  Entoniolosy,  i/o  DepailMu-iit  ol  KnlonioloyN ,  Hntisli  Museum  (Natural  Hist()r\),  Cromwell  hu.id,  London.  SVV7.5BD.  UK. 


324 


1988 


WiLsoN:  Genus  Afhonisia 


325 


MNHN  Museum  National  d'Histoin-  NaturelU-,   Paris, 

France 
MRAC    Musee  Royal  de  I'Africiue  Centrale,  Tervuren, 

Belgium 
NCI         National    Insect    Collection,     Pretoria,     South 

Africa 
RL  R.  Linnavuori  Collection,  Raisio,  Finland 

Key  to  African  Genera  of  Kermesiinae 

1.  Clypeus  rounded,  without  lateral  carinae. 
Forewing  with  only  one  branch  to  M  apically  .  . 
Nisid 

—  Clypeus  with  lateral  carinae.  Forewing  with  two 
branches  of  M  at  apex  (Fig.  4)    2 

2(1).  Eyes  large,  twice  as  long  as  wide.  Forewing  with 
clavus  more  than  half  total  length.  Hind  tarsal 
segments  with  segment  1  approximately  ecjual- 
ing  in  length  segments  2  +  3   ...  Afronmii  gen.  n. 

—  Eyes  smaller,  only  slightly  longer  than  wide. 
Forewing  with  clavus  one-third  total  length. 

Hind  tarsal  segment  1  longer  than  2  +  3 

Kcrmesia 

Afronisia,  gen.  n. 

Type-species. — Eponisia  albovittata  Fen- 
nah. 

Description. — Small,  slender  meenoplids, 
length  overall  3.5-5.9  mm.  Head  with  vertex 
longer  than  wide,  merging  smoothly  into 
frons  with  profile  strongly  rounded,  postero- 
lateral areolets  widely  spaced.  Frons  over 
twice  as  long  as  broad;  lateral  carinae  parallel 
for  most  of  their  length,  diverging  slightly  on 
clypeus,  interrupted  at  frontoclypeal  suture. 
Clypeus  with  lateral  carinae  extending  from 
lateral  carinae  of  frons;  median  carinae  weakly 
developed  but  present  toward  rostrum.  Ros- 
trum long,  extending  beyond  hind  tro- 
chanters. Pronotum  with  well-defined  ante- 
rior carina  parallel  to  the  anterior  margin, 
carina  also  present  on  shoulder  of  pronotum. 
Mesonotum  with  median  carina  present  but 
not  sharply  defined,  lateral  carinae  obscure. 

Forewing:  Relatively  long  and  narrow,  2.5- 
3.0  times  longer  than  maximum  width,  clavus 
longer  than  half  total  length.  Sc  +  R  united  for 
almost  half  wing  length,  R  bifurcate  at  apex, 
M  bifurcate  at  apex.  Claval  veins  appear  to 
join  inner  margin  before  apex  of  clavus  but 
actually  enter  the  apex. 

Hindwing:  r-m  crossvein  at  approximately 
half  length  of  wing,  R  and  M  bifurcate  at  apex. 

Hindlegs;  Hind  tibiae  with  lateral  spines. 
Posttibia  with  8  spines  grouped  as  3  +  5.  Apex 
of  segment  1  with  approximately  8  spines, 
median  ones  less  sclerotized  than  other  ones. 


Apex  of  segment  2  with  similar  arrangement 
of  spines.  Tarsal  segment  1  approximately 
ecjual  in  length  to  segments  2  +  3. 

Male  (;ENiTAi.iA. — Pygofer  dorsoventrally 
elongated,  in  lateral  view  with  dorsolateral 
angle  slightly  or  strongly  produced,  either 
rounded  or  pointed.  Anal  segment  with  lat- 
eral margins  diverging  apically,  posterolateral 
angles  produced  or  rounded.  Aedeagus  (phal- 
lus) tubular,  enlarged  at  apex,  more  or  less 
trumpet-shaped.  Periandrium  (phallobase) 
with  several  pairs  of  appendages  arising  from 
base,  inner  pairs  more  slender  than  outer 
pairs.  Parameres  long  and  narrow,  apical  por- 
tion upcurved. 

Female  genitalia. — Of  the  "reduced" 
meenoplid  form. 

Distribution. — Africa  south  of  Sahara. 

Diagnosis. — The  genus  may  be  distin- 
guished from  Nisia  and  Kermesia  by  the  fol- 
lowing characters.  In  ISlisia  the  lateral  carinae 
of  the  clypeus  are  absent;  they  are  present  in 
Afronisia  and  Kermesia.  In  Kermesia  the  cari- 
nae extend  smoothly  from  the  frons;  in  Afro- 
nisia they  are  disjunct.  The  eyes  oi Afronisia 
species  are  large,  twice  as  long  as  wide,  while 
in  Kermesia  and  ISlisia  they  are  more  globular. 
The  forewing  venation  in  Afronisia  differs, 
with  the  clavus  more  than  half  the  length  of 
the  forewing  and  M  bifurcate  at  its  apex.  In 
Kermesia  M  is  also  bifurcate,  but  the  clavus  is 
short,  only  one-third  the  length  of  the  fore- 
wing. In  Nisia  M  is  single  at  its  apex,  with  the 
clavus  more  or  less  half  the  length  of  the 
forewing.  The  hind  tarsal  segment  1  is  longer 
than  2  +  3  in  both  Nisia  and  Kermesia  and  is 
more  or  less  equal  to  the  length  of  2  +  3  in 
Afronisia.  All  of  these  features  are  at  present 
only  diagnostic.  However,  the  appendages  of 
the  aedeagus  are  characteristic  and  are  not 
present  in  other  meenoplid  genera  but  are 
here  regarded  as  apomorphic  for  the  genus. 
Oriental  species  oi Eponisia  resemble  Kerme- 
sia in  some  respects  and  differ  markedly  from 
Afronisia  in  the  form  of  the  male  genitalia 
(Tsaur  et  al.  1987). 

The  biology  and  host  plants  are  little 
known.  Linnavuori  (1973)  records  A.  albovit- 
tata (Fennah)  and  A.  brunnescens  (Synave)  as 
being  collected  "in  a  swamp." 

Key  to  Species  (Males)  oi Afronisia 

1.  Forewing  almost  entirely  dark  brown,  contrast- 
ing strongly  with  yellow,  dorsal  stripe  from  ver- 
tex to  pronotum  and  mesonotum.    Aedeagus 


326 


Great  Basin  Naturalist  Memoirs 


No.  12 


with  3  pairs  of  processes  (Fig.  7)    

albovittata  (Fennah) 

—  Forewing  variable  in  color  but  never  entirely 
dark  brown.  Aedeagus  with  between  2  and  4 
pairs  of  processes 2 

2(1).  Aedeagus  with  2  pairs  of  large,  sclerotized  pro- 
cesses       3 

—  Aedeagus  with  3  or  4  pairs  of  processes  (Figs.  22, 
35,  45) 4 

3(2).  Forewings  overall  pale  whitish  gray.  Aedeagus 
with  outer  processes  very  large,  curved,  extend- 
ing beyond  phallus  (Fig.  56),  short,  central  pro- 
cess present bredoi,  n.  sp. 

—  Forewing  with  white  veins,  membranes  pale 
brown  (Fig.  15).  Aedeagus  with  outer  processes 
shorter  than  phallus,  short,  central  process  ab- 
sent   muiri  (Metcalf) 

4(2).  Aedeagus  with  4  pairs  of  processes  (Fig.  22)  .  .  . 

flavescens  (Synave) 

—  Aedeagus  with  3  pairs  of  processes    5 

5(3).  Aedeagus  with  outer  pair  of  processes  longer 
than  inner  pairs,  curved  inward  and  dorsad 
(Figs.  33-35).  Pygofer  with  laterodorsal  angles 
pointed  (Fig.  31)   hninnescens  (Synave) 

—  Aedeagus  with  outer  pair  of  processes  shorter 
than  inner  pairs,  pygofer  with  laterodorsal  an- 
gles rounded   6 

6(5).  Outer  pair  of  aedeagal  processes  short  and  bifur- 
cate (Fig.  50)    gemhuensis,  n.  sp. 

—  Outer  pair  longer,  about  two-thirds  length  of 
inner  pairs  (Fig.  45) pallida  (Linnavuori) 

Afronisia  albovittata  (Fennah),  comb.  n. 

Figs.  1-12 

Eponisia  albovittata  Fennah  1955.   Ann.    Mus.   Congo 
Tervuren  in  ser.  8.  Zool.  40;  434. 

Length. — Male  4.2-4.4  mm,  female 
4.5-4.8  mm. 

Description. — Coloration:  Overall  dark 
brown  species.  Vertex  and  face  with  lateral 
edges  of  carinae  dark  brown,  face  between 
carinae  brown  or  light  yellow  in  some  speci- 
mens, becoming  yellow  at  junction  of  face  and 
vertex.  Antennae  dark  brown,  eyes  dark.  Ver- 
tex pale  yellow.  Pronotum  and  mesonotum 
with  central,  yellow  stripe  (Fig.  1)  with  orange 
pigmentation  following  median  carina.  Fore- 
wing dark  brown,  inner  margin  of  clavus  and 
subapical  line  of  transverse  veins  between  R 
and  Cu  and  between  apex  of  clavus  and  Cuj 
white  (Fig.  4).  Legs  brown.  Abdomen  and 
genital  capsule  dark  brown. 

Male  genitalia  (Figs.  7-12). — Pygofer 
longer  than  broad,  laterodorsal  angles  acute 
(Fig.  10).  Anal  segment  with  apical  (posterior) 
margin  concave  between  rounded  posterolat- 


eral margins  (Fig.  11),  slightly  deflexed  in 
lateral  view  (Fig.  10).  Aedeagus  with  perian- 
drium  bearing  three  pairs  of  sclerotized  pro- 
cesses from  ventral  portion  (Fig.  7),  the  outer 
pair  broader  and  slightly  shorter  than  the  in- 
ner pair  and  central  pair  very  short.  Long, 
trumpet-shaped  central  portion  (phallus)  ex- 
tending beyond  processes.  Parameres  long, 
with  upcurved,  subspinose,  apical  process 
(Fig.  9),  slightly  widened  before  apical  por- 
tion in  dorsal  view  (Fig.  12). 

Distribution, — East  Africa:  Rwanda  (type 
locality),  Sudan  (Equatoria  Prov.)  (Linnavuori 
1973),  Uganda,  Kenya,  Zaire. 

Material  examined. — Males  and  females 
from  Kenya,  Uganda,  Sudan  (Equatoria 
Prov.)(BMMH). 

Remarks. — Fennah's  original  description 
(1955)  states  that  the  phallobase  (perian- 
drium)  possesses  two  pairs  of  sinuate  spines, 
not  three  as  actually  present.  However,  Fen- 
nah's drawing  was  made  from  a  lateral  view, 
and  the  short,  central  (ventral)  pair  may  not 
have  been  visible.  The  male  genitalia  were 
figured  correctly  by  Synave  (1957b)  from  ma- 
terial from  the  type  locality,  and  for  this  rea- 
son the  type-specimen  was  not  reexamined. 

Diagnosis. — The  dark  coloration  of  the 
species  and  the  pale  yellow,  dorsal  stripe  dis- 
tinguish it  from  all  others  in  the  genus. 

Afronisia  muiri  (Metcalf),  comb.  n. 

Figs.  13-21 

Nisia  albovenosa  Muir  1927.  Ann.  Mag.  Nat.  Hist.  (9)19: 
200.  [Preoccupied  by  Nisia  albovenosa  Distant, 
1906]  Nisia  muiri  Metcalf  1945:  228  replacement 
name  for  Nisia  albovenosa  Muir. 

Length. — Male  3.5-3.8  mm,  female  4.5- 
4.6  mm. 

Description. — Coloration:  Lateral  carinae 
of  head  and  vertex  dark  brown.  Face  and 
clypeus  brown,  lateral  carinae  of  clypeus  pale. 
Vertex  pale  yellow-white.  Pronotum  and  mes- 
onotum with  lateral  margins  brown  with  cen- 
tral, pale  stripe  (Fig.  13),  median  carina  nar- 
rowly orange.  Forewing  with  veins  white, 
membranes  light  brown,  darker  adjacent  to 
veins  (Fig.  15),  granulations  dark  brown  on 
white  background.  Abdomen  dark  brown,  in- 
tersegmental membranes  white.  Legs  light 
brown. 

Male  genitalia  (Figs.  16-21). — Pygofer 
longer  than  broad,  laterodorsal  angles  almost 
rectangular  (Fig.  18).  Anal  segment  posteri- 
orly deflexed  in  lateral  view  (Fig.  18),  slightly 


1988 


WiLSON:  Genus  Afronisia 


327 


Figs.  1-12.  Afronisia  allwvittata  (Fennah):  1,  head  and  thorax,  dorsal  view;  2,  head,  face;  3,  head,  lateral  view;  4, 
forewing;  5,  hindwing;  6,  hindleg;  7,  aedeagus,  ventral  view;  S,  aedeagus,  dorsal  view;  9,  aedeagus  and  parameres, 
lateral  view;  10,  pygofer  and  anal  segment,  lateral  view;  11,  anal  segment,  dorsal  view;  12,  parameres,  ventral  view. 


328 


Great  Basin  Naturalist  Memoirs 


1988 


Figs.  13-28.  Figs.  13-21,  Afronisia  muiri  (Metcalf);  13,  head  and  thorax,  dorsal  view;  14,  head,  lateral  view;  15, 
forewing;  16,  aedeagus,  ventral  view;  17,  aedeagus,  lateral  view;  18,  pygofer  and  anal  segment,  lateral  view;  19,  anal 
segment,  dorsal  view;  20,  paramere,  ventral  view;  21,  paramere,  lateral  view.  Figs.  22-28,  Afronisia  flavescens 
(Synave):  22,  aedeagus,  ventral  view;  23,  aedeagus,  lateral  view;  24,  pygofer,  lateral  view;  25,  anal  segment,  lateral 
view;  26,  anal  segment,  dorsal  view;  27,  paramere,  ventral  view;  28,  paramere,  lateral  view. 


1988 


WiLsoN:  Genus  Afronisia 


329 


produced  posterior  processes  (Fig.  19).  Ae- 
deagus  with  periandrium  bearing  two  pairs  of 
sclerotized  processes  (Fig.  16),  outer  pair 
large,  curved,  inner  pair  longer,  thin  (Fig. 
17).  Parameres  with  upturned  apical  portion 
(Fig.  21),  subapical  lobe  in  dorsal  view  (Fig. 
20). 

Distribution. — South  Africa. 

Material E.XAMiNED. — Holotype  9,  South 
Africa,  Port  St.  John,  Pondoland,  R.  E. 
Turner,  iv.l923  (BMNH);  1  9,  same  data  as 
holotype;  1  c?  6  9 ,  Rustenburg,  Transvaal, 
10-13. xii.  1961  (Capener);  1  9,  Trazaneen, 
Transvaal,  10. xii.  1918;  1  9,  Umtentweni,  Na- 
tal, 9-14.iii.1961  (Capener)  (NCI,  2  9 
BMNH);  1  d,  Ladvsmith,  29. i.  1981  (Theron); 
1  9,  Dundee,  21. i.  1981  (Theron);  1  9,  Dur- 
ban, 7. i.  1971  (Kluge)  (BMNH). 

Remarks. — The  species  was  originally  de- 
scribed from  one  female.  A  second  female 
from  the  same  locality  (Port  St.  John,  Pon- 
doland) is  also  present  in  the  BMNH  collec- 
tion. Only  two  males  were  present  among  the 
specimens  from  South  Africa.  No  further  spe- 
cies have  been  found  in  South  Africa.  Com- 
parison of  wing  pigmentation  supports  the  as- 
sociation of  the  males  with  the  type  female. 
Synave  (1957b)  repeated  Muir  s  original  de- 
scription but  did  not  examine  the  type-speci- 
men. 

Diagnosis. — This  is  the  only  species  known 
at  present  from  South  Africa.  The  forewing 
coloration  is  distinctive,  and  the  male  aedea- 
gus  is  characteristic  with  the  two  pairs  of  pro- 
cesses. 

Afronisia  flavescens  {Sy nave),  comb.  n. 

Figs.  22-28 

Eponisia  flavescens  Svwdve  1971.  Bull.  Inst.  r.  Sci.  nat. 
Belg.  47(39):  10. 

Length. — Male  4.2  mm,  female  4.8  mm. 

Description. — Coloration:  Male  head  and 
body  orange,  pale  yellow  stripe  on  pronotum 
and  mesonotum  with  orange  median  pigmen- 
tation following  median  carina.  Lateral  cari- 
nae  of  head  edged  dark  brown.  Legs  orange. 

Forewing  with  veins  white,  apices  pale 
brown.  Membranes  suffused,  very  pale 
brown.  Granulations  with  black  points  on 
white  background. 

Female  much  paler  than  male.  Overall  stra- 
mineous. 

Male  genitalia  (Figs.  22-28). — Pygofer 
with  laterodorsal  angles  acute  (Fig.  24).  Anal 


segment  deflexed  posteriorly  in  lateral  view 
(Fig.  25),  posterior  margin  scarcely  produced 
(Fig.  26).  Aedeagus  with  periandrium  with 
four  pairs  of  sclerotized  processes  (P'ig.  22), 
outer  pair  thickest,  two  inner  pairs  thinner 
but  more  or  less  equal  in  length,  central  pair 
about  half  the  length.  Central  portion  (phal- 
lus) longer  than  processes  (Fig.  22).  Par- 
ameres as  in  other  species  with  upturned  api- 
cal portion  (Figs.  27,  28). 

Distribution. — Nigeria. 

Material  examined. — Holotype  d,  Nige- 
ria, Badeggi,  N.W.  State  (IRSNB).  Paratype 
9  (not  male  as  stated  in  original  description), 
Nigeria,  Yankar,  N.E.  State  (IRSNB). 

Diagnosis. — A  pale  species,  similar  in  col- 
oration to  A.  hredoi  n.  sp.  The  only  species  to 
have  four  distinct  pairs  of  aedeagal  processes. 

Remarks. — Known  only  from  the  type  ma- 
terial. 

Afronisia  brnnnescens  (Synave),  comb.  n. 

Figs.  29-37 

Eponisia  hninnescens  Synave  1957a.  Expl.  Pare  National 
de  rUpemba.  Fasc.  43(2):  7. 

Length. — Overall  male  4.2-4.4  mm,  fe- 
male 4.6-4.9  mm. 

Description. — Coloration:  Male  body  and 
legs  pale  orange.  Vertex  and  mesonotum  with 
distinct,  pale  yellow  stripe,  darker  orange  lat- 
erally. Edges  of  lateral  carinae  of  head  dark 
brown  extending  to  lateral  ocellus,  inner  mar- 
gin (sensory  pits)  whitish.  Antennae  pale  yel- 
low brown. 

Forewing:  Membranes  pale,  stramineous, 
light  brown,  darker  adjacent  to  veins.  Veins 
white,  with  apices  darker.  Claval  vein  with 
sensory  pits  dark  brown.  Forewing  narrow, 
parallel-sided,  rounded  at  apex. 

Hindwing:  Whitish,  veins  pale  brown. 

Female  paler  generally  than  male. 

Male  genitalia  (Figs.  31-37). — Pygofer 
with  laterodorsal  angles  extended,  acute  in 
profile  (Fig.  31).  Anal  segment  with  postero- 
lateral apical  lobes  reflexed  (Figs.  31,  32). 
Aedeagus  with  periandrium  bearing  three 
pairs  of  sclerotized  processes,  the  outer  pair 
longer  and  thicker  than  two  inner  pairs, 
curved  inward  and  dorsad  apically  (Figs. 
33-35),  inner  pair  diverging  apically  (Figs. 
33,  35),  central  pair  short,  about  half  the 
length  of  outer  pairs.  Phallus  longer  than  pro- 
cesses, trumpet-shaped.  Parameres  slender, 
apical  quarter  directed  dorsad  (Fig.  36).  Some 


330 


Great  Basin  Naturalist  Memoirs 


No.  12 


variation  appears  in  length  of  processes  (Fig. 
33  illustrates  the  paratype.  Fig.  35  is  a  speci- 
men from  Cameroon). 

Distribution. — Central  and  west  Africa: 
Cameroon,  Sudan  (Equatoria  Prov.)  (Lin- 
navuori  1973),  Zaire  (Katanga  Prov.,  type  lo- 
cality, Orientale  Prov.),  Central  African  Re- 
public (Bossangoa). 

Material  EXAMINED. — Holotype  6 ,  Zaire, 
Kaziba  (MRAC).  Paratype  d,  Zaire,  Kabwe 
(MRAC).  Zaire,  numerous  specimens  from 
Pare  National  de  la  Garamba  (northeast  Zaire) 
(IRSNB,  MRAC);  1  6,  Cameroon  (Banyo) 
(BMNH);  1  6,  Central  African  Republic 
(Bossangoa)  (RL). 

Diagnosis — The  male  genitalia  are  very 
characteristic  with  the  curved  outer  pair  of 
processes.  The  species  was  found  mixed  in 
series  with  A.  pallida  and  may  be  separated  by 
the  narrow,  parallel-sided  forewings  as  well  as 
by  the  male  genitalia. 

Afronisia  pallida  (Linnavuori),  comb.  n. 

Figs.  38-45 

Eponisia  pallida  Linnavuori  1973.  Notiil.  ent.  .53:  111. 

Length. — Overall  male  4.5-5.0  mm,  fe- 
male 5.2-5.8  mm. 

Description. — Coloration:  Male/female, 
lateral  carinae  of  head  with  edges  dark 
brown,  body  and  head  pale  yellow,  mesono- 
tum  orange  laterally,  pale,  median  stripe  in- 
terrupted by  orange,  median  carina.  Legs 
pale  yellow.  Abdomen  and  genital  capsule 
stramineous.  Forewing  with  pale  yellow 
membrane,  white  veins,  apices  of  veins 
marked  with  brown  (Fig.  38),  apical  cells 
darkened  adjacent  to  veins,  black  tips  to  white 
sensory  pits  on  claval  veins.  Forewing  becom- 
ing broader  toward  apex. 

Male  genitalia  (Figs.  40-45). — Pygofer 
with  laterodorsal  angles  broadly  rounded  in 
profile  (Fig.  41),  anal  segment  with  postero- 
lateral lobes  (Fig.  40)  reflexed  (Fig.  41). 
Aedeagus  (Figs.  45,  46)  with  periandrium 
bearing  three  pairs  of  sclerotized  processes 
and  one  central  (ventral)  process,  outer  pair 
shorter  than  inner  pairs  (Fig.  45),  the  more 
dorsal  of  the  inner  pairs  converging  apically 
(Fig.  45),  the  inner  pair  parallel  for  half  their 
length  before  diverging  (Fig.  45),  single,  cen- 
tral spine  short.  Two  small  spines  on  dorsal 
surface  of  phallus.  Parameres  with  distinct 
ventral  lobe  one-third  distance  from  apex  with 


apical  third  curved  upward  and  twisted  (Figs. 
42,  43). 

Distribution. — Central  Africa:  Sudan 
(Equatoria  Prov.,  type  locality)  (Linnavuori 
1973),  Central  African  Republic,  Zaire. 

Material  e.xamined. — Holotype  d,  Su- 
dan, Equatoria,  Yambio  (RL).  Paratype  9 , 
Sudan,  Equatoria,  Yambio  (BMNH).  Cen- 
tral African  Republic  (Bossangoa)  (IRSNB). 
Zaire,  numerous  specimens  from  Pare  Na- 
tional de  la  Caramba  (northeast  Zaire) 
(IRSNB,  MRAC). 

Remarks. — Found  in  same  localities  as  A. 
bninnescens  since  it  is  mixed  in  collections 
from  Zaire. 

Diagnosis. — A  pale  species,  externally 
similar  to  A.  bninnescens  but  slightly  larger 
and  with  forewings  broadened  rather  than 
parallel-sided.  Male  genitalia  very  distinct 
with  three  pairs  of  long  processes. 

Afronisia  gembuensis,  n.  sp. 

Figs.  46-51 

Length. — Male  4.7  mm,  female  5.2  mm. 

Description. — Body  color  pale  orange,  re- 
sembling A.  mtiiri,  especially  in  darker  pig- 
mentation of  forewings. 

Male  genitalia  (Figs.  46-51). — Pygofer 
with  laterodorsal  angles  rounded  (Fig.  46). 
Anal  segment  with  lateral  margins  diverging 
apically,  posterolateral  angles  acute  (Fig.  47), 
with  apical  margin  concave.  Aedeagus  with 
periandrium  with  three  pairs  of  sclerotized 
processes  and  one  short,  central  process, 
outer  pair  short  and  bifurcate  (Fig.  50),  inner 
pairs  long,  with  outer  of  these  diverging,  in- 
ner one  almost  parallel  (Fig.  50),  central  pro- 
cess short.  Parameres  with  pronounced  ven- 
tral lobe  two-thirds  as  long  as  its  length,  apical 
third  thinner  and  directed  dorsad,  slightly 
twisted  (Figs.  48,  49). 

Distribution. — Nigeria. 

Type  material. — Holotype  6,  Nigeria, 
N.E.  State,  Gembu-Yelwa,' 22.viii.1973,  Lin- 
navuori (RL).  Paratvpes,  3  (5  6  9,  Nigeria, 
N.E.  State,  Gembu,  21-22. viii.  1973,  Lin- 
navuori (RL,  16  19  deposited  in  each 
BMNH,  MRAC). 

Diagnosis. — Somewhat  similar  to  A.  pall- 
ida in  aspects  of  the  male  genitalia  and  partic- 
ularly in  the  shape  of  the  laterodorsal  angles  of 
the  pygofer  and  the  shape  of  parameres.  The 
aedeagal  (periandrium)  processes  are,  how- 
ever, diagnostic,  particularly  the  bifurcated 
outer  pair. 


1988 


WILSON:  Genus  Afronisia 


331 


Figs.  29-45.  Figs.  29-37,  Afronisia  bnmnescens  (Synave);  29,  head,  lateral  view;  30,  forewing;  31,  male  pygofer  and 
anal  segment,  lateral  view;  32,  anal  segment,  dorsal  view;  33,  aedeagus,  ventral  view,  paratype;  34,  aedeagus,  lateral 
view;  35,  aedeagus,  ventral  view;  36,  paramere,  lateral  view;  37,  paramere,  ventral  view.  Figs.  38-45,  Afronisia  pallida 
(Linnavuori):  38,  forewing;  39,  head,  lateral  view;  40,  male  anal  segment,  dorsal  view;  41,  pygofer  and  anal  segment 
lateral  view;  42,  paramere,  ventral  view;  43,  paramere,  lateral  view;  44,  aedeagus,  lateral  view;  45,  aedeagus,  ventral 


332 


Great  Basin  Naturalist  Memoirs 


No.  12 


Figs.  46-51.  Afronisia  gemhuensis ,  n.  .sp. :  46,  male  pygofer  and  anal  segment,  lateral  view;  47,  anal  segment,  dorsal 
view;  48,  paramere,  ventral  view;  49,  paramere,  lateral  view;  50,  aedeagus,  ventral  view;  51,  aedeagus,  lateral  view. 


Afronisia  bredoi,  n.  sp. 

Figs.  52-57 

Length. — Male  5.2  mm,  female  5.7-5.9 
mm. 

Description. — Coloration  male/female: 
body  and  legs  light  orange  dusted  with  wax. 
Head  with  lateral  carinae  of  vertex  and  face 
edged  with  dark  brown,  face  suffused  with 
light  brown.  Mesonotum/pronotum  light  or- 
ange laterally,  central,  pale  yellow  stripe  with 
median  carina  light  orange.  Antennal  scape 
very  pale  brown,  pedicel  darker. 

Forewing:  Overall  pale  grey-white,  mem- 
branes darker,  particularly  adjacent  to  veins. 
Apices  of  veins  at  margin  darker,  especially  in 
female.  Forewing  distinctly  wider  toward 
apex. 

Male  genitalia  (Figs.  52-57). — Pygofer 
with  laterodorsal  angles  rounded  (Fig.  52). 
Anal  segment  with  lateral  margins  diverg- 
ing apically,  posterolateral  angles  produced 
(Fig.  53).  Aedeagus  with  periandrium  bearing 
two  pairs  of  sclerotized  processes,  and  a  sin- 
gle, central  one,  outer  processes  very  large, 


curved  inward  beyond  the  phallus,  with  short 
processes  arising  laterally  at  base.  These  are 
asymmetric  in  specimen  examined  (Fig.  56). 
Inner  processes  shorter  than  phallus,  more  or 
less  parallel,  central  spine  short.  Parameres 
with  apical  third  curved  smoothly  upward  in 
lateral  view  (Fig.  57),  twisted  apicallv  (Fig. 
54). 

Distribution. — Zaire  (Elisabethville). 

Type  material. — Holotype  6,  Zaire,  la- 
beled Congo  Beige,  Elisabethville,  iv.l939, 
H.  J.  Bredo  (MRAC).  Paratypes:  2  9,  same 
data  as  holotype  (MRAC). 

Dia(;n()SIS — A  pale-colored  species  resem- 
bling A.  pallida.  The  long,  curved,  outer  ap- 
pendages of  the  male  aedeagus  are  diagnostic 
and  resemble  no  other  species. 

Remarks  on  Eponisia  alhiiicriosa  Muir  1934 

This  species  is  considered  the  junior  syn- 
onym of  Kermesia  alhinervosa  Muir  1927 
(type  locality.  Sierra  Leone).  The  male  holo- 
type and  female  allotype  were  very  badly 
damaged  in  transit  from  Honolulu  to  London 


1988 


WiLSON:  Genus  Afronisia 


333 


Figs.  52-57.  Afronisia  brecJoi,  n.  sp. ;  52,  male  pygofer  and  segment,  lateral  view;  53,  anal  segment,  dorsal  view;  54, 
paramere,  ventral  view;  55,  paramere,  lateral  view;  56,  aedeagus,  lateral  view;  57,  aedeagus,  ventral  view. 


in  1927  (together  with  other  African  meen- 
oplid  types  described  in  the  same  paper).  For- 
tunately, the  male  genitalia  of  the  holotype 
were  intact  and  other  specimens  have  been 
associated.  It  is  with  these  specimens  that  the 
female  holotype  o(  Eponisia  albinervosa  from 
Uganda  has  been  compared. 
The  synonymy  is  as  follows: 

Kermesia  albinervosa  Muir  1927.  Ann.  Mag.  Nat.  Hist. 

(9)19:  203. 
Eponisia  albinervosa  Muir  1934.  Ann.  Mag.  Nat.  Hist. 

(10)14:  564.  syn.  n. 


Acknowledgments 

I  am  pleased  to  thank  the  following  for  the 
loan  of  material  used  in  this  study:  Dr.  Jan  Van 
Stalle  (IRSNB,  Brussels,  Belgium),  Dr.  H. 
Andre  (MRAC,  Tervuren,  Belgium),  Dr.  R. 
Linnavuori  (Raisio,  Finland),  the  late  Dr.  J. 
Theron,  deceased  January  1988  (Stellenbosch 
University,  South  Africa),  and  Dr.  1.  M.  Mil- 
lar (NCI,  Pretoria,  South  Africa).  I  am  very 
grateful  to  Mr.   W.   R.   Dolling,   Dr.   W.  J. 


Knight,  and  Mr.  M.  D.  Webb  for  making 
useful  comments  and  criticism  of  the  manu- 
script. 

Literature  Cited 

Distant,  W  L  1906.  Fauna  of  British  India  including 
Ceylon  and  Burma.  Rhynchota.  Heteroptera- 
Homoptera  3.  Taylor  and  Francis,  London,  UK. 
503  pp. 

Emeljanov.  a.  F  1984.  [A  contribution  of  knowledge  of 
the  families  Kinnaridae  and  Meenoplidae  (Ho- 
moptera,  Fulgoroidea).]  Ent.  Obozreniye  63(3): 
468-483.  [Translated  in  Ent.  Rev.  64:  49-65, 
1985.] 

Fennah,  R  G  1955.  Contributions  a  I'etude  de  la  faune 
entomologique  du  Ruanda-Urundi  (Mission  P. 
Basilewsky  1953).  LXXVIII.  Homoptera,  Ful- 
goroidea. Ann.  Mus.  Congo  Tervuren  in  ser.  8 
Zool.  40:  427-446. 

1957.  Results  from  the  Danish  expedition  to  the 

French  Cameroons  1949-50.  XXIV  Fulgoroidea. 
Bull,  de  IT. F.A.N,  ser.  A.  19:  1274-1311. 

1958.  Fulgoroidea  (Homoptera)  from  West  Africa. 

Bull,  de  ri. F.A.N,  ser.  A.  20:  460-538. 

KiRKALDY,  G  W  1906.  Leaf-hoppers  and  their  natural 
enemies,  Pt.  IX.  Leafhoppers-Hemiptera.  Bull. 
Hawaiian  Sugar  Planters  Assoc,  Div.  Ent.  1: 
271-479. 


334 


Great  Basin  Naturalist  M  emoirs 


No.  12 


1907.    Leaf-hopper.s — supplement    (Hemiptera). 

Bull.  Hawaiian  Sugar  Planters  A.SSOC.,  Div.  Ent.  3: 
1-186. 

LiNNAVUORl,  R  1973.  Hemiptera  of  the  Sudan,  with  re- 
marks on  some  species  of  the  adjacent  countries  2. 
Homoptera  Auchenorrhyncha:  Cicadidae,  Cer- 
copidae,  Machaerotidae,  and  Fulgoroidea.  Notul. 
Ent.  53:65-137. 

Metcalf.  Z.  P  1945.  Meenoplidae.  Pages  219-2.38  in 
General  catalogue  of  the  Hemiptera.  Ease.  IV.  Pt. 
6.  Smith  College,  Northampton,  Massachusetts, 
USA. 

Mum,  F  1927.  New  species  of  African  Meenoplidae  (Eul- 
goroidea,  Homoptera).  Ann.  Mag.  Nat.  Hist. 
(9)19;  197-208. 

1934.  New  and  little-known  Fulgoroidea  (Ho- 
moptera). Ann.  Mag.  Nat.  Hist.  (10)14;  .561-586. 


Synave,  H  1957a.  E.xploration  de  Pare  National  de 
I'Upemba.  Mission  G.  F.  de  Witte.  2  Meenopli- 
dae. Ease.  43;  79-81. 

1957b.  Pare  National  Albert  1.  Mission  G.  E.  de 

Witte  1933-35,  Meenoplidae  (Homoptera,  Eul- 
goroidea).  Ease.  90(2);  7-.30. 

1971.    Contribution   a  la  connaissance   des   ful- 

gorides  du  Nigeria  (Homoptera)  (recoltes  J.  T. 
Medler).  Bull.  Inst.  r.  Sci.  nat.  Belg.  47:  1-34. 

TsAUR.  S  C ,  C  T  Yang,  and  M  R  Wilson  1987. 
Meenoplidae  of  Taiwan  (Homoptera:  Eul- 
goroidea).  Monograph  of  Taiwan  Museum  No.  6; 
81-117.  December  1986  (1987), 

Wilson,  M  R  1987.  African  Derbidae  (Homoptera,  Ful- 
goroidea); taxonomic  notes  with  descriptions  of 
new  species  collected  mainly  from  coconut.  J.  Nat. 
Hist.  21:567-595. 


DELPHACIDAE  OF  ALASKA  (HOMOPTERA:  FULGOROIDEA) 

Stephen  W.  Wilson' 

Abstract. — Fifteen  species,  in  10  genera,  of  Delphacidae  are  recorded  from  Alaska.  One,  Kusnezoviella  niacleani,  is 
described  as  new  to  science.  Acanthodelphax  analis  (Crawford),  Chilodelphax  magnifrons  (Crawford),  Javesella 
arcanastyla  (Beanier),  and  J.  atrata  (Osborn)  are  new  combinations.  Delphacudes  saileri  Beamer  is  a  synonym  of 
Javesella  sinullima  (Linnavuori). 

Of  the  15  species,  1  is  recorded  only  from  Alaska,  3  are  Holarctic,  4  are  Nearctic,  and  7  are  Palearctic  and  restricted 
to  tundra  habitats,  with  5  of  these  apparently  found  only  north  of  the  Brooks  Range. 


Little  information  is  available  on  species 
diversity  and  distribution  of  much  of  the 
Alaskan  insect  fauna  (MacLean  and  Hodkin- 
son  1980).  For  example,  only  seven  species  of 
Delphacidae  have  been  described  from 
Alaska.  Delphacodes  stejnegeri  (Ashmead) 
was  described  by  Ashmead  (1904)  and  re- 
described  and  illustrated  by  Dozier  (1926). 
Vilbaste  (1980)  synonymized  this  species  with 
Javesella  pelhicida  (Fabricius).  Delphacodes 
atrata  Osborn  was  recorded  from  Alaska  by 
Beamer  (1951).  Delphacodes  saileri  Beamer 
(1952b)  was  described  from  Alaska  but,  as  in- 
dicated below,  is  a  synonym  oi Javesella  simil- 
lima  (Linnavuori).  DuBose  (1960)  recorded 
the  Holarctic  D.  pelhicida  from  Alaska.  Del- 
phacodes uniformis  (Walker)  was  listed  from 
Alaska  by  Metcalf  (1943);  however,  Muir 
(1919)  stated  that  this  insect  is  not  a  delphacid. 
Megamelus  flavus  Crawford  was  recorded 
from  Alaska  by  Beamer  (1955). 

The  present  paper  provides  a  key  and  infor- 
mation on  distribution  for  the  15  delphacid 
species  found  in  the  Canadian  National  Col- 
lection, Ottawa,  and  material  collected  by 
MacLean  in  the  northern  part  of  the  state. 
The  name  used  by  the  first  describer  and  the 
first  placement  in  the  current  genus  are  given 
for  each  species.  Because  the  species  have 
been  described  in  detail  elsewhere,  only  the 
new  species  o( Kusnezoviella  is  described.  Ac- 
curate identification  of  delphacids  relies  on 
use  of  characters  of  the  male  genitalia;  for  this 
reason,  all  species  treated  here  are  illustrated. 


2(1) 


3(1). 


4(3). 


5(3). 


6(5). 


Key  to  the  Alaskan  Delphacidae 

Frons  with  2  longitudinal,   median  carinae 

(Fig.  1) 2 

Frons  with  1  longitudinal,  median  carina  ....    3 

Large  pits  present  on  frons,  pronota, 
mesonota,  and  abdomen  (Figs.  1,  2);  aedeagus 
strongly    decurved   with    poorly   developed 

teeth  (Figs.  2,  4)    

Achorotile  sitbarctica  Scudder 

Pits  absent;  aedeagus  straight  with  well- 
developed  teeth  (Figs.  6,  7)  

....    C riomorphiis  ivilhelmi  Anufriev  &  Averkin 

Pronotal  lateral  carinae  straight,  extending  to, 
or  almost  to,  posterior  margin  of  pronotum 
(Fig.  12);  metatibial  spur  with  or  without 
black-tipped  teeth 4 

Pronotal  lateral  carinae  curving  laterad,  not 
extending  to  posterior  margin  of  pronotum; 
metatibial  spur  with  black-tipped  teeth,  these 
sometimes  weak    5 

Metatibial  spur  with  well-developed,  black- 
tipped  teeth;  pygofer  with  lateral,  inflated 
lobes  (Figs.  9,  11)    .  Megamelus  flavus  CravAord 

Metatibial  spur  lacking  teeth;  pygofer  not  in- 
flated laterally  (Fig.  13) ' 

....   N othodelphax  ebiirneocarinatiis  (Anuiriev) 

Broad,  pale  stripe  usually  bordering  longitu- 
dinal, median  carinae  on  vertex,  pronotum, 
and/or  niesonotum;  pygofer  with  broad  lobe 
laterally  on  caudal  margin  (Figs.  16,  21) 6 

Broad,  pale  stripe  not  present;  pygofer  with- 
out broad  lobe  7 


Styles  each  with  irregular  teeth  on  broad  apex 
(Fig.  20)   Unkanodes  excisa  (Melichar) 

Styles  each  with  acute  apex  (Fig.  23) 

Chilodelphax  magnifrons  (Crawford) 


'Department  of  Biology,  Central  Missouri  State  University,  Warrensburg,  Missouri  64093 


335 


336 


Great  Basin  Naturalist  Memoirs 


No.  12 


Figs.  1-20.  Figs.  1-5,  Achorotile  subarctica  Scudder;  1,  head,  frontal  view;  2,  male  genitalia,  lateral  view;  3,  anal 
tube,  dorsal  view;  4,  aedeagus,  lateral  view;  5,  styles,  caudal  view.  Figs.  6-8,  Criomorphiis  wilhelmi  Anufriev  & 
Averkin;  6,  male  genitalia,  lateral  view,  7,  aedeagus,  lateral  view;  8,  styles,  caudal  view.  Figs.  9-11,  Megamelus  flavus 
Crawford:  9,  male  genitalia,  lateral  view;  10,  aedeagus,  lateral  view;  11,  male  genitalia,  caudal  view.  Figs.  12-15, 
Nothodelphax  ehurneocarinatus  (Anufriev):  12,  head  and  thorax,  dorsal  view;  13,  male  genitalia,  lateral  view;  14, 
aedeagus,  lateral  view;  15,  styles,  caudal  view.  Figs.  16-20,  t/n/ca norfe.s  excisa  (Melichar):  16,  pyfoger  and  style,  lateral 
view;  17,  anal  tube,  lateral  view;  18,  19,  aedeagus,  lateral  view;  20,  styles,  caudal  view. 


7(5).       Anal  tube  with  spines  crossing  (Fig.  27)   .... 

Ribautodelphax  albostriatus  (Fieber) 

—  Anal  tube  with  spines  parallel  or  diverging  .  .      8 

8(7).  Pygofer  with  caudally  directed,  median  pro- 
jection ventral  to  base  of  styles  (Fig.  29)  .... 
Acanthodelphax  analis  (Crawford) 

—  Pygofer  without  median  projection 9 

9(8).       Apices  of  styles  converging  (Fig.  34) 

Kusnezoviella  rnacleani,  n.  sp. 

—  Apices  of  styles  diverging 10 

10(9).     Aedeagus  strongly  decurved  (Fig.  36) 

Javesella  pellucida  (Fabricius) 

—  Aedeagus  recurved  or  straight  (Figs.  40,  43, 
47,50,53) 11 

11(10).  Aedeagus  evenly  forked  (Fig.  40) 

javesella  obscurella  (Boheman) 

—  Aedeagus  not  forked  (Figs.  43,  47,  50,  53)   .  .    12 

12(11).  Aedeagus  with  projection  in  basal  half  on  ven- 
tral aspect  (Figs.  50,  53)  or  with  broad,  weakly 


toothed  region  in  basal  half  on  ventral  aspect 

(Fig.  47)   13 

—  Aedeagus  not  as  above  (Fig.  43) 

Javesella  atrata  (Osborn) 

13(12).   Aedeagus  with  large,  ventral,  basal  projection 

(Figs.  50,  53)   14 

—  Aedeagus  without  large,  basal  projection 
(P'ig.  47)   Javesella  simillima  (Linnavuori) 

14(13).  Aedeagal,  ventral,  basal  projection  acute, 
aedeagal  dorsal  margin  slightly  cur\'ed  (Fig. 
50) Javesella  discolor  (Boheman) 

—  Aedeagal,  ventral,  basal  projection  rounded, 
aedeagal  dorsal  margin  constricted  (Fig.  53) 
Javesella  areanastijla  (Beamer) 

Achorotile  subarctica  Scudder 

Figs.  1-5 

Achrotile  [sic]  subarctica  Scudder,  1963:169 

Distribution  records  for  specimens  used  in 
this  study  are:  Alaska:  Brooks  Range,  Spruce 


1988 


WiLsoN;  Delphacidae  OF  Alaska 


337 


Tree  Mine,  68°03'N,  149°W,  800  m,  4  July 
1982,  Seppo  Keponen,  ex.  B.  <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. 

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Dlabola.  J  1965.  Ergebnisse  der  zoologisehen 
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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- 
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Emeljanov,  A  F.  1982.  Fulgoroidea  (Homoptera)  col- 
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entomofaunistical  group  of  the  Soviet-Mongolian 
complex  biological  expedition  in  1970-1975.  Ins. 
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Fabricius,  J  C  1794.  Ryngota.  Entomologia  systematica 
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Fennah,  R.  G.  1963.  New  genera  of  Delphaeidae  (Ho- 
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KWON,  Y.  J  1982.  New  and  little  known  planthoppers  of 
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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  bocanus  Oman  19.34. 

Texas. 
Laevicephahis  cartwrighti  Oman  1932. 

CaHfornia. 
Laevicephahis  excavatus  Oman  1931. 

California,  Oregon. 
Laevicephahis  incongruus  Oman  1937. 

California. 
Laevicephahis  joaqiiinus  Oman  1937. 

California. 
Laevicephahis  pacificus  1937.  Cahfornia. 
Laevicephahis  siskiyoii  Oman  1937.  USA. 
Laevicephahis  iihleri  Oman  1931.  USA. 
Laevicephahis  wilsoni  Oman  1932. 

California. 
Lystridea  nuda  Oman  1938.  California. 
Neocoehdia  beameri  Oman  1931.  Texas. 
Neocoehcha  distincta  Oman  1931. 

Arizona,  Texas. 
Neocoehdia  fiiscovittata  Oman  1931. 

Arizona. 


Neocoehdia  penehneata  Oman  1931. 

California,  Nevada. 
Neocoehdia  texana  Oman  1931.  Texas. 
Neonirvana  hyahna  Oman  1936.  Costa 

Rica. 
Onura  ehurneola  Oman  1938.  Argentina. 
Pagaronia  confiisa  Oman  1938.  California. 
P agar onia  f areata  Oman  1938.  California. 
Paracoehdia  fiisconeura  Oman  1930. 

Arizona. 
Phlepsiiis  divergens  Oman  1931.  Texas. 
Phlepsius  obviiis  Oman  1931.  Texas. 
Protalebra  brunnea  Oman  1937.  Puerto 

Rico. 
Sanctanus  tectus  Oman  1934.  Virginia, 

South  Carolina 
Scaphoidula  dentata  Oman  1937.  St. 

Lucia. 
Scaphoidula  incisa  Oman  1937.  Central 

America. 
Scaphoidula  unica  Oman  1937.  Brazil. 
Tiaja  arenaria  Oman  1972.  California. 
Tiaja  cruzensis  Gill  &  Oman  1982. 

California. 
Tiaja  montara  Oman  1941.  California. 
Tiaja  ventura  Oman  1941.  California. 
Thatuna  gilletti  Oman  1938.  Idaho. 
Xerophloea  carinata  Oman  1936. 

Argentina. 
Xerophloea  carinata  gigas  Oman  1936. 

Argentina. 
Xerophloea  cephalica  Oman  1936. 

Argentina. 
Xerophloea  diffonnis  Oman  1936. 

Argentina. 
Xerophloea  elegans  Oman  1936. 

Argentina. 
Xerophloea  elongata  Oman  1936. 

Argentina. 
Xerophloea  magna  Oman  1936. 

Argentina. 


INDEX 

New  names  are  indicated  by  boldface  type,  invalid  names  l)y  italics. 


abbreviata,  244 

aberrans,  45,  52 

abrupta,  68,  76 

Acanthodelphax,  336,  338 

Achorotile,  335,  336 

Acurhinus,  224,  225 

actis,  117 

acuta,  79 

Aetalion,  61 

Aflexia,  224 

Afronisia,  324,  325 

Alapus,  224 

albida,  255 

albifrons,  132 

albocinctus,  105 

albostriatus,  336,  338 

albovittata,  326 

amabilis,  141,  142 

Amerzana,  139,  140 

Amycle,  135,  139,  141 

anademus,  105,  125 

analis,  336,  338 

Angulanus,  133 

Anigrus,  324 

annularis,  149 

apertus.  132 

aphoda, 18,  19,  25 

Aphrodisias,  139,  144 

arcanastyla,  336,  341 

arenicola,  260,  264 

areolata,  273 

arizonensis,  258,  261 

Artacie,  137,  146 

aspersa,  18,  19,  25 

Athysanella,  18,  229 

atlantica,  277,  280 

atrata  (Delphacodes),  335 

atrataQavesella),  336,  341 

auromaculatus,  160,  162 

Austroagalloides,  65 

bakeri  (Idiodonus),  134 

bakeri  (Sophonia)  198,  203 

Balbillini,  182,  183 

Balbillus,  184 

Balclutha,  86,  90 

balli,  82,  84 

Balolina,  86,  90,  100 

bandarita,  234,  239 

barbara  lusitanica,  171 

beameri  (Colladonus),  105,  122 

beameri  (Flexamia),  258,  261,  270 

beamerellus,  122 


belli,  105,  107 

bengalensis,  213 

bicinctus,  105,  113 

bidentata  (Flexamia),  283 

bidentatiis  (Ileopeltus),  45 

bifida  (Athvsanella),  18,  20,  34 

bifida  (Sophonia),  198,  210 

bifurcatus,  16 

bispinosa,  188,  191 

blockeri,  44,  51 

bohemani,  153 

boliviana,  68,  71 

Bonneyana,  104,  132 

brachialis,  157 

bredoi,  326,  332 

brevis,  141,  142 

brunnea  (Rhabdocephala),  135,  165 

brunnea  (Stalubra),  167 

brunnescens,  326,  329 

brunneus  (Calliscarta),  80 

cacica,  144 

Caladonus,  104 

caldwelli,  132 

Calliscarta,  67 

Calotartessus,  62 

candelaria,  135 

canyonensis,  246,  247 

capicola,  10 

carrenoi,  159 

castresii,  157 

caternaulti,  63 

Cathedra,  136,  137,  147 

cearensis,  157 

celata,  258,  265 

Chiasmus,  13,  63 

Chilobia,  137,  147 

Chilodelphax,  335,  337 

Chlorotettix,  43 

Chudania,  182,  188,  194 

Cicada,  171 

Cicadelloidea,  61 

Cicadidae,  171 

cicadina,  156 

cinxia,  148 

Circulifer,  4 

clathrus,  105,  120 

claustrus,  105,  115 

clavatus,  44 

clayi,  277,  285 

coanaco,  148 

Colladonus,  103 

collorum,  234,  241 


349 


350 

Coloborrhis,  63,  64 

Columbiana,  68,  77 

complexa,  198,  206 

complicata,  199,  210 

conspersa,  149,  150 

Copidocephala,  139,  148 

copulus,  134 

Cornutipo,  61,  63 

corticina,  65 

corvenda,  68,  77 

costatus,  133 

Crioniorphus,  335,  337 

Crispina,  182 

crocodilia,  157 

cruzanus.  52 

cuneus,  45,  54 

cursa,  18,  20,  30 

curvata,  251,  252 

Cyclops,  44,  54 

Cvrpoptus,  136 

dakota,  276,  277,  283 

dampfi,  105,  112 

Darthula,  61,  65 

decora  (Calliscarta),  68 

decora  (Flexamia),  258,  261,  267 

decora  (Kana),  188,  194 

delecta,  182,  196 

delongi,  276,  277,  284 

delicata  (Calliscarta),  68,  72 

delicatus  (Phrictus),  160,  162 

Delphacidae,  335 

Delphacodes,  335 

Deltocephalinae,  82 

Deltocephalus,  224 

deserta,  18,  20,  35 

diadema,  135,  162 

Diareusa,  139,  149 

dichopteroides,  147 

Didius,  221 

diligens,  160,  162 

dilutus,  45 

discolor,  336,  341 

diserus,  109 

doeringae,  234,  240 

Doleranus,  43 

dorsalus,  44,  46 

dufourii,  146 

eburneocarinatus,  335,  337 

ecuadoria,  147 

edentulus,  134 

edulis,  182 

elongata,  68,  70 

erninenta,  155 

Enantiocephalus,  224 

Enchophora,  135,  139,  150,  151 

Enhydria,  137,  156 

Eoscartoides,  61 

Eponisia,  324 

Errhomus,  5 

Euacanthella,  63 

Eurymela,  62 

Eurymeloides,  62 

Eurymelops,  62 

evansi,  63 

Evansiola,  65 

excavatus,  134 


Great  Basin  Naturalist  Memoirs 

excisa,  335,  337 
Exitianus,  10 
expanda,  68,  75 
fasciata  (Calliscarta).  67,  80 
fasciata  (Kana),  188,  194 
fasciaticollis,  105,  109 
fenestrata,  62 
festivus,  68 
flavescens,  326,  329 
flavus,  335,  337 
Flexamia,  224,  225 
tlexulosa,  258,  261,  263 
florens,  154 
foeda,  18,  20,  27 
fowleri,  182 
fredonia,  18,  20,  29 
fri^ida,  19,  21 
Fulgora,  135,  137,  157 
Fulgoridae,  135 
Fulgoroidea,  324,  335 
fiinbuliis,  254 
furnaca,  18,  20, 34 
fuscifrons,  91 
gardenia,  19,  21 
gembuensis,  326,  330 
generosa,  62 
gila,  234,  238 
gillettei,  107 
Gladionura,  18,  229 
globosa,  19,  23 
Goniagnathus,  16 
goodi,  118 
graciliceps,  157 
graminea,  276,  277,  285 
grammica,  250 
grandis  (Amycle),  141,  142 
grandis(Balclutha),  91,93 
granulosus,  184 
greeni,  182 
guttata,  148 
haemoptera,  146 
haplus,  44,  49 
hardwicki,  61,  62 
hastulus,  45,  49 
Hebecephalus,  82 
hemijona,  19,  20,  30 
Hododoecus,  224 
hoffmannsi,  160,  163 
Homaloscytina,  63 
hospes,  91,  96 
Idiodonus,  103 
IdiotcUix,  67,  68 
Ileopeltus,  43 
illaborata,  188,  194 
iUuminata,  198,  204 
imitatrix,  149,  150 
imputans,  272 
incerta,  19,  20,  32 
incidus,  105,  126 
incisurus,  133 
incongrua,  19,  20,  38 
indicus,  184,  185 
inflata,  258,  261,  270 
inocarpi,  182 
Inocarpus,  182 
insculptus,  127 


No.  12 


1988 


Index 


351 


insignis,  198,  203 

invita,  68,  71 

Itzalana,  135 

Jaacunga,  104,  129 

jacala,  234,  241 

Jassiis,  224 

Javesella,  335,  336,  339 

kadokana,  19,  20,  32 

Kana,  188 

kanabana,  19,  20,  33 

karachiensis,  13 

kenineri,  149,  150 

keralica,  198,  203 

Kermesia,  324 

Kermesiinae,  324 

kilaueae,  100 

kinonantis,  45 

Kirkaldiella,  86 

Koebelia,  61 

krameri,  19,  23 

kuscheli,  65 

Kuznezoviella,  335,  336 

laeta,  19,  20,  30 

Laevicephalus,  224 

lampetis,  159 

laternaria,  135,  159 

latidens,  128 

lautus,  80 

linealis,  198,  199 

lineata,  98 

Listrophora,  63 

lobata(Balclutha),  91,93 

lobata  (Chiasmus),  14 

longicephala  (Quercinirvana),  182 

longicephala  (Sophonia)  198,  201 

longicornuta,  156 

longirostris  (Enchophora),  154 

longirostris  (Rhabdocephala),  135 

longitudinalis,  198,  201 

lora,  68,  71 

lucifera,  159 

macleani,  336,  339 

Macrostelini,  86 

maculata,  151 

magdalena,  19,  20,  37 

magna,  68,  75 

magnificus,  68 

magnifrons,  335,  337 

major,  10 

mankinsi,  141,  142 

marginata  (Calliscarta),  78 

/Jiargtnafu.s  (Colladonus),  122 

marthae,  19,  20,  29 

Meenoplidae,  324 

Meenoplinae,  324 

Meenoplus,  324 

Megamelus,  335,  337 

melanoptera,  149 

Membracoidea,  61 

merula,  148 

mescalero,  234,  242 

mexicana  (Calliscarta),  80 

mexicana  (Scolopsella),  165 

mexicanus  (CoWadonus),  117 

minima,  234 

minor,  12 


minutacanthis,  162,  163 
mitrii,  159 
moebiusi,  160,  163 
modesta,  198,  207 
modica,  260,  261,269 
Mohunia,  182 
monoiyo,  82 

montana  (Athysanella),  21 
montana  (Taslopa),  63,  65 
Monteithia,  65 
muiri,  326 
nanocanthus,  44,  45 
Narecho,  182 
Neobalinae,  67 
Nesolina,  86,  90,  96 
Nesophrosyne,  86 
Nesophryne,  86 
nigridens,  106 
Nigridonus,  104 
nigrifrons,  132 
nigriventris,  91,  93 
nigromaculata,  151 
nigropicta,  188,  192 
Nirvana,  182,  188,211 
Nirvaninae,  182,  183 
Nirvanini,  182,  183,  187 
Nisia,  324 

Nothodelphax,  335,  337 
obscurella,  336,  340 
Occinirvanini,  182,  183,  219 
ocellatus,  160,  163 
Odontoptera,  139,  159 
Ollarianus,  104,  132 
Omaranus,  221 
Ophiuchus,  188,  217 
ordinata,  189 
ornanda,  149 
ornata,  68,  73 
orni,  171 
Orosius,  86 
orthocephala,  157 
pallida  (Afronisia),  326,  330 
pallida  (Nirvana),  182,  212,  213 
pallidipunctata,  135,  151,  153 
pallidus  (Colladonus),  126 
Paracolladonus,  104,  127 
Paracrassana,  104,  132 
Paranurenus,  104,  127 
Paratanus,  43,  104,  133 
parca,  19,  20,  29 
parvipennis,  155 
pastora,  19,  21 
Pazu,  82 

pectinata,  234,  236 
peculiaris,  213,  216 
pellucida,  335,  336,  339 
peregrina,  91,  95 
peruana,  141 
peshawarensis,  10 
Phaconeura,  324 
phosphorea,  135 
phoxocephala,  91,  96 
Phrictus,  135,  139,  160 
picta,  253 
pinyonae,  135,  142 
plana  (Athysanella),  19,  25 


352 


Great  Basin  Naturalist  Memoirs 


No.  12 


plana  (Homaloscytina),  63 

planata,  19,  23 

Platyeurymela,  63 

plummeri,  134 

plutonis,  91,  95 

Poiocera,  136 

Polyamia,  224 

prairiana,  275,  277 

prasina,  151,  153 

pravus,  109 

princeps,  219 

Pristiopsis,  135 

producta  (Flexamia),  275,  277,  281 

productus  (Jassus),  224 

pseudommatos,  213 

Pseudonirvana,  182,  196 

punctata  (Eurymeloides),  62 

punctatus  (Phrictus),  162,  164 

Pyrops,  135 

pyrops  (Flexamia),  253,  254 

pyrrhocrypta,  151,  153 

Quercinirvana,  182,  198,  213 

quinquepartitus  (Phrictus),  160,  164 

quinquepartitus  (Rhabdocephala),  135 

ramificata,  189,  190 

recurva,  151,  153 

reflexa,  275,  277,  279 

regalis,  160,  164 

reticulata,  165,  166 

Rhabdocephala,  135,  139,  165 

Ribautdelphax,  336,  338 

riograndensis,  159 

ritana,  256 

robusta,  19,  21 

rosacea,  151,  154 

rosea,  65 

rostrata,  19,  20,  29 

rubella,  131 

rugosa,  78 

saileri,  335 

salsa,  19,  20,  32 

saltuella,  91,  96 

sandersi,  276,  277,  283 

sanguinea,  151,  154 

satilla,  277,  286 

saxatilis,  141,  142 

Secopennis,  224 

semifascia,  63 

semipullatus ,  107 

serrata  (Cathedra),  147 

serrata  (Flexamia),  257 

serratus  (Colladonus),  105,  108 

serratus  (Pristiopsis),  135 

servillei,  159 

sexpunctatiis,  106 

sexualis,  221 

schmidti,  166 

Scolopsella,  139,  165 

shaman,  144,  146 

sif^nata,  204 

silena,  148 

simillima,  335,  336,  341 

singularius,  105,  119 

Sinuala,  139,  166 

slossonae,  255 

smaragdina,  148 


sodalis,  141,  144 

sonorae,  107 

Sophonia,  188,  196,  198 

sordidus,  163 

Spartopyge,  224 

spatulata,  131 

spectabilis,  159 

spinosus,  44,  48 

stall  (Sinuala),  166 

stalii  (Calotartessus),  62 

Stalubra,  137 

stejnegere,  335 

Stenotortor,  182,  184,  186 

stigmata,  68,  76 

stillifera,  151,  154 

Stirellus,  43 

striata,  213,  216 

strobila,  19,  27 

stylata,  258,  260,  266 

stylosa,  19,  20,  37 

subarctica,  335,  336 

subhimalaya,  186 

submaculata,  135 

subviridis,  151,  154 

supina,  19,  27 

surcula,  251 

suturalis,  182,  213 

Suva,  324 

Tahura,  182 

tartessoides,  68,  70 

Taslopa,  63,  65 

tecomariae,  182 

tenellus,  4 

tenera,  19,  20,  38 

terebrans,  19,  20,  40 

tessellata  (Enhydria),  156,  157 

tessellatus  (Colladonus),  105,  124 

tethys,  44,  45 

Tettigarcta,  61 

texana,  258,  261,  271 

thoracica,  189,  190 

timberlakei,  91,  95 

tinga,  71 

titulus,  105,  118 

tolucensis,  105,  111 

trabilis,  105,  119 

tricornis,  63 

tripartitus  (Ollarianus),  132 

tripartitus  (Phrictus),  160,  164 

tuba,  151,  155 

tuberculata  (Enchophora),  151,  155 

tuberculata  (Sinuala),  166 

tubtdu-s.  109 

tumacacoriae,  141,  144 

turpiter,  120 

typicus,  221 

ultimus,  105,  HI 

ultihra,  156 

uniformis  (Delphacodes),  335 

uniformis  (Enchophora),  151,  155 

Unkanodes,  335,  337 

usitata,  91,  95 

utahna,  19,  20,  35 

varicolor,  63 

vativala,  19,  20,  38 

ventriculus,  44,  48 


1988  Index  353 

verecundus,  105,  116  wilhelmi,  335,  337 

vernalis,  141,  144  Wolfella.  63 

vincula,  130  xanthopterus,  160,  164 

viridiguttata,  148  youngi,  258,  267 
viridipennis,  151,  155  in  on  oc 

J       OC1  yumana,  19,  20,  35 

volcanicola,  91,  93  zacate,  246,  248 

whitcombi,  19,  25  zamora,  234,  235