Skip to main content

Full text of "The University of Kansas science bulletin"

See other formats 


^  --/V/*/  -U 


HARVARD  UNIVERSITY 


LIBRARY 


OF  THE 


Museum  of  Comparative  Zoology 


Nuv  1  6  lyo** 


BULLETIN  of  1Z 

The  University  oi  Kansas 


ia,'-is6' 

SCIENCE  BULLETIN 


(Continuation  of  Kansas  University   Quarterly) 


Vol.  XXI 


LAWRENCE,  KANSAS 

Published  Semimonthly  from  January  to  June 
and  Moniltly  from  July  to  December 


Vol.  34  MARCH  15,  1933  No.  5 

Entered  as  second-class  matter  December  29,  1910,  at  the  post  ofiBce  at 

Lawrence,   Kansas,   under  act   of   July   16,   1894. 

15-3482 


^ 


NOTICE  TO  EXCHANGES 

The  attention  of  learned  societies  and  other  institutions  which 
exchange  scientific  publications  with  the  University  of  Kansas  is 
called  to  the  list  of  publications  of  this  University  on  the  third  and 
fourth  pages  of  the  cover  of  this  issue. 

Those  marked  "Supply  exhausted"  cannot  be  furnished  at  all;  as 
far  as  the  supply  permits  the  remaining  numbers  will  glady  be  fur- 
nished to  any  of  our  exchanges  who  may  need  them  to  complete  their 
files. 

Back  numbers  of  the  Kansas  University  Quarterly,  as  far  as  pos- 
sible, will  be  sent  to  those  of  our  newer  correspondents  who  are  able 
and  willing  to  reciprocate.    Separates  are  available  to  specialists. 


ANNOUNCEMENT 

The  Kansas  University  Science  Bulletin  (continuation  of  the 
Kansas  University  Quarterly)  is  issued  in  parts  at  irregular  inter- 
vals. Each  volume  contains  from  300  to  400  pages  of  reading  mat- 
ter, with  necessary  illustrations.  Exchanges  with  other  institutions 
and  learned  societies  everywhere  are  solicited.  All  exchanges  should 
be  addressed  to  the  Library  of  the  University  of  Kansas. 

The  Kansas  University  Science  Bulletin, 
Library  of  the  University  of  Kansas, 
Lawrence,  Kan. 


EDITORIAL  BOARD 

H.  B.  HuNGERFORD,  Chairman.  C.  M.  Baker. 

E.  H.  Taylor,  Secretary.  0.  0.  Stoland. 

J.  D.  Stranathan.  R.  C.  Moore. 
A.  W.  Davidson. 


THE 

KANSAS  UNIVERSITY 

Science  Bulletin 


DEVOTED   TO 

THE  PUBLICATION  OF  THE  RESULTS  OF 

RESEARCH  BY  MEMBERS  OF  THE 

UNIVERSITY  OF  KANSAS 


Vol.  XXI 

(Whole  Series,  Vol.  34) 


PUBLISHED  BY  THE  UNIV-ERSITT 
LAWRENCE,   KANSAS 

1933 


PR[NTED    BY    KANSAS    STATE    PRINTING    PLANT 
W.     C.     AUSTIN.     STATE     PRINTER 

TOPEKA      1934 

15-3482 


^~/^7i   U 


CONTENTS  OF  VOLUME  XXI 

No.  PAGE 

1.  The  Genus  Notonecta  of  the  World.    H.  B.  Hunger j or d.  ...       5 

2.  The  Giant  Water  Bugs  (Belostomatidse,  Hemiptera).    Carl 

Cicmmings 197 

3.  A  Study  of  the  Genus  Brachymctra  (Hemiptera,  Gerridse). 

J.  Gilbert  Shaw 221 

4.  The  Genus  Chlorotettix  in  America  North  of  Mexico 

(Homoptera,  Cicadellidse).    Win.  F.  Brown 235 

5.  A  New  Species  of  Lizard  from  Mexico.    E.  H.  Taylor 257 

6.  Observations  on  the  Courtship  of  Turtles.    E.  H.  Taylor.  .   2G9 

7.  A  New  Bassariscus  from  the  Lower  Pliocene  of  Nebraska. 

Claude  W.  Hibbard 273 

8.  Two  New'  Species  of  Coelacanthus  from  the  Middle  Penn- 

sylvanian  of  Anderson  County.    Claude  W.  Hibbard.  .  .  .   279 

9.  Cephalopods  of  the  Pierre  Formation  of  W^allace  County. 

M.  K.  Elias 289 

10.  A  New  Megasecopteron  from  the  Carboniferous  of  Kansas. 

F.  M.  Carpenter 365 

11.  Comparative  Anatomical  Research  within  the  Genus  Ribes. 

James  C.  Bates 369 

12.  Morphology  and  Anatomy  of  Mollugo  verticillata  L. 

Sister  M.  Anthony  Payne 399 

13.  Morphology  and  Anatomy  of  Lygodesmia  juncea  (Pursh) 

D.  Don.    Liza  Spann 421 

14.  Comparative  Anatomy  of  Mentzelia  oligosperma  and  M. 

decapetala.     W.  E.  Booth 439 

15.  Some  Studies  on  the  Transference  of  Ions  in  Anhydrous 

Acetic  Acid  Solution.    Arthur  W.  Davidson  and  Vernoji 
Holm    463 

16.  The  Effects  of  a  Minimal  Dose  of  X-Rays  upon  Chick 

Embryos.    H.  Ernest  Crow 479 

(3) 


THE  UNIVEESITY  OF  KANSAS 

SCIENCE  BULLETIN 

Vol.  XXL]  March,  1933.  [No.  1. 


The  Genus  Notonecta  of  the  World 
(Notonectida3-Hemiptcra) 

H.   B.   HUNGERFORD, 
llcvul   cif  Depaitment   of  Entomologj-,  University  of  Kansas,  Lawivnoe.* 

CONTENTS 

PAGE 

I.  lulioduction  6 

II.  Family  Characteristics  Including  Key  to  Genera 9 

III.  Anatomical  Notes,  Technique  and  Terminology 12 

IV.  Biology  of  the  Notonectida^ Ki 

V.  Relations  to  Other  Aquatic  Life 17 

VI.   Geographical  Distribution 18 

VII.    Geological  Distribution  21 

VIII.    Taxonomy  of  the  Notonecta 23 

A.  Key  to  Subgenera 23 

B.  Notonecta  of  the  Eastern  Hemisphere 25 

C.  Notonecta  of  Western  Hemisphere 65 

1.  North,  Central  and  Insular  America 65 

2.  South   America    125 

D.  Check  List  of  Species 145 

IX     Acknowledgments   147 

X.    Bibliography   151 

XI.   Addenda  I55 

XII.   Index I57 

XIII.    Plates    r.  162 


Abstract:  This  paper  is  the  result  of  studies  covering  a  number  of  years 
and  visitations  to  many  important  museums  of  Europe  and  North  America. 
All  species  are  redescribed  and  most  of  them  are  illustrated  in  color.  Besides 
five  color  plates  there  are  twelve  plates  in  black  and  white,  illustrating  struc- 
tural parts  of  taxonomic  interest.  Keys  to  the  species  of  each  hemisphere  are 
included. 


*  Contribution  from  the  Department  of  Entomology,  University  of  Kansas  witli  acknowl- 
edgments to  the  University  of  Michigan  Biological  Station  for  opportunity  to  pursue  this 
study  during  several  summers  there. 

(5) 
2—3482 


(3  The  University  Science  Bulletin 

INTRODUCTION 

IT  IS  now  twenty-nine  years  since  the  appearance  of  G.  W.  Kir- 
kaldy's^^  "Uber  Notonectiden,"  which  concluded  his  studies  on 
the  Notonectidse  of  the  world,  and  thirty-six  years  since  his  "Re- 
vision of  the  Notonectidse,  Part  I."  ^-^  was  published.  It  was  in  this 
last-named  paper  that  Doctor  Kirkaldy  reported  his  studies  upon 
the  genus  Notonecta.  In  the  years  that  have  intervened  between 
that  time  and  the  present  some  species  have  been  described  as 
new  and  structural  characters  of  specific  value  discovered  that  are 
much  more  precise  than  those  known  to  him.  I  believe,  therefore, 
that  it  is  time  to  assemble  our  knowledge  of  this  genus,  and,  if 
possible,  stimulate  a  wider  interest  in  this  group.  Surely  there  are 
species  not  yet  taken  or  recognized  as  new,  and  more  intensive  col- 
lecting and  closer  scrutiny  will  prove  fruitful  indeed.  There  are 
vast  areas  where  the  terrestrial  insect  fauna  is  quite  well  known 
from  which  the  aquatic  collections  are  very  meager.  This  is  due 
in  large  measure  to  the  fact  that  the  equipment  necessary  for 
thorough  water  collecting  is  somewhat  different  from  that  employed 
in  the  capture  of  terrestrial  insects.  While  much  interesting  material 
could  be  taken  with  the  ordinary  land  nets,  collectors  are  loath  to 
dampen  their  equipment.  A  little  more  attention  to  water  insects 
by  the  general  collector  would  be  amply  repaid. 

In  passing  in  review  the  literature  that  has  dealt  with  the  Noto- 
necta of  the  world  we  find  just  two  papers  of  length — Fieber's 
"Rhynchotographieen,"  ^^  1851,  and  Kirkaldy 's  "Revision  of  the 
Notonectidffi,"  •*'  1897.  These  papers  were  preceded  by  the  follow- 
ing, which  paved  the  way:  Linnaus  in  his  tenth  edition  of  "Systema 
Naturae,"  1758,  erected  the  genus  Notonecta  and  placed  in  it  three 
species.  .V.  glauca,  N.  striata  and  N.  minutissima.  The  last  two 
were  later  removed  to  other  genera.  Leach  in  his  "Classification  of 
Notonectides,"  ^^  1817,  proposed  a  new  genus  Plea,  and  Spinola  in 
his  Essai,^^  in  1837,  brought  out  the  genera  Anisops  and  Enithares. 

In  Fieber's  paper  a  number  of  new  species  were  described  and 
many  varieties  proposed  on  the  basis  of  size  and  color  alone.  Thus 
only  in  those  cases  where  the  types  are  extant  can  Fieber's  species 
be  fixed  with  satisfaction.  His  casual  disposition  of  previously 
described  species  is  well  illustrated  by  his  treatment  of  Amyot  and 
Servilles  Notonecta  viexicana.  After  describing  A',  klugii  he  re- 
marks that  N.  mexicana  "diirfte  wohl  nur  eine  varietlit  der  oben 
beschriebenen  N.  Klugii  sein,  oder  zur  folgenden  gehoren"  (his  own 
N.  ijyipressa).    A  further  problem  in  nomenclature  arises  from  this 


Hungerford:    Genus  Notonecta  7 

author's  treatment  of  specific  names.  For  illustration:  He  de- 
scribes Notonecta  variabilis  in  three  lines  and  then  names  and 
describes  four  varieties,  citing  the  habitat  of  each;  the  first  variety, 
A',  scutellaris,  from  Brazil  and  Porto  Rico,  the  second,  N.  maculata, 
from  Baltimore  (U.  S.  A.),  and  the  third  and  fourth  varieties  from 
Brazil.  Since  two  or  more  species  are  involved,  it  is  necessary  to 
consider  his  first-named  variety  as  his  A' .  variabilis  and  its  locality 
as  the  type  locality. 

In  Kirkaldy's  paper,  which  appeared  nearly  fifty  years  later, 
the  author  made  an  honest  and  worthy  efTort  to  set  in  order  the 
taxonomy  of  the  genus  Notonecta.  He  appears  to  have  made  every 
attempt  to  locate  and  examine  types  and  to  gather  together  all  the 
material  he  could  command.  His  types  have  been  designated  and 
preserved.  He  introduced  some  structural  characters  and  brought 
some  order  out  of  the  chaos.  His  allowance  for  wide  variability  in 
some  species,  has,  in  some  cases,  been  too  great,  giving  an  un- 
warranted distributional  range  for  such  species.  For  example,  he 
did  not  note  the  specific  differences  between  the  common  North 
American  Notonecta  umhdata  Say  and  a  much  smaller  species  in 
Chile. 

In  this,  the  third  general  study  of  the  genus,  I  have  gone  to  the 
sources  both  for  descriptions  and  types  as  far  as  they  have  been 
available  to  me.  I  have  tried  to  account  for  every  named  species, 
but  there  are  some  descriptions  without  existing  types  or  without 
material  from  type  localities  that  fit  the  descriptions  which  are 
often  too  brief.  Whether  the  descriptions  are  inaccurate,  or  whether 
the  species  in  question  have  not  been  seen  by  me,  1  cannot  say. 
Most  of  the  older  descriptions  are  based  on  color  and  it  is  almost 
impossible  to  appropriate  them  with  any  degree  of  certainty.  This 
paper  has  been  in  progress  for  twelve  years  and  publication  with- 
held in  the  hope  that  every  case  might  be  clarified.  I  find  that  the 
time  has  come  when  I  must  present  my  results,  such  as  they  are, 
hoping  that  the  future  may  uncover  points  now  obscure  to  me.  The 
colored  figures  so  faithfully  drawn  for  me  by  Miss  Kathleen  Doer- 
ing  will  enable  anyone  to  determine  the  specimens  in  the  catch  that 
are  fully  and  typically  colored.  The  tcneral  forms  and  the  bi- 
chromatic  species  may  be  perplexing.  In  such  cases  note  the  size, 
shape  of  the  head,  thorax,  mesotrochanters,  abdominal  sternites  and 
genitalia.  Kirkaldy  ^'^  made  a  significant  contribution  to  the  tax- 
onomy of  the  Notonecta  when  he  introduced  the  more  or  less  precise 
measurements  of  the  head.     Champion,  1901,  in  the  Biologia  Cen- 


8  The  University  Science  Bulletin 

trali  Americana  (Hem.-Het.,  Vol.  II)  noted  the  fact  that  certain 
species  have  the  ventral  abdominal  keel  bare,  while  others  have  this 
keel  covered  with  hair.  Delcourt's  -  study  of  the  biology  and  pig- 
mental variations  of  the  forms  of  his  region  have  been  helpful; 
indeed,  he  demonstrated  the  value  of  biological  facts  in  establishing 
his  A^.  viridis  as  a  distinct  species.  Edwards  ^^  in  describing  his  N. 
halophila  (now  known  to  be  N.  viridis  Delcourt)  called  attention  to 
the  anterolateral  angles  of  the  prothorax. 

In  1917,  while  studying  the  biology  of  some  species  of  Notonecta 
in  New  York  state,  I  discovered  that  the  female  gonapophyses  have 
some  specific  characteristics.^^  A  little  later  I^^  investigated  the  value 
of  the  male  genital  capsules  and  found  them  to  be  quite  precise  and 
satisfactory  in  specific  determination.  An  examination  of  long  series 
of  a  given  species  taken  from  widely  separated  places  in  the  United 
States  has  demonstrated  the  fixed  characters  Of  the  male  genitalia. 
Since  that  time  I  have  submitted  drawings  of  the  genital  capsules  of 
the  species  that  I  have  described,  and  other  workers  have  used  the 
male  genital  characters  in  taxonomic  problems  with  success. 

In  1922  R.  Despax  ^^  made  a  study  of  Notonecta  maculata  Fahr. 
and  Notonecta  glauca  L.  and  figured  the  genitalia.  In  1924  Ray- 
mond Poisson  '^■"'  figured  the  male  claspers  of  Notonecta  furcata, 
Notonecta  glauca,  Notonecta  maculata  and  Notonecta  viridis  Del- 
court  and  demonstrated  their  specific  differences.  He  writes: 
".  .  .  les  differentes  pieces  de  I'armature  genitale,  dans  chacune 
des  especes,  conservant  une  structure  constante."  More  recently 
this  last  author^"  has  compared  the  three  species  common  to  his 
country  and  Asia  Minor.  His  drawings  show  general  resemblance 
in  each  case.  Esaki  (1927),  in  describing  his  N.  horvathi  {=  N. 
pallidida  Poisson,  1926),  compared  the  direction  and  relative  length 
of  the  lateral  margin  of  the  pronotum  of  this  species  with  that  of 
A^.  glauca  Linn.,  and  in  his  recent  description  of  A'',  saramao  Esaki 
he  has  attempted  to  state  precisely  in  degrees  the  anterolateral 
angles  of  the  pronotum. 

Another  character  that  I  have  found  to  be  of  value  in  separating 
certain  confusing  species  is  that  of  the  mesotrochanter.''^  In  some 
species,  the  majority,  indeed,  it  is  rounded,  but  in  others  it  is  angu- 
late  or  even  spurred.  The  shape  of  the  sternites  of  the  abdomen  are 
often  specific,  and  the  last  abdominal  sternites  of  the  females  are 
especially  useful.  The  characteristic  shape  of  the  head  and  thorax 
often  defy  a  word  description,  yet  are  sufficiently  distinctive  to 
identify  a  species.    The  sexes  are  sometimes  of  different  color,  as, 


Hungerford:    Genus  Notonecta  9 

for  example,  the  males  of  Notonecta  lobata  Hungerford,  which  are 
usually  black,  while  the  females  are  red.  They  may  also  be  of 
somewhat  different  shape.  The  head  and  thorax,  for  example,  of 
Kirkaldy's  Notonecta  ceres  are  cjuite  unlike  in  the  two  sexes  (see 
Color  Plate  III,  figures  2  and  3).  The  males  are,  in  many  species, 
slightly  smaller  and  the  distance  between  the  eyes  slightly  less, 
relatively,  than  in  the  females.  In  every  case  of  doubt  an  examina- 
tion of  the  quite  precise  characters  of  the  male  genital  capsule  will 
settle  the  question  of  specific  identity. 

FAMILY  CHARACTERISTICS 

The  Notonectidffi  embrace  perfectly  aquatic  forms  differing  from 
all  others  (except  the  Pleidse  and  Helotrephidse  formerly  included  in 
this  family*)  in  the  persistent  habit  of  swimming  on  their  backs. 
They  are  deep-bodied,  convex  dorsally  and  more  or  less  triangular 
in  cross  section.  The  eyes  are  large,  reniform,  and  twice  sinuated 
on  the  outer  side.  Ocelli  are  absent.  Antennae  with  three  or  four 
well-defined  segments  and  partly  concealed  between  the  head  and 
thorax.  Beak  four-segmented.  Front  and  middle  legs  adapted  for 
grasping  and  hind  legs  more  or  less  flattened  and  fringed  for  swim- 
ming. Tarsi  two  or  three  segmented  (counting  a  small  basal  seg- 
ment). Two  tarsal  claws  present  in  all,  but  inconspicuous  on  the 
tarsi  of  the  hind  legs.  Scutellum  visible.  Abdominal  venter  with 
median  longitudinal  keel  provided  along  the  lateral  margins  at  least 
with  guard  hairs,  which,  together  with  movable  rows  of  similar  hairs 
from  the  sides  of  the  venter,  are  capable  of  closing  over  the  two 
longitudinal  troughs  thus  forming  air  chambers. 

I  propose  to  arrange  the  gensra  of  the  family  as  now  restricted 
as  indicated  in  the  key  which  follows: 

Key  to  Genera  of  Ngtonectid.e 

A.     HTnelytral  commissure  without  pit  at  anterior  end (Subf.    Notonrctin.s.) 

B.    Intermediate  tarsus  with  two  well-developed  segments  and  a  very  small  basal 
one.     Intermediate  femur  stout  with   anteapical   pointed  protuberance. 

(Tribe  Notonectini.) 

C.    Anterolateral   narg'ns  of  prothorax   not   foveate Notonecta. 

CC.    Anterolateral  marg-ns  of  prothorax  foveate.     (See  PI.   VI,  fig.   Z).  .  .Enit hares. 
BB.    Intermediate   tarsus   with    one   well -developed    segment    and    the    middle    femur 

without  anteapical   pointrd   protuberance (Tribe   Nychini.) 

C.    Antenna   4   segmented Martarega. 

CC.    Ant  nna   3  s'^gniented Nychia. 

AA.     Hemelytral  commissiue  with  pit  at  anterior  end.f      (See  PI.   VI,  fig.   7.) 

(Subf.   Anisopin.?;.) 

*  These  two  families  fonnin";  the  superfamily  Pleoideae  Esaki  and  China  are  distinguished 
from  the  Notontctoideas  by  having  hemelytra  thick  and  unittd  apically.  Ventral  abdominal 
segments  may  be  medianly  tuberculate  but  not  carinate. 

t  Poisson"'  1926  considers  this  a  sensorial  organ  in  Anisop.'i  producta  Fieb. 


10  The  University  Science  Bulletin 

B.     Ventral  keel  not  extending  onto  last  abdominal   segment.      Male  genital  capsule 
cleft  behind.     Males  without  stridulator  protuberance  on   front  tibia.     Females 

with    short    gonapophyses Paranisops. 

BB.  Ventral  abdominal  keel  extending  onto  last  abdominal  segment.  Male  genital 
capsule  closed  behind.  Males  with  stridulatory  protuberance  on  front  tibia. 
Females  with  elongate  subspatulate  gonapophyses. 

C.    Male  with   anterior  tarsus   1    segmented Atjisops. 

CC.    Male  with  anterior  tarsus  2   segmented Buenoa. 

The  above  arrangement  is  new,  but  appears  to  me  to  be  more 
logical  than  any  proposed  thus  far.  Kirkaldy,^-'^  1897,  divided  the 
family  into  Notonectinse  and  Pleinae.  Hale,  1923^^  and  1924,^^ 
followed  Kirkaldy,  as  did  Hutchinson,*^  1929.  Esaki  and  China, -^ 
1928,  removed  Plea  from  the  Notonectidse  and  revived  the  family 
Pleidae  first  erected  by  Fieber  under  the  name  Plese.  They  made  the 
following  disposition  of  the  groups  formerly  embraced  by  the  family 
Notonectidae: 

Superfamily  Notonectoidese  for  the  family  Notonectidse. 
Superfamily  Pleoidese  for  the  families  Pleidae  and  Helotrephidse. 

Hutchinson,'*^  1929,  proposed  to  divide  his  Notonectinae  into  the 
tribes  Notonectini  and  Anisopini,  the  first  embracing  Notonecta  and 
Enithares  and  the  second  Nychia,  Anisops,  Buenoa,  and  Paranisops. 
He  did  not  undertake  to  place  Martarega  B.  White  and  Signoretiella 
Berg,  but  suggested  that  a  third  tribe  might  have  to  be  instituted 
for  them. 

There  seems  to  be  no  good  reason  for  separating  Nychia  from 
]\Iartarega.  These  two  genera  belong  in  the  same  tribe.  They  agree 
in  general  facies,  both  have  eyes  contiguous  at  base;  both  have  the 
face  transversally  depressed;  both  have  a  curious  curve  in  the 
costal  margin  of  the  hemelytra;  both  have  male  genital  capsules 
that  are  cleft Ibehind.*  In  every  way  save  one  they  are  quite  alike. 
Nychia  has  a  three-segmented  antenna  and  Martarega  a  four- 
segmented  antenna.  However,  it  is  conceivable  that  Nychia  has 
lost  the  fourth  segment  by  atrophy  since  it  is  small  and  loosely 
joined  in  Martarega.  They  are  indeed  closely  related  genera  and 
belong  in  the  same  tribe. 

Whether  Enithares  is  closer  to  Notonecta  than  to  the  Nychia- 
]Martarega  group  is  a  debatable  question.f  It  shares  some  charac- 
ters with  one  and  some  with  the  other.  It  is  like  Notonecta  in 
having  the  eyes  well  separated,  in  having  middle  femora  stout  and 

*  Doctor  Jaczewski,''-  192S,  in  his  "Notonectidie  from  the  .State  of  Parana,"  p.  132,  describes 
the  genital  capsule  of  Martarega. 

t  Doctor  Jaczewski,"  in  Notes  on  Some  West-African  Heteroptera,"  1926,  p.  TS,  places 
it  with  Notonecta,  and  I  conci.r  with  him  in  this  opinion. 


Hun'gerford:    Genus  Xotoxecta  11 

equipped  with  an  anteapical  proturbanee,  in  having  the  claspers  of 
the  male  genital  capsule  in  most  species  alikcj  in  having  the  same 
number  of  tarsal  segments  (two  well-developed  segments  and  a  very 
small  basal  one  in  the  front  and  middle  legs). 

On  the  other  hand  it  has  the  anterolateral  margins  of  the  pro- 
thorax  foveate  like  Nychia  and  the  shiny  surface  and  texture  of  the 
hemelytra.    The  aedeagus  is  unlike  Notonecta. 

As  a  matter  of  fact  it  would  seem  to  me  that  a  cytological  study 
of  these  various  genera  might  throw  some  light  upon  the  problem. 
A  little  work  has  been  done  on  Notonecta  and  has  been  of  real  value 
in  understanding  relationships  within  the  genus. 

In  the  above  key  the  genera  are  arranged  in  phylogenetic  order 
beginning  with  Notonecta,  the  most  primitive  genus,  and  progressing 
to  the  highest.  No  linear  arrangement  is  consistent  in  all  regardsl 
The  best  one  can.  do  is  to  use  the  most  fundamental  differences  and 
allow  for  parallel  and  independent  modifications.  For  example,  the 
ovipositors  of  Notonecta  females  may  be  short  and  weak  or  long  and 
strong. 

The  genus  Notonecta  embraces  63  species,  14  subspecies,  and  is 
widespread  over  the  earth.  There  is  one  species  in  Australia,  one 
in  Africa  south  of  the  Sahara,  several  in  Asia,  several  in  Europe  and 
North  Africa,  25  in  North  and  Central  America  and  20  in  South 
America.  The  genus  Enithares,  with  a  single  exception  {E.  brasi- 
liensis  Spinola  from  S.  A.)  is  found  in  the  Eastern  Hemisphere  only. 
Nychia  belongs  to  the  Eastern  Hemisphere  and  has  its  counterpart 
Martarega  in  South  America.  Anisops  is  widely  distributed  over 
the  Eastern  Hemisphere  and  is  replaced  by  Buenoa  in  the  AVestern 
Hemisphere.    Paranisops  is  confined  to  Australia. 

From  the  standpoint  of  distribution  alone,  Notonecta  might  be 
considered  the  oldest  genus.  This  consideration,  however,  is  sup- 
ported by  the  four  segmented  antennae,  by  the  three  segmented  tarsi 
and  by  the  lack  of  asymmetry  in  the  male  genitalia.  Enithares,  as 
stated  above,  comes  next.  It  is  the  only  other  genus  represented  in 
both  Hemispheres.  Its  only  representative  in  the  Western  Hemi- 
sphere is  a  very  large  species  which  has  the  male  genital  claspers 
unlike.  So  far  as  I  know  all  of  the  other  species  in  the  genus  hick 
this  asymmetry. 

In  some  regards  Enithares  may  be  more  primitive  than  Notonecta. 
The  interocular  space  shows  less  variability  than  in  Notonecta.  The 
female  gonapophyses  are  uniformly  weak,  and  Hale^-  has  reported 

%  Enithares  braziliensis  Spinola  has  the  right   and  U'ft  claspers  unlike.      (See  PI.   ^'III.) 


12  The  University  Science  Bulletin 

that  "Enithares  commonly  anchors  itself  to  submerged  objects,  even 
walking  up  a  plant  stem  to  the  surface  for  a  renewal  of  its  air  sup- 
ply." While  species  of  the  genus  Notonecta  anchor  themselves  to 
submerged  objects,  I  do  not  recall  having  seen  them  climb  supports 
to  the  surface.  Nevertheless  the  foveate  anterolateral  margins  of 
the  prothorax,  the  shiny  surface  and  the  beginnings  of  asymmetry  in 
the  male  genitalia  anticipate  the  Nychini. 

Nychia  and  Martarega  have  the  male  claspers  unlike  on  the  two 
sides.  In  IMartarega  the  claspers  are  the  less  differentiated.  Nychia 
has  lost  the  terminal  antennal  segment.  The  increasing  unlikeness 
of  the  male  claspers  and  the  three  segmented  antennae  of  Nychia 
approach  the  condition  in  the  Anisopinae.  The  Nychini  usually  have 
the  flight  wings  reduced,  accompanied  by  a  small  scutellum.  When 
fully  developed  flight  wings  are  present,  the  scutellum  is  large.  This 
is  true  also  of  Paranisops,  which  belongs  in  the  Anisopinae,  and  is 
the  most  primitive  of  that  subfamily. 

I  consider  the  Anisopinae  at  the  top  of  the  series  because:  (1)  They 
have  oxyhsemoglobin  and  a  pit  on  the  anterior  end  of  the  commissure 
of  the  hemelytra;  (2)  the  males  possess  stridulatory  organs  and  a 
courting  ritual  (excepting  Paranisops) ;  (3)  the  antennae  are  three- 
segmented;  (4)  in  Buenoa  and  Anisops  the  female  gonapophyses 
are  uniform  and  long.  The  male  genital  capsule  is  closed  behind 
and  the  claspers  decidedly  unlike  on  the  two  sides.  The  genitalia 
in  these  last  two  genera  have  reached  such  a  stability  and  uniformity 
that  they  are  of  little  value  for  purposes  of  specific  determination. 

The  Anisopinae  are  more  at  home  in  the  open  water,  cruising  about 
gracefully  and  in  perfect  equilibrium  at  some  distance  beneath  the 
surface. 

Besides  the  differences  stated  in  the  key  the  Notonectinse  and 
Anisopinae  are  unlike  in  the  character  of  the  labrum.* 

ANATOMICAL  NOTES,  TECHNIQUE  AND  TERMINOLOGY 

A  number  of  contributions  have  been  made  to  our  knowledge  of 
the  anatomy'  of  the  backswimming  water  bugs.  Wefelscheid,^^  1912, 
produced  a  study,  "Uber  die  Biologic  und  Anatomic  von  Plea,"  that 
is  monographic  in  nature.  Esaki  ^^  and  China  investigated  the 
Helotrephidao.  In  the  Notonectidae  as  now  restricted  Buenoa  has 
been  studied  by  Bare  ^  and  Anisops  by  Poisson."^  Notonecta  has 
been  the  object  of  many  studies,  for  the  most  part  in  connection  with 
some  general  problem  dealing  with  mouthparts,  wings,  respiration, 

*  Except  Paranisops  which  has  labium  like  Notonecta.  Paranisops  also  has  infracoxal 
plates  hairy  like  Notonecta. 


Hungerford:    Genus  Notonecta  J  3 

or  genitalia.  Some  studies  on  embryology  and  cytology  are  also 
available.  Several  of  these  have  considerable  value  to  the  sys- 
tematist.  Hutchinson,'*^  in  his  "Revision  of  the  Notonectidse  and 
Corixidse  of  South  Africa,"  figures  the  venation  of  the  hind  wings  of 
Notonecta,  Enithares,  Nychia,  Anisops  and  Plea.  Specific  differ- 
ences in  the  ovipositors  of  American  Notonecta  (1918)^^  and  in  the 
genital  capsules  of  the  males  (1919)  were  reported  by  Hungerford.^" 
Since  then  the  genitalia  have  been  used  in  systematic  papers  by 
Hale,  Jaczewski,  Bare,  Hutchinson,  Despax,  Poisson  and  Esaki. 

Hutchinson,  in  the  paper  cited  above,  employed  the  cytological 
evidence  in  chromosome  studies  made  by  Browne,^''  in  1916  and  by 
Poisson  ^"  in  1927  in  his  division  of  the  genus  Notonecta  into  the 
subgenera  Notonecta  S.  Str.  and  Paranecta  (new) ;  the  latter  em- 
bracing Notonecta  lactitans  Kirk,  and  all  the  other  species  without 
digitiform  extension  of  the  male  genital  capsule  and  with  short 
ovipositors  in  the  female;  the  former,  with  Notonecta  glauca  L.  as 
the  type,  and  including  species  in  which  the  males  have  the  digiti- 
form extension  of  the  male  capsules  and  elongate  ovipositors. 
Hutchinson  believes  that  the  subgenus  Paranecta  is  the  older  group 
because  of  the  less  specialized  condition  of  the  ovipostor,  the  simpler 
genital  capsule  and  the  larger  number  of  chromosomes.  Browne 
found  that  in  N.  undulata  Say,  A^.  indica  Linn.  (=  N.  unifasciata 
Guer.  probably)  and  in  N.  shooteri  Uhler,  the  diploid  number  of 
chromosomes  is  26,  the  first  spermatocyte  division  showing  14  (the 
sex  chormosomes  conjugate  late)  and  the  second  13. 

In  both  N.  Glauca  Linn,  of  Europe  and  N.  irrorata  Uhler  of  North 
America  the  diploid  number  is  24,  the  first  spermatocyte  division 
showing  13  and  the  second  12.  It  would  be  very  helpful  if  further 
cytological  studies  in  the  Notonectidse  were  made. 

The  Notonecta  mexicana  A.  &  S.  series,  for  instance,  could  be 
studied  with  profit.  This  group  is  in  some  respects  quite  distinct, 
and  certain  species  such  as  Notonecta  lobata  Hungerford  could  be 
secured  in  large  numbers  in  the  southwestern  United  States.  A 
study  of  Notonecta  handlirschi  Kirkaldy,  which  was  placed  in  the 
new  subgenus  Enitharonecta  by  me  in  1928,"'"  should  be  made  and 
compared  with  a  study  of  some  species  of  Enithares.  Unfortunately 
we  must  first  discover  the  breeding  place  of  this  unique  Australian 
representative  of  the  genus  Notonecta.  So  far  we  know  it  only  from 
the  two  specimens  of  the  type  series  in  the  Vienna  Museum  which 
are  labeled  "Post.,  A.  Fischer,  1878,"  and  one  other  specimen. 

In  this  paper  it  is  necessary  to  present  only  such  gross  anatomical 


14  The  University  Science  Bulletin 

features  as  may  be  useful  in  the  separation  of  the  species.  The 
illustrations  on  Plates  VI,  IX  and  X  will  give  an  idea  of  the  chief 
anatomical  characteristics  of  the  genus  Notonecta. 

In  understanding  the  descriptions  given  in  this  paper  it  is  im- 
portant to  note  that  the  measurements  have  been  made  from  above 
the  dorsal  side  of  the  insect  when  held  in  a  horizontal  position. 
Care  must  be  taken  to  have  the  longitudinal  and  transverse  axes 
exactly  horizontal.  The  length  of  the  head,  compared  to  median 
length  of  pronotum,  has  been  determined  by  projection  and  not  by 
tilting  the  insect  out  of  position;  so,  also,  in  comparing  length  of 
vertex  with  its  anterior  width  and  the  convexity  of  the  anterior 
margin  of  the  vertex  with  that  of  the  frontal  margin  of  the  eye.  The 
ratio  of  the  anterior  breadth  of  vertex  to  the  synthlipsis  is  secured 
by  measuring  the  distance  between  the  eyes  in  front  and  the  nar- 
rowest interocular  space,  the  latter  being  called  "synthlipsis." 

The  lateral  margins  of  the  prothorax  are  always  more  or  less  com- 
pressed dorsoventrally  into  a  more  or  less  thin  edge,  or  ledge,  which 
in  some  species  is  quite  broad  along  the  anterior  portion.  By  view- 
ing this  edge  from  the  side  it  is  usually  seen  to  be  somewhat  sig- 
moid, and,  compared  with  the  long  axis  of  the  insect,  is  either  oblique 
or  nearly  parallel  with  it.  The  shape  of  this  ledge  may  be  ciuite 
different  in  the  sexes  of  the  same  species. 

The  abdominal  sternites  are  often  of  interest  (see  Plate  X).  In 
some  species  the  edge  of  the  mid-ventral  keel  is  bare  on  one  or  more 
segments.  The  so-called  "fourth  abdominal  sternite"  is  so  desig- 
nated for  the  convenience  of  the  taxonomist.  It  is  really  the  sternite 
of  the  fifth  segment.  In  the  females  the  shape  of  the  last  two  ab- 
dominal sternites*  afford  excellent  characters  (see  Plate  X).  In 
the  A^.  mexicana  A.  &  S.  series  the  penultimate  sternite  is  very  nar- 
row, in  A^.  montezuma  Kirk,  it  is  long,  broad  and  keel  is  bare;  in 
A",  undulata  Say  it  is  relatively  long  and  cleft  behind,  while  in  A^. 
irrorata  Uhler  it  is  broader  than  long.  The  last  abdominal  sternite 
is  no  less  interesting.  On  Plate  X  I  have  figured  four  distinct  types. 
Figure  1  illustrates  the  characteristic  shape  of  the  A",  mexicana 
A.  &  S.  series,  figure  3  the  usual  shape  of  the  subgenus  Paranccta 
Hutchinson,  and  figure  4  that  of  the  subgenus  Notonecta.  In  Para- 
necta  the  first  pair  of  gonapophyses  are  short  and  in  Notonecta  they 
are  elongate. 

*  Also  referring  to  those  visible  as  the  fifth  and  sixth.  Of  course  they  are  actually  the 
sternites  of  the  sixth  and  seventh  and  the  three  pairs  of  gonapophyses  are  attached  to  the 
eighth  and  ninth.     The  genital  capsule  of  the  male  belongs  to  the  ninth  segment. 


Huxgerford:    Gexus  Xotoxecta  15 

The  male  genital  capsules  with  their  claspers  or  paramcres  afford 
good  characters.  There  are  three  distinct  groups  of  the  genus  based 
upon  the  shape  of  the  capsule  itself.  The  subgenus  Enitharonecta, 
with  its  caudal  projection  and  reduced  lobes  behind  the  claspers 
(see  Plate  VIII,  fig.  4) ;  the  subgenus  Paranecta,  which  lacks  a 
digitate  process  (see  Plate  XIII) ;  and  the  subgenus  Notonecta, 
which  possesses  such  a  process  (see  Plates  XVI  and  XVII).  The 
male  genital  capsule  is  hidden  within  the  last  abdominal  segment 
and  must  be  drawn  out  into  view,  and  for  careful  study  must  be 
cleared.  Both  operations  are  quite  simple  after  a  little  practice. 
To  remove  the  capsule  moisten  the  insect  with  5  per  cent  alcohol  to 
relax  it.  After  a  few  minutes  hold  the  insect  between  thumb  and 
finger  of  one  hand  and  with  the  other,  by  means  of  a  dissecting 
needle  bent  at  the  tip,  reach  in  and  bring  out  the  capsule.  It  is 
often  best  to  slip  the  needle  in  between  the  body  wall  and  the 
capsule  on  both  sides  to  loosen  the  connections  before  undertaking 
to  draw  out  the  capsule.  To  clear  the  capsule  place  it  in  caustic 
potash  solution  and  either  boil  it  for  a  time  till  clear  or  let  it  stand 
in  the  cold  solution  two  or  three  days.  I  prefer  the  latter  method. 
After  washing  the  cleared'  capsule  in  water  it  can  be  studied  under 
the  binocular  and  then  stored  in  a  vial  of  alcohol  (70  per  cent)  or 
dehydrated  with  alcohol,  placed  for  a  few  minutes  in  xylol  and 
mounted  in  a  drop  of  balsam  on  a  card  or  celluloid  slip  to  be  placed 
on  the  pin  beneath  the  insect.  Another  method  that  is  even  better 
is  to  clear  a  little,  wash  in  water,  then  place  in  a  little  dish  of  acetic 
acid.  After  ten  minutes  add  a  drop  of  clove  oil,  adding  clove  oil 
drop  by  drop  every  few  minutes,  and  drawing  off  the  excess  with  a 
pipette.  After  a  time  the  capsule  will  be  in  pure  clove  oil.  from 
which  it  can  be  mounted  directly  in  a  drop  of  balsam  on  a  celluloid 
card  for  pinning  beneath  the  insect.  If  the  insect  is  then  placed  in 
a  dust-free  chamber  until  the  mount  hardens,  this  method  will  be 
found  most  satisfactory. 

The  aedeagusf  also  should  be  examined  and  the  amount  of  clear- 
ing of  the  genital  capsule  must  be  enough  to  enable  a  study  of  the 
claspers  and  yet  not  enough  to  destroy  the  characters  of  the  aedeagus, 
Avhich  in  certain  cases  are  useful.  The  difficulty  in  securing  uniform 
clearing  so  that  precise  comparisons  of  scleratized  parts  can  be 
made  and  the  wide  differences  in  the  relative  positions  of  the  parts 
of  the  aedeagus  in  different  individuals  has  led  me  to  omit  specific 
descriptions  of  this  structure.     The  structure,  called  the  "internal 

t  Phallic  organ,  according  to  Snodgrass. 


16  The  University  Science  Bulletin 

•stay"  by  Hutchinson''^  (see  his  Plate  XXVIII,  fig.  2),  appears  to 
be  most  developed  in  the  N.  mexicana  A.  &  S.  group  and  but  slightly 
developed  in  the  N.  lutea  Miiller  group. 

THE  BIOLOGY  OF  THE  NOTONECTID.E 

The  backswimmers  are  among  the  best  known  of  the  water  bugs. 
They  inhabit  the  ponds  and  pools  of  nearly  every  land.  The  fact 
that  they  swim  on  their  backs  readily  distinguishes  them  in  their 
native  element.  They  are  predaceous  creatures,  the  larger  species 
often  attacking  insects  and  other  animals,  even  small  fish,  larger 
than  themselves.  The  small  nymphs  of  Notonecta  and  probably 
all  of  the  Anisopina?,  even  in  the  adult  stage,  feed  to  a  large  extent 
upon  the  small  Entomostracan  life  of  the  water.!  The  Anisopinse 
have  the  two  anterior  pairs  of  legs  margined  with  rather  long  spines, 
which  form,  when  flexed,  a  splendid  crib  for  the  retention  of  ostra- 
cods,  cladocera,  etc.,  many  of  which  may  be  held  captive  at  one 
time. 

Members  of  the  genus  Notonecta,  and  no  doubt  of  the  Enithares§ 
also,  can  inflict  a  severe  "sting"  with  their  beaks  when  incautiously 
handled,  a  fact  which  led  to  the  use  of  the  name  "Wasser-bienen" 
by  the  early  German  writers.  This  disagreeable  behavior  has  given 
them  a  place  in  the  literature  of  Medical  Entomology  (see  Riley 
and  Johannsen  Handbook  of  Medical  Entomology).  No  doubt  most 
of  the  observations  in  this  regard  have  been  made  by  collectors,  but 
on  several  occasions  I  have  been  impressed  by  their  reputation  as 
"stingers"  and  their  eccncmic  importance  to  the  owners  of  com- 
mercial swimming  pools.  Instances  have  been  reported  to  me  in 
which  the  swimming  pools  have  been  so  infested  with  thes?  insects 
and  the  complaints  so  numerous  among  bathers  that  the  elimination 
of  the  backswimmers  became  necessars^  to  restore  the  popularity 
of  the  pools. 

Most  species  winter  as  adults  either  hidden  in  the  mud  and  other 
debris  of  the  pool  or  remain  more  or  less  active  as  conditions  may 
permit.  They  have  been  observed  swimming  beneath  the  ice  in 
spring-fed  pools  in  midwinter.  There  is  some  evidence,  mostly  un- 
published, to  indicate  that  certain  species  may  also  overwinter  in  the 
egg  stage.*    Some  species  have  been  observed  to  leave  the  shallow 

t  Anisops  and  Buenoa  can  be  anil  Iiave  b^'cn  reared  in  rapti\  ily  upon  mosquito 
wriggl  rs. 

§  H.  M.  Hale  in  the  Records  of  the  South  .•\ustralian  M\iseuni.  Vol.  H,  No.  3,  1923, 
reports  the  "sting"   of  Enithares  as  similar  to  tl'at   of  a  bee  sting. 

*  See  under  .V.  lutra  Mijller  and  A^.  borealis  Bueno  and  Hussey. 


Hungerford:    Genus  Notonecta  17 

temporary  pools  of  summer  to  fly  to  deeper  waters  in  which  to 
winter. 

There  are  some  interesting  records  of  countless  swarms  of  Noto- 
necta that  for  some  reason  have  been  precipitated  from  their  flight  in 
mid-air  to  drop  like  hail  upon  the  ground.  One  such  "rain  of  insects" 
recorded  in  the  northern  United  States  in  1846  covered  an  extent  of 
at  least  twenty-five  miles  in  one  direction. 

Sometimes  Notonectida3  fly  to  trap  lights,  and  Mr.  R.  A.  Stirton, 
who  collected  these  insects  for  me  in  Central  America,  drew  them 
within  reach  of  his  net  by  the  use  of  a  flash  light  over  the  water. 

Mating  takes  place  in  the  water,  and  the  courting  maneuver  ol 
Buenoa,  and  probably  mostf  other  Anisopinse,$  includes  stridulation 
of  the  males,  which  make  a  chirping  sound  by  rubbing  the  front 
legs  against  the  base  of  the  beak  as  they  approach  the  females. 
The  eggs  of  some  of  the  Notonecta  (the  majority)  are  affixed  to 
aquatic  vegetation  and  other  supports  in  the  water,  while  others 
insert  their  eggs  more  or  less  deeply  into  plant  tissues.  The  oviposi- 
tion  of  only  one  species  of  Enithares  has  been  published,  and  this 
one  affixed  its  eggs  to  aquatic  vegetation.  So  far  as  known,  all  of 
them  have  the  first  pair  of  female  gonapophyses  short  and  weak 
and  not  adapted  for  piercing  plant  tissue.  The  Anisopinse,  on  the 
other  hand,  with  the  exception  of  Paranisops,  have  well-developed 
gonapophyses  and  must  insert  their  eggs  as  has  been  observed  for 
some  of  the  Buenoa.  So  far  as  known  there  are  five  nymphal  in- 
stars,  and  there  may  be  one  or  more  generations,  depending  upon 
the  species  and  the  climate. 

RELATIONS  TO  OTHER  AQUATIC  LIFE 

The  Notonecta  are  powerful  swimmers,  but  not  as  fleet  as  the 
Corixids.  Ordinarily  they  swim  short  distances,  coming  to  rest 
back  downward  just  beneath  the  surface  film,  the  tip  of  the  abdo- 
men in  contact  with  the  surface,  the  body  and  head  downward  at 
such  an  angle  that  just  the  claws  of  the  intermediate  limbs  may 
touch  the  surface  and  the  hind  limbs  directed  well  forward  of  right 
angles  to  the  body,  poised  for  a  sudden  rowing  stroke,  or  else  they 
dive  to  cling  to  some  submerged  object.  Certain  species  prefer 
open  water,  others  the  shelter  of  aquatic  vegetation,  and  still  others 
the  dark,  shadowy  places  beneath  overhanging  bushes.     Some  will 

t  Paranisops  is  doubtless  mule. 

t  Hale  in  Records  of  the  South  Australian  Museum,  Vol.  II,  No.  3,  1923,  p.  406,  describes 
the  courtship  and  stridulation  of  Anisops  hyperion  Kirk,  which  is  very  similar  to  observations 
made  by  Hungerford  and  by  Bare  of  Buenoa. 


18  The  University  Science  Bulletin 

live  in  muddy,  stagnant  little  ponds,  others  only  in  cool,  clear 
waters.  For  the  most  part  the  Notonecta  live  in  standing  bodies 
of  water  or  in  quiet  pools  of  a  watercourse.  One  species  will  tolerate 
the  brackish  waters  by  the  sea,  and  some  have  been  taken  in  the 
acrid  coffee-colored  waters  of  some  peat-bog  pool.  As  a  group  they 
are  probably  indifferent  to  a  wide  range  of  pH. 

Their  relation  to  the  other  small  creatures  of  the  water  is  that  of 
predator.  In  their  early  stages  they  feed  upon  the  small  Crustacea 
(Entomostraca  mostly)  and  small  insect  larvse;  in  their  later  stages 
upon  larger  forms,  even  small  tadpoles  and  little  fishes.  In  turn 
they  may  be  taken  and  devoured  by  the  naiads  of  dragon  and  dam- 
sel flies,  by  Belostomatids  and  Nepids  and  by  predaceous  water 
beetles.  Now  and  then  they  are  captured  by  some  surface-dwelling 
insect  or  other  animal.  They  are  seldom  eaten  by  the  fishes,  at 
least  much  less  frequently  than  are  the  Corixids,  which  often  dwell 
with  them  in  large  numbers. 

Their  contention  for  the  same  food*  that  small  fishes  eat  and 
their  destruction  of  the  earty  stages  of  some  of  the  fishes  make  them 
of  some  economic  importance  in  fish  culture.  The  economic  relation 
of  the  Notonecta  to  the  plant  life  of  the  water  is  trivial.  Some 
species  make  incisions  in  the  stems  of  plants  for  the  deposition  of 
their  eggs,  others  merely  attach  them  to  the  leaves  and  stems. 
While  chlorophyl  from  Spirogyra  has  been  found  by  Bare^  in  the 
stomachs  of  some  Buenoa,  it  is  not  probable  that  the  Notonecta 
feed  directly  upon  plant  life. 

GEOGRAPHICAL  DISTRIBUTION 

The  family  Notonectidse  is  represented  by  one  or  more  species  in 
nearly  every  land.  Only  one  genus,  however,  has  anything  like  a 
world-wide  distribution.  This  is  the  genus  Notonecta,  and  it  has  not 
been  reported  for  the  East  Indian  Archipelago,  Philippine  Islands, 
New  Zealand,  Tasmania,  Madagascar,  Hawaii  or  other  Pacific  in- 
sular groups.  Furthermore  there  is  but  one  species  found  in  Austra- 
lia, and  this  one  is  a  curious  species  found  only  in  west  Australia  (X. 
handlirschi  Kirkaldy,  known  only  from  three  specimens).  It  is  the 
type  of  the  subgenus  Enitharonecta  Hungerford  and,  as  the  name 
implies,  is  in  some  respects  intermediate  between  Notonecta  and 
Enithares.  The  other  subgenera  of  the  genus  Notonecta  are  Para- 
NECTA  Hutchinson,  Bichromonecta  Hungerford,  Erythronecta 
Hungerford  and  Notonecta  Linnaeus. 

*  G.  C.   Embody-^  records  them  killing  amphlipods  as  large  as  themselves. 


Hl'ngerford:    Genus  Xotonecta  19 

The  subgenus  Paranecta  appears  to  me  to  be  the  oldest.  It  is 
widely  distributed  in  the  Western  Hemisphere,  having  15  species 
in  South  America  and  13  species  covering  Insular  America,  Central 
and  Xorth  America.  It  is  then  the  dominant  subgenus  of  the  West- 
ern Hemisphere.  In  the  Eastern  Hemisphere  we  find  a  species,  .V. 
lactitans  Kirkaldy  in  Africa  south  of  the  Sahara,  but  not  one  in 
Europe!  In  eastern  Asia  we  find  3  in  China,  1  in  eastern  Siberia, 
1  in  Japan  and  1  in  Formosa.  The  distribution  suggests  that  the 
subgenus  may  have  arisen  in  the  new  world,  and  that  the  old-world 
species,  namely  the  one  in  south  Africa,  those  in  eastern  Asia,  Japan 
and  Formosa,  represent  the  terminals  of  an  early  dispersal.  Even 
the  subgenus  Enith.aronecta  in  remote  Australia  may  have  de- 
veloped from  some  ancestral  Paranecta. 

In  Xorth  America  we  have  N.  insulata  Kirby  and  N.  kirbyi  Hun- 
gerford  that  represent  a  link  between  this  subgenus  and  subgenus 
Notonecta.  The  fomier  species  occurs  in  northern  United  States 
and  Canada  east  of  100th  meridian  and  the  latter  in  western  Canada 
and  western  United  States,  ranging  southward  along  tlie  mountain 
chain.  These  two  species,  together  with  A^.  mipressa  Fieber  (A'. 
montezuma  Kirkaldy)  from  JMexico  and  north  into  Texas  form  a 
distinct  group  of  the  subgenus  Paranecta. 

Other  groups  in  this  subgenus  can  be  distinguished,  such  as  the 
N.  bifasciata  Guerin  group,  the  N.  undidata  Say  group,  and  the  N. 
chinensis  group. 

■Most  of  the  South  American  species  are  small  in  size  and  much 
alike  and  belong  to  the  X.  bifasciata  Guerin  group.  The  A^  un- 
dulaia  Say  group  consists  of  species  of  moderate  size  and  is  repre- 
sented in  South  America  by  A^  fazi  Hungerford  in  Chile,  its  close 
relative  A',  vereertbruggheni  Hungerford  just  over  the  mountains  to 
the  east,  and  N.  variabilis  Fieber  in  Brazil.  N.  undidata  Say  is 
widely  distributed  over  North  America  from  coast  to  coast.  In  the 
northern  half  of  the  United  States  and  Canada  it  is  readily  recog- 
nized in  the  field,  but  as  we  proceed  southward  into  the  Lower 
Austral  Zone  we  find  it  more  and  more  replaced  by  N.  indica  Lin- 
naeus, which  is  an  abundant  species,  not  only  in  the  southern  states 
but  in  Cuba  and  Jamaica  and  extending  southward  from  Mexico  to 
Colombia.  N.  variabilis  Fieber  in  Brazil  is  almost  indistinguishable 
from  A',  indica  Linnaeus. 

The  N.  imifasciata  Guerin  group,  possessing  angulate  mesotro- 
chanters  and  a  black  scutellum  margined  with  flavous,  ranges  from 
Mexico  northward  into  British   Columbia.     The   species,  N.   uni- 


20  The  University  Science  Bulletin 

fasciata  Guerin  embraces  several  subspecies  which  in  the  northern 
ranges  is  often  abundant  in  warm  or  so-called  "hot  springs."  While 
no  records  of  this  species  have  been  made  from  South  America  in 
recent  years,  there  are  two  specimens  in  Kirkaldy's  collection  labeled 
"S.  Amer."  and  called  by  him  A^.  americana.  There  are  also  a  few 
specimens  in  Vienna  labeled  'Trfld.  Am."  and  "Frfld.  1874."  Where 
they  came  from  is  uncertain. f 

The  subgenus  Bichromonecta  Hungerford  may  have  arisen  from 
the  Paranecta,  but  has  developed  a  striking  reduction  of  the  penul- 
timate sternite  in  the  female  (see  text  figures,  p.  101),  a  char- 
acter anticipating  the  great  reduction  of  this  sternite  in  the  next 
subgenus.  The  subgenus  is  typified  by  N.  shooteri  Uhler  and  oc- 
cupies a  restricted  range  from  Colombia,  South  America,  on  the 
south  to  California  on  the  north. 

The  subgenus  Erythbonecta  Hungerford,  of  which  N.  mexicana 
Amyot  and  Serville  is  characteristic,  occupies  the  same  range  as 
that  of  Bichromonecta  Hungerford. 

The  subgenus  Notonecta  Linnaeus:  This  subgenus  represented  by 
N.  glauca  Linnaeus  is  characterized  by  having  a  digitate  process  on 
the  male  genital  capsule.  In  most  of  the  species  the  females  have 
long  ovipositors  for  inserting  the  eggs  in  the  tissues  of  plants,  a 
character  not  known  in  any  of  the  other  subgenera.  It  probably 
developed  from  the  Paranecta  through  species  much  like  N.  insulata 
Kirby.  The  most  primitive  species  are  N.  maculata  Fabr.  and  N. 
pallidula  Poisson  etc.,  which  have  short  ovipositors  and  merely 
attach  their  eggs  to  supports  in  the  water,  as  do  all  the  Paranecta 
known.  Moreover  the  last  abdominal  sternite  of  the  female  is  no 
more  developed  than  in  some  species  of  Paranecta.  This  subgenus 
is  dominant  in  Europe  and  North  Africa,  a  territory,  it  will  be  re- 
membered, in  which  Paranecta  is  absent.  From  southern  Eurcps  it 
spreads  eastward  to  India,  and  from  northern  Europe  we  find  a 
circumpolar  group  typified  by  N.  lutea  Miiller  in  northern  Europe, 
a  species  which  follows  the  mountains  southward  to  France,  and  I  y 
N.  borealis  Bueno  and  Hussey,  a  boreal  species  of  North  America.:/ 
The  subgenus  is  represented  also  by  a  species  in  Burma,  one  in 
China,  one  in  southwestern  Siberia,  and  by  N.  irrorata  Uhler  in 
eastern  North  America.  Thus  there  are  2  species  in  North  America, 
1  of  them  confined  to  the  Boreal;  3  confined  to  Eastern  Asia,  1  to 

t  It  was  supposed  that  those  had  been  roUectrd  by  Frau'-nWd  when  Entomologist  on 
the  Novara  which  touched  South  America  at  Rio  Janeiro,  Brazil,  and  ^'alparais^,  Chile. 
This  cannot  be  true,  however,  since  the  Novara  made  its  trip  around  the  world  in  1857-1859. 

t  We  must  bear  in  mind  also  A^.  nigra  Fieber  described  from  Brazil,  S.  A.  See  discussion 
under  this  species. 


Hungerford:    Genus  Notonecta  2] 

Arabia,  and  9  species  and  their  subspecies  which  are  found  in 
Europe,  the  Islands  adjacent  thereto,  and  in  North  Africa.  A''. 
maculata  Fabr.  ranges  east  to  India  and  A',  viridis  kashtniriaria 
Hungerford  represents  the  eastward  distribution  of  this  species. 

Summarizing  the  distribution  of  the  genus  Notonecta  Linnaeus  we 
find  it  well  represented  in  the  Western  Hemisphere,  in  Europe, 
North  Africa  and  Asia.§  It  appears  to  be  lacking  entirely  in  the 
East  Indian  Archipelago,  etc.,  where  it  is  replaced  by  the  genus 
Enithares.  The  genus  is  divided  into  five  subgenera:  Paraxecta, 
abundant  in  the  Western  Hemisphere,  is  represented  in  the  Eastern 
Hemisphere  by  a  few  species  in  the  Eastern  Palaearctic  and  one  in 
the  Ethiopian.  Bichromonecta  and  Erythronecta  are  confined 
to  the  region  between  Colombia,  S.  A.,  and  California  and  New 
Mexico.  U.  S.  A.  The  subgenus  Notonecta  is  common  in  Western 
Palsearctic.  has  a  few  species  in  Eastern  Palaearctic,  only  two  species 
in  Nearctic  and  possibly  one  in  the  Neotropical.*  Finally  the  sub- 
genus Enitharonecta,  which  is  represented  by  a  single  species  in 
the  West  Australian  realm. 

The  above  gives  us  a  picture  of  the  present  distribution  of  the 
subgenera  of  Notonecta.  Unfortunately  there  is  little  evidence  in 
the  fossil  remains  to  help  us  locate  the  centers  of  dispersal,  as  wull  be 
seen  in  the  following  chapter.  Most  of  our  fossils  are  from  the  Ter- 
tiary and  far  too  recent  and  too  few  to  assist  us.  The  one  from  the 
Jurassic  suggests  that  the  Notonectinae  and  the  Anisopinae  were  al- 
ready separated.  The  relation  of  the  various  genera  and  subgenera 
must  be  based,  therefore,  upon  the  morphology  of  present-day  forms 
and  what  we  may  conjecture  from  their  present  distribution. 

GEOLOGICAL  DISTRIBUTION 

Only  a  few  fossil  species  of  the  family  have  been  described,  and 
these  all  appear  to  belong  to  the  Anisopinae. 

The  oldest  fossil  is  described  under  the  name  Notonecta  elterleini 
Deichmiiller.^^  and  comes  from  the  lithographic  chalk  in  Bayern 
(Jurassic).  Kirkaldy^'^  says  it  must  belong  to  a  new  genus  and 
Handlirsch^'*  says  it  cannot  belong  to  Notonecta  S.  Str. 

All  the  other  fossils  are  from  the  Tertiary  and  all  but  one  from 
the  old  world.  Six  species  from  the  upper  Oligocene,  one  from 
lower  Miocene,  and  one  from  the  Florissant  in  Colorado  (Miocene). 

The  curiously  short  and  thickened  front  legs  of  Notonecta  jubata 

§  Records   are   lacking   over   vast    stretches    of   fliis    great    continent,    due    probably    to    the 
few  collections  that  have  been  made. 
*  A'',  nigra  Fieber. 

3—3482 


22 


The  University  Science  Bulletin 


Schlechtendal  and  apparently  also  of  N.  comata  Schlechtendal  and 
A'^.  deichmuellari  Schlechtendal  make  them  unlike  anything  living 
to-day.  While  Kirkaldy'*^  places  A',  hornacki  Schlechtendal  as  a 
true  Anisops,  there  is  no  indication  of  a  stridulatory  protuberance 
on  the  tibia  and  the  thickened  femur  suggests  that  the  specimen  is 
a  male. 

The  other  Europt.,."  -pccies  Kirkaldy  either  assigns  a  position 
near  Buenoa  or  fi-  undescribed  genera.  To  my  mind  all  of  them 
belong  in  the  second  category. 

Scudder's'^^  N.  emersoni  from  the  Florissant  is  also  impossible  to 
place.  If  his  interpretation  that  "the  median  forked  line  on  its 
posterior  portion  seems  to  indicate  the  hemelytral  suture  of  the 
upper  surface  seen  through  the  body"  is  true,  then  the  insect  is  a 
true  Anisopinae,  because  the  claval  orifice  is  present.  I  supposed  from 
the  drawing  this  was  the  midventral  keel  present  in  all  Notonectidaj. 
The  evidence  is  not  clear  that  any  of  the  fossils  belong  to  existing 
genera,  although  the  general  shape  is  that  of  Anisopinse. 


^ 
.<^^ 


X^ 


A^ 


i^ 


.X> 


'^^  ^^^ 


> 
1/ 


-r/\>^v- 


Text  Figure  1.     Diagram  illustrating  the  possible  phylogeny  of  the 
LT>iiera  and  subgenera   of  the  Notonectidae. 


Hungerford:    Genus  Notonecta  23 

TAXONOMY  OF  THE  NOTONECTA 

The  Genus  Notonecta  Linn^us 
(Logotype  A'',  glauca  Linnaeus) 

1758.  Notonecta  Linna?us ;  Systema  Naturs,  10th  Edn.,  p.  439.  (For  otlier  references  see 
Van  Duzee's  Catalogue  of  the  Hemiptera,  pp.   449-450.) 

1801.  Notonecta  Linnieus;  Lamarck,  Syst.  Nat.  Anim.  s.  Vert.,  p.  29G,  names  A',  glauca 
Linn,  as  type. 

Besides  the  diagnosis  given  in  the  key,  the  following  character- 
istics may  be  noted :  * 

The  head  shorter  than  the  pronotum.  Lateral  margin  of  pronotum 
compressed  dorsoventrally  into  an  edge.  Hemelytra  clothed  more  or 
less  with  short,  flattened  hairs.  Sides  of  thorax  densely  clothed  with 
rather  long,  flattened  hairs.  The  infracoxal  plates  covering  the 
bases  of  the  hind  coxae  not  longitudinally  carinate  but  densely 
clothed  with  long  hairs. 

Male  genital  capsule  with  or  without  a  ventral  digitifonn  process. 
Claspers  f  or  parameres  symmetrical. 

Key  to  Subgenera  of  Notonecta 

A.    Last  two  abdominal  sternites  of  male  normal  (not  long  and  slender).     Male  genital 
capsule  not  produced  caudally  on  its  ventral  line,  caudal  lobes  behind  rfaspers  not 
greatly  reduced.     Claspers  not  large  thin  plates. 
B.    Genital    capsule    of    male    without    digitiform    process.      The    last    abdominal 
stemite  of  female  not  enlarged. 
C.    Keel    of    fourth    abdominal    sternite    usually    not    bare.      Eyes    normal    in 
size,  or  if  broad  not  as  broad   as   length   of   the  lateral   margin   of   the 
pronotum.     Penultimate  abdominal  sternite  of  female  not  slender.      (See 
Plate  X.) 
D.    Anterior  trochanter  of  male  with  or  without  small  hook  and  with  no 
basal  protuberance.     Penultimate  abdominal  sternite  of  female  more 

or   less   rectangular Paranecta    (Hutchinson). 

DD.  Anterior  trochanter  of  male  with  large,  stout  hook  and  a  basal 
protuberance.  Penultimate  abdominal  sternite  of  female  triangular, 
its  tip  overlapping  base  of  last  segment.     (See  text  figure  7.) 

Bichromonecta  (Hungerford). 
CO.    Keel   of   fourth   abdominal   sternite   always   bare.      Eyes   unusually   broad, 
about  as  broad  or  broader  than  the  short  lateral  margin  of  the  pronotum. 
Penultimate  abdominal  sternite  of  female  slender.  .Erythronecta  (Hungerford). 
BB.    Genital    capsule    of    male    with    a    digitiform    process.      The    last    abdominal 
sternite  of  female  enlarged  and  usually  constricted  just  before  the  tip. 

Notonecta    (Linnaeus). 
AA.    Last  two  abdominal  sternites  of  male  long  and  slender.     Male  genital  capsule   pro- 
duced caudally  on  its  ventral   line,   caudal   lobes   behind   claspers   reduced.      Claspers 
large  thin  plates.     Known  only  from  Australia Enitharonecta  (Hungerford). 

*  Mr.  Hutchinson  "  adds  statements  regarding  the  venation  of  the  hind  wings ;  they  are 
not  correct  for  the  genus.  Using  his  nomenclature  Notonecta  lobata  Hungerford  has  M-Cu 
strongly  "chitinized"  and  many  species  show  Cui  and  Cuo  not  only  "chitinized"  at  base,  but 
complete  throughout  and  united  distally  as  in  his  figure  of  the  wing  of  Enithares  sohrina  Stal. 
(Plate  XXVII,  fig.  5.) 

t  Snodgrass  proposes  the  name  harpagones  for  these. 


24  The  University  Science  Bulletin 

Subgenus  Paeanecta  Hutchinson  1929 
(Type  species  Notonecta  lactitans  Kirkaldy) 

Hutchinson,  G.  E.,  Annals  South  African  Museum,  Vol.  XXV,  Pt.  3,  p.  363. 

Notonecta  in  which  the  male  genital  capsule  lacks  a  digitiform 
process  and  in  which  the  females  have  the  first  pair  of  gonapophyses 
short  and  weak.  The  females  have  the  penultimate  abdominal 
sternite  rectagular.  The  last  abdominal  sternite  of  female  is  not 
large  and  is  usually  incised  at  tip  (see  Plate  X,  fig.  3).  The  diploid 
number  of  chromosomes  for  those  known  is  26. 

Subgenus  Bichromonecta  Hungerford  1934 
(Type  species  Notonecta  shooteri  Uhler) 

Hungerford,  H.  B.,  Jl.  Kans.  Ento.  Soc,  Vol.   VII,  No.  3,  p.   98. 

Notonecta  of  compact  form  wdth  two  color  phases,  one  luteous  and 
the  other  pigmented.  The  luteous  form  has  reduced  flight  wing, 
reduced  posterior  lobe  of  membrane,  smaller  scutellum  and  less 
prominent  humeri,  resulting  in  more  nearly  parallel  lateral  margins  of 
pronotum.  The  pigmented  form  has  normal  wings  and  more  promi- 
nent humeri.  Eyes  are  flattened  and  anterior  margins  oblique;  syn- 
thlipsis  relatively  broad.  Anterolateral  angles  of  pronotum  embrac- 
ing the  eyes  and  lateral  margins  straight  to  convex.  The  females 
have  the  penultimate  abdominal  sternite  triangular,  its  tip  notched 
and  overlapping  the  base  of  the  last  segment  (see  text  figures,  p. 
101).  The  males  have  a  protuberance  at  the  angle  and  a  broad 
stout  hook  on  the  anterior  trochanter  (see  Plate  IX,  fig.  4) .  The 
"internal  stay"  of  the  sedeagus  is  recurved  at  tip. 

Subgenus  Erythronecta  Hungerford  1934 
(Type  species  Notonecta  lobata  Hungerford) 

Hungerford,  H.  B.,  Jl.  Kans.  Ento,  Soc,  Vol.  VII,  No.   3,  p.   98. 

Notonecta  of  compact  form,  usually  red  and  black  in  color.  The 
genital  capsule  of  the  male  lacks  a  digitiform  process  and  the  "in- 
ternal stay"  of  sedeagus  is  heavily  sclerotized.  The  first  pair  of 
gonapophyses  of  the  female  is  short.  The  so-called  fourth  abdomi- 
nal sternite  is  broad  and  short  in  both  sexes  and  the  keel  is  bare. 
The  penultimate  abdominal  sternite  of  the  female  is  very  slender 
and  the  last  one  is  narrow  and  usually  emarginate  at  tip  (see  Plate 
X,  fig.  1). 


Huxgerford:    Genus  Notonecta  25 

Subgenus  Notonecta  Hutchinson  1929 
(Type  species  Notonecta  glauca  Linn.) 

Hutchinson,  G.  E.,  Annals  South  African  Museum,  \o\.  XX\',  p.   303. 

Notonecta  in  which  the  male  genital  capsule  is  provided  with  a 
digitiform  process  and  in  which  the  females  have  the  first  pair  of 
gonapophyses  well  developed.  The  last  abdominal  sternite  of 
female  is  large  and  usually  constricted  just  before  the  tip,  which  is 
not  incised.  The  diploid  number  of  chromosomes  for  those  known 
is  24  (see  Plate  X,  fig.  4.) 

Subgenus  Enitharonecta  Hungerford  1928 
(Type  species  Notonecta  handlirschi  Kivkaldy.) 

Hungt'rford,  H.  B.,  Annals  Ento.  See.  America  XXI,  p.   143. 

Resembles  other  Notonecta  in  general  appearance,  but  decidedly 
unlike  them  in  shape  of  last  two  abdominal  sternites  of  male,  which 
arc  long  and  slender,  and  in  the  shape  of  the  genital  capsule  of  the 
male,  which  is  produced  cadually  on  its  ventral  line.  The  claspers 
or  parameres  are  large,  thin  plates  convex  without  and  concave 
within.  The  caudal  lobes  of  the  capsule  are  greatly  reduced.  (See 
PlateVIII,  fig.  4.) 

NOTONECTA  OF  EASTERN  HEMISPHERE 
Key  to  Notonecta  of  Eastern  Hemisphere 

1.  Small  species,  not  to  exceed  12  mm.  long  or  4  mm.  across  humeri* (.2) 

Species  larger  than  above (3) 

2.  Species  plump.      Middle   femur  with   distinct  spiniferous   tubercle   near   base   of   rear 

margin.     Last  two  abdominal  sternites   of  male  slender A'',   handlirschi. 

(Aastralia,  p.  27.) 

Species  slender,   at   least   three  times  as  long  as  broad.      Middle   femur  without   the 

distinct    tubercle    mentioned     above.       Last    two    abdominal     sternites    of     n^ale 

normal   .' -V-  lactilans. 

(S.  Africa,  p.  28.) 

3.  Male    genitale    capsule    without    digitate    prolongation;    last    abdominal    sternite    of 

female  not  large  and  constricted  near  the  tip.     (See  Plate  X,  fig.  3) (.i) 

Male  genital   capsule  with   digitate   prolongation;    females   of  most   species   with    last 

abdominal  sternite  large  and  constricted  near  the  tip.     (See  Plate  X,  fig.   4)....      (9) 

4.  Mesotrochanter  right   angulate -V.    kirkaldni. 

(Chma,  p.  30.) 
Mesotrochanter  with  angle  produced (5) 

5.  Typical  color,  red  and  black (6) 

Typical  color  flavous  and  black (7) 

6.  Hair  on  ventral  side  of  body  silvery  gray  to  light  brnwn.     Lateral  margin  of  prono- 

tum    not     sharp-edg.d.       Anterior     margin     of     vertex,     seen     from    above,     not 

flattentKl A',  chinensis. 

(China,  p.  31.) 
Hair    on    ventral    side    of    body    black.      Lateral    margin    of    pronotum    sharp-edged. 

Anterior  margin  of  \ertex,  seen  from  above,  flattened A',  saramoa. 

(Formosa,  p.  32.) 

*  Poisson  described  his  A^.  paUidula  as  11.5  mm.  to  12  mm.  long;  all  specimens  I  have 
seen  are  at  least  broader  than  4  mm.  and  most  of  them  longer  than  12  mm. 


26  The  University  Science  Bulletin 

7.    Mesotrochanter  produced,  but  not  into  a  long,  sharp,   thornlike   process (8) 

Mesotrochanter  produced  into  long,  sharp,  thomlike  process.     (See  Plate  IX.) 

A^.   kiangsis. 
(China,  p.  33.) 

S.    Connexivum  black  beneath A^.   immediata. 

(E.  Siberia,  p.  35.) 

Connexivum  green  to  flavous A',   triguttata. 

(Japan,  p.  36.) 

9.    Mesotrochanter    right    angulate.      General    facies    reddish-orange,    unevenly    marked 

with  black    A^.   montandoni. 

(China,  p.  37.) 
Mesotrochanter  rounded    (10) 

10.  Last  abdominal  sternite  of  female  not  constricted  near  the  tip.     (See  Plate  X,  fig.  3),   (11) 
Last  abdominal  sternite  of  female  constricted  near  the  tip   (see  Plate  X,   fig.    4)   or 

large  and  depressed  on  sides  before  the  tip (14) 

11.  Scutellum  pale,  sordid  brown A^.   pallidula. 

(N.  Africa,  p.  38.) 
Scutellum   black (12) 

12.  Anteapical  tooth  of  mesofemur  of  normal  size.     (See  Plate  VI,   fig.   5.)     Metanotum 

and  first  abdominal  tergite  orange  or  yellow (13) 

Anteapical  tooth  of  mesofemur  unusually  long.     Metanotum  black.  ..  .A',   cavariensis. 

(Canary  Islands,  p.  40.) 

13.  Ant.  rolateral  angles   of  pronotum  slightly  produced A^.   maculata. 

(Western  and  Southern  Europe,  North  Africa,  and  eastward  to  India,  p.  41.) 

Anteri .lateral  angles  of  pronotum  obtuse A',    meinertzhagem. 

(Hoggar  Mountains  in  the  Sahara,  p.  44.) 

14.  Last  abdominal  st«rnite  of  female  large  and  slightly   depressed   on   the  sides   before 

the  tip TV.  arabievsis. 

(Arabia,  p.  44.) 
Last  abdominal  sternite  of  female  strongly  depressed  on  the  sides  before  the  tip   or 

strongly   constricted   before   the   tip (15) 

15.  Anterior  angles  of  pronotum   not   acute   or   closely   embracing   the   eyes.      (See   Plate 

VIII,   fig.    8) (16) 

Anterior    angles    of    pronotum    acute    and    closely    embracing    the    eyes.      (See    Plate 

XVII,  fig.  4) (26) 

16.  Scutellum   tjpically   entirely   black (17) 

Scutellum  not   entirely  black (25) 

17.  Large  reddish  species,  more  than  6  mm.  across  humeri A^.  violacea. 

(Burma,  p.  45.) 
Species  not  surpassing  5.4  mm.  across  humeri (18) 

18.  Hemelytra  typically  yellowish  or  flavous,  lacking  oblique  stripes (19) 

Hemelytra  reddish  or  brownish,  maculated  or  not (20) 

19.  Hemelytra    typically    yellowish    or    flavous,    with    brownish    or    blackish    maculations 

(sometimes  confluent)  along  costal  margin,  and  often  a  transverse  spot  at  apex 

of  clavus  and  internal  angle  of  corium A^.  glauca  glauca. 

(Northern  Europe  and  southward  to  Central  France,  p.  46.) 

Hemelytra  yellowish  or  flavous  with  a  broad  transverse  black  band  covering  distal 

third  of  corium,  tip  of  clavus  and  basal  portion  of  membrane.  .A'^.  glauca  poissoni. 

(Asia  Minor,  p.  49.) 

20.  Hemelytra  reddish  or  brownish,  maculated  or  not (21) 

Hemelytra  largely  velvety  black  with  one  or  two  flavous  stripes  at  the  base (22) 

21.  Hemelytra  more  or  less  strongly  maculated,  presenting  at  the  base  either  one  pale 

longitudinal  stripe  on  the  clavus,  or  two  such  stripes,  one  on  the  clavus  and  the 

other  on  the  corium N.  glauca  hybrida. 

(Southern  France  to  North  Africa,  p.  50.) 
Hemelytra  generally  clear  reddish,  sometimes  with  some  vague  markings  of  dull  black 
in  the  marginal  canal  and  some  drop -shaped  spots  on  the  disc. 

A'^.  glauca  rufescens*    (N.  glauca  fulva?) 
(Southern  Europe,  N.  Africa,  p.  51.) 

*  Doctor  Poisson  mentions  a  new  subsoecies.     A',  glauca  kirvillei,  from  Asia  Minor,  which 
he  says  is  related  to  this. 


Hungerford:    Genus  Notonecta  27 

22.  Size  usually   15  to   16  mm (23) 

Size  usually  13  to  14  mm.     (If  larger  with  only  one  pale  stripe  on  hemelytron) (24) 

23.  Hemelytra  with  two  very  clear  flavous  stripes  at  the  base,  the  one  on  corium  a  little 

shorter  than  the  oni?  on  the  clavus N.  obliqua  obliqua. 

(Northern  Europe  to  Central  France,  p.  51.) 
Hemelytra   as   above,   plus   one   large   flavous   spot    beyond    distal    end    of   the   stripe 

on  corium    N.   obliqua  delcourti* 

(Belgium  and  Northwestern  France,  p.  54.) 

24.  Hemelytra  with  two  flavous  stripes,   but  the  one  on  corium   more  or  less   restricted 

or  only  the  claval  stripe  present,   and   perhaps  some  yellowish   spots  in   place   of 

the  corial  stripe    A^.  obliqua  meridionalis. 

(About  the  Mediterranean,  p.  54.) 

25.  Size   very   large:     17    mm.    to    18.3    mm.    long.      Last   abdominal    sternite   of    female 

greatly  and  abruptly  constricted  before  the  tip N.  amplifica. 

(E.  Siberia,  p.  55.) 
Size  seldom  over  15  mm.  long.     Last  abdominal  sternite  of  female  only  moderately 

and  not  abruptly  constricted  before  the  tip N.  lutea. 

(N.  Europe,  p.  56.) 

26.  Scutellum   seldom    entirely    black,    sometimes    entirely    pale.      Anterior   trochanter    of 

male  with   a   hook (27) 

Scutellum  solid  black.     Anterior  trochanter  of  male  with  slight  elevation (28) 

27.  Synthlipsis  usually  not  over  half  the  width  of  the  eye.     Last  abdominal  sternite  of 

female  only  moderately  and  not  abruptly  constricted  before  the  tip.  Male 
clasper  as  on  Plato  XVII,  fig.  2 N.  lutea. 

(N.  Europe,  p.  56.) 
Synthlipsis    over    half   the    width    of    the   eye.      Last    abdominal    sternite    of    female 

abruptly  constricted  before  the  tip.     Male  clasper  as  on  Plate  XVII,  fig.  1.  .A^.  reuteri. 

(N.  Europe,  p.  59.) 

28.  Hemelytra   flavous   or   greenish    flavous,    costal    margin    often   maculated    and   a    dark 

X-shaped  figure  at  tip  of  henielytral  commissure N.  viridis  viridis. 

(Western  Emope  from  France  southward,  north  Africa,  p.  60.) 
Hemelytra  largely  maculated (29) 

29.  Anterior  margin   of   head   as  seen   from   above   slightly   convex.      Genital   claspers   of 

male  like  above A',  viridis  mediterranea. 

(Mediterranean  region  and  eastward,  p.  63.) 
Anterior  margin  of  head  as  seen  from  above  flattened.     Genital  claspers  of  male  as 

shown  on  Plate  XVII,  fig.  5 N.  viridis  kashmiriana. 

(Kashmir,  p.  65.) 

Notonecta  handlirschi  Kirkaldy  1897 

(Color  Plate  IV,  fig.  2,  Plates  VIII,  fig.  4,  and  IX,  fig.  7.) 

1897.    A'',  handlirschi  Kirkaldy,  Trans.  Ento.  Soc.  London,  p.  408. 

1904.    A'^.  handlirschi  Kirkaldy,  Wien.  Ento.  Zeit.  xxiii,  p.   132. 

1923.    N.  handlirschi  Kirkaldy;  Hale,  Rec.  S.  Aust.  Mus.,  Vol.  II,  No.  3,  June  30,  p.  418. 

1928.  A^.  handlirschi  Kirkaldy;  Hungerford,  Annals  Ento.  Soc.  Amer.  xxi,  p.  143,  PI.  ix, 
fig.  6.     (Fig.   genital  capsule.)     (New  subgenus  Enitharonecta.) 

1929.  N.  handlirschi  Kirkaldy;  Hutchinson,  Annals  of  the  South  African  Museum,  Vol. 
XXV,  Part  3,  p.   363. 

Size.  Length  10  mm.  to  10.5  mm.;  width  of  pronotum  3.5  mm.  to 
4  mm. 

Color.  As  shown  on  Color  Plate  IV,  fig.  2.  A  species  with  reddish 
hemelytra  sometimes  nearly  black,  with  clavus  dark  reddish  brown. 
Metanotum  and  dorsum  of  abdomen  nifotestaceous.    Venter  black. 

Structural  Characteristics.    Anterior  outline  of  head  viewed  from 

*  A  variety,  not  a  subspecies. 


28  The  University  Science  Bulletin 

above  flattened;  vertex  slightly  shorter  than  its  anterior  width; 
margin  of  A'ertex  less  convex  than  the  margin  of  the  eye;  anterior 
breadth  of  vertex :  synthlipsis :: 22: 13.*  Pronotum  twice  the  length 
of  the  head;  lateral  margins  straight  and  moderately  divergent; 
anterolateral  angles  produced  forward  beyond  the  caudolateral  mar- 
gins of  the  eyes;  lateral  ledge  as  seen  from  the  side  shghtly  curved, 
oblique  and  about  as  Iqng  as  rear  margin  of  the  eye  below  it. 
Scutellum  longer  than  the  pronotum.  Hemelytral  commissure 
shorter  than  pronotum;  posterior  lobe  of  membrane  well  developed. 
Anterior  trochanter  of  the  male  without  hook  or  elevation.  Meso- 
trochanter  rounded  or  faintly  angulate;  mesofemur  with  distinct 
spiniferous  tubercle  on  base  of  caudal  margin;  the  distal  meso- 
femoral  tooth  not  prominent.  The  last  two  abdominal  sternites  long 
and  slender  in  the  male;  the  lobe  of  ninth  tergite  slender  and 
pointed.  The  male  genital  capsule  as  shown  on  Plate  VIII,  fig.  4. 
The  claspers  are  thin  plates,  convex  without  and  concave  within; 
as  one  views  the  clasper  from  the  rear  there  is  a  recurved,  short  sub- 
marginal  fold  parallel  with  the  upper  margin;  a  much  lesser  thick- 
ening occurs  before  the  tip  below  the  upper  margin;  neither  of  these 
is  visible  from  the  lateral  view. 

Location  oj  Type.  Two  males  in  the  Vienna  Museum.  They  bear 
the  label  "Fischer,  Austr.  Post  1,  1878."  One  of  them  now  (1928) 
has  the  abdomen  damaged  by  dermestids. 

Comparative  Notes.  This  species,  the  only  one  in  the  genus  re- 
ported for  Australia,  appears  to  stand  quite  alone.  For  this  reason 
I  have  placed  it  in  a  subgenus  by  itself. 

Data  on  Distribution.  There  is  a  male  specimen  in  the  British 
Museum  labeled  "Western  Australia,  1922-1923,  B.  R.  Lucas."  In 
this  specimen  the  scutellum  is  black  and  the  hemelytra  nearly  black, 
with  the  clavus  dark  reddish  brown. 

Notonecta  lactitans  Kirkaldy  1897 

(Color  Plates  IV,  fig.  3  and  V,  fig.  12;   Plates  Vlll,  fig.  9  and  IX,  fig.  S.) 

1897.    N.  lactitans  Kirkaldy,  Ann.   Mag.  Nat.  Hist.,  (6)  xx,  p.   58. 

1897.    N.  lactitans  Kirkaldy,  Trans.  Ento.   See.   London,  p.   405. 

1904.    N.  lactitans  Kirkaldy,  Wien.   Ento.  Zeit.,  xxiii,   p.    132. 

1925.    A',  lactitans  Kirkaldy;    Hungerford,  Annals  Ento.   Soc.  Am.,  xviii,  ]i.   417. 

1929.  N.  lactitans  Kirk.;  Hutchinson,  .A.nnals  of  South  African  Museum,  xxv,  Pt.  3,  pp.3C3 
and  365-368.  (Type  of  Paranecta  subg.  n.)  PI.  xxvii,  figs.  1,  2  and  3  (Head  and  pronotum, 
mesofemur  and  hind  wing);  PI.  xxviii,  figs.  1,  2  and  3  (Genital  capsule,  aedeagus  and  clasper); 
PI.  xxix,  figs.   10  and  11  (Head  and  pronotum  of   ?  3rd  and  5th  instars). 

*  Kirkaldy  says:  "Vertex  not  quite  as  wide  as  synthlipsis."  This  is  not  true  in  the  types 
which  I  have  examined  on  two  occasions. 


Hungerford:    Genus  Notonecta  29 

Referring  to  this  species,  also: 

1897.  A'',  lactitans  var.  sfygica  Kirkaldy,  Trans.  Eiito.  &oc.  I>ondoii,  p.  406.  This  is  the 
male  of  A'^.  lactitans  Kirk. 

1904.  jV.  lactitans  var.  sti/f/ica  Kirkaldy.  Wifii.  Ento.  Zeit..  xxiii,  p.  132.  (Name  listed 
only.) 

Size.  Length  10.5  mm.  to  12  mm.  The  females  longer  than  the 
males.  A  slender  species  that  is  at  least  three  times  as  long  as 
broad. 

Color.  A  yellowish  gray  and  black  species  as  shown  on  Color 
Plates  IV  and  V.  The  males  are  darker  than  the  females.  Scutellum 
is  black.  Scutellmn  and  hemelytra  covered  sparcely  with  short,  fine, 
golden  pubescence.    Abdominal  dorsum  and  venter  dark. 

Structural  Characteristics.  Head  small,  notocephalon  very  faintly 
longitudinally  carinate  on  posterior  half.  Vertex  broad  at  base. 
Anterior  breadth  of  vertex  :  synthlipsis  : :  5.6:3.5.  Pronotum  twice  as 
long  as  the  head,  anterior  angles  obtuse,  lateral  margins  slightly 
divergent  and  sinuate;  lateral  marginal  ledge  nearly  straight  termi- 
nating caudally  below  the  humeral  elevation.  Scutellum  slightly 
longer  than  the  pronotum.  Outer  lobe  of  membrane  very  slightly 
longer  than  inner.  Anterior  trochanters  of  male  without  hook. 
Mesotrochanters  slender,  rounded,  a  small  tuft  of  long  hairs  at  the 
base.  Mesofemur  with  tuft  of  bristle-like  hairs  on  basal  angle  of 
rear  margin.  The  subapical  tooth  directed  slightly  toward  apex. 
Small  tuft  of  hairs  on  median  line  of  the  short  third  ventral  abdo- 
minal segment,  fourth  ventral  abdominal  segment  longest,  terminal 
one  in  female  deeply  notched.  Male  genital  capsule  as  shown  in 
Plate  VIII,  fig.  9.  •  First  pair  of  gonapophyses  of  female  short. 

Location  of  Types.  Kirkaldy  recorded  the  type  as  being  in  his 
own  collection  and  listed:  "Africa,  Guinea  (my  coll.);  Gaboon 
(Vienna  Mus.) ;  Cape  of  Good  Hope  (Paris  Mus.)".  The  type,  a 
female,  broken  and  dismembered  is  in  the  U.  S.  N.  M.  with  the 
Kirkaldy  collection.  In  the  Vienna  Museum  there  is  a  female 
labeled  cotype  and  "Gaboon  Coll.  Signoret."  Thorax  on  the  left 
side  has  been  eaten  by  dermestids  (1928).  In  the  Paris  Museum 
there  is  also  a  female  bearing  a  red  label  "Comp.  with  type  by 
Kirkaldy  in  Kirkaldy  Coll."  This  is  the  specimen  mentioned  by 
Kirkaldy  as  from  Cape  of  Good  Hope.  It  carries  the  label  "Museum 
Paris.  Cap  de  Bonne  Esperance.  Delalande  1820."  The  type  of 
V.  lactitans  var.  stygica  Kirk,  is  in  the  British  Museum.  It  is  only 
the  dark  male  of  A^.  lactitans  Kirkaldy. 

Comparative  Notes.  The  most  slender  Notonecta  known  and  the 
only  representative  of  the  Genus  in  Africa  south  of  the  Sahara. 


30  The  University  Science  Bulletin 

The  males  are  usually  darker  than  the  females,  the  hemelytra  being 
blackish  with  the  outer  margin  of  the  embolium,  a  streak  on  outer 
edge  of  clavus  and  a  spot  on  the  inner  part  of  the  posterior  margin 
of  the  corium  dull  yellow.  As  Mr.  Hutchinson  has  pointed  out,  this 
species  is  quite  distinct,  but  more  nearly  related  to  the  Western 
Hemisphere  forms  than  to  the  European.  It  is  the  type  of  Hutchin- 
son's subgenus  Paranecta. 

Data  on  Distribution.  Besides  the  type  localities  listed  by  Kirk- 
aldy,  Mr.  Hutchinson  has  recorded  some  thirteen  collections  in  the 
Cape  and  believes  that:  "In  the  Cape  Peninsula  there  is  a  migration 
from  mountain  pools  in  the  autumn  to  low-lying  waters  on  the  Cape 
Flats  and  elsewhere  in  which  the  insects  breed,  the  newly  emerged 
adults  then  ascending  and  populating  the  localities  left  by  their 
parents."  I  have  specimens  kindly  sent  to  me  by  F.  W.  Pettey  from 
Elsenbourg,  S.  Africa,  and  from  Muilder's  Vlei,  Cap.  Prov.  There 
are  four  specimens  in  Dr.  Herbert  Osborn's  collection  "Vlei  Su  Pr. 
Common  June  6,  1909." 

Character  of  Habitat.  The  specimens  sent  to  me  by  Mr.  Pettey 
are  from  a  clear  mountain  stream  pool.  Hutchinson  records  collec- 
tions from  reservoir,  bog,  pond,  weedy  pool  and  pools  in  streams. 
The  elevations  range  from  low  fiats  to  a  pool  near  the  top  of  a  pass 
in  the  mountains.    The  highest  elevation  he  gives  is  4,200  feet. 

Life  Histonj  Notes.  Hutchinson  says  the  eggs  are  unknown,  but 
presumably  are  laid  on  submerged  objects  without  being  imbedded. 
He  describes  what  he  believes  to  be  the  third,  fourth  and  fifth 
instars. 

Notonecta  kirkaldyi  Martin  1902 

(Color  Plate  V,  fig.   9;    Plate  XV,  fig.  3.) 

1902.  A',  kirkaldyi  Martin;  Bull.  Mus.  d'Hist.  Nat.  (Paris),  viii,  p.   336. 

1904.  iV.  kirkaldyi  Martin;  Kirkaldy,  Wien.  Ento.  Zeit.,  xxiii,  pp.  95  and  132. 

1925.  A'',  kirkaldyi  Martin;  Esaki,  Notula  Entoniologicas,  v.  p.   14. 

1925.  A',  kirkaldyi  Martin;  Hungerford,  Annals.  Ento.  Soc.  Am.,  xviii,  p.   418. 

Size.    Length,  13  mm.;  width  across  thorax,  4.6  mm. 

Color.  A  reddish  orange  and  black  species.  Head  and  limbs  of 
usual  testaceous  color.  Scutellum  black.  Hemelytra  reddish  orange 
and  black;  the  distal  end  of  clavus,  a  stripe  on  corium  bordering 
clavus  and  a  broad  irregular  band  of  black  crossing  distal  half  of 
corium;  the  membrane  is  dark  brown  to  black. 

Structural  characteristics.  Anterior  breadth  of  vertex  :  synthlip- 
sis  ::  7:3;  head  appearing  fairly  long;  its  length  to  that  of  prono- 
tum  ::  2.5:4.2.    Lateral  margins  of  pronotum  diverging  and  nearly 


Hungerford:    Genus  Notonecta  31 

straight;  anterior  angles  normal;  lateral  ledge,  seen  from  the  side, 
straight  with  rear  end  turned  upward  beneath  the  humeral  eleva- 
tion. The  scutellum  a  little  longer  than  pronotum  (nearly  5  to  4). 
Posterior  lobe  of  membrane  slightly  longer  than  anterior.  Anterior 
trochanter  of  the  male  without  hook  or  tooth.  IMesotrochanter 
right  angulate.  Mesotibia  with  slight  prominence  on  rear  margin. 
Female  not  examined  by  me.  Male  genital  capsule  as  shown  on 
Plate  XV,  fig.  3. 

Location  of  Type.  Paris  Museum,  ootypes  in  U.  S.  N.  M.  and  in 
my  collection. 

Comparative  Notes.  A  smaller  species  than  A-",  chinensis  Fallou 
and  with  the  trochanter  of  middle  leg  right  angulate  instead  of 
acutely  angulate. 

Data  on  Distribution.  Described  from  "Yun-Nan-Nansen  (Mgr. 
Excofiier)  R.  Oberthur.  1898."  There  were  31  specimens  in  the  type 
series  in  the  Paris  Museum,  all  labeled  as  above. 

Notonecta  chinensis  Fallou  1887 

(Color  Plate  V,  fig.   8;    Plate  XV,  fig.   8.) 

1887.  iV.   chinensis  Fallou;    Le  Naturaliste  (Deyrolle  Paris),   viii,   p.   413. 

1894.  .V.  chinensis  Fallou;  Bergroth,  Rev.  Ento.  franc,  xiii,  p.  164.  Redescribed  at 
length.) 

1897.  A",  chinensis  Fallou;   Kirkaldy,  Trans.  Ento.  Soc.  London,  1897,  pp.  415,  416. 

1904.  N.   chinensis  Fallou;    Kirkaldy,  Wien.   Ento.  Zeit.,  xxiii,  pp.   95  and   132. 

1912.  A',   chineyxsis  Fallou ;    Oshanin,   Katalog  der  palaarktischen  Hemipteren,  p.   91. 

1925.  N.   chinensis  Fallou ;    Esaki,   Entomolog.    Mitteilungen,   xiv,    1925,   Nr.    5/6,   p.    312. 

1925.  A^.   chinensis  Fallou;    Hungerford,  Annals  Ento.  Soc.   Am.,  xviii,  p.   418. 

1930.  A^.   chinensis  Fallou;    Hungerford,  Bull.   Brook.  Ento.   Soc,  xxv..  No.   3,  p.   138. 

1930.  N.  chinensis  Fallou;  Kiritshenko,  Annuaire  du  Musee  Zoologique  de  I'Acad^mie  des 
Sciences  de  I'll.  R.  S.  S.,  p.  434. 

Referring  to  this  species,  also : 

1873.    N.  sinica  Walker,  Cat.  Hem.-Het.   Brit.   Mus.,  viii,  p.   204.     (Kirkaldy.) 

Size.  Length,  13-14  mm.;  width  across  thorax,  4.5  mm.  to  5.1 
mm.,  the  females  averaging  a  little  larger  than  the  males. 

Color.  An  orange-red  and  black  species.  Head,  pronotum  and 
limbs  light  testaceous.  Face,  limbs  and  connexivum  beneath  may 
be  greenish.  Scutellum  black.  Hemelytra  orange-red  to  red-brown 
with  an  undulating  bluish-black  band  (occasionally  represented  by 
scattered  spots)  extending  from  suture  to  lateral  margin,  near  the 
apical  margin  of  the  corium,  but  diverging  from  that  as  it  approaches 
the  lateral  margin;  membrane  bluish-black. 

Structural  Characteristics.  Eyes  close  together  at  synthlipsis; 
anterior  breadth  of  vertex  :  synthlipsis  :  :  26  :  7;  head  about  five- 


32  The  University  Science  Bulletin 

eights  as  long  as  pronotum.  Lateral  margins  of  pronotum  divergent 
and  sinuate;  anterior  angles  somewhat  produced;  lateral  ledge  as 
seen  from  the  side  slightly  sigmoid.  Scutellum  a  little  longer  than 
pronotum.  Lobes  of  membrane  about  equal  in  length.  Anterior 
trochanter  of  male  with  a  median  raised  dentation  in  place  of  hook; 
mesotrochanter  acutely  angulate,  the  angle  produced.  Terminal 
abdominal  sternite  of  female  with  lateral  margins  concave  and 
apex  shallowly  notched.  Female  gonapophyses  of  moderate  length. 
Male  genital  capsule  as  shown  on  Plate  XV,  fig.  8. 

Location  of  Type.  Fallou  collection  in  Paris  Museum.  The  male 
type  is  from  Fo-Kien. 

Comparative  Notes.  This  species  is  readily  separated  from  other 
oriental  Notonecta  (except  N.  kiangsis  Kirk,  and  N.  saramoa  Esaki) 
by  the  very  narrow  interocular  space  (synthlipsis).  N.  kiangsis 
Kirk  is  very  different  in  color,  more  slender  in  form,  and  the  angle 
of  the  mesotrochanter  is  produced  into  a  long,  sharp  thorn-like 
process. 

Data  on  Distribution.  Kirkaldy  says,  "Well  distributed  over  the 
Chinese  Empire"  and  gives  the  following:  "Pekin,  Kian-Si,  Se- 
Tchouen,  Chen-Si  (Paris  Mus.)  Kin  Kiang  (?  Chin-Kiang) ,  Foo- 
Chan  (Brit.  Mus.)  Fo.-Kien  (Fallou  and  Bergroth  Co.).  Ngan 
Hoei,  (Montandon  and  my  colls.)." 

The  following  additions  I  have  noted:  "Prov.  Fo  Kien  (China) 
G.  Siemssen  vend.  31-V-1904  (Hamburg  Mus.)";  ''Chin  Kiang, 
China  V,  20,  1924,  E.  Suenson"  (my  coll.) ;  Peking,  China.  Oct., 
1925,  P.  W.  Claassen,  (my  coll.  and  Cornell)  ;  Hweisin,  Kansu, 
China  (my  coll.). 

Notonecta  saramoa  Esaki  1933 

(Plate  XV,  figure  7) 
1933.    N.  saramoa  Esaki;   Trans.  Nat.  Hist.  Soc.  Formosa,  Vol.  XXII,  pp.  493-495. 

Size.  Length,  13.5  mm.  to  14  mm.;  width  across  thorax,  4.5  mm. 
to  5  mm. 

Color.  An  orange-red  and  black  species  resembling  very  closely 
A",  chinensis  Fallou.  The  broad,  black  band  commonly  present 
across  the  end  of  the  clavus  and  corium  in  N.  chinensis  Fallou  is  re- 
placed by  three  angular  spots  in  the  four  paratypes  before  me. 
However,  the  species  from  China  sometimes  has  identically  these 
markings.    Hair  on  ventral  side  of  body  black. 

Structural  Characteristics.  Eyes  close  together  at  synthlipsis; 
anterior  margin  of  vertex,  seen  from  above  flattened ;  anterior  breadth 
of  vertex  :  synthlipsis  : :  29:9.  Head  about  half  as  long  as  pronotum; 


Hungerford:    Genus  Notonecta  33 

lateral  margins  of  pronotum  divergent,  slightly  sinuate  in  the  male, 
more  so  in  the  female;  anterior  angles  somewhat  produced;  lateral 
ledge  as  seen  from  the  side  thin  and  sigmoid.  Scutellum  a  little 
longer  than  pronotum.  Lobes  of  membrane  about  equal  in  length. 
Anterior  trochanter  of  male  with  a  median  raised  dentation  in  place 
of  a  hook;  mesotrochanter  acutely  angulate,  the  angle  produced. 
Terminal  abdominal  sternite  of  female  with  lateral  margins  concave 
and  apex  faintly  incised.  Male  genital  capsule  as  shown  on  Plate 
XV,  iig.7. 

Location  of  Type.  In  collection  of  Professor  Teiso  Esaki.  Four 
paratypes  in  the  Francis  Huntington  Snow  Entomological  Museum. 
These  paratypes  are  labeled:  "Formosa,"  "Meoto-ike  (Taichu-shu) 
near  Kunugigaoka,  8,000  ft.,  17  -  VII,  1932,  Teiso  Esaki."  and  were 
collected,  in  a  small  pond  with  the  holotype  in  central  Formosa. 
Other  paratypes  were  collected  in  Mururoafu-no-ike,  a  larger  pond 
7,500  ft.  in  Taiheizan  district,  Northern  Formosa,  on  July  23,  1932, 
by  Professor  Esaki.  Paratypes  will  also  be  found  in  the  collections 
of  the  Government  Research  Institute  of  Fomiosa,  Taihoku  and  in 
the  British  Museum. 

Comparative  Notes.  This  species  is  near  A',  chinensis  Fallou,  from 
which  it  is  distinguished  by  its  more  truncate  head,  by  the  thin 
lateral  edge  of  pronotum,  by  the  black  hair  on  the  ventral  side  of  the 
body,  by  shape  of  genital  capsule  and  mesotrochanter. 

Data  on  Distribution.  Known  only  by  the  type  series  from 
Formosa. 

Notonecta  kiangsis  Kirkaldy  1897 

(Color  Plate  V,  fig.   7;   Plate  XV,  fig.   5.) 

1897.    .V.  chineiisis  Fallou  var.  kiangsis  Kirkaldy,  Trans.  Ento.  Soc.  London,  1897,  p.   416. 
1904.    X.   chinensis   Fallou   var   kiangds   Kirkaldy,    Wien.    Ento.    Zeit.,    .\xiii,    p.    132    (lists 
name  onlj). 

1925.    .V.  kiangsis  Kirkaldy;   Esaki,  Notul.  Entom.,  Helsingfors  v,  p.   15. 

1929.  .V.   kiangsis  Kirkaldy;    Hutchinson,  Annals  S.   African  Museum,  xxv,  pt.   3,   p.    3G3. 

1930.  A',   kiangsis  Kirkaldy;    Hungerford,  Bull.   Brooklyn  Ento.   Soc,  xxv,  p.   138. 

1930.  X.  kiangsis  Kirkaldy;  Kiritchenko,  Annuaire  du  Musee  Zool.  de  I'Academie  des 
Sciences  de  I'URSS'.,  pp.   436  and  440. 

Referring  to  this  species,  also: 

1925.    A',   hergrothi  Esaki,  Notul.  Entom.  Helsingfors  v,  p.   14. 

1925.    K.  hergrothi  Esaki,  Entom.   Mitt.,  Berlin-Dahlem,  xiv^  p.   313  (fig.   1). 

1925.  A",   siietuoni  Hungerford,  Annals  Ento.   Soc.   America,   xviii,   p.    417. 

1926.  iV.  suensoni  Hungerford,  Annals  Ento.  Soc.  America,  xix,  p.  92.  (Syn.  note  ;=  A^ 
hergrothi  Esaki). 

Size.  Length  of  male,  12.5  mm.;  of  female,  13.8  mm.;  width 
across  the  eyes,  male,  2.28  mm.;  female,  3  mm.;  width  across  widest 
part  of  pronotum,  male,  4  mm.,  female,  4.5  mm. 


34  The  University  Science  Bulletin 

Color.  General  color  luteous.  Scutellura  black.  Sparse  silvery 
pubescence  on  hemelytra.  Brownish-black  band  along  apical  margin 
of  clavus  (caudal  half  of  elytral  commissure) ;  slender,  brown  line 
on  base  of  clavocorial  suture,  and  another  longitudinal  line  on  disk 
of  corium;  margins  and  distal  third  of  embolium  more  or  less  brown- 
ish; distal  portion  of  corium  marked  with  two  nearly  quadrate 
brownish-black  areas,  one  before  the  other,  the  posterior  one  larger 
and  both  covered  with  conspicuous  silvery  hairs;  inner  angle  of 
corium  and  inner  base  of  membrane  embrowned.  Venter  black. 
The  ventral  side  of  the  connexivum  greenish. 

Structural  Characteristics.  Head  long;  eyes  close  together  at 
synthlipsis;  vertex  :  synthlipsis  : :  5:1.  Lateral  margins  of  pronotum 
divergent,  sinuate,  and  narrowly  explanate;  anterior  angles  acute; 
lateral  ledge,  as  seen  from  the  side,  faintly  sigmoid  and  ending 
caudally  just  beneath  the  humeral  elevations,  which  are  not  marked. 
Scutellum  slightly  longer  than  pronotum.  Anterior  lobe  of  hemely- 
tral  membrane  slightly  longer  than  posterior  lobe.  Anterior  trochan- 
ter of  male  with  a  small  tubercle,  two-fifths  of  distance  from  base 
to  apex.  The  angle  of  the  mesotrochanter  produced  into  a  long, 
slender,  pointed  process.  Small  hair  tuft  on  median  line  of  third 
ventral  abdominal  segment,  fourth  ventral  abdominal  segment  long- 
est. Last  ventral  segment  of  female  but  shallowly  notched.  First 
pair  of  gonapophyses  of  female  short.  Male  genital  capsule  as 
shown  on  Plate  XV,  fig.  5. 

Location  of  Types.  In  Paris  Museum.  At  the  end  of  his  dis- 
cussion of  A^.  chinensis  Fallou,  Kirkaldy  says:  "To  this  species  1 
have  referred  four  specimens  in  the  Paris  Museum  (from  Chen-Si, 
Se-Tchouen,  and  Kiang-Si)  which  seem  sufficiently  distinct  to  bear 
a  varietal  name — kiangsis  var.  nov."  This  was  followed  by  a  short 
description  of  color.  Since  Kirkaldy  gave  it  as  a  variety  name,  and 
gave  only  a  color  description,  it  was  assumed  by  both  Professor 
Esaki  and  me  that  his  variety  was  structurally  identical  with  N. 
chinensis  Fallou.  We,  therefore,  independently,  but  the  same  year, 
1925,  described  as  new,  specimens  that  came  to  our  hands.  Some- 
time later  Esaki  found  three  specimens  in  the  Paris  Museum  labeled 
by  Kirkaldy  '^Notonecta  chinensis  Fallou  var.  1897"  that  bear  the 
locality  labels  mentioned  above.  While  not  bearing  the  name 
kiangsis  nor  marked  types,  they  certainly  are  to  be  so  considered. 
In  1928  I  also  examined  these  specimens.  How  anyone  of  Dr. 
Kirkaldy 's  experience  could  have  thought  them  to  be  a  variety  of 
N.  chinensis  Fallou  is  a  puzzle  to  me. 


Hungerford:    Genus  Notonecta  35 

Comparative  Notes.  This  species  is  very  distinct  from  any  others 
of  the  orient.  It  differs  from  all  of  them  by  its  pale  color  and  more 
slender  shape.  It  differs  from  N.  montandoni  Kirkaldy,  A'',  violacea 
Kirkaldy,  N.  Kirkaldyi  Martin  and  the  A^.  glauca  group  in  having 
the  angle  of  the  mesotrochanter  plainly  produced.  In  this  respect  it  is 
like  N.  triguttata  Motsch.  and  A^.  chinensis  Fallou,  from  which  it 
is  readily  separated  by  this  process  being  longer  and  sharper.  The 
claspers  of  the  male  are  also  different. 

Data  on  Distribution.  Esaki  described  his  specimens  from  Amur 
and  Ussuri  and  says  in  his  1927  paper:  "This  species  is  now  known  to 
be  widely  distributed  in  Central  and  Northeastern  China  and  East 
Siberia."  There  are  also  many  specimens  from  South  Ussuri  in 
the  Zoological  Museum  of  the  Academy  of  Sciences,  St.  Petersburg 
(Leningrad)  and  a  single  specimen  from  "China  meridionalis  in 
the  Hungarian  National  Museum,  Budapest."  My  specimens  are  as 
follows:  "Shanghai,  China,  April  22,  1923;  E.  Suenson.  (Holotype 
and  allotype  of  A'',  suensoni  Hungerford";  "Tsing  Hua  Col.  Peking, 
China,  11-8-1924,  P.  W.  Claassen";  another  collection  in  1925  at 
same  place  by  Doctor  Claassen;  Mr.  Suenson  also  sent  me  some 
collected  April  18,  1926,  from  Shanghai;  Hweisin,  Kansu,  China 
(my  coll.) ;  Dr.  Kiritshenko  gives  southeastern  Siberia,  Alanchuria, 
Korea,  North  China  and  the  oriental  region. 

Character  of  Habitat.    I  have  no  data  on  this  point, 

Notonecta  inimediata  Kiritshenko  1930 

(Color  plate  V,   fig.    6.) 

19.30.  N.  immediata  Kiritshenko,  Anniaire  du  Musee  Zoologique  de  TAcademie  des  Sciences 
de  \'V.  R.  S.  S.,  p.  438.     (Figs.  5,  6,  9  and  10.) 

Thanks  to  the  kindness  of  Doctor  Kiritshenko,  I  possess  a  female 
paratype  of  this  species.  Aside  from  the  difference  mentioned  in 
the  key,  I  cannot  distinguish  it  from  A^.  triguttata  Alotsch.  from 
Japan.  It  may  be  that  the  male  shows  some  specific  structural 
difference.  The  punctations  on  the  partype  in  my  possession  do  not 
clearly  separate  it  from  N .  triguttata.  I  quote  below  Doctor  Kirit- 
shenko's  description  and  notes: 

"  5 ,  ? .  Corpus  elongatum,  sat  angustum,  supenie  et  inferne  fere  totum 
nigrum.  Caput,  fronte,  viridi  excepta,  et  pronotum  superficie  superna  tota 
nee  non  inferne  limbis  lateralibus  sat  latis  albida.  Hemelytra  nigra,  clavo 
dimidio  basali  toto,  plaga  communi  clavi  et  corii,  comissuram  clavi  occupante 
et  apicem  clavi  baud  attingente,  corii  plaga  propter  venam  radialem  nee  non 
macula  parva  ad  medium  suturae  membranae  sita  luteseentibus ;  exocorio  mar- 
gine  externo  etiam  tenuiter  lutescenti-limbato,  clavo  ad  dimidium  apicalem 


36  The  University  Science  Bulletin 

marginis  scutellaris  vitta  angusta  submarginali  arcuata  lutescenti,  propter  comis- 
suram  clavi  breviter  prolongata  praedito;  membrana  et  ventre  toto  nigris. 
pedibus  sordide  flavescenti-virescentibus. 

"Caput  ante  oculos  late  rotundatum,  vertice  synthlipsi  triple  latiore.  Pro- 
notum  latitudini  suae  anticse  aequilongum,  latitudine  postica  duplo  brevius, 
marginibus  lateralibus  a  supero  visis  subrectis,  angulis  anticis  rectangularibus, 
superficie  superna  nitida,  tenuiter  punctulata;  metanotum  totum  nigrum. 
Scutellum  totum  nigrum,  opacum,  densissime  punctatum,  apice  acuminatum 
et  longe  attenuatum,  latitudine  sua  basali  paullo  brevius.  Hemelytra  densissime 
punctata,  pilis  incumbentibus  aureis  parce  tecta,  angulo  apicali  corii  multo 
densius  piloso,  comissura  clavi  margine  scutellari  aequilonga,  lobis  membranse 
aequalibus  et  aequilongis.  Abdomen  supra  nigrum,  segmentis  4  ultimis  margine 
extemo  flavescenti-maculatis. 

"Long,  corporis  13.5-14  mm.,  long  pronoti  2.6-2.8  mm.,  lat.  hemelytrorum 
4.4-5.1  mm.,  lat.  synthlipsis  0.7-0.8  mm. 

"Habitat  in  prov.  Primorskaja  Sibirise  orientalis:  .Novokievskoje  in  litore 
sinus  Posjet  (27-28  IX  1928,  Prinada  leg.,  Posjet  (11  V  1928,  Volk  leg.). 

"Species  supra  descripta  ad  sectionem  specierum  generis  orientali-asiati- 
carum:  N.  triguitata  Motsch.,  N.  chinensis  Fallou,  A^.  kirkaldyi  Martin,  N. 
kiangsis  Kirk,  pertinet,  ex  quibus  A'',  triguttafa  Motsch.,  specie  japonica  valde 
affinis  et  similis,  differt  statura  angustiore  et  longiore,  connexivo,  ventre  et 
maxima  parte  hemelytrorum  nigris,  hemelytris  praecipue  clavo  et  exocorio 
distinctius  punctatis;  a  N.  kiangsis  Kirk.,  specie  alia  ussuriensi,  magnitudine 
majore  et  latiore,  colore,  vertice  latiore  facile  cognoscitur." 

Notonecta  triguttata  Motsch.  1861 

(Color  Plate  IV,  fig.  4;   Plate  XV,  fig.   6.) 

18(il.    N.  triguttata  Motschulsky,  Etudes  Ento.,  x,  p.  24. 

189V.    N.  triguttata  Motsch. ;  Kirkaldy,  Trans.   Ento.  Soc.  London,  p.   417. 

1904.    N.  triguttata  Motsch.;  Kirkaldy,  Wicn.   Ento.  Zeit.,  xxiii,  p.  95,  132. 

1912.    A'^.   triguttata  Motsch. ;  Oshanin,  Katalog  der  palaarktisclien  Hemiptercn,  p.  91. 

1925.  A'',  triguttata  Motiich. :  Esaki,  Notulac  Entomologies,  v,  p.  14,  fig.  2.  (Figures, 
head  and  pronotum.) 

1925.    N.  triguttata   Motsch.;  Hungerford,  Annals  Ento.  Soc.  Am.,  xviii,  p.  418.     (Notes.) 

1928.  A'',  triguttata  Motsch.;  Hutchinson,  Ento.  Mo.  Mag.,  I.xiv,  pp.  36,  37.  (Figures 
male  genital  capsule.) 

1929.  A',  triguttata  Mutsch.  :  Hutchinson,  Annals  South  .\frii-an  Museum  xxv,  Pt.  3,  p. 
363.     (Subg.  Paranecta.) 

1930.  A^  triguttata  Motsch.  ;  Kiritshenko,  Annuaire  du  Mu.s(e  V.  R.  S.  S.  Zoologique 
de  I'Academic  des  Sciences  de  I'U.  R.  S.  S.,  p.  436. 

Size.  Length,  13-14  mm. ;  width  across  thorax,  4.5  mm.  to  5.1  mm. 
The  females  appear  to  be  Larger  than  the  males. 

Color.  Black  anci  flavous.  Head  and  limbs  of  usual  color;  scutel- 
lum  black;  hemelytra  black  or  bluish  black  with  t-wo  oblique  flavous 
stripes  on  basal  half  and  spot  on  distal  margin  of  corium  midway 
between  lateral  margins ;  sometimes  the  flavous  bands  are  broad  and 
coalesce,  again  they  may  be  reduced  to  narrow  lines.  The  gencial 
pattern  is  that  of  A^.  obliqua  Gallen.  The  venter  and  the  dorsum  of 
abdomen  are  dark  in  color. 


Huxgerford:    Genus  Xotoxecta  37 

Structural  Characteristics.  Head  relatively  smaller  than  in  N. 
chinensis  Fallon;  anterior  breadth  of  vertex  :  ?;ynthlipsis  ::  13:5; 
head  a  little  more  than  half  as  long  as  pronotum.  Lateral  margins 
of  pronotum  divergent  and  but  slightly  sinuate  or  incurved,  nearly 
straight,  especially  in  the  males;  anterior  angles  of  jM-onotum  normal; 
lateral  ledge  as  seen  from  the  side,  faintly  sigmoid,  the  rear  end 
turned  up  beneath  the  humeral  elevations.  Scutellum  longer  than 
pronotum.  Posterior  lobe  of  membrane  a  trifle  longer  than  anterior 
lobe.  Anterior  trochanter  of  male  with  a  small  dentation  instead  of 
a  hook;  mesotrochanter  "with  angle  sharply  produced.  Terminal 
abdominal  sternite  in  female  notched  at  tip;  female  with  first  pair 
of  gonapophyses  short;  male  gential  capsule  as  shown  on  PI.  XV, 
fig.  6. 

Location  of  Type.  Kirkaldy  wrote  '".'Moscow."  In  1915  Kirit- 
shenko  examined  INIotschulsky's  types  and  reported  that  there  were 
two  specimens,  one  labeled  "Japonia''  and  one  without  locality  label. 
They  are  at  Moscow. 

Data  on  Distribution.  Japan.  Kirkaldy.  in  his  ''Uber  Xotonecti- 
den,"  adds,  ''Shantung,  China — Distant  collection." 

Notonccta  montandoni  Kirkaldy  1897 

(Color  I^late  IV,  fig.  I;    Plate  XVII,  fig.  G) 

1897.    N.   montandoni  Kirkaldy,   Ann.   and   Mag.   Xat.   Hist.    (6),   .\x,   p.    56. 
1897.    A',  montandoni  Kirkaldy,  Trans.   Ento.  Soc.  London,  1897,  pp.   417,  418. 

1904.  A',   montandoni  Kirkaldy,  Wien.   Ento.,  xxiii,  pp.   95  and   132. 

1906.  V.  montandoni  Kirkaldy;  Distant,  Faun.  Brit.  Ind.  Rhynchota,  vol.  iii,  p.  41, 
fig.   25. 

1912.  A',    montandoni  Kirkaldy;    Cshanin,   Katalog   der  palaarktisrhen  Hemipteren,   p.    91. 

1925.  A',   montandoni  Kirkaldy;   Hungerford,  Annals  Ento.  Soc.  Am.,  .xviii,  p.  418  (notes). 

1925.  A'^.   montandoni  Kirkald\';    Esaki,  Notulje  Entomologis,  v,  p.   13. 

1930.  N.  montandoni  Kirkaldy;  Kiritshenko,  Annuaire  du  Musee  Zocllogique  de  I'Academie 
des  Sciences  de  I'U.  R.  S.  S.,  p.  436. 

Referring  to  this  species,  also: 

1905.  A',  bivittata  Matsumura,  Jl.  Sapporo  Agric.  Cull.,  vol.  ii,  p.  59,  PI.  1,  fig.  9  0, 
1905   (Esaki). 

1925.  A'',  bivittata  Matsumura;  Esaki,  Xotuls  Entomoligicje,  p.  14,  footnote  says 
examined  the  type  and  finds  it  to  be  A',   montandoni  Kirkaldy. 

Size.  Length,  15  mm.  to  16  mm.;  width  across  protliorax,  5  mm. 
to  5.5  mm. 

Color.  Head  and  limbs  of  usual  color;  scutellum  black;  hemelytra 
reddish-orange  unevenly  marked  with  black,  the  black  figures  often 
coalescing  into  an  irregular  transverse  band  across  eorium,  mem- 
brane black  with  basal  part  of  anterior  lobe  somewhat  mottled  with 
the  ground  color  of  the  eorium. 

4—3482 


38  The  University  Science  Bulletin 

Structural  Characteristics.  Head  with  the  anterior  breadth  of 
vertex  :  synthlipsis  : :  15:6.  Some  specimens  have  synthlipsis  a  little 
narrower  than  above.  Head  half  as  long  as  pronotum.  Lateral 
margins  of  pronotum  divergent  and  nearly  straight,  but  slightly 
convex  on  anterior  half;  anterior  angles  nomial;  lateral  ledge,  seen 
from  the  side,  slightly  oblique  and  up-curved  beneath  the  humeral 
prominence.  Scutellum  slightly  longer  than  pronotum  (6:5).  Lobes 
of  hemelytral  membrane  about  equal  in  length.  Anterior  trochanter 
of  male  without  hook  or  tooth.  Mesotrochanter  nearly  right  angu- 
late.  Mesofemur  with  anteapical  tooth  of  usual  form.  Terminal 
abdominal  sternite  of  female  somewhat  constricted  near  apex,  which 
is  not  notched.  Female  gonapophyses  of  moderate  length.  Male 
genital  capsule  as  shown  on  Plate  XVII,  fig.  6. 

Location  of  Type.    "Montandon  collections." 

Comparative  Notes.  This  species  is  a  little  larger  than  A',  chinen- 
sis  Fallou  and  has  some  maculations  on  basal  half  of  hemelytra.  It 
also  has  a  mesotrochanter  that  is  right  angulate  instead  of  acutely 
angulate  as  in  N.  chinensis  Fallou,  N.  triguttata  Motsch.,  N.  kiang- 
sis  Kirkaldy,  and  N.  immediata  Kiritch. 

Data  on  Distribution.  Given  in  Kirkaldy's  Revision  of  the  No- 
tonectidae  as  follows:  "China  Kiang-Si  (Paris  Mus.),  Ngan-Hoei 
(Montandon  and  my  collns.),  Mou  Piu,  Tibet.  (Paris  Mus.)" 
Paris  Museum  also  has  specimens  labeled  "Museum  Paris  Kouy- 
Tcheou,  Kouy-Yang,  P.  P.  Cavalerie  et  Fortunat  1906."  The  U.  S. 
N.  M.  has  some  labeled:  "Shin  Kai  Si,  Mt.  Omei,  Szechuen, 
China,  4,000-6,000  ft.,  D.  C.  Graham,  collector";  "Suifu,  Szechuen, 
China,  1920.  D.  C.  Graham  Coll.";  "4,400  ft.  Shin  Kai  Si  Mt. 
Omei,  near  Kianting,  Szechuen,  China,  D.  C.  Graham;"  Kiritshenko 
lists  Japan,  China  and  oriental  region. 

Notonecta  pallidula  Poisson  1926 

(Color  Plate  V,   fig.   2;    Plate  XVII,   fig.    3) 

1926.  A''  pallidula  Poisson,  Bull.  Soc.  Hist.  Nat.  Afr.  Nord,  xvii,  p.  238,  figs.  1  and  2. 
(Figs,  of  head  and  pronotum  and  genital  capsule.) 

1928.    N.  pallidula  Hutchinson,  Ento.   Mont.   Mag.,  Ixiv,  p.  35. 

1928.  N.  pallidula  Poisson,  Bull.  Soc.  Ento.  France,  No.  6,  pp.  105,  lOfi. 

1929.  N.  pallidula  Poisson ;  Hutchinson,  Annals  of  South  African  Museum,  xxv,  pt.  3, 
p.   363. 

1933.    A^.   pallidula  Poisson,  Annals  de  la  Soc.  Ento.   de  France,  Janvier,   1933. 
1933.    N.   pallidula  Poisson,    Bull,    de    la    Soc.    Scien.    de    Bretagne    X,    Fasc.    Ill    et    IV. 
(Reprint  p.  1.) 

Referring  to  this  species,  also : 

1897.  N.  glauca  var.  maculata  "leucochroic  form"  Kirkaldy,  Trans.  Ento.  Soc.  Loadon. 
1897,  pp.  422,   423. 


Hungerford:    Genus  Notonecta 


39 


1927.  N.  horvathi  Esaki,  Ann.  &  Mag.  Nat.  Hist.  (9),  xx,  p.  284,  figs.  1,  2,  4.  (Figs, 
dorsal  view  of  insect,  lateral  vdew  of  head  and  thorax  and  tips  of  hemelytral  membranes.) 

Size.  Length,  11.5  mm.  to  13  mm.;  breadth  of  pronotum,  4  mm. 
to  4.5  mm.    Females  slightly  larger  than  the  males. 

Color.  Body  concolorous,  pale  sordid  brown,  including  scutellum. 
Irregular  obscure  dark-brown  markings  along  the  costal  margin  of 
hemelytra  and  along  the  line  between  corium  and  clavus.  These 
may  be  reduced  in  the  male.  Metanotum  and  dorsum  of  abdomen 
yellowish  with  a  dark  marking  on  the  second,  third  and  fourth  and 
part  of  the  fifth  segments.  Underside  of  body  black  with  pale- 
brown  hairs.  In  some  specimens  the  abdominal  venter  may  be 
green  in  color. 

Structural  Characteristics.  Body  comparatively  stout.  Head 
rather  large  with  large  eyes ;  vertex  narrow  at  base ;  anterior  breadth 
of  vertex  :  synthlipsis  ::  8:3.  Pronotum  transverse,  lateral  margins 
divergent,  the  lateral  ledges  more  oblique  and  shorter  than  in  N. 
glauca  Linne.  Scutellum  much  shorter  than  in  A^.  glauca  Linne  and 
more  strongly  narrowed  toward  the  apex.  Membrane  divided  into 
two  nearly  equal  parts  along  the  median  longitudinal  line,  the  apex 
of  the  basal  portion  is  more  angularly  and  more  posteriorly  pro- 
jecting than  that  of  the  apical  half.  Mesotrochanter  rounded.  The 
male  genital  capsule  provided  with  a  finger-like  projection.  The 
clasper  as  shown  in  figure  8,  Plate  XVII.  (See,  also,  figures  3 
and  14.) 

Location  of  Types.  The  author  does  not  state  the  depository  of 
the  types. 

Data  on  Distribution.    Poisson  described  this  species  from  ma- 


B. 


Text  Figure  2.  A.  Notonecta  pallidida  Poisscn  (=  N.  horvathi 
Esaki).  B.  Notonecta  glauca  Linn.  Lateral  view  of  the  head  and  pro- 
thorax.  L.  =  lateral  ledge  of  pronotum.  E  =  rear  margin  of  the  eye 
below  the  lateral  ledge  of  pronotum.  Compare  lengths  of  E.  &  L.  in 
figures  A.  &  B.     (Redrawn  from  Esaki.) 


40  The  University  Science  Bulletin 

terial  taken  by  M.  H.  Gauthier  in  North  Africa,  who  found  it  living 
alone  in  the  ''oiicd  Reghaia,"  but  in  company  with  A',  maculata 
and  y.  glauca  in  the  "oued  Safsaf."" 

Professor  Teiso  Esaki,  who  described  this  species  under  the  name 
A',  hori'othi,  studied  a  male  and  two  females  from  Tangier  that  were 
in  the  National  Museum  of  Natural  History,  Paris.  Esaki  says 
they  were  determined  by  Kirkaldy  as  A^.  glauca  var.  maculata.  Pois- 
son  bclives  that  Delcourt  (1909)  had  seen  this  species  from  Spain, 
Portugal,  Italy  and  North  Africa,  but  had  considered  it  to  be  N. 
macvlata.  Doctor  Poisson  informs  me  that  Algeria  and  Morocco 
are  the  only  records  confirmed. 

Notonccta  canariensis  Kirkaldy  1897 

Color  Plate  IV,  fig.  S;   Plato  XVII,  fig.  10.) 
1S9T.    -V.    (jlcnca  var.    canariensis  Kirkaldy,  Trans.  Ento.  Soc.  London.  1897,  pp.   420,   4-22 

and  4-23. 

1904.     -V.   glauca  var.   caiuirhnsis  Kirkaldy,  Wien.   Ento.   Zeit,  xiii,  p.   1.S2. 

Size.    Length,  14  mm.  to  15  mm.;  width  of  pronotum,  4.85  mm.  to 

5  mm. 

Color.  General  color  effect  dark.  Head,  thorax  and  legs  dark 
testaceous;  an  indefinitely  V-shaped  dark  spot  on  the  pronotum 
behind  the  vertex  of  the  head.  Femur  with  more  or  less  definite 
longitudinal  dark  stripe.  Venter,  except  the  connexivum,  nearly 
black.  Scutellum  black.  Hemelytra  brown  almost  completely  sub- 
merged by  irrorations  (often  suffused)  that  are  veiy  dark  brown 
or  bluish-black;  membrane  dark  brown  to  bluish-black. 

Structural  Characteristics.  Anterior  outline  of  the  head  viewed 
from  above  flattened;  vertex  shorter  than  its  anterior  width;  margin 
of  vertex  less  convex  and  longer  than  the  margin  of  an  eye ;  anterior 
breadth  of  vertex  :  synthlipsis  ::  2  +  :  1.  Pronotum  slightly  more 
than  twice  as  long  as  the  head;  lateral  ledge  as  seen  from  the  side 
sigmoid  and  oblifiuc,  shorter  than  the  rear  margin  of  the  eye  below 
it.  Scutellum  longer  than  the  pronotum.  The  ridge  of  the  hemely- 
tral  suture  :  length  of  scutellum  : :  13:16.  Anterior  lobe  of  membrane 
distinctly  longer  than  the  posterior  lobe.  Anterior  trochanter  of 
male  with  tubercle.  Mesotrochanter  not  angulate.  Anteapical  tooth 
of  fenuir  large.  Terminal  abdominal  sternite  of  female  with  lateral 
margins  not  concave  and  tip  not  plainly  incised.  First  pair  of 
gonapophyses  probably  moderately  short.  Male  genital  capsule  as 
shown  on  Plate  XVII. 


*Oued— A.n  arable  word  signifying  running  water  and  which  designates  more  particularly 
the  courses  of  temporary  streams  in  the  Sahara.  It  becomes  a  part  of  the  name  of  a  great 
niuiibcr  of  courses  of  water  and  localities. 


Hungerford:    Genus  Kotoxecta  41 

Location  of  Types.  In  the  Vienna  ]\Iuseum  I  found  four  speci- 
mens, three  males  and  one  female,  labeled  ''O.  Simony,  1888  Tene- 
riffa"  and,  in  addition,  on  one  label  is  written  '"Barenco  Nedalgo." 
I  placed  red  cotype  labels  on  these  specimens,  since  they  are  the 
specimens  studied  and  recorded  by  Kirkaldy. 

Comparative  Notes.  Kirkaldy  in  his  description  wrote:  "Noto- 
cephalon  a  trifle  narrower  than  in  the  other  forms.  Hemelytra 
purple-black,  obscurely  irrorated  with  dark,  rich  castaneous,  mar- 
gins of  hemielytral  areas  narrowly  the  same  colour;  anterior  half  of 
exocorium  rich  castaneous  blotched  with  black,  posterior  half  black. 
JNlembrane  black.    Pedes  rufotestaceous,  suffused  with  rich  green." 

This  sturdy  species  is  closely  related  to  A^  maculata  Fabr.  The 
mesofemural  tooth  is  longer  and  stouter,  the  tibia  bears  a  conspicuous 
elevation.  The  margin  of  the  pronotum  is  more  strongly  ledged, 
especially  on  the  front  half;  the  abdomen  without  the  orange-yellow 
band  at  base.  The  clasper  of  the  male  genital  capsule  slightly  dif- 
ferent. 

Data  on  Distributio7is.  The  Canary  Islands.  Besides  the  cotypes 
tlie  Vienna  ^Museum  has  ten  specimens  labeled  "Gomera"  ''Canar. 
Ins.  Polatzek,"  and  seventeen  specimens  labeled  "Tenerife"  "Canar. 
Ins.  Polatzek." 

Notonecta  maculata  Fabricius  1794 

(Color  Plate  V,  fig.   10;    Plate  XVI,  fig.    T.) 

1794.    A',    macxdata  Fabricius,  Ento.  Syst.,  iv,  p.   58. 

1799.    A'',   maculata  Coquebert,  III.  Icon.  Ins.,  p.  38,  tab.  x,  fig.   1. 

180.3.    A',   maculata  Fabricius,  Syst.  Rhyng.,  p.   103. 

1817.    A',   maculata  Leach,  Trans.   Linn.   Soc.   London,  xii,  pt.   1,  p.   12. 

1824.  N.  maculata  Curtis,  Brit.  Ento.  (ed.  1.),  I,  pi.  x.  (Esaki  says  plate  is  in  different 
volunje   in   later  editions.) 

1843.    A',   glauca  var.   maculata  Amyot  et  Serville,  Hem.,  p.  453. 

1845.  A^  maculata  Herrich-Scliaffer  (fide  Curtis)  =  marmorea  H.  S.,  Wanz.  Ins.  viii,  p. 
23,  tab.  cclvi,  fig.  797. 

1848.    A',   maculata  Amyot,  Ento.  Franc,  Rhynch.,  p.  338.    (Name  invalid.) 

1805.    A^.   nmculata  Douglas  &  Scott,  Brit.  Hem.  I,  p.  588. 

1888.  A',  glauca  var.  maculata  Fabr.,  Renter,  Synonymica  Heteropteroruin  palaearcticorum 
in:  Acta  Societatis  Scientiaruni  Fennicie  Tomus  XV,  p.  727. 

1892.    AT.   glauca  var.   maculata   Saunders,  Hem.  Heter.  Brit.  Is.,  p.  329. 

1897.    N.   glauca  var.   maculata  Kirkaldy,  Rev.  d'Ento.,  xvi,  pp.  222,  224. 

1897.    A',   glauca  var.   maculata  Kirkaldy,  Trans.   Ento.  Soc.  London,   1897,  p.  422. 

1905.  A',  glauca  \3.r.  maculata  Hiieber,  Jahresh.  Ver.  Vaterl.  Natur.  Wurttemberg,  Ixi 
117. 

1909.  A',  maculata  Fabricius;  Delcourt,  Bull.  S<'ientifique  Fr.  Belg.  (7)  xliii,  p.  378  and 
pp.    389   and  beyond. 

1909.  A'^.   glauca  var.   maculata   Oshanin,   Verz.  Pala?.  Hem.   I,  p.   97."). 

1910.  AT.  glauca  var.  maculata  ^Vliittaker,  Entomologist,  xviii,  p.  97.  London.  (Mating 
in  Jan.   near  Winscombe,   Somerset.) 

1922.  A',  maculata  Despax,  Bull.  Soc.  Hist.  Nat.  Toulouse  L.,  1922,  pp.  97-112,  19  figs, 
(morphology  of  genitalia.) 

1923.  A',   glauca  var.   vmculata  Lundblad,   Ento.   Tidsk.,   1922-23,   p.   72. 


42  The  University  Science  Bulletin 

1023.    A'',   maculata  Butler,  Biol.   Brit.  Hem.   Heter.,  p.   5.'^9. 

1921.  A^.  maculata  Poi.sson,  Bull.  Biol,  de  France  et  Belgique,  T.  Iviii  (1924),  Fasc.  1. 
(Many   ref.    and   figs.) 

1924.  N.   maculata  Hale,  Proc.  Lin.  Soc,  New  South  Wales,  xlix,  pt.   4,  p.  462.    (Notes.) 

1925.  N.  maculata  Jaczewski,  Ann.  Zool.  Mus.  Polon.  Hist.  Nat.,  iv,  p.  128,  fig.  2. 

1926.  maculata  Poisson,  Bull,  de  la  Soc.  d'Hist.  Nat.  de  I'Afrique  du  Nord.,  T.  Dix- 
Septieme,   p.   247,   Nov.,    192G.      (Figures,   head   and   protrorax.)      (Also,   p.    269,   Dec,   1926.) 

1926.  A^.  maculata  Fabr. ;  Poisson,  (3pt.  rend.  Soc.  Biol.  Paris,  Bd.  94,  p.  562  and  1007 
(Cytology.) 

1927.  A^.  maculata  Fabr.;  Poisson,  Archives  de  Zoologie  Experimentale  et  General..., 
Tome  66,  Fasc.  2.  pp.  23-70.  (Cytological  study.) 

1928.  N.  maculata  Esaki,  Annals  &  Mag.  Nat.  Hist.,  Ser.   10  ii,  pp.  69,  70,  75. 

1929.  N.  ma-culata  Fabr.;  Poisson,  Bull.  Soc.  d'Hist.  Nat.  de  I'Afr.  du  Nord,  xx,  pp.  112, 
113. 

1929.    N.  matulata  Hutchinson,  Annals  of  South  African  Mus.,  vol.  xxv,  pt.  3,  p.  363. 

1929.  N.  maculata  Lindberg,  Soc.  Scient.  Fennica  Comment.  Biologies?,  iii,  4,  pp.  11,12. 
(Took  species  m  Spain  and  Morrocco,  and  says  in  both  flowing  and  standing  water.) 

1930.  N.  maculata  Larsen,  Entomologisk  Tidskrift  1930,  pp.  219-241  (fig.  2a,  chorion  of 
Pgg.) 

1930.  A'',  maculata  Fabr. ;  Jones,  H.  P.,  reprint  from  The  Entomologists  Record,  Vols. 
XL-XLII,  p.  69. 

1933.    A^.   maculata  Fabr.;   Poisson,  Annales  de  la  Soc.  Ento.  de  France.  Janvier,  1933. 

1933.  N.  maculata  Fabr.,  Poisson,  Bull,  de  la  Soc.  Scien.  de  Bretagne,  X.,  Fasc.  Ill  et  IV 
(Reprint  p.  2). 

Referring  to  this  species,  also: 

1800.  N.  glauca  Schellenberg,  Cimic.   Helvet.,  p.   21,  tab.  x. 

1803.  N.  marmorea  Fabricius,  Systema  Rhyng.,  p.   103. 

1826.  A^.  variegate  Leach  et  Risso,  Hist.  Princ.  Prod.  Europ.  Merid.  (Risso),  v,  p.  216. 

1835.  N.  marmorea  Herrich-Schaffer,  Nomencl.  Ento.,  I,  p.  63.     (See  Esaki,  1928.) 

1840.  N.  glauca  var.  marmorea  Blanchard,  Hist.  Nat.  Ins.,  iii,  p.  89. 

1851.  N.  jnbricii  var.   umhrina  Fieber,  Abd.  Bohm.   Gesel.  Wis.   (5),  vii,  p.  474. 

1861.  A'',  fabricii  var.  umbrina  Fieber,  Eur.  Hem.,  p.  101. 

1880.  A^.  glauca  var.  umbrina  Puton,  Synop.  Hem.  Heter.  France,  iii,  p.  218. 

1881.  A^.  glauca  var.  umbrina  Puton,  Synopsis  H  m.-Heteroptera  France  in:  Memoires  de 
la  Societe  des  Sciences  de  I'Agriculture  et  des  Arts,  4  Serie,  Tom.  ix,  p.  122. 

1905.  N.  glauca  var.  marmorea  Hiieber,  Jahresh.  Ver.  Vaterl.  Natur.  Wiirttemberg,  Ixi, 
p.  116. 

1907.  N.  umbrina  Dtelcourt,  Comptes  Rendus  des  Seances  et  Memoires  de  la  Societe  de 
Fr.  Biologie,  Ixii,  pp.  11-13.     (Eggs  in  October  to  January  on  supports.) 

1909.    A^.  marmorea  Kuhlgatz,  Siisswasserf.  Deutschl.  (Brauer),  vii,  p.  81. 

Size.  Length,  12.6  mm.  to  15.6  mm.;  wicith  of  pronotum,  4.5  mm. 
to  5.4  mm.,  an  astonishing  variation!  My  smallest  specimen  is  a 
male  from  Jericho,  and  the  largest  a  female  from  Oran  (Algeria) .  I 
have  a  specimen  from  Paris  that  attains  the  maximum  length  but 
not  quite  the  width. 

Color.  An  orange-red  or  orange-yellow  species  irrorated  with 
brownish-red  and  blackish-brown.  Anterior  half  of  pronotum,  head, 
limbs  and  connexivum  of  usual  color,  often  suffused  with  green. 
Scutellum  black.  Metanotum  orange-yellow  with  a  large  blackish 
blotch  bordering  the  basal  half  of  the  scutellar  margins.  First  ab- 
dominal dorsum  orange-yellow,  second  to  fourth  black  (the  fourth 
narrowly  orange-yellow  apically)  fifth  and  beyond,  orange-yellow. 


Hungerford:    Genus  Notonecta  43 

The  liemelytral  markings  variable,  the  dark  areas  often  coalescing 
at  the  distal  inner  angle  of  the  corium. 

Structural  Characteristics.  Anterior  outline  of  head,  viewed  from 
above,  flattened;  vertex  shorter  than  its  anterior  width;  margin  of 
vertex  less  convex  and  longer  than  the  margin  of  an  eye;  anterior 
breadth  of  vertex  :  synthlipsis  : :  2  +  :  1.  Pronotum  slightly  more 
than  twice  as  long  as  the  head;  lateral  ledge,  as  seen  from  the  side, 
sigmoid  and  oblique,  shorter  than  the  margin  of  the  eye  below  it. 
Scutellum  longer  than  the  pronotum.  The  ridge  of  the  hemelytral 
suture  :  length  of  scutellum  : :  13 :  18 — .  Anterior  lobe  of  membrane 
a  little  longer  than  the  posterior  lobe.  Anterior  trochanter  of  male 
with  very  slight  inconspicuous  tubercle.  Mesotrochanter  not  angu- 
late.  Anteapical  tooth  of  femur  of  normal  size  (not  large  as  in  A^. 
canariensis  Kirk.).  Tenninal  abdominal  sternite  of  female  with 
lateral  margins  slightly  concave  and  tip  shallowly  incised.  First 
pair  of  gonapophyses  moderately  short.  Male  genital  capsule  as 
shown  on  Plate  XVI. 

Location  of  Types.  Delcourt  says  the  types  are  in  the  Paris 
Museum. 

Comparative  Notes.  This  species  differs  from  other  common 
European  species  in  having  a  head  that  is  broad  and  truncate  in 
front,  in  having  a  mottled  brown  color  and  in  having  the  last  ventral 
abdominal  sternite  of  the  female  not  compressed  just  before  the  tip. 
In  this  regard  it  agrees  with  N.  paUidula  Poisson,  from  which  it  is 
distinguished  as  shown  by  Poisson  in  his  description. 

Biological  Notes.  Delcourt,  1907,  says  this  species  attaches  its 
eggs  to  the  surface  of  some  support  in  the  water.  The  egg-laying 
period  is  from  October  to  January.  By  keeping  some  eggs  at  a 
temperature  of  from  0  to  8  degrees  they  were  held  from  October  to 
following  September.  At  18  degrees  they  hatched  in  20  days.  Whit- 
taker  records  the  mating  on  January  14  in  Summerset,  England. 

Data  on  Distribution.  Doctor  Poisson  has  been  kind  enough  to 
give  me  the  following  information:  Denmark,  Holland,  Belgium, 
England  (West  and  South),  Iberian  Peninsula,  North  Africa  (Algeria 
and  Morocco),  Egypt,  Macedonia,  Greece,  Italy,  Corsica,  Central 
Europe,  West  and  Southwest  of  Persia.  My  own  records  include 
the  following:  Paris,  France;  La  Galle,  Le  Boul;  Algeria;  Phillippe- 
ville,  Le  Boul;  Brindisi,  J.  Sahib;  Egypt,  G.  Fallou;  Sardinia,  A.  H. 
Krausse;  Morocco,  G.  Buchet,  1903  (Paris  Mus.),  Pushire,  Persia; 
Karab,  23,  1931,  VII,  Alban.  Exp.  1918.     (Vienna  Mus.) ;  Baluchi- 


44  The  University  Science  Bulletin 

Stan,  Qiietta,  Fund  in  Res.  Garden,  Sta.  No.  3,  11-10-18.  B.  S.;  Shisba 
R.  Pond  6,640  ft.,  8  II,  96  Pamir  Exp.  (Indian  Mus.). 

Notonecta  meincrtzhageni  Poisson  1934 

.V.  meinertzhageni  Polxson,  Annals  &  Mae..  Nat.  Hist.  (10)  xiii  p.  170-17.3  (with  text 
figures). 

Size.    Length,  15  mm.;  width  of  pronotum,  5.7  mm.* 

Color.  An  orange-red  and  black  species  with  pattern  same  as 
N.  chinensis  Fallou.    (See  Color  Plate  IV,  fig.  1.) 

Structural  Characteristics.  Anterior  outline  of  head,  viewed  from 
above,  flattened;  the  anterior  outline  of  vertex  slightly  produced 
medianly;  vertex  shorter  than  its  anterior  width;  margin  of  vertex 
less  convex  and  about  equal  to  anterior  margin  of  the  eye ;  anterior 
breadth  of  vertex  :  synthlipsis  ::  37:17.  Pronotum  slightly  more 
than  twice  as  long  as  the  head;  lateral  ledge,  as  seen  from  the  side, 
sigmoid  and  oblicjue,  shorter  than  the  margin  of  the  eye  below  it; 
anterolateral  angles  obtuse.  Anterior  trochanter  of  male  with  very 
slight  inconspicuous  tubercle.  Mesotrochanter  rounded.  Male  geni- 
tal capsule  resembles  that  of  N.  macidata  Fabr.  The  clasper  has  a 
slight  angle  or  projection  on  caudal  margin.  Poisson's  drawing 
shows  lateral  edge  of  pronotum  straight. 

Location  of  Types.  In  British  Museum  and  in  collection  of  Doc- 
tor Poisson. 

Data  on  Distribution.  Described  from  siiecimens  labeled  "Ahag- 
gar  ]Mts.,  1931,  col.  Mcinertzhagen." 

Notonecta  arabien,sis  Hungerford  1926 

(Color  Plate  V,  fig.  1) 
1926.    A',   arabiensis  Hungerford,  Annals  Ento.  Soc.  of  Amer.,  xix.  No.  .3,  p.   280. 

Size.  Length,  16  mm. ;  width  across  the  eyes,  4  mm. ;  width  across 
the  thorax,  6  mm. 

Shape.    Robust. 

Color.  Head,  pronotum,  conncxivum  and  limbs  yellow.  Scutellum 
black,  rear  margin  of  pronotum  somewhat  darkened.  Apical  por- 
tion of  wing  membrane  dark,  remainder  of  membrane,  corium,  and 
clavus  brick  red  with  some  faint  irregular  blackish  blotches.  Venter 
blackish,  midventral  keel  slightly  and  abdominal  margins  plainly 
yellow. 

Structural  Characteristics.     Head  short.    Anterior  outline  of  the 

*  This  short  description  made  from  a  paratype  belonging  to  the  British   Museum. 


Huxgerford:    Gexus  Notonecta  45 

head,  viewed  from  above,  flattened.  Vertex  considerably  shorter 
tlian  its  anterior  width,  margin  of  vertex  nearly  straight,  curving  at 
its  lateral  margins  and  nearly  equal  in  length  to  the  frontal  margin  of 
the  eye;  anterior  breadth  of  vertex  :  synthlipsis  : :  19:10.  Pronotum 
arched  and  at  least  three  times  as  long  as  the  head;  lateral  margins 
strongly  divergent  and  straight;  anterior  angles  normal;  lateral 
ledge,  as  seen  from  the  side,  slightly  sigmoid  and  oblique  and  a 
trifle  shorter  than  the  rear  margin  of  the  eye  below  it.  Scutellum  a 
little  longer  than  the  pronotum  or  the  ridge  of  the  hemelytral  suture. 
Lobes  of  membrane  about  equal.  Legs  stout.  Mesotrochanter 
rounded.  Terminal  abdominal  sternite  of  female  quite  large  and 
but  slightly  constricted  before  the  tip,  which  is  not  notched. 

Location  of  Types.  Described  from  two  females  taken  in  Arabia 
and  belonging  to  British  jVIuseum.  These  are  designated  as  holo- 
type  and  paratype.  It  is  to  be  hoped  that  male  specimens  may  be 
found  to  enable  us  to  figure  the  male  genital  capsule. 

Comparative  Notes.  These  specimens  are  of  the  same  size  and 
color  as  A'.  violac£a  Kirkaldy.  From  this  species  they  differ,  how- 
ever, in  having  a  broader,  shorter  head,  wider  synthlipsis,  and  less 
flattened  lateral  margins  of  pronotum  as  seen  from  above.  Shorter, 
blunter  mesofemoral  tooth.  No  prominence  on  caudal  margin  of 
mcsotibia  just  distad  to  femoral  tooth  when  the  limb  is  flexed. 
Notonecta  violacea  Kirkaldy  is  provided  with  one.  The  last  ab- 
dominal sternite  of  female  is  not  conspicuously  constricted  near  its 
tip  as  in  N.  violacea  Kirkaldy.  I  have  before  me  Kirkaldy's  type,  a 
male  from  Catcin  Cauri,  Birmania. 

Notonecta  violacea  Kirkaldy  lb!)7 

(Color  Plate  IV,  fig  11;   Plate  XVII,  fig.   11.) 
1S97.    A',   vxontnndoni  Kiik.    var.   violacea   Kirkaldy,    Trans.    Ento.    Soc.    London,   1S97,   p. 


41S. 


1004.  .v.  viontandoni  Kirk.  var.  violacea  Kirkaldy,  Wien.  Ento.  Zeit,  xxiii,  p.  132  (lists 
only). 

ISlo.    .V.   violacea  Kirk.;   Hungerford,  Annals  Ento.  Soe.  Am.,  .xviii,  p.  418  (notes). 

Size.    Length,  17.2  mm.;  width  across  pronotum,  6.5-7  mm. 

Shape.    Large,  plump  species. 

Color.  Head,  anterior  part  of  pronotum  and  limbs  dark  tes- 
taceous ;  front  of  head  and  limbs  may  be  greenish ;  scutellum  black ; 
hemelytra  including  basal  two-thirds  of  anterior  lobe  of  membrane 
reddish,  faint-violet  in  background,  rest  of  membrane  brownish- 
black.    Base  of  hind  wing  and  marginal  vein  red. 

Structural  Characteristics.    Anterior  breadth  of  vertex  :  synthlip- 


46  The  University  Science  Bulletin 

sis  ::  9:3;  lateral  margins  of  pronotum  nearly  straight  but  faintly- 
sinuate;  anterior  angles  not  acute.  Scutellum  longer  than  pronotum 
7 — :  5).  Lobes  of  membrane  equal  in  length.  Anterior  trochanter 
of  male  without  hook  or  dentation.  Mesotrochanter  rounded  (per- 
haps faintly  angulate).  Mesofemur  with  usual  anteapical  tooth. 
Mesotibia  with  denticulate  prominence  on  rear  margin.  Terminal 
abdominal  sternite  of  female  large  and  strongly  constricted  near 
tip,  which  is  entire.  First  pair  of  gonapophyses  long.  Male  genital 
capsule  as  shown  on  Plate  XVII,  fig.  11. 

Location  of  Types.  Described  by  Kirkaldy  as  a  variety  of  N. 
montandoni  Kirk.  He  gives  the  habitat  as:  "Burma:  'Catcin 
Cauri,'  "  (Fea  Ag.  Nov.,  1886)  and  specimens  as  being  in  Montan- 
don's,  Hamburg  Mus.  and  his  own  collection.  I  have  found  speci- 
mens in  the  Kirkaldy  collection  at  the  U.  S.  N.  M.  labeled  "Catcin 
Cauri  Birmania  Fea,  Ag.  Nov.,  1886"  and  determined  by  Kirkaldy 
as  N.  montandoni  var. 

Comparative  Notes.  Although  described  as  a  variety  of  N.  mon- 
tandoni, it  is  a  distinct  species  which  is  related  to  N.  arabiensis 
Hungerford  and  N.  amplijica  Kirit.,  from  which  it  may  be  distin- 
guished by  the  key  presented  below. 

A.    Mesotibia  with  denticulate  prominence  on  rear  margin A',  violacea. 

AA.    Mesotibia  without  denticulate  prominence  on  rear  margin. 

B.    Rear  margin  of  middle  tibia  at   base  eniarginate  and   detinculate.     Last  ab- 
dominal sternite  of  female  strongly  constricted A^.  amplifica. 

BB.    Rear  margin  of  middle  tibia  at  base  straight  and  not  denticulate.     Last  ab- 
dominal sternite  of  female  not  stronglj   constrict «'d A^.  arabiensis. 

Notonecta  glaiica  glauca  Linne  1758 

(Color  Plate  IV,  fig.  10;   Plate  XVII  fig.  7.) 

1758.  N.  glauca  Linne,  Syst.  Nat.,  ed.  x.,  p.  439. 

1778.  N.  glauca  Goeze,  Ento.  Beytr.,  ii.,  p.  166. 

1789.  N.  glauca  Gmelin,  Syst.  Nat.  (Linne),  ed.  xiii,  p.  2118. 

1794.  A',  glauca  Fabricius,  Ento.  Syst.,  iv,  p.  57. 

1803.  N.  glauca  Fabricius,  Syst.  Rliyng.,  p.  102. 

1804.  N.  glauca  Panzer,  Schafferi  Icon.  Ins.  Ratisbon.,  Enum.  Syst.  i.,  p.   48. 
1817.  N.   glauca  Leach.  Trans.  Linn.  Soc.  London,  xii,  pt.  i.,  p.   13. 

1829.  N.  glauca  Fallen,  Hem.  Svec,  p.   177. 

1835.  A^.  glauca  Burmeister,  Handb.   Ento.,  ii,  p.   190. 

1835.  A^.  glauca  Herrich-Schiiffer,  Nomencl.  Ento.  i,  p.  63. 

1840.  N.  glauca  Spinola,  Essai  Ins.  Hem.,  p.  59. 

1840.  N.  glauca  Blanchard,  Hist.  Nat.  Ins.,  iii,  p.  88,  tab.  Hem.  i,  fig.  2. 

1843.  A'',  glauca  Amyot  et  Serville,  Hem.,  p.  452. 

1845.  A^.  glauca  Herrich-Schaffer,  Wanz.  Ins.  viii,  p.  23. 

18C0.  A^.  glauca  Fior,  Rhynch.  Livl.  i,  p.  772. 

1865.  A^.  glauca  Douglas  et  Scott,  Brit.  Htm.  i,  p.  587,  tab.  xx.,  fig.   4. 

1869.  N.  glauca  Linne;    Stal,  Hemiptera  Fabriciana,  p.   136. 

1875.  A^.  glauca  Regimbart,  Ann.   Soc.   Ento.   Fr.   (5),   pp.   204,   205,  pi.   IV   (oviposition). 

1878.  A^.  glauca  Snellen  van  Vollenhoven,  Hem.  Heter.  Neerland,  p.  347,  tab.  xx,  figs.  2,  2a. 

1880.  A^.  glauca  var.  glauca  Puton,  Synop.  Hem.  Heter.  France,  iii,  p.  217. 


Hungerford:    Genus  Notonecta  47 

1881.  N.  glauca  var.  glauca  Linn.  Puton.  Synopsis,  Hem.-Heter.  France  in:  Memoires  de 
la  Societe  des  Sciences  de  I'Agnculture  et  des  Arts,  4°  Serie,  Tom.  ix,  p.  121. 

1888.  A^.  glauca  Reuter,  Rev.  Synomymica  Heter.  Palaearcticorum  ...  in  Acta  Soc. 
Sclent.  Fennica  Tom  xv,  p.  725. 

1890.    A^.  glauca  Pedaschenco,  Rev.  Sci.  Nat.  St.  Petersb.  I,  pp.  358-362  (embrj'ology). 

1892.    A^.  glauca  Saunders,  Hem.  Heter.  Brit.  Is.,  p.  329. 

1897.    A^.  glauca  var.  glauca  Kirkaldy,  Trans.  Ento.  Soc.  London,  1897,  p.  421. 

1897.    A^.   glauca  Linne;   Kirkaldy,  Rev.  Ento.  France,  XVI,  pp.  222-224. 

1897.  A',   glauca  Linne;   Horvath,  Rev.  Ento.  France,  1897,  p.  91. 

1898.  N.  glauca  Lnme;   Kirkaldy,  Ento.  Mo.  Mag.,  ix,  (2d),  p.   173  (ovipositioii). 
1900.    A',  glauca  Linne ;   Puton,  Revue  D'Entomologie,  xviii,  p.  80. 

1902.    A^.  glaura  Pantel  and  de  Sincty,  Compt.   Rend.  Ac.   Sc,  Paris,  cxxxv. 

1904.  A^.  glauca  Linne;   Kirkaldj',  Wien.  Ento.  Zeit,  xxiii,  p.   132. 

1905.  N.  glauca  Prowazek,  Zutschr.   Insbiol.,  I,  p.   63  (migration). 

1905.  A',   glauca  Hueber,  Jahresh.   Ver.  Vaterl.   Naturk.   Wiirttemberg,  Ixi,  p.   113. 

1906.  A^.  glauca  Pantel  and  de  Sinety,  La  Cellule,  xxiii,  pp.   87-303,  8  plates  (cytology). 

1906.  A'^.  glauca  Poulton,  Trans.  Ento.  Soc,  1906,  pp.  403-408  (Preying  on  Donacia). 

1907.  N.  glauca  Linne;  Delcourt,  Comptes  Rendus  des  Seances  et  Memoirs  de  la  Societe 
de  Fr.  Biologic,  Ixii,  pp.  11-13.     (Mates  and  oviposits  in  spring  time). 

1909.  N.  glauca  Linne;  Delcourt,  Comptes  Rendus  des  Seances  et  Mem.  9oc.  de  Biologie, 
Paris,  Ixvi,  pp.  589-591. 

1909.    N.  glauca  Oshanin,  Verz.  False,  Hem.  i,  p.  974. 

1909.    A',  glatica  Kuhlgatz,  Siisswasserf.     Deutschl.   (Brauer),  vii,  p.   79,  fig.  68. 

1912.    A^.  glauca  Jensen-Haarup,  Danmarks  Fauna,  xii  (Taeger),  p.  31,  fig.   10. 

1912.    N.   glauca  Wefelscheid,  Zool.  Jahrb.   (Jena),  xxxii,  pp.  391  and  407. 

1912.  N.  glauca  Hoppe,  Zool.  Jahrb.  Jena  Abt.  f.  allg.  Zool.,  xxxi,  pp.  189-244,  2,  tablfs 
(respiration). 

1915.  N.  glauca  Linne;  Wesenberg-Lund.,  Insektlivet  I.  Ferske  Vande,  pp.  118-125. 

1916.  N.   glauca  Browne,  Jl.  of  Morph,  xxvii,  p.  123  (Cytology). 

1917.  N.  glauca  Linne;   Hungerford,  Ento.  News,  xxviii,  p.  268. 

1917.  A'',  glauca  Linne;   Hungerford,  Ento.  News,  xxviii,  p.  174. 

1918.  N.  glauca  Linne;  Hungerford,  Ento.  News,  xxix,  p.  241,  pi.  xv,  fig.  6  (figures 
ovipositor). 

1919.  N.  glauca  Linne;  Hungerford,  K.  U.  Sci.  Bull.,  xi,  pp.  43,  166,  169,  179,  180,  181. 
186,  190,  255,  256,  258,  259,  262,  263,  265. 

1919.    A^.  glauca  Lmne;    Hungerford,  K.  U.   Sci.   Bull.,  xi,  p.    329.     (Error  in  drawing). 
1922.    A^.  glauca  Linne;   Lehmann,  Zool.  Jahrb.  Jena,  xlvi,  pp.  121-158,  2  pL  (Biology  cO- 

1922.  N.  glauca  Linne;  Despax,  Bull.  Soc.  His.  Nat.  Toulouse,  vol.  L,  pp.  97-112,  figs. 
7-12  and  17-19.     (Male  and  female  genitalia.) 

1923.  A^.   glauca  Linne;    Lundhlad,  Ento.  Tidsk.,  p.   71. 

1923.    N.  glauca  Linne;   Ha'e,  Rec.  S.  Aust.  Mus.,  ii.  No.  3,  p.  418. 

1923.  N.  glauca  Butler,  Biol.  Brit.  Hem.   Heter.,  pp.   558,  5.59. 

1924.  N.  glauca  Linne;  Poisson,  Bull.  Biol,  de  France  et  Belgique,  Iviii,  pp.  58,  62-65. 
69-71,  73-76,  79,  110,  112,  113,  114,  185,  186,  188,  191,  276-278,  290,  295,  298,  300;  pi. 
1,  fig.  1,  pi.  3,  figs.  3-5. 

1925.  A^.   glauca  Poisson,  Bull.  Soc.  Ento.  France,  p.  328,  fig.  2. 

1925.  A'',   glauca  Linne;    Hungerford,   Annals  Ento.   Soc.  Am.,  xviii,  p.   418. 

1926.  A^.  glauca  Linne;  Poisson,  Bull,  de  la  Soc.  d'Hist.  Nat.  de  I'Afrique  du  Nord.,  vol. 
xvii,  p.   269. 

1926.    A^.  glauca  Kerkis,  Revue  Russe  d'Ento.,   xx  p.   300   (internal  sex  organs.) 

1928.  A^.   glauca  Linne;    Esaki,  Annals  and   Mag.  Nat.  Hist.,  ser.   10,  vol.   II,  p.  67. 

1929.  A^.  glauca  Linne;  Lindberg,  Soc.  Scient.  Fennica  Comment.  Biologicae  III,  4, 
p.  11   (Spain). 

1929.  A',   glauca  Linne;    Hutchinson,  Annals  of  South  African  Mus.,  xxv,  pt.  3,  p.  303. 

1930.  N.   glauca  Linne;    Lars.n,  Ento.   Tidsk.,   1930,  p.   219. 

1930.  A^.  glauca  Linne;  Jones  in  Reprint  from  The  Entomologists  Record,  vols.  XL- 
XLII,  p.  69. 

1933.    N.  glauca  glauca  Linne;  Poisson,  Annales  de  la  Soc.  Ento.  de  France.     Janvier,  1933. 

1933.  N.  glauca  glauca  Linne;  Poisson,  Bull,  de  la  Soc.  Scien.  de  Bretagne  X.  (Reprint 
p.  2.) 


48  The  University  Science  Bulletin 

Referring  to  this  species,  also: 

1766.  N.  prima  Schaffer,  Icon.  Ins.  Ratisb,  I,  pt.  1,  tab.  xxxiii,  figs.  5  and  6.  (Esaki 
says  name  "invaliduni  ct  nudum.") 

1818.  A',  var.  glauca  Amyot,  Ento.  Fiance,  Rliynch,  p.  337,  tab.  iii,  fig.  .54.  (Esaki  says 
"nom.  invaliduni.") 

1851.  A',  fabricii  var  glauca  Fieber,  Abh.  Bohm.  Gesel.  Wis.  (5),  vii,  p.  474.  (=  Rhyn- 
chotographien). 

Size.  Length,  14  mm.  to  16  mm.;  width  of  thorax,  4.8  mm.  to 
5.1  mm. 

Color.  Typically  yellowish-brown  with  black  scutellum  and 
brownish-black  maculations  along  the  costal  margin  of  the  hemely- 
tra ;  venter  and  abdominal  dorsum  more  or  less  dark. 

Structural  Characteristics.  Head  moderately  convex,  vertex 
shorter  than  its  anterior  width;  margin" of  vertex  less  convex  than 
margin  of  the  eye ;  margin  of  vertex  nearly  as  long  as  frontal  mar- 
gin of  the  eye;  anterior  breadth  of  vertex  :  synthlipsis  ::  2:1;  head 
less  than  half  as  long  as  pronotum.  Pronotum  with  lateral  margins 
moderately  divergent  and  slightly  concave,  more  so  in  female  than 
in  the  male;  anterior  angles  normal;  lateral  ledge,  as  seen  from  the 
side,  sigmoid,  caudal  end  turned  up  beneath  the  humeral  angle,  wdiich 
is  not  prominent,  somewhat  oblique  and  as  long  as  the  rear  margin 
of  the  eye  below  it.  Scutellum  longer  than  pronotum.  Anterior 
lobe  of  membrane  longer  than  the  posterior.  Anterior  trochanter  of 
male  wdth  slight  protuberance.  Mesotrochanter  rounded.  Terminal 
abdominal  sternite  of  female  constricted  near  the  tip,  which  is 
bluntly  pointed.  First  pair  of  gonapophyses  of  female  moderately 
long.    Male  genital  capsule  as  shown  on  plate  XVII,  fig.  7. 

Location  of  Type.  Kirkaldy  gives  the  location  of  the  type  ''? 
Upsala."  I  did  not  find  it  there.  I  found,  however,  a  female,  in  the 
Linnean  Society's  collection  in  London.  Delcourt  --  reported  the 
type  as  being  in  London. 

Discussion.  This  species  name  has  been  applied  to  so  many  forms 
that  one  must  take  the  references  with  some  reservation.  Kirkaldy 
placed  as  varieties  of  this  species  the  following:  N.  marginata 
Miiller,  A',  camiriensis  Kirk.,  A",  marmorea  Fabr.  and  N.  macidata 
Fabr.  I  have  examined  most  of  the  material  mentioned  by  Doctor 
Kirkaldy  in  his  "Revision  of  the  Notonectidae"  and  feel  that  his 
conception  of  this  species  was  quite  confused.  Doctor  Delcourt  and 
others  have  attempted  to  clear  up  the  specific  relationships  of  the 
European  forms. 

Under  "Notonecta  var.  1  glauca  L.,"  Kirkaldy  included  specimens 
from  brackish  water  near  Gravesend  (England).    These  specimens 


Hungerford:    Gexls  Xotoxecta  49 

belong  to  N.  vindis  Delcoiirt.  The  specimens  in  the  Kirkaldy  col- 
lection bearing  the  label  Gravesend  are  certainly  Delcourt's  species. 

Doctor  Kirkaldy,  under  this  variety  (p.  421  of  his  Revision); 
says:  "From  jMaie-Nousky,  Erzerum  (Vienna  Mus.)  I  have  seen 
two  specimens  resembling  in  color  and  disposition  of  the  apical 
hemielytral  fascia,  the  typical  form  of  A^.  americana  Fabr."  I  have 
these  two  specimens  before  me;  they  are  from  Mali,  not  "Male," 
and  I  have  seen  eight  more  in  the  Vienna  Musevmi.  They  are,  it 
seems  to  me,  a  little  larger  than  typical  N.  glauca  Linne.  The  broad, 
black  band  across  the  hemelytra  is  a  striking  characteristic  of  all 
of  them.  (See  Color  Plate  V,  fig.  3.)  A  specimen  like  these  labeled 
"Monts  Baktyaris,  1500  a  4500  m.  J.  de  Morgan  1904"  is  in  the 
Paris  Museum.* 

Data  on  Distribution.  Since  Doctor  Kirkaldy's  conception  of  this 
species  was  rather  vague,  his  idea  of  distribution  is  not  acceptable. 
Doctor  Poisson  has  been  kind  enough  to  write  me  concerning  his 
knowledge  of  this  species.  He  says  it  is  widely  distributed  over 
Europe  and  is  common  in  France,  but  that  its  southern  distribution 
should  be  more  precisely  ascertained.  This,  I  think,  applies  also 
to  the  Asiatic  records.  The  specimens  in  our  own  collection  came 
from  England,  France,  Finland  and  Germany. 

Biological  Notes.  This  species  has  been  studied  by  various 
European  workers.  Regimbart,"-  1875,  reported  that  the  eggs  are 
inserted  in  the  stems  of  aquatic  plants.  Wesenberg-Lund  ^*  says 
the  eggs  are  inserted  in  plant  tissues,  air  filled,  not  water-soaked 
plant  parts.  He  further  says  that  eggs  laid  in  April  and  May  are 
hatched  in  the  course  of  three  to  six  weeks.  The  nymphs  pass 
through  five  moults,  and  after  five  or  six  weeks  they  are  fully 
developed.  Mating  has  been  observed  in  late  summer,  but  the 
usual  time  is  early  spring.  These  insects  are  good  fliers  and  in  the 
fall  seek  ponds  rich  in  plant  life  and  overwinter  there  as  adults. 
Delcourt  --  has  reared  this  species  and  studied  the  results  of  crosses 
of  forms  of  different  color  i^atterns. 

Notonecta  glauca  poissoni  new  subsp. 

■(Color  Plate  V,  fig.   3) 

This  form  is  larger  than  the  average  .V.  glauca  glauca  Linn, 
and  has  a  large  transverse  band  across  the  hemelytra,  as  shown  in 
the  colored  figure.  Doctor  Kirkaldy  mentions  specimens  belonging 
to  the  Vienna  Museum  (see  his  Revision,  p.  421)  that  came  from 
Mali-Nousky,  Erzerum,  and  says  "they  resemble  in  color  and  dis- 

*  I  have  named  these  N.  Glauca  poinsoni. 


50  The  University  Science  Bulletin 

position  of  the  apical  hemielytral  fascia  the  typical  form  of  N. 
americana  Fabr."  The  two  specimens  mentioned  by  Kirkaldy  are 
before  me.  I  have  seen  eight  more  in  Vienna.  Paris  Museum  has 
a  specimen  labeled  "Monts  Baktyaris,  1500  a  4500  m.  J.  de  Morgan 
1904." 

I  propose  to  make  the  series  in  Vienna  the  types  and  will  mark  the 
holotype,  allotype  and  paratypes. 

Notonecta  glauca  hybrida  Poisson*  1933 

1933.    N.   glauca  hybrida  Poisson,   Annales  de  la  Soc.  Ento.   de  France.  Janvier,   1933. 
1933.    N.   glauca  hybrida  Poisson,  Bull,  de  la  Soc.  Scien.  de  Bretagne  X,  Fasc.  Ill  et.  JV. 
(Reprint,  p.   2.) 

This  subspecies  is,  in  color,  intennediate  between  N.  glauca  Linn, 
and  N.  maculata  Fabr.  The  hemelytra  are  reddish  or  brownish, 
more  or  less  strongly  maculated,  presenting  at  the  base  either  one 
pale  longitudinal  band  on  the  clavus  or  two  bands,  one  on  clavus 
and  the  other  on  the  corium.  Structurally  this  form  is  like  A'',  glauca 
Linn,  and  the  females  may  be  placed  at  once  by  examining  the  last 
abdominal  sternite,  which  is  strongly  constricted  before  the  tip,  a 
character  like  A^.  glauca  Linn,  and  not  like  N.  maculata  Fabr. 
Doctor  Poisson  gives  the  distribution  as  Southern  France,  Iberian 
Peninsula  and  North  Africa.  I  have  two  species  sent  me  by  Doctor 
Poisson.    One  is  labeled  "Banyuls"  and  the  other  'Touade." 

Notonecta  glauca  fulva  de  la  Fuente  1898 

(Color  Plate  V,  fig.  11;   Plate  XVI,  fig.  6.) 

1898.  A'^.  glauca  var.  fulva,  de  la  Fuente,  Actas  Esp.  Hist.  Nat.,  xxvi,  p.   130. 

1927.  A^.  maculata  var.  fulva,  Hutchinson,  Annals  &  Mag.  Nat.  Hist.  (9),  xix,  p.  375. 

1928.  N.  glauca  var.  fulva,  Poisson,  Bull.  Soc.  Ento.  France,  No.  6,  p.  106. 
1928.  A',  maculata  var.  fulva,  Hutchinson,  Ento.  Mont.  Mag.,  \xiv,  p.  35. 

1928.  N.  macuJafa  var.  fulva,  Esaki,  Annals  &  Mag.  Nat.  Hist.  (Ser.  10),  vol.  ii,  pp.  70 
and  75. 

J.  M.  de  la  Fuente,  in  a  paper  entitled  ''Datos  para  la  fauna  de  la 
provincia  de  Cindadreal  in  Actas  of  the  "Anales  de  la  Sociedad 
Espafiola  de  Historia  Natural,  1897,  p.  130,"  described  this  variety 
as  follows:  "Notonecta  glauca  L.  v.  fulva:  Elytra  fulva,  sine 
maculis,  una  tantum  levissima  excepta  in  basi  suture  locata." 

The  type  of  this  variety  has  not  been  reexamined  since  it  was 
described,  and  I  do  not  know  its  location.  It  has  been  assumed  by 
Hutchinson  that  it  is  an  immaculate  variety  of  A^.  maculata  Fabr. 
Mr.  Hutchinson  says  there  are  specimens  from  Naples  and  Southern 
Tunisia  that  fit  Fuente's  fulva  and  which  are  a  variety  of  A^. 

*  This  may  be  A'^.  octopunctata  Goeze,  1878,  Ento.  Beytr.,  ii,  p,  169.  See  Esaki,  Annals 
and  Mag.  Nat.  Hist.  (Ser.  10),  vol.  ii,  p.  76,  for  comment  on  A^  secunda  Schaffer. 


Hungerford:    Genus  Notonecta  51 

maculata  Fabr.  and  not  N.  pallidula  Poisson.  Esaki  reports  speci- 
mens taken  by  W.  E.  China  in  Guernsey,  Channel  Islands,  August, 
1927,  that  should  be  assigned  to  N.  maculata  var.  fulva  Fuente. 

Mr.  China,  in  1928,  gave  me  seventeen  specimens  labeled  "Guern- 
sey, August,  1927,  W.  E.  China."  All  but  one  of  these  are  typical 
N.  maculata  Fabr.  There  is  one  male,  however,  that  I  have  taken 
to  be  N.  maculata  fulva  Fuente,  and  figured  in  Color  Plate  V.  This 
specimen,  besides  having  a  little  differently  shaped  head  and 
straighter  lateral  margins  of  prothorax,  has  the  dorsum  of  the  abdo- 
men black  (lacking  the  orange  band  characteristic  of  all  the  other 
specimens)  and  the  anteapical  tooth  of  middle  femur  small.  I  have 
no  female  specimens  from  Guernsey.  An  examination  of  the  last 
abdominal  sternite*  of  female  would  show  at  once  whether  this  form 
should  be  assigned  as  subspecies  of  N.  maculata  Fabr.,  as  Hutchin- 
son and  Esaki  assume,  or  to  N.  glauca  Linn.,  as  Poisson  believes. 
Doctor  Poisson  has  recently  written  me  that  he  has  attempted  to 
locate  Fuente's  type  without  success  and  considers  the  specimens 
from  Guernsey  as  belonging  to  N.  glauca  rufescens  Poisson.  It  is 
in  deference  to  his  opinion  that  I  transfer  Fuente's  species  to  N. 
glauca  fulva,  but  strongly  suspect  that  A^.  glauca  rufescens  Poisson 
will  prove  to  be  N.  glauca  fulva  Fuente. 

Notonecta  glauca  rufescens  Poisson  1933 

(Color  Plate  V,  fig.   11.) 

1933.    N.  glauca  rufescens  Poisson,  Annales  de  la  Soc.  Ento.  de  France.     Janvier,  1933. 
1933.    N.  glauca  rufescens  Poisson,  Bull,  de  la  Soc.  Scien.  de  Bretagne  X,  Fasc.  Ill  et  IV, 
1933.     (Reprint,  p.  3.) 

This  subspecies  has  the  structural  characters  of  A^.  glauca  Linn, 
and  is  distinguished  by  the  almost  immaculate  reddish  hemelytra 
(see  key  to  species).  Since  N.  glauca  fulva  Fuente  must  have  a 
similar  appearance  it  may  be  the  same  thing.  Doctor  Poisson  has 
most  kindly  sent  me  specimens  from  Banyuls  and  Salses. 

Notonecta  obliqua  obliqua  Gallen  1787 

(Color  Plate  IV,  fig.  6;  Plate  XVII,  fig.  9) 

1787.    N.  obliqua  Gallen,  Mus.  Nat.  Acad.  Upsal.,  pt.  v,  p.  61  (Esaki). 
1897.    N.  obliqua  Thunb. ;    Kirka'dy,   Trans.    Ento.    Soc.    London,   p.    419.      (According   to 
Esaki,  Kirkaldy  credited  wrong  author.) 

1928.  N.  obliqua  var.   obliqua  Esaki,  Ann.  &  Mag.  Nat.  Hist.,  ser.  10,  vol.  ii,  p.  68. 

1929.  N.  obliqua  var.  obliqua  Hutchinson,  Ann.  South  African  Museum,  XXV,  pt.  3, 
p.  363. 

*  I  have  asked  Mr.  China,  of  the  British  Museum,  to  examine  the  females  from  Guernsey 
with   reference  to  this  point.     He  replies;     "No   females  are  present." 


52  The  University  Science  Bulletin 

1933.    N.   obliqua  Gallen;   Poisson,  Annales  de  la  Soc.  Euto.  de  France.     Janvier,  1933. 
1933.    A',   obliqua   obliqua   Gallen;    Poisson,   Bull,   de  la   Soc.   Scien.   de  Bretagne  X,   Ease. 
Ill  et  IV  (Reprint  p.   3).     (Also,  Ease.  I  et  II,  Reprint,  p.   29,  egg  of). 

Referring  to  this  species,  also: 

1794.  A',   furcata  Fabricius,  Ento.  Syst.,  iv,  p.   58. 

1799.  A',   furcata  Coquebert,  111.  Icon.  Ins.,  p.   38,  tab.  x,  fig.   2. 

1803.  A'',  furcata  Fabricious,  Syst.  Rhyng.,  p.  102. 

1807.  A'^.  furcata  Haworth,  Trans.   Ento.   Soc.   London,  i,  p.   98. 

1817.  A',   furcata  Leach,  Trans.  Linn.  Soc.  London,  xii,  pt.  1,  p.  12. 

1826.  A^.  melanota  Leach  and  Risso,  Hist.  Princ.  Prod.  Eur.  Merid.  (Risso),  v,  p.  2I.'> 
(Esaki). 

1829.  A^.   furcata  Fallen,  Hem.  Svec,  p.   178. 

1835.  A',   glauca  var.   a  Burmeister,   Handb.   Ento.,   ii,   p.    190. 

1835.  A'',   furcata  Herrich-Schaffer,  Nomtncl.   Ent.,   i,   p.   63. 

1840.  A'^.   fuscata  (!)  Spinola,  Essai,  Ins.  Hem.,  p.   59. 

1840.  A^   glauca  var.   furcata  Blanchard,  Hist.  Nat;  Ins.,  iii,  p.  89. 

1843.  A'^.   glauca  var.  furcata  Amyot  et  Serville,  Hem.,  p.  453. 

1845.  (sine  nom.)    Herrich-Schaffer,  Wanz.  Ins.,  viii,   p.   24,   No.    4. 

1848.  A',   var.  furcata  Amyot,  Ento.  Franc,  Rhynch.  p.   337,  nom.   invalid. 

1851.  A^  fabricii  var.  furcata  Fieber,  Abh.   Bohni.  Ciesel.   Wis.   (5),  vii.,  p.   474. 

1861.  A',  fabricii  var.  furcata  Fieber,  Eur.  Hem.,  p.  101. 

1865.  A',  glauca  var.  furcata  D'ouglas  et  Scott,  Brit.  Hem.,  i,  p.   58S. 

1880.  A'',  glauca  var.  furcata  Puton,  &j-nop.  Hem.  Heter.  France,  iii,  p.  217. 

1881.  A^.  glauca  var.  furcata  Fabr. ;  Puton,  Synopsis  Hem.  Heter.  France  in :  Memoires  de 
la  Societe  des  Science  de  I'Agriculturei  et  des  Arts,  4°   Serie,  Tom.  IX,  p.   121. 

1888.  A'',  glauca  var.  furcata  Reuter,  Syn.  Heterop.  pal.Tarcticorum  in:  Acta  Soc.  Seient. 
Fennicff,  Tomus  XV,  p.   727. 

1892.    A',  glauca  var.  furcata  Saunders,  Hem.  Heter.  Brit.  Is.,  p.  329,  tab.  xxxi,  fig.  2. 

1897.  A',  glauca  var.  marginata  Miiller;  Kirkaldy,  Trans.  Ento.  Soc.  London,  1897,  p.  421 
(Esaki  says  name  applied  to  a  corixid). 

1904.  A^.  fabricii  var.  fasciata  ( ! ),  Kirkaldy  ("fide  Fieber,  1861"),  Wien.  Ent.  Zeit.,  xxiii, 
l.p.   95,  132. 

1905.  A',  glauca  var.  furcata  Hueber,  Jahresh.  Ver.  Vaterl.  Naturk.  Wiirttemberg,  Ixi, 
p.   117. 

1906.  A^  glauca  var.  marginata  Distant,  Faun.  Brit.  Ind.,  Rhynch.  iii,  p.   41. 

1907.  N.  furcata  Fabr. ;  Delcourt,  Comptes  Rendus  des  Seances  et  Memoires  de  la  Societe 
de  Fr.  Biologie,  Ixii,  pp.   11-13. 

1909.  A'^.  furcata  Fabr. ;  Delcourt,  Bull.  Seient.  Fr.  et  Beige,  xliii,  pp.  373,  461  (numerous 
references). 

1909.    A',   glauca  var.   furcata  Oshanin,  Verz.   Pala?.   Hem.,   i,  p.   975. 

1909.     N.  glauca  var.  furcata  Kuhlgatz,   Siisswasserf.  Deut.schl.   (Brauer),  vii,  p.   81. 

1915.  A',  glauca  var.  furcata  Fabr. ;  Bollweg,  Verhandlungen  des  Natiu-historischen  Vereins 
der  Preussischen  Rheinlande  und  Westfalens  71.     Jahrgang   1914,   p.    174. 

1922.  A^.  glauca  var.  furcata  Fabr.;  Despax,  Bull.  Soc.  His-t.  Nat.  Toulouse  1922, 
pp.  97-112. 

1923.  A',   glauca  var.   furcata  Fabr.;    Lundblad,  Ento.   Tidskrift   1922-1923,  p.   72. 
1923.    N.  furcata  Butler,  Biol.  Brit.  Hem.  Heter.,  p.   559. 

1929.  A'',  furcata  Hutchinson,  Annals  of  South  African  Museum,  xxv,  pt.   3,  p.   363. 

1930.  N.  furcata  Jones,  in  Reprint  from  the  Entomologists  Record,  XL-XLII,  p.  69. 
1930.    A',  furcata  Larsen,  Entomologisk  Tidskrift,  1930,  pp.   219-221. 

Size.  Length,  16  mm.;  width  of  pronotum,  5.1  mm.  to  5.4  mm. 
A  large  species  that  may  vary  both  ways  from  above  measurements. 

Color.  A  black  and  luteous  species.  Head  and  limbs  of  usual 
color;  scutellum  black;  hemelytra  black  or  blue-black  with  two 
oblique  luteous  stripes  on  basal  half;  hemelytral  suture  pale;  venter 
and  abdominal  dorsum  dark. 


Hungerford:    Genus  Notonecta  53 

Structural  Characteristics.  Head  not  prominent.  Anterior  out- 
line of  head  moderately  convex;  vertex  shorter  than  anterior  width; 
margin  of  vertex  less  convex  than  margin  of  the  eye;  margin  of 
vertex  longer  than  the  frontal  margin  of  the  eye;  anterior  breadth 
of  vertex  :  synthlipsis  : :  2:1.  Pronotum  more  than  twice  as  long  as 
the  head;  lateral  margins  moderately  divergent  and  slightly  concave; 
anterior  angles  normal;  lateral  ledge,  as  seen  from  the  side,  slightly 
sigmoid,  nearly  horizontal  and  longer  than  rear  margin  of  eye  below 
it.  Sciitellum  longer  than  the  pronotum.  Anterior  lobe  of  membrane 
slightly  longer  than  posterior.  Anterior  trochanter  of  male  with 
slight  elevation  that  is  obscured  by  a  more  or  less  dense  pilosity. 
Mesotrochanter  rounded.  Terminal  abdominal  stemite  of  female 
large,  constricted  near  the  tip,  which  is  bluntly  pointed.  First  pair 
of  gonapophyses  moderately  long. 

Location  of  Type.  Unknown  to  me.  I  did  not  see  it  in  the 
Museum  at  Upsala.  Delcourt  reports  type  of  N.  furcata  Fabr.  to 
be  in  Paris. 

Discm-<^io}i.  This  species  is  better  known  under  the  names  N. 
furcata  Fabr.;  A',  marginata  ■Nliiller,  or  treated  as  a  variety  of  N. 
glaiica  Linn.  Doctor  Esaki,  1928,  says,  "Kirkaldy  identified  this 
species.  Notonecta  furcata,  auct.,  with  Notonecta  marginata  Miiller." 
Miiller  writes:  ''N.  marginata  elytris  nigris:  margine  suturaque 
luteis"  [Zool.  Dan.  Prodr.  p.  104  (1776)  ].  This  description  does  not 
agree  with  this  species,  and  although  "furcata"  is  mentioned  by 
Jensen-Haarup  in  Danmarks  Fauna,  I  have  no  verification  of  its 
occurrence;  in  any  case  it  is  not  a  common  form  in  Denmark,  so 
that  I  do  not  agree  with  Kirkaldy's  opinion.  On  the  other  hand, 
marginata,  ]Muller,  is  most  probably  identical  with  one  of  the  Corixa 
(sensu  lat.)  species,  as  the  description  covers  all  the  forms  which 
are  common  in  Denmark,  and  marginata  was  placed  by  Aliiller  be- 
tween Notonecta  striata  and  N.  minutissima,  both  of  which  are 
Corixids.  The  British  Museum  copy  of  Miiller 's  work  originally 
belonged  to  J.  C.  Schiodte,  the  well-known  Danish  hemipterist,  and 
among  his  annotations  he  writes  of  Notonecta  marginata  "=  Cori.va 
hellensi.  Fallen,  183,"  so  that  there  is  little  doubt  as  to  the  generic 
identity  of  Miiller's  species," 

Doctor  Esaki,  1928,  also  says:  "The  name  ohliqua  was  referred 
to  "Thunberg  DD.  Ac.  Mus.  Upsal.,  p.  61"  by  Kirkaldy.  The  cor- 
rect title  of  this  book  is:  "Museum  Naturalium  Academiae  Upsalien- 
sis  (praeside  C.  P.  Thunberg) ,"  and  different  parts  were  worked  out 
by  different  authors.    Part  V  (pp.  59-68),  "Donation  Thunbergiana) 

5—3482 


54  The  University  Science  Bulletin 

1785,  Continiiat.  Ill,"  in  which  this  species  is  included,  was  written 
by  01a^als  Gallen,  not  by  Thunberg,  and  was  published  in  1787 
(according  to  Hagen,  Bibl.  Ent.,  on  December  5).  His  description 
is  as  follows:  "Hemelytra  nigris  lineis  duabus  obliquis  abbreviatis 
cinereis."  This  agrees  quite  well  with  Notonecta  jurcata  Fabricius, 
and  therefore  I  have  synonymized  the  latter  with  N.  obliqua  Gallen. 
While  Delcourt  ^^  found  all  color  pattern  gradations  between  this 
species  and  N.  glauca  Linn.,  and  Despax  ^"^  concludes  that  it  is  only 
a  variety  of  A^.  glauca  Linn.,  I  am  retaining  them  as  distinct  though 
closely  related  species  and  await  the  conclusions  of  Doctor  Poisson, 
who  has  at  his  command  much  material  and  the  advantage  of  recent 
morphological  studies. 

Data  on  Distribution.  Northern  and  middle  palsearctic.  Doctor 
Poisson  writes  me  that  it  is  to  be  found  in  the  east,  north,  north- 
west, west  and  central  France,  and  that  most  typical  form  appears 
to  be  that  from  Normandy. 

Notonecta  obliqua  delcourtV^  Poisson  1933 

1933.    N.  obliqua  delcourti  Poisson,  Annales  de  la  Soc.  Ento.  de  Fiance.     Janvier,  1933. 
1933.    A',  obliqua  delcourti  Poisson,  Bull,  de  la  Soc.  Scien.  de  Bretagne  X,  Fasc.  Ill  et  IV. 
(Reprint,  p,  3.) 

This  variety,  Doctor  Poisson  says,  is  distinguished  from  N.  obliqua 
obliqua  Gallen  by  having  in  addition  to  the  two  typical  bands  on 
the  hemelytra  a  large  flavous  spot  beyond  the  corial  band.  I  have 
not  seen  this  variety. 

Notonecta  obliqua  meridionalis  Poisson  1926 

(Color  Plate  V,  fig.   5;   Plate  XVI,  fig.   5) 

192C.  N.  jurcata  Fahr.  var.  meridional'!,  Poisson,  Bull.  Soc.  Hist.  Nat.  Afnque  Nord, 
xvii,  p.  240,  November,  1926.     (also  p.  2G9,  December). 

1928.    A'^.  furcata  Fabr.   var.   meridionalis  Poisson,  Bull.  Soc.  Ento.  France,  No.   6,  p.   106. 

1928.  A^.  obliqua  Gallen  var.  viiridionalis  Poisson;  Esaki,  AnnaLs  and  Mag.  Xat.  Hist., 
ser.  10,  vol.  ii,  p.  09. 

1929.  A',  obliqua  Gallen  var.  meridionalis  Poisson;  Hutchinson,  .\nnal>  South  .\frican 
Museum,  xxv,  part  3,  p.  303. 

1933.    N.   oblicjua  meridionalis  Poisson,  Annales  de  la  Soc.  Ento.  de  France,  Janvier,  1033. 
1933.    A',  obliqua  meridionalis  Poisson,  Bull,   de  la   Soc.   Scien.    de  Bretagne   X,   Fasc.   Ill 
and  IV.     (Reprint,  p.  3.) 

Referring  to  this  species,  also: 

1897.    .V.  marginata  Kirkaldy,  Trans.  Ento.  Soc.   London,  p.    421    (Sind  Valley,  Kashmir). 
1925.    A^.  furcata  Poisson,  Bull.  Soc.  Ento.  France,  p.  270. 
1925.    N.  jurcata  Poisson,  Bull.  Soc.  Ento.  France,  p.  328,  fig.  3. 

Size.  Length,  18.2  mm.  to  15  mm.;  width  of  pronotum,  4.8  mm.  to 
5.4  mm. 


*  Poisson  describes  as  a  N-ariety,  not  a  subspecies. 


Hungerford:    Genus  Notonecta  55 

Color.  General  facies  darker  than  N.  obliqua  Gallen.  Hemelytra 
nearly  black  except  for  a  broad  flavous  line  on  the  basal  half  of 
clavus.  Some  specimens  may  show  a  small  flavous  stripe  or  some 
maculations  on  the  corium. 

Structural  Characteristics.  While  I  cannot  fix  upon  any  very 
good  external  characters  to  separate  this  from  N.  obliqua  Gallen, 
tlic  lateral  margins  of  the  pronotum  seem  straighter  and  more  di- 
vergent and  as  viewed  from  the  side  the  pronotum  more  elevated 
in  the  rear.  The  claspers  of  the  two  males  I  have  examined  differ 
from  those  of  A^.  obliqua  as  indicated  on  Plate  XVI. 

Location  of  Type.    Not  indicated  in  Poisson's  description. 

Comparative  Notes.  In  naming  this  variety  Doctor  Poisson  says 
that  it  is  a  little  smaller  than  the  typical  A'',  obliqua  Gallen  and 
usually  only  presents  one  flavous  line  on  the  hemelytron,  this  is  on 
the  clavus.  In  A^.  obliqua  Gallen  there  are  two  slightly  diverging 
longitudinal  flavous  lines  on  the  base  of  the  dark  hemelytra,  one 
on  the  corium  and  the  other  on  the  clavus. 

Data  on  Distnbution.  Doctor  Poisson  writes  me  that  this  species 
is  around  the  Mediterranean  region.  Southern  France.  Iberian  Pe- 
ninsula, North  Africa,  Asia  Minor,  Greece,  Macedonia.  Albania  and 
Italy.  I  have  specimens  before  me  labeled  as  follows:  Tunisia, 
Tabarca  and  Ain  Draham  (Budapest  Mus.) ;  Sind  Valley.  Kashmir 
Det.  by  Kirkaldy  as  N.  glauca  L. ;  Mor.  Sidi  Dris  de  Barcin  G. 
Buchet  1903  (Paris  Mus.) ;  Constantine,  J.  Sahib.  Det.  by  Kirkaldy 
as  N.  glauca  var.  marginata  Mull. ;  Zaguan,  J.  Sahib. ;  Philippeville, 
Algerie,  A.  Thery. 

Notonecta  amplifica  Kiritshenko  1930 

(Color  Plate  V,  fig.   4  ;    Plate  XVI,  fig.   4) 

1930.  N.  amplifica  Kiritshenko,  Annuaire  du  Musee  Zoologique  de  rAcadeniie  des 
Sciences  de  I'U.   R.  S.   S.,   1930,  pp.   434  and  desc.   on  pp.   437-438,  figs.   1,   2,   3,   4. 

Size.  Length,  17  mm.  to  18.3  mm.;  width  across  thorax,  6  mm. 
to  6.3  mm. 

Shape.  Robust. 

Color.  Head,  pronotum,  limbs  and  connexivum  dark  testaceous. 
Rear  half  of  pronotum  darkened  by  black  beneath.  Scutellum  with 
basal  two-thirds  black  mottled  with  pale  yellowisii  spots.  Tip  pale. 
Hemelytra  pale  brown. 

Structural  Characteristics.  Male  head  a  little  i)roader  than  that 
of  female;  anterior  margin  of  vertex  :  synthlipsis  :  :  9  :  4.1.  Pro- 
notum somewhat  arched,  lateral  margins  slightly  sinuate.    Scutc^llum 


5(3  The  University  Science  Bulletin 

and  elytral  suture  about  equal  in  length.  Anterior  lobe  of  mem- 
brane very  slightly  longer  than  posterior.  Anterior  trochanter  of 
male  without  hook  or  marked  dentation.  Middle  leg  as  in  A^. 
arabiensis,  but  distal  femoral  tooth  stouter  and  rear  margin  of  tibia 
slightly  emarginate  and  minutely  denticulate.  Terminal  abdominal 
sternite  of  female  strongly  constricted.  Male  genital  capsule  as 
shown  on  Plate,  XVI,  fig.  4. 

Location  of  Types.  Described  from  southeastern  Siberia.  Types 
in  Pctrograd,  paratypes  in  University  of  Kansas  Collection. 

Comparative^Notes.  A  large  robust  species  related  to  A^.  violacca 
Kirkaldy  and  A^.  arabiensis  Hunger  ford,  see  key  under  A^.  violacea 
Kirkaldy.  Three  specimens  more  slender  and  of  luteous  color  from 
"Amur,"  in  British  Museum,  were  labeled  by  Doctor  Kiriishenko 
as  A^.  lutca  var.  scuteUaris.  The  male  genitalia  shows  them  to  be 
A^  ampUfica  Kiritshenko. 

Notes  on  Distribution.  "Habitat  in  prov.  Primorskaja  Sibiris 
Orientalis:  baud  procul  a  Vladivostok  (Sedanka,  Tshernaja 
Rjetshka,  Oke'anskaja)  et  Novakievskojc  in  litore  sinus  Posjet. 
(27-27  ix  1928,  jPrinada  leg.) " 

Doctor  Kiritshenko  was  kind  enougli  to  send  me  two  specimens 
from  Vladivostok. 

Notonecta  lutea  Miillcr  1776 

(Color   Plate   IV,   fig.    7;    Plate   XVII,   fig.    2) 


177C.    A'.    Intra   Miiller,   Zocil.   Dan  Prodr.,   r 


103. 


Ooeze,  Ento.    Beytr.,   11,   p.    170. 

AVeber  and  Mohr,  Naturh.  Reise  de.   Schwed.,  p.   66   (Fieber). 

Fallen,  Hydroc.  Nauc.  Suec,  p.  6   (Van  Duzee). 

Fallen,  Hem.   Svec,   p.   178. 

Fieber,   Abh.   Bohm.    Gesel.    Wiss.    (5),   vii,   p.    473. 

Flor,  Rhynch.  Livl.,   1,  p.  774. 

Fieber,  Eur.   Hem.,  p.   100. 

Sahlberg,  Not.  Sallsk.  Fauna  Fenn.  Forh.,  xiv,  p.  274   (Kirkaldy). 

Puton,  Synop.  Heter,  Fr.,  p.   218.     (From  A'an  Duzee,  pagination 


1778.  A',  liiica  Miiller; 
1804.  A'.  Ivtea  Miiller; 
1814.  A',  lutea  Miiller; 
1829.  A',  lutea  Miiller; 
1851.    A',   lutea  Miiller; 

1860.  .V.   lutea   Miiller; 

1861.  A',  lutea  Miiller; 
1875.  N.  lutea  Miiller; 
1879.    -V.   lutea  Miiller; 

of  Separate.) 

1881.  A',  lutea  Miiller;  Puton,  Synopsis  des  Hemipt.-Heteropt.  de  France  3°  Partie. 
Memoires  de  la  Societe  des  Sciences  de  I'Agri.  et  des  Arts  de  Lille,  4°  Serie,  Tome  ix,  p.  122. 

1888.  V.  lutea  MuUer;  Reuter,  Rev.  Synom.,  p.  374,  No.  351.  (From  Van  Duzee— 
Pagination  of  Separate.) 

1888.  N.  lutca  Miiller;  Reuter,  Revisio  Synonymioa  Heteropterorum  Pala-arcticorum  .  .  . 
in:    Acta  Societatis  Scientiarum  Fennica?,  Tom.  XV,  p.   728. 

1891.    A',   lutea  Miiller;   Duda,  Klub.  prirod.    Praze,  p.   13,  pi.  iv,  fig.   1.      (Kirkaldy). 

1897.    A',    lutea  MiiUer;   Kirkaldy,  Trans.  Ento.  Soc.   London,   1897,  p.   425. 

1905.    A',   lutea  Mtiller;    Hueber,  Jahresh.  Ver.   Vaterl.   Naturk.   Wiirttiiiiberg,  Ixi,   p.    118. 

1909.    A',    lutea  MiiUer;    O-shanin,  Verz.  Pala>.   Hem.,  I,  p.   97(",. 

1909.    X.   lutca  Miiller;    Kuhlgatz,   Susswasserf.   Deutschl.    (Brauer)    VII,   p.    82. 

1912.    A',   lutea   Miiller;    Jensin-Harrup.    Danmarks   Fauna,   xii    (Ta-ger),    p.    31. 

1912.    .V.    lutea   Muller;    Oshanin,   Katalng   der  Palaarktischen   Hemipteren,   p.    91. 

lOir,.     A',    lutca    Miiller;    Lundbhul.   Ento.   Tidsk.,   p.    220. 


Hungerford:    Genus  Notonecta  57 

1922.  .V.   tut(a  Miiller;    Lundlilad.  Ento.  Ticls-k.,  i\   73. 

1923.  A'^.   lutea  Miiller;    Biieno  &  Hussey,  Bull.   Brooklyn  Ento.  See,  xviii,  p.    lOfi. 

1924.  .V.  lutea  Muller;  Hale,  Proc.  Lin.  Soc.  New  South  Wale?,  vol.  xliSfJ;  pt.  4,  p.  4ii2. 
(Notes  concerning.) 

1925.  A^.  lutea  Miiller;  Schmidt,  Jahrb.  des  Nassauischen  X'eriens  fiir  Natiiikinide, 
Jahrgang  77,  p.  78  ("neu  fiir  die  Rheinprovinz"). 

1928.  A^.  lutea  Miiller;  Hutchinson,  Ento.  Mo.  Mag.,  Ixiv,  p.  35,  3ii.  (Figs,  gi-nital 
capsule). 

1929.  A'',   lutea  Miiller;    Hutchinson,  Annal.s  of  South  African  Museum,  xxv,  pt.  3,  p.   303. 

Referring  to  this  species,  also: 

1835.    A^.   vnicolor  Herrich-Schiiffer,  Nomcncl.  Ento.,  1,  p.  03. 

1845.    N.    unicolor  Herrich-SchaffiT,  Wanz.   Ins.,  viii,  p.   23. 

1851.     A^   tumida   Germar  Ms.,   I'^ieber  mentions  in  his  Rhynrhotographieen,  p.    473. 

Size.    Length,  14-16  mm.;  width  of  pronotum,  4.8-5  mm. 

Shape.    Fairly  plump-bodied  insects. 

Color.  Typical  color  is  luteous  with  brown  ventor  and  legs  of 
intermediate  color.  The  costal  margin  of  hemelytra  often  brown. 
Rarely  there  may  occur  melanistic  specimens  having  black  scutcllum 
and  dark  hemelytra. 

Structural  Characteristics.  x\nterior  outline  of  head,  viewed  from 
above,  flattened;  vertex  shorter  than  its  anterior  width;  anterior 
margin  of  vertex  less  convex  than  the  margin  of  an  eye;  anterior 
breadth  of  vertex  :  synthlipsis  : :  2.2  ±  :  1.  Pronotum  at  least  twice 
as  long  as  the  head  as  seen  from  above;  lateral  margins  of  pronotum 
concave  and  only  moderately  divergent;  anterior  angles  slightly 
produced,  blunt;  lateral  ledge  as  seen  from  the  side  nearly  straight, 
but  slightly  oblique,  and  subeciual  in  length  to  the  rear  margin  of 
the  eye  below  it.  Scutellum  subecjual  in  length  to  the  pronotum. 
Ridge  of  hemelytral  suture  longer  than  scutellum.  Anterior  lobe  of 
membrane  longer  than  posterior  one.  Anterior  trochanter  of  male 
with  hook;  mesotrachanter  rounded,  middle  femur  with  a  long  stout 
anteapical  tooth.  Terminal  abdominal  sternite  of  female  broad, 
somewhat  constricted  before  the  tip,  which  is  blunt.  First  pair  of 
gonapophyses  in  the  female  stout,  broad  and  moderately  long.  The 
male  genital  capsule  with  short  digitate  process  and  clasper  as  shown 
on  Plate  XVII,  fig.  2. 

Location  of  Type.  Kirkaldy  reports  that  the  type  has  been  lost. 
The  species,  however,  is  well  known. 

Comparative  Notes.  This  large  pale  species  has  its  counterpart 
in  A^.  borealis  Bueno  and  Hussey  of  Northern  North  America  which, 
for  a  long  time,  was  confused  with  the  European  species  and  led 
to  the  erroneous  record  of  "Nearctic"  in  Oshanin's  catalogue.  It  is, 
however,  structurally  different  from  the  American  species,  as  shown 
by  a  study  of  the  genitalia  of  both  sexes.     It  is  also  specifically 


58  The  University  Science  Bulletin 

different  from  the  species  named  N.  hitca  var.  scutellaris  by  Reuter 
(see  Plate  XVII,  figm-es  1  and  2. 

Data  on  Distribution.  This  is  a  northern  species.  Reported  by 
Oshanin  in  his  catalogue  as  present  in  middle  and  northern  Europe, 
Crimea  and  Siberia.  Schmidt  has  reported  it  from  the  Rhine  prov- 
ince, and  I  have  specimens  taken  by  Dr.  0.  Lundblad  in  Sweden; 
by  H.  Lindberg,  in  Finland;  and  others  from  Bohemia  and  Kossau. 
Doctor  Poisson  has  kindly  written  me  that  this  species  is  found  in 
the  northern  part  of  eastern  and  central  Europe,  its  western  limits 
passing  from  Hanover  southward  through  Alsace,  the  Jura  and  the 
Alps.  He  reports  finding  this  species  at  one  place,  however,  farther 
west  (Lac  d'Estivadoux  near  Basse  in  the  department  Puy-de-D6me, 
France) .  From  the  map  before  me  this  appears  to  be  in  the  Auvergne 
mountain  district.  Since  the  specimens  from  this  place  in  France 
are  structurally  like  A'',  reuteri  Hungerford  instead  of  like  what  I 
believe  to  be  A',  lutea  Midler,  the  distributional  records  of  these  two 
species  must  be  reviewed  by  the  reexamination  of  all  available 
material. 

Biological  Notes.  Delcourt,--  1909,  gives  considerable  informa- 
tion about  the  distribution  of  this  species  and  reports  upon  catches 
he  has  made.  In  collecting  at  Ulm,  on  August  25, 1907,  he  discovered 
that  this  species  was  fully  developed  sexually,  while  A'',  glauca  L. 
was  plainly  of  the  new  generation.  Wesenberg-Lund,^^  1915,  reports 
that  A^.  lutea  Miiller  inserts  its  eggs  into  the  tissues  of  plants  and 
winters  in  the  egg  stage,  its  young  hatching  in  the  spring  along 
with  those  of  N.  glauca  L.  While  the  egg  stage  of  the  latter  lasts 
from  four  to  six  weeks,  that  of  A",  lutea  Miiller  from  eight  to  nine 
months. 

Notonecta  lutea  Miiller  var.  schumacheri,  Schirmer,  1911 

1911.  .V.  lutea  Miiller  var.  schwnacheri,  Schirmer,   Deutsch.  Ento.  Zeit.,  p.   680. 

1912.  .V.  lutea  Miiller  var.  schumacheri,  Schirmer;  Oshanin,  Katalog  der  paliiarkt.  Hem., 
p.  91. 

1928.  .V.  lutea  var.  schumacheri,  Schirmer;  Esaki,  .\nnals  and  Mag.  Nat.  Hist.  (10), 
II,  p.  71. 

Mr.  Schumacher  describes  this  variety  as  follows : 
"Var.  schumacheri  Schirmer  nov.  In  Grosse  iind  Form  der  Stammart 
gleich,  aber  ganz  Schwarz,  nur  der  Kopf,  das  Pronotum  und  ein  schmaler 
FlugcLseitenrand  gelb.  Diese  aufallige  Varietiit  erlaube  ich  mir  dem  ausserst 
strebsamen  jmigen  Forscher  auf  dem  stiefmiitterlich  behandelten  Gebiete  der 
Rhynchoten,  Herren  F.  Schumacher,  zu  widmen.  In  2  Ex.  wurde  diese 
Varietiit  von  mir  mit  der  Stammform  zusammen  gefangen." 

I  do  not  agree  that  this  should  receive  a  varietal  name.    It  is,  no 


Hungp:rford:    Genus  Notonecta  59 

doubt,  nothing  more  than  an  example  of  melanchroism  in  N.  lutea 
Miiller,  such  as  I  described  in  A^  borealis  Bueno  and  Hussey  (1928). 

Notonecta  reuteri  Hungerford  1928 

(Color  Plate  IV,  fig.  9;    Piatt-  XVII,  fig.   1.) 

192S.  X.  lutea  Miiller  var.  reuteri  Hungerford,  Bull.  Brooklyn  Ento.  See,  xxiii,  June, 
1!>28.  No.  3,  p.  128.  (New  name  for  iV.  lutea  var.  scute.llaris  Reuter,  1886,  which  was 
preoccupied  hy  .V.  variahilis  var.  sciitellaris  Yieher  1S51.) 

Referring  to  this  species,  also: 

1886.    A',  lutea  Miiller  var.  sculellaris  Reuter,   Medd.   Soc.   Faun.   Flor.  Fenn.,  xiii,  p.   234. 

1897.  .V.  lutea  Miiller  var.  scittellaris  Kirkaldy,  Trans.  Ento  Soc.  London,  1897,  p.  425 
(credits  variety  name  to  "Sahib.  S."). 

1899.  A^.  hitca  Miiller  var.  scutellaris  Puton,  Revue  D'Entomologie,  Tome  xviii,  p.  80 
(credits  var.  name  to  "Sahib.  S."). 

1904.  N.  lutea  Miiller  var.  scutellaris  Kirkaldy,  Wiin.  Ento.  Zi'it.,  xxiii,  p.  132  'credits 
var.  name  to  "Sahib.   S."). 

1909.    A',  lutea  Miiller  var.  scutellaris  Reuter;    Oslianin,  ^'erz.   Pah-e.   Hem.   1,   p.   97G. 

1912.  A',  lutia  Miiller  var.  scutellaris  Reuter;  Oslianin,  Catalog,  der  Paliiarkt.  Hem., 
p.  91. 

1927.  A',  lutea  Miiller  var.  scuteliaris  Reuter;  Esaki,  Ann.  &  Mag.  Nat.  Hist.  ''9),  xx. 
p.  280  (Footnote). 

1928.  A",  lutea  Miiller  var.  scutellaris  Reuter:  Esaki,  Ann.  .t  Mag.  Nat.  Hist.  (10), 
vol.  II,  p.   TO. 

1933.    A",   lutea  Poisson,   Annales  de  la   Soc.   Ento.    de  France.   Janvier,    1933. 
1933.    A',   lutea  Poisson,  Bull,  de  ia  Soc.  Scien.  de  Bretagne,  X,  Fasc.   Ill  et  IV  (Reprint, 
p.   1). 

Size.  Length,  14-15  mm.;'"  width  of  pronotum,  4.8  mm. 

Shape.  Slightly  more  slender  and  less  elevated  on  the  back  than 
N.  lutea  Miiller. 

Color.  Typical  color  is  luteous  with  a  dark  median  spot  on  the 
anterior  portion  of  scutellum  and  a  brown  band  along  costal  area 
of  hemelytra,  which  is  broader  in  rear  than  in  front. 

Structural  Characteristics.  Anterior  outline  of  head,  viewed  from 
above,  flattened;  vertex  a  little  shorter  than  its  anterior  width;  an- 
terior margin  of  vertex  less  convex  than  the  margin  of  an  eye; 
anterior  margin  of  vertex  :  synthlipsis  :  :  2  :  1.  Pronotum  usually 
less  than  twice  as  long  as  the  head  as  seen  from  above ;  lateral  mar- 
gins of  pronotum  concave  and  only  moderately  divergent;  anterior 
angles  more  sharply  produced  than  in  N.  lutea  Miiller;  lateral  ledge 
as  seen  from  the  side  nearly  straight,  but  slightly  oblique  and  a 
little  longer  than  the  rear  margin  of  the  eye  below  it.  Scutellum 
subequal  in  length  to  pronotum.  Ridge  of  hemelytral  suture  longer 
than  scutellum.  (Anterior  lobe  of  membrane  longer  than  posterior 
one.)     Anterior  trochanter  of  male  with  a  hook;  mesotrochanter 

*  The  specimens  so  kindly  sent  to  me  from  France  by  Doctor  Poisson  are  only  slightly 
more  than  12  mm.  long  and  have  the  scutellum  pale.  See  my  comments  under  N.  lutea 
Miiller. 


GO  The  University  Science  Bulletin 

rounded.  Middle  femur  with  a  stout  anteapical  tooth.  Terminal 
abdominal  sternite  of  female  broad,  more  strongly  constricted  be- 
fore the  tip  than  in  N.  lutea  Miiller.  First  pair  of  gonapophyses  in 
the  female  strong  and  moderately  long.  Male  genital  capsule  with 
short  digitate  process  and  clasper  as  shown  on  Plate  XVII,  fig.  1. 

Location  of  Type.  Kirkaldy  referred  to  N.  lutea  var.  scutellaris 
J.  Sahib,  and  said  it  was  exhibited  to  the  Societas  Fennica  and  a 
specimen  given  to  him.  I  have  this  specimen  before  me.  Doctor 
Kirkaldy  was  in  error  in  assigning  this  species  to  J.  Sahlberg,  since 
Doctor  Renter  had  named  it  in  1886.  The  type  ought,  therefore,  to 
be  in  Helsingfors,  Finland.  Since  the  name  N.  scutellaris  was  used 
by  Fieber,  1851,  for  a  Notonecta  from  South  America,  a  change  in 
name  was  necessary  and,  therefore,  in  1928,  A^.  reuteri  was  proposed. 

Comparative  Notes.  This  will  be  seen  to  be  a  species  distinct 
from  N.  lutea  Miiller.  Aside  from  a  difference  in  the  typical  coloring, 
the  shape  of  this  species  is  more  slender,  the  head  and  thorax  are 
longer,  and  the  male  genitalia  are  different,  as  shown  on  Plate  XVII, 
figures  1  and  2. 

Data  on  Distnbution.  The  J.  Sahlberg  specimen  before  me  is 
from  'Maakima  vii-20-1881."  It  is  this  specimen  that  is  figured  in 
Color  Plate  IV.  Other  specimens  are  recorded  from  Finland,  Tam- 
mati ;  Jaakima,  Finland  (on  the  northw^est  shore  of  lake  Ladoga  and 
almost  directly  north  of  Petrograd) .  Doctor  Kiritshenko  has  kindly 
given  me  specimens  labeled  "Lake  Tsheznoje,  near  the  rise  of  the 
river  Oredezh,  distr.  Luga,  Petrograd."  Doctor  Poisson  writes  me 
that  the  distribution  of  this  species  as  he  knows  it  is  Finland,  north- 
ern Russia  and  eastern  Siberia. 

There  are  in  the  British  Museum  three  large,  pale  specimens, 
labeled  by  Kiritshenko  A^.  lutea  var.  scutellaris,  coming  from 
"Amur"  An  examination  of  a  male  shows  them  to  be  smaller  and 
paler  than  A^.  amplijica,  but  the  male  possesses  a  genital  capsule  and 
claspers  like  A^.  amplifica  Kirit.  1930. 

Notonecta  vindis  viridis  Delcourt  1909 

(Color  Plate  V,  lig.   U;   Plate  XVII,  fig.  12) 

1909.  A',  viridis  Delcourt,  Bull,  Sci.  France  de  Belgique,  xliii,  p.  379;  Tab.  iv.,  lin.  2; 
Tab.  V,  lin.  18. 

1924.  A',  viridis  Poisson,  Bull.  Biol.  Sci.  France  et  Beliquc,  Iviii,  Fasc.  1  (Life  history  and 
figures). 

1925.  A',  viridis  Poisson,  Bull.  Soc.  Ento.  France,  1925,  p.  256.  (Says  A^.  halophila  Edw. 
is  a  synonym.) 

1928.    N.  viridis  Hungerford,  Annals  Ento.  Soc.  Am.,  xxi,  p.  143,  pi.  ix,  fig.  1. 
1928.    N.  viridis  Esaki,  Annals  and  Mag.  Nat.  Hist.,  sen  10,  vol.  ii,  pp.  68  and  74. 
1933.    A^.   Hindis  viridis  Poisson,  Annales  de  la  Soc.  Ento.  de  France.     Janvier,  1933. 


Hungerford:    Gexus  Notoxecta  61 

1933.  A'^.  viridis  viridis  Delcourt;  Poisson,  Bull,  cle  la  Soc.  Scien.  tie  Bretagne,  Tom.  X, 
Fasc.  Ill  et  IV.     (Reprint,  p.  1.) 

Referring  to  this  species,  also: 

1848.  ;'  N.  var.  pallida  Amyot,  Ento.  Franc.  Rhynch.,  p.  337  (nom  invalid,  note  by  Esaki). 

1897.  A'',   glauca  var.  glaxica  Kirk.,  Trans.  Ento.  Soc.  London,  1897,  p.   421  (in  part). 

1918.  A'',   halophila  Edwards,  Ento.  Men.  Mag.,  liv,  p.  58. 

1923.  N.  halophila  Butler,  Biol.  Brit.  Hem.  Heter.,  p.  559. 

1930.  N.  halophila  Jones,  !>;  Reprint  from  The  Entonjologists  Record,  vols.  XL-XLII,  )i.  09. 

Size.  Length,  14.4  mm.  to  15  mm.;  width  of  pronotum,  4.5  mm. 
to  4.8  mm. 

Coloi'.  General  facies  light.  Head  and  limbs  of  usual  color,  often 
suffused  with  green.  Scutellum  black.  Hemclytra  when  fully  pig- 
mented as  shown  in  Color  Plate  V.  Usually  the  hemelytra  are  less 
pigmented,  the  dark  pigment  confined  to  a  row  of  spots  along  costal 
area  and  an  X-shaped  figure,  the  center  of  which  is  just  behind  the 
tip  of  the  clavus. 

Structural  Characteristics.  Head  not  prominent.  Anterior  out- 
line of  head  very  slightly  convex;  vertex  shorter  than  anterior  width; 
margin  of  vertex  less  convex  than  margin  of  the  eye;  anterior 
breadtli  of  vertex  :  synthlipsis  : :  2  +  :  1.  Pronotum  more  than  twice 
as  long  as  the  head;  lateral  margins  only  slightly  divergent  and 
plainly  concave  near  the  anterior  end;  anterior  angles  acutely  pro- 
duced and  strongly  embracing  the  eyes;  lateral  ledge  distinctly 
sigmoid,  nearly  horizontal  and  a  little  longer  than  the  rear  margin 
of  the  eye  below  it;  other  characters  like  subspecies  below. 

Location  of  Type.  In  Delcourt's  collection.  I  am  very  unhappy 
to  differ  with  Doctor  Esaki  concerning  this  nomenclatorial  problem. 
He  gives  some  quite  convincing  arguments  why  this  species  should 
be  called  A'',  marmorea  viridis  Dele,  and  the  next  one  N.  marmorea 
marmorea  Fabr. 

At  the  time  I  labeled  my  colored  plates  I  w^as  inclined  to  accept 
Doctor  Esaki's  conclusions.  After  further  reflection,  however,  I  am 
forced  to  reject  my  former  stand  in  the  matter.  Fabricius  described 
N.  maculata  in  1794.  This  species  he  described  from  specimens 
taken  near  Paris  and  is  a  readily  recognized  species  with  "abdomen 
atrum,  fascia  baseos  apiceque  fulvis."  Although  he  wrote  ''Scutellum 
album,"  it  is  generally  conceded  that  this  was  a  typographical  error 
for  ''Scutellum  atrum."  The  identity  of  this  species  is  not  disputed. 
Then,  in  1803,  he  described  specimens  from  Algeria  as  N.  marmorea 
as  follows:  "A^  glauca  elytris  testaceis  fusco  maculatis.  Habitat  in 
Algier.  Dom.  Stubb.  Mus.  Dom.  de  Sestestedt.  Statura  omnino  A'. 
glaucae.  Caput  et  thorax  glauca,  immaculata.  Scutellum  atrum. 
Elytra  testacea,  fusco  maculata.  Corpus  nigrum  pedibus  glaucis." 


62  The  Uxivkksity  Sciknce  Bulletin 

When  he  says  "Corpus  atrum  pedibus  glaucis"  his  attention  may 
have  been  focused  upon  the  underside  of  the  entire  body.  To  con- 
sider "Corpus  atrum"  as  meaning  the  abdominal  dorsum  would  not 
be  true  for  N.  viridis  Dele,  any  more  than  for  A",  maaidata  Fabr., 
because  Delcourt's  species  has  the  distal  part  of  the  abdomen  pale 
above. 

I  agree  that  the  remainder  of  the  description  seems  to  fit  A''. 
viridis  Dele.  However,  there  are  no  specimens  of  N.  viridis  coming 
down  fi-oni  Fabricius'  time,  and  there  are  two  specimens  in  the 
Museum  at  Copenhagen  that  are  now  labeled  types  and  which  were 
named  by  Fabricius.  One  is  a  male  labeled  "Mus.  Seh.  &  T.  Lund" 
and  ".V.  marmorea  ex.  Algier  Stub."  The  other  is  a  female  labeled 
"Mus.  Seh.  &  T.  Lund."  The  male  now  has  the  hemelytra  open  just 
enough  to  show  the  yellow  base  of  the  abdominal  dorsum. 

Delcourt,  1909  (p.  377),  noted  this,  for  he  wrote  "Les  caracteres 
t€ut  a  fait  particuliers  de  coloration  jaune  d'une  pai-tie  des  tergites 
et  tout  I'ensemble  ne  permettent  pas  d'en  douter."  He,  too,  had 
seen  these  types  and  declared  them  to  be  N.  maculata  Fabr.  Esaki, 
1928  (p.  72),  writes:  "Thus  Delcourt  synonymized  marmorea '-with 
macidata,  but  he  does  not  actually  mention  whether  or  not  he  ex- 
amined the  dorsal  surface  of  the  Fabrician  specimens,  and  he  was 
ignorant  of  the  pronotal  differences  between  viridis  and  maculata. 
I  am  of  the  opinion  that  the  type  specimens  of  authors  writing  at 
the  time,  when  the  conception  of  the  type  specimen  was  not  yet 
established,  cannot  be  accepted  as  the  standard  for  the  species,  unless 
they  well  agree  with  the  description  of  the  author,  becavLse,  in  those 
days,  so  many  specimens  were  not,  or  insufficiently,  labeled  that  there 
were  a  great  many  possibilities  of  confusion.  .  ."  I  must  report 
that  in  these  two  specimens  the  anterior  lateral  angles  of  the  pro- 
notum  are  not  produced  as  in  A',  viridis  Dele.  The  specimens  are 
the  same  as  N.  maculata  Fabr.  I  believe  them  to  be  the  types  of 
A',  marmorea  Fabr.  and  in  spite  of  the  description  should  be  recog- 
nized as  such.  While  Doctor  Esaki's  arguments  would  be  acceptable 
if  no  historical  specimens  were  at  hand,  it  would  be  unwise  to  ignore 
the  specimens  in  Copenhagen  and  apply  the  name  to  some  other 
species  which  also  fails  to  meet  the  requirements  of  the  description 
in  some  particular. 

Comparative  Notes.  Distinguished  from  other  European  species 
at  once  by  the  acutely  produced  anterior  angles  of  pronotum  which 
embrace  the  eyes  (except  N.  reuteri  Hungerford). 

Data  on  Distribution.    For  the  most  part  at  least  this  form  in- 


Hungerford:    Genus  Notonecta  68 

habits  brackish  water,  although  Delcourt  took  it  in  fresh  water  near 
Paris  and  Poisson  records  it  from  Caen  and  Rennes.  Doctor  Poisson 
writes  me  that  the  distribution  as  known  includes  England,  the  west 
and  northwest  coast  of  France  and  the  fresh-water  records  in 
France  as  given  above.    My  own  specimens  are  from  England. 

Biological  Notes.  Doctor  Delcourt  in  establishing  this  species 
based  his  conclusions  regarding  its  specific  distinctness  more  upon 
biological  than  structural  evidence.  He  studied  its  natural  habitat, 
its  refusal  to  mate  with  N.  glauca  Linn.,  its  less  early  development, 
its  relative  resistance  to  environmental  adversity,  and  noted  that  its 
eggs  and  nymphs  are  smaller  than  those  of  A',  glauca  Linn.  It  re- 
mained for  Edwards  to  point  out  a  structural  detail  useful  in  deter- 
mining this  species.  He  pointed  out  the  acute  anterolateral  angles 
of  the  pronotum  of  his  A^.  halophila,  which  is  now  recognized  to  be 
A^.  viridis  Delcourt.  Like  most  other  European  species,  this  one 
deposits  its  eggs  in  the  tissues  of  plants  and  Avinters  as  an  adult 
insect. 

Notonecta  viridis  meditcrranea  Hutchinson  1928 

(Color  Plate  TV,  fig.   5.) 

1928.  .V.  viridis  mediterranea  Hutchinson,  Ento.  Mo.  Mag.,  ixi\-,  p.  35  (new  name  for 
N.  viridis  meridionalis  Hutch.). 

1933.  N.  viridis  mediterranea  Hutchinson;  Poisson.  Annales  de  la  Soc.  Ento.  de  France. 
Janvier,  1933. 

1933.  .V.  viridis  mediterranea  Hutchinson ;  Poisson,  Bull,  de  la  Soc.  Scien.  de  Bretagne  X. 
(reprint,  p.  2). 

Referring  to  this  species,  also: 

1835.  .V.   glauca  var.  P  Burmeister,  Handb.  Ent.  II,  p.  190. 

1840.  f.  X.  marmorata  Spinola  fide  Fabr.     Essai  Ins.  Hem.,  p.  59. 

1840.  N.  glauca  var.  maculata  Blanchard,  Hist.  Nat.  Ins.,  iii,  p.  89. 

1843.  N.  glauca  var.  marmorea  Amyot  and  Ser\-ille,  Hem.,  p.  453. 

1848.  N.  var.   marmorea  Ams'ot,  Ento.  Franc.  Rhynch,  p.  337  (Esaki  says  "nom.  invalid"). 

1851.  A',  fabricii  var.   marmorea  Fieber,  .\bh.  Bohm.  Gesel.  Wis.  (5),  vii,  p.  474. 

1861.  N.  fabricii  var.  marmorea  Fieber,  Eur.  Hem.,  p.  101. 

1863.  N.  glauca  var.  marmorea  Stal,  Svensk.  Vet.  Akad,  Handl.,  vii.  No.  11  (Hem.  Fabr.), 
p.  136. 

1880.  A',   glauca  var.  marmorea  Puton,  Synop.  Hem.  Heter.  France,  iii,  p.  217. 

1881.  AT.  glauca  var.  marmorea  Fabr.,  Puton,  Synopsis,  Hem.  Heter.  France  in:  Memoires 
de  la  Societe  des  Sciences  de  I'Agriculture  et  des  Arts,  4°  Serie,  Tom.  ix,  p.  121. 

1888.  N.  glauca  var.  marmorea  Reuter;  Rev.  Syn.  Heter.  Palsearcticorum  in:  Acta  Soc. 
Scient.  Fennicje,  Tom.  xv,  p.  726. 

1897.    N.  glauca  var.   marmorea  Kirk.,  Rev.  d'Ento.,  xvi,  pp.  223,  224. 

1897.    N.  glauca  var.  marmorea  Kirk.,  Trans.  Ento.,  Soc.  London,  1897,  p.  421. 

1906.  A'^.  glauca  var.  marmorea  Distant,  Faun.  Brit.  Ind.  Rhynch,  iii,  p.  42. 

1907.  N.  vujrmorea  Fabr.  ;  Delcourt,  Comptes  Rendus  des  Seances  et  Mem.  de  la  Soc.  de 
Fr.  Biologic,  vol.  62,  pp.  11-13. 

1909.    N.   viridis  Delcourt,  Bull.  Soc.  France  et  Belgique,  xliii,  p.   379  (in  part). 
1909.    A'',  glauca  var.  marmorea  Oshanin,  Verz.  Palse.  Hem.,  I,  p.  975. 
1915.    A^.  glauca  var.  marmorea  Bollweg,    Verh.    des    Naturhist.    Vereins   der   Preussischen 
Rheinland  und  Westfalens  71,  Jahrg.   1914,  Mai,  1915,  p.  174. 


64  The  University  Science  Bulletin 

1922.    iV.   glauca  var.   marmorea  Despax,  Bull.  Soc.  Hist.  Nat.  Toulouse,  1922,  pp.  97-112. 

1924.  iV.   marmorea  Poisson,  Bull.  Biol,  de  la  France  et  de  la  Belg.  T.  Iviii,  Fasc.  1,  p.  UO. 

1925.  N.  viridis  Poisson,  Bull,  Soc.  Ento.  France,  p.  328,  fig.  1. 

1927.  A',  viridis  meridionalis  Hutchinson,  Ann.  and  Mag.  Nat.  Hist.  (9),  xx.  p.  S75 
(description  given,  name  later  changed.). 

1928.  A'',  marmorea  viarmorea  Esaki,  Annals  and  Mag.  Nat.  Hist.,  aer.  10,  vol.  ii,  p.  67. 

1928.  A'^.   viridis  meridionalis  Poisson,  Bull.  Soc.  Ento.  France,  No.  6,  p.   107. 

1929.  A^  marmorea  mormorca  Hutchinson,  Annals  of  South  African  Mus.,  vol.  xxv,  pt.  3, 
p.  303. 

Size.  Length,  13.2  mm.  to  14.4  mm.;  width  of  pronotmu,  4.2  mm. 
to  5  mm. 

Color.  Some  specimens  are  colored  as  in  the  illustration  on  Color 
Plate  IV,  others  show  the  rear  half  of  pronotum  dark,  due  to  the 
black  mesonotum  beneath,  and  have  the  lighter  markings  of  the 
hemelytra  tan  to  pale  orange.  Scutellum  black.  The  costal  area 
of  the  hemelytra  nearly  always  checkered  with  a  row  of  roughly 
rectangular  spots.  Venter  black  except  connexivum  and  tip  of  ab- 
domen. 

Structural  Char  act  enstics.  Head  not  prominent.  Anterior  out- 
line of  head  moderately  convex;  vertex  shorter  than  anterior  width; 
margin  of  vertex  less  convex  than  margin  of  the  eye;  margin  of 
vertex  longer  than  frontal  margin  of  the  eye;  anterior  breadth  of 
vertex  :  synthlipsis  : :  2  +  :  1.  Pronotum  more  than  twice  as  long  as 
the  head;  lateral  margins  only  moderately  divergent  and  slightly 
concave;  anterior  angles  acutely  produced  and  embracing  the  eyes; 
lateral  ledge,  as  seen  from  the  side,  distinctly  sigmoid,  nearly  hori- 
zontal and  a  little  longer  than  rear  margin  of  the  eye  below  it. 
Scutellum  longer  than  the  pronotum.  Anterior  lobe  of  membrane 
longer  than  the  posterior.  Anterior  trochanter  of  male  with  slight 
obscure  elevation.  Mesotrochanter  rounded.  Terminal  abdominal 
sternite  of  female  large,  constricted  near  the  tip,  which  is  bluntly 
pointed.    First  pair  of  gonapophyses  moderately  long. 

Location  of  Type.  Now  in  British  Museum.  Type  labeled  "As- 
troni  near  Naples,  Italy,  19,  IV,  1926." 

Data  on  Distribution.  I  consider  the  species  to  include  3  sub- 
species, one  of  which  is  N.  viridis  mediterranea  Hutch.,  and  have 
seen  specimens  labeled  as  follows:  "Egypte,  Bir-Hooker,  J.  Dewitz, 
1904;"  "Milano,  Italy;  Tyrolia,  J.  Sahib;"  "Bethlehem,  J.  Sahib;" 
"Kotschy,  Ost  Medit."  (Vienna  Mus.);  "Baluchistan,  Kushdil, 
Khan  Pishin,  India";  "Plateau  Persan  occid.  De  Hamadan  a 
Zendjan  J.  de  Morgan,  1904  (Paris  Mus.)."  Doctor  Poisson  has 
kindly  supplied  me  with  the  following  distributional  notes:  "South- 
em  France,  Spain,  North  Africa,  Italy,  Sardinia,  Sicily,  Corsica, 
Hungary,  Greece,  Cypinis,  Macedonia,  Asia-minor,  Palestine,  Cri- 


Hungerford:    Genus  Notonecta  65 

mea,  Transcaspia,  Persia,  Turkestan.  About  the  Mediterranian, 
but  showing  a  tendency  toward  a  vast  dispersion  east  and  south- 
eastward." My  own  observations  extend  this  eastward  distribution 
to  India,  and  Hutchinson  lists  Daulatabad,  Seistan. 

Notonecta  viridis  kashinlriana  new  subsp. 

(Color  Plate  V,  fig.   13;   Plate  XVII,  fig.   5) 

Size.    A  little  smaller  than  the  others. 

Color.    Typical  color  as  shown  on  Color  Plate  V. 

Structural  Characteristics.  Closely  related  to  N.  viridis  mcditer- 
ranea  Hutch.  The  synthlipsis  relatively  a  trifle  wider,  the  lateral 
margins  of  the  prothorax  more  divergent  and  the  genital  claspers 
of  the  male  as  figured  on  Plate  XVII,  fig.  5. 

Location  of  Types.  Holotype  and  allotype  in  Indian  Museum, 
Calcutta;  paratypes  in  Francis  Huntington  Snow  Entomological 
Museum,  University  of  Kansas.  Described  from  five  specimens 
labeled:  "Kashmir  Survey,  Pond  on  roadside  near  Missionary  Hos- 
pital, Sprinagar,  Kashmir,  N.  W.  Himalayas,  Sta.  3,  10,  VI,  21." 

Discussion.  There  is  before  me  also  a  series  labeled:  "Daulata- 
bad, Seistan  20.  XII,  18.  Sta.  29,  B.  S.  Village  Pond."  In  these  the 
male  genital  clasper  is  not  quite  like  the  above,  but  intermediate 
between  it  and  A',  viridis  mediterranea  Hutch.  These  groups  appear 
to  involve  the  same  problem  encountered  with  N.  unijasciata  Guerin 
in  America. 

NOTONECTA  OF  WESTERN  HEMISPHERE 

Key  to  Xotonect.v  of  North,  Cextr.^l,  .\nd  Insul.'Vr  America! 

1.  Keel  of  fourth  abdominal  stemite  bare,  the  hairs  confined  to  the  sides.     (See  Plate 

X,   figs.    1   and    2) (.j) 

Keel  of  fourth  -sternite  nut  liare  (except  iu  N.  borealis).     (See  Plate  X,  figs.  3  and  •},   (14) 

2.  Head  broad;    eyes   large  and   broad,   rear  width   usually   greater  than   length   of   the 

lateral    margin    of    pronotvim;    scutelluni    conspicuously    broader    than    long;    last 
abdominal  sternite   of   female   narrow,   side   margins   usually   concave.      (See   Plate 

X,    fig.    1) (3) 

Head  not  conspicuously  broad  and  short.  Eyes  not  transverse,  rear  width  not  as 
great  as  length  of  lateral  margin  of  pronotum.  Scutellum  not  conspicuously 
broader  than  long.  The  last  abdominal  sternite  of  female  not  narrow  with  con- 
cave side  margins.     (See  Plate  X,  figs.  2  and  3) (12) 

3.  Lateral  margins  of  pronotum  of  males  straight*  and  nearly  so  in  female.     Penultimate 

abdominal  sternite  of  female  slender  at  tip (4) 

Lateral  margins  of  pronotum  of  males  at  least  slightly  conca\e,  females  plainly  so. 
Penultimate  abdominal  sternite  of  female  fairly  broad,  explanate  near  caudal  end 
or,  if  slender,  plainly  running  to  this  section  by  the  concave  lateral  margins  of 
pronotum (6) 

t  Read  Section  III  beginning  p.  12. 

*  M^les  of  -V.  ceres  stirtoni  have  lateral  margins  of  pronotum  almost  straight,  but  the 
females  have  the  margins  plainly  concave. 


66  The  Uxiversity  Science  Bulletin 

4.  Length    not    more    than    12    mm.      Last    abdominal    sternite    of    male   not    umisually 

broad.     Penultimate  abdominal  sternite,  of  female,  pointed  at  tip.     Last  sternite 

slender.     (See  text  figure  3) iV.   hoffmanni. 

(California,  Arizona  and  Lower  California,  p.  68.) 
Length    more    than    12    mm.      Last    abdominal    sternite    of    male    unusually    broad. 
Penultimate  abdominal  sternite   of  female  not   pointed,   but  more   or   less   incised 
at  tip  and  last  sternite  rather  broad.     (Plate  X,  fig.   1) (5) 

5.  Genital  capsule  of  male  with  large  lobe  in  front  of  clasper.     PenuHimate  abdominal 

sternite  of  female  plainly  notched  at  tip -V.   lobata. 

(New  Mexico,  Texas,  Mexico,  p.   70.) 
Genital    capsule    without    large    lobe    in    front    of    clasper.      Penultimate    abdominal 

sternite  of  female  faintly  notched  at  tip -V.  hintoni. 

(Mexico,  p.  72.) 

6.  Anterolateral    angles    of    pronotum    acute    and    snugly    embracing    tlie    eyes.       (Plate 

XVH,    fig.    4) (7) 

Anterolateral  angles  of  pronotum  not  as  above  (See  Plate  VIII,  fig.  8) (9) 

7.  Lateral    margins    of    pronotum    abruptly    constricted,    greatly    so    in    female.      Last 

abdominal   sternite  of   female  broad.      (Text  figure   5) N.    compacta. 

(Mexico,  p.  73.) 
Lateral    margins    of    pronotum    concave,    not   abruptly    constricted.      Last    abdominal 

sternite  of  female  narrow.     (Text  figure  5) (8) 

8.  Length   more  than   12   mm -V.    mcxjrnna. 

(Mexico,  p.  75.) 

Length   12  mm.  or  less N.   m.  creaseri. 

(Mexico,  p.  77.) 

0.    Large   and    robust.      Male    clasper    without    sharp    basal    projection.      Females    with 

lateral    margins    of    pronotum    constricted    just    behind    the    anterolateral    angles, 

which  flare  outward -V-  robusta. 

(Mexico,  p.  7S.) 
Size  moderate  to  small   for  tliis  subgenus.     Male  clasper  with   a  sharp  basal  projec- 
tion.    Females  not   as  above (10) 

10.  Lateral    marg-ns    of    pronotum    of    male    almost    straight.      Penultimate    abdominal 

sternite   of   female   fairly   broad   and    expanded   before   the  tip.      (Text    figure    6) 

A',  ceres  stirtovi. 

(Mexico,  p.  80.) 

Lateral    margins    of    )ironotinn    of    male    plainly    concave.      Penultimate    abdominal 

sternite  of  female  narrow  and  slightly  expanded  before  the  tip.     (Text  figure  6),   (11) 

11.  Length  12  mm.  or  less.     The  lateral  margins  of  the  pronotum  of  female  constricted 

near  the  middle A',    ceres   ceres. 

(Costa  Rica,  p.   79.) 
Length   more   than   12   mm.     The  lateral   margins    of   the   pronotum    of   female   con- 
stricted in  front  of  the  middle N.  c.  rogersi. 

(Co.sta  Rica,  p.  81.) 

12.  Keel  of  fourth  and  fifth   abdominal  sternites  bare,  the  hairs  confined  to  the 

sides   A^.  impressa  and  N.  montezuma. 

(Mexico  and  Texas,  pp.  81  and  82.) 
Keel  of  fourth  abdominal  sternite  only  bare,  the  hairs  confined  to  its  sides (13) 

13.  Synthlipsis  usually  less  than   one-half  width   of  eye.      Metaxyphus   typically  yellow. 

Head  one-half  as  long  as  pronotum A^.  insulata. 

(Eastern  Canada  and  Northern  United  States  east  of  100th  meridian,  p.   84.) 
Synthlipsis  usually  fully  one-half  to  two-thirds  width  of  eye.     Metaxyphus  typically 

black.     Head  less  than  one-half  as  long  as  pronotum A^.  kirbyi. 

(Western  Canada,  Northern  United  States,  west  of  100th  meridian  and  south 
to  Texas,  p.   88.) 

14.  Last  abdominal  sternite  of  female  large  but  strongly  constricted  just  before  the  tip. 

Male  with  digitate  prolongation  on  genital  capsule (15) 

Last  abdominal  sternite  of  female  not   large  nor  strongly  constricted   just  before  the 

tip.     Male  without  digitate  prolongation  on  the  genital  capsule (16) 

15.  Color    dark,    scutellum    black,    hemelytra    irrorated    with     brown     and    blue-black. 

Synthlipsis    narrow,    only    one-third     anterior    margin    of    vertex     viewed     from 
above A",  irrorata. 


Hungerford:    Genus  Notonecta  67 

(Eastern    Canada    and    Eastern    United    States    south    to    Gulf    and    west    to 
Kansas,  p.  92.) 
Culur    light,    scutellum    pale,    hemelytra    pale.*      Synthlipsis    greater    than    one-third 

anterior  margin  of  vertex .V.   borcalis 

(Across  Canada  and  in  Michigan  and  Minnesota,  U.  S.  A.,  p.  0.5.) 

16.  Pronotum  broad  in  front,  embracing  the  eyes,  which  are  flattened  and  receding  from 

the  anterior  margin  of  the  vertex.  Synthlipsis  broad,  on^'-half  anterior  margin 
of  vertex  as  seen  from  above.  Scutellum  plainly  broader  than  long.  Males  with 
stout  tubercle  at  angle  of  front  trochanter  and  a  very  stout,  broad  hook  as  shown 

on  Plate  IX,  fig.   4 (17 , 

Pronotum  not  broad  in  front.  Eyes  not  as  above.  Scutellum  not  plainly  transverse. 
Males  without  stout  tubercle  at  angle  of  front  trochanter  and  with  a  small  hook 
on   its   anterior   surface (21) 

17.  Lrngth  usually  not  more  than  12  mm " (^g) 

Length  usually  more  than   12   mm (19) 

18.  Pronotum   steeply   declivant  as   viewed    from    the   side.      Dark    form   with    hemelytra 

brown    to    nearly    black    without    luteous    stripes.      Luteous    form    with    tinge    of 

orange.     Male  genital  capsule  as  on  Plate  XIII N.   melaena. 

(Mexico,  p.  97.) 
Pronotum  not   steeply  declivant.      Dark   form   unknown   to  me.      Luteous    form   j  ale 
luteous.     Male  genital  capsule  with  a  thickened  protuberance  on  keel... 

N.   ochrothoe. 
(Southern  California  to  Colombia,  S.  A.,  p.  99.) 

19.  Anterior  end  of  lateral   ledge  of  pronotum  broadly  flattened,  the  ledge  plainly  wider 

beneath  than  the  diameter  of  the  second  antennal  segment.  Penultimate  ab- 
dominal sternite  of  female  broadly  triangular.      (Text  figure  7) A'^.  repanda. 

(Arizona  to  Mexico,  p.  100.) 
Anterior  end   of   lateral   ledge   of   pronotum   not   broadly   flattened,    scarcely,    if   any, 
wider  than  the  diameter  of  the  second  antennal  segment.     Penultimate  abdominal 
sternite  of   female  slenderly  triangular.      (Text   figure   7) (20) 

20.  Caudal    half    of    lateral    margin    of    pronotum    plainly    undulate    as    seen    from    side. 

^'■ertex  of  ftmale  not  as  below.     Male  genital  capsule  as  on  Plate  XIII,  fig.  8. 

A'^.  shoot eri. 

(California,  p.  101.) 

Caudal    half    of    lateral    margin    of    pronotum    of    male    nearly    straight.       An'erior 

margin    of    vertex    of    female    considerably    produced    beyond    the    very    receding 

eyes A^.   colombiana. 

(Colombia,  S.  A.,  p.  128.) 

21.  Mesotrochanter  plainly  angulate  or  produced  into  a  tooth  or  stout  spinelike  process,  (22) 
Mesotrochanter  rounded   or  nearly  so (26) 

22.  Synthlipsis  very  narrow,  not  more  than  one-sixth  the  anterior  margin  of  the  vertex 

as   seen    from    above.      Hemelytra    brick    red    to    orange    with    black    blotch    on 

corium   A'^.   uhleri. 

(Eastern  United  States  from  Massachusetts  to  Florida  westward  to  Mississippi 
river,  p.  103.) 
Synthlipsis  usually  broader  than  above.     Hemelytra  not  red  or  orange (23) 

23.  Scutellum  typically  pale (24) 

Scutellum  typically  black  with   flavous  lateral  margins (25) 

24.  Synthlipsis  about  one-fourth  width   of  eye A',   raleighi. 

(Southeastern  United  States,  p.  10.").) 

Synthlipsis  about  one-third  width  of  eye A',  lunata. 

(Eastern  Canada  and  northern  states  west  to  Wisconsin,  p.  107.) 

25.  SjTithlipsis   at   least   one-half   anterior   margin   of   vertex.      Mesotrochanter   not   pro- 

duced into  a  long  spinose  process A'^.  unifasciata  and  subspecies. 

(Mexico  north  to  western  Canada,  pp.  109  to  111.) 
Synthlipsis   less   than    one-half   anterior   margin    of    vertex.      Mesotrochanter    produced 

into  a  long  spinose  process A',  spinosa. 

(Western  Canada  and  northwestern  IT.  S.  A.,  p.   112.) 

*  Often   there    is   a    dark    submarginal    streak,    and    rarely    the    scutellum    is    black    and    the 
hemelytra  marked  nearly  as  in  A',  irrorata  Uhler. 


68  The  University  Science  Bulletin 

20.    Synthlipsis  distinctly  less  tlian  one-half  anterior  margin  of  vertex,   head  prominent, 

hemelytra  usually  black  and  white (2^) 

Synthlipsis  nearly   one-half  anterior  margin   of  vertex.      Head   not   prominent.      Pale 

or  marked  with  black (28) 

27.  Anterior  margin   of  vertex   (seen   from   above)   straight   and   width   less   than   length 

of  vertex.     Male  clasper  broad,  shallowly  incised  at  tip.     Last  abdominal  sternite 
of  female  with   apical   notch  broader  than   deep   if   present.      Typical   color  with 

broad  black  band  across  distal  end  of  hemelytra A',   indica. 

(North  America  south  of  37°  latitude.  Insular  America,  S.  .^,^  p.  113.) 

Anterior  margin  of  vertex  rounded  and   width   sul)equal   to  length   of  vertex.      Male 

clasper    deeply    incised.      Last    abdominal    sternite    of    female    with    apical    notch 

deeper    than    broad.      Typical    color    with    narrow    undulate    band    across    liem- 

glytra    ^-   undulata. 

(A.-ross  Canada,  widespread  in  United  States  to  Mexico,  p.  117.) 

28.  Male   clasper   bifurcate (29) 

Male  clasper  not  bifurcate.      (Plate  XIV,  fig.   8) N.  indicoidea. 

(Mexico,  p.  123.) 

29.  Clasper  broad  and  deeply  furcate.     (Plate  XIII,  fig.   7) .V-   coiifusa. 

(South  America,  p.  130.) 

Clasper  plump  and  shallowly  notched.     (Plate  XIII,   fig.   9) -V.    distwctoidea. 

(Mexico,  p.  124.) 

Notoriecta  hoffmanni''  Hungcrford  1925 

(Color  Plate  III,  fig.  12;  Plate  XI,  fig.  G;   text  figure  3.) 
1925.    .V.   hoffmani  Hungerford,  Can.  Ento.  Ivii,  p.  241,  pi.  vi,  fig.   6. 

Referring  to  this  species  also: 

1894.  N.  7ticxicana  Lhler,  Proc.   Calif.   Acad.   Sci.    (2),  iv,  p.   202. 

1697.  N.  mcxicava  Kirkaldy,  Trans.  Ento.  Soc.  London,  1897,  p.  4ri2.    (Bassee  Californie, 

Diguet  1895,  Paris  Mus.) 

1901.  N.  mexicava  Champion,    Biol.   Centr.   Amer.   Hem.-Het.    ii,   p.    368. 

1905.  N.  mexicana  Bueno,  Jl.   N.  Y.  Ento.   Soc.  xiii,   p.   158   (in  part). 

1907.  N.  mexicana  Kirkaldy  and  Bueno,  Proc.  Ento.  Soc.  Wash,   x,  p.   198   (in  part). 

1914.  .V.  mericana  Van  Duzee,  Trans.   San  Diego  Soc.   Nat.   Hist,  ii,   p.    33. 

1917.  .V.  mexicana  Van  Duzee,  Catalog  of  Hem.   p.   454   (in  part). 

1919.  X.  mexicana  Hungerford,  Kans.   I'niv.   Sci.   Bull,   xi,   p.   170   (in  part). 

Size.  Length,  10  mm.  to  11.5  mm.;  width  of  pronotiim.  3.9  mm. 
to  4.5  mm. 

Color.  Orange  red  and  black  to  horn  and  black.  Head,  pronotiim 
and  limbs  yellow.  Scutellum  black.  Membrane  of  hemelytra  from 
dusky  to  brownish  black.  Trochanter  and  femur  of  middle  and 
hind  legs  with  the  longitudinal  dark  stripe  characteristic  of  the 
N.  nicvicann  A.  &  S.  groui). 

Structural  Characteristics.  Head  large,  anterior  outline,  as  viewed 
from  above,  flattened;  vertex  a  little  longer  than  its  anterior  width; 
anterior  margin  of  vertex  less  convex  and  plainly  shorter  than  the 
frontal  margin  of  the  eye;  anterior  breadth  of  vertex  :  synthlipsis  : : 
3:1,  sometimes  a  little  broader.  Pronotum  about  one  and  eight- 
tenths  the  length  of  the  head;  lateral  margins  moderately  divergent 
and  nearly  straight,  somewhat  explanate  on  anterior  two-thirds, 

*  Name  in  honor  of  W.  E.  Hoffmann  and  one  "n"  omitted  in  error. 


Hungerford:    Genus  Notoxecta  69 

anterior  lateral  angles  rounded;  lateral  ledge,  as  seen  from  the  side, 
nearly  straight.  Anterior  lobe  of  membrane  of  hemelytron  larger  and 
longer  than  the  posterior  one.  Anterior  trochanter  of  the  male  with 
a  short,  inconspicuous  hook.  Mesotrochanters  rounded.  Carina  of 
fourth  abdominal  sternite  bare;  last  abdominal  sternite  of  female 
slender,  boarder  at  tip  than  middle  and  incised  at  tip.  First  pair  of 
gonapophyses  short.    INIale  genital  capsule  as  shown  on  Plate  XI. 

Location  of  Types.  In  the  Francis  Huntington  Snow  Entomo- 
logical Museum  of  the  University  of  Kansas.  Described  from  a 
series  of  insects  taken  at  Laguna  Beach,*  California,  C.  T.  Dodds. 
collector.    Other  paratypes  from  California  and  Lower  California. 


Text  Figure  3.  A'',  hojfmaymi 
Hungerford,  showing  the  venter  of 
the  abdomen  of  a  female.  Com- 
pare with  text  figures  5  and  6. 
(Subgenus  Erythronecta.) 


N.  hoffmanni. 

Comparative  Notes.  This  species  belongs  to  the  N.  mexicana  A. 
&  S.  group  and  until  1925  was  confused  with  N.  mexicana  A.  &  S. 
It  is  a  trifle  smaller  than  N.  ceres  Kirkaldy,  from  which  it  differs  in 
having  almost  straight,  not  sinuate,  lateral  margins  of  the  prothorax. 

Data  on  Distribution: 

Californu.  Laguna  Beach,  C.  T.  Dodds;  Laguna  Mts.,  July  6,  1929,  L.  D. 
Anderson;  Alpine,  July  9,  1929,  R.  H.  Beamer;  San  Diego  Co.,  July  4,  1929, 
L.  D.  Anderson;  San  Diego  Co.,  August  5,  1913,  E.  P.  VanDuzee  (Drake  Coll.) ; 
San  Diego  Co.,  April  9,  1930,  C.  &  D.  Martin;  Indio,  July  24,  1929,  L.  D. 
Anderson;  Claremont,  C.  F.  Baker  (Uhler  Coll.) ;  Monrovia  Canyon,  March  2, 
1930,  Mrs.  C.  H.  Martin;  Mission  Creek,  Santa  Barbara,  1915,  C.  H.  Kennedy; 
California,  L.  Lethierry  (Det.  by  him  as  A'',  mexicana) ;  Winters,  August  6, 
1929,  R.  H.  Beamer. 

*  Not  Long  Beach,  as  stated  in  original  description. 
6—3482 


70  The  University  Science  Bulletin 

Arizona.  Huachuca  Mts.,  5,200  ft.,  May  20,  1919,  R.  D.  Champ.;  Salome 
Creek,  Sierra  Ancha  Mts.,  Gila  Co.,  September  1,  1932,  D.  K.  Duncan;  Chiri- 
cahua  Mts.,  Pinerey  Canyon,  Hands  Ranch,  June  16,  1932,  D.  K.  Duncan. 

Lower  California,  Mexico.  El  Paraiso,  May,  1889,  Chas.  D.  Haines.  (Det. 
by  Kirkaldy  as  N.  mexicana).  Santa  Maria,  May,  1889,  Chas.  D.  Haines 
(Uhler  Coll.). 

Notonecta  lobata  Hungerford  1925 

(Color  Plate  III,  iigs.   5,  10,   11,   13;    Plates  X,  fig.   1,  and  XI,  fig   2.) 

1925.  N.  lobata  Hungerford,  Can.  Ento.  Ivii,  p.  239,  PI.  vi,  fig.  3. 
1928.  JV.  lobata  Hungerford,  Annals  Ento.  &oc.  Amer.  xxi,  p.  142. 
1931.    A^.  lobata  Bueno,  Bull.  Brooklyn  Ento.  Soc.  xxvi.  No.   3,  p.  138. 

Referring  to  this  species,  also: 

1884.  N.  mexicana  Uhler,  Standard  Nat.  Hist.  II,  p.  252. 

1897.  N.  mexicana  Kirkaldy,  Trans.  Ento.  Soc.  London  for  1897,  pp.  401,  402  (in  part). 

1901.  N.  mexicana  Champion,  Biol.  Centr.  Amer.  Hem.-Het.  II,  p.  368  (in  part). 

1905.  A'^.  mexicana  Bueno,  Jl.  N.  Y.  Ento.  Soc.  xiii,  p.  158  (in  part). 

1906.  N.  mexicana  Snow,  Trans.  Kans.  Acad.  Sci.  xx,  pt.  1,  p.  181. 

1907.  A'^.  mexicana  Snow,  Trans.  Kans.  Acad.   &ci.   xx,  pt.   2,  p.   160. 
1909.  N.  mexicana  Kirkaldy  and  Bueno,  Proc.  Ento.  Soc.  Wash,  x,  p.  198. 

1918.  N.  mexicana  Hungerford,  Ento.  News  xxix,  p.   245,  pt.  xv,  fig.  7. 

1919.  JV.  mexicana  Hungerford,  Kans.  Univ.  Sci.  Bull,  xi,  p.   170  (in  part). 

1919.  N.  mexicana  Hungerford,  Kans.  Univ.  Sci.  Bull,  xi,  pi.  xxxii,  fig.  4  (male  genital 
capsule). 

Size.  Length,  13  mm.  to  14  mm.;  width  of  pronotum.  4.3  mm.  to 
5  mm. 

Color.  Typically  red  and  black.  Some  specimens  are  brownish 
black  with  beige  trimmings  and  others  are  greenish-tan.  All  have 
the  head,  pronotum  and  limbs  yellow  or  gray  and  scutellum  and 
membrane  of  hemelytron  black.  The  hemelytra,  excepting  the  mem- 
brane, are  typically  red.  The  males  of  the  red  forms  frequently 
have  the  black  of  the  membrane  extending  foi-ward  upon  the  distal 
portion  of  clavus  and  corium.  The  pale  and  dark  forms  have  only 
the  slightest  suggestion  of  red.  Middle  and  hind  trochanters  and 
femora  with  longitudinal  median  streaks  of  brown  or  black. 

Structural  Characteristics.  Head  large,  anterior  outline,  as  viewed 
from  above,  flattened;  vertex  slightly  longer  than  its  anterior  width; 
anterior  margin  of  vertex  less  convex  and  plainly  shorter  than  the 
frontal  margin  of  the  eye ;  anterior  breadth  of  vertex  :  synthlipsis  : : 
19:7.  Pronotum  about  one  and  two-thirds  length  of  the  head,  lateral 
margins  divergent,  straight  in  male  and  nearly  so  in  female.  An- 
terior angles  normal;  lateral  ledge  as  seen  from  the  side  sinuate 
and  oblique  and  shorter  than  the  rear  margin  of  eye  below  it;  an- 
terior half  moderately  explanate.  Anterior  lobe  of  membrane  larger 
than  the  posterior  one.  Anterior  trochanter  of  the  male  with  short 
inconspicuous  hook.  Mesotrochanters  rounded.  The  keel  of  fourth 
abdominal  sternite  bare.    Last  abdominal  sternite  of  female  slender. 


Hungerford:    Genus  Notonecta  71 

not,  or  little,  broader  at  tip  than  middle,  tip  incised.  First  pair  of 
gonapophyses  short.  Last  abdominal  sternite  of  male  unusually 
broad  and  rounded;  male  genital  capsule  with  large  lobe  in  front 
of  clasper.    (See  Plate  XI,  fig.  2.) 

Location  of  Types.  In  the  Francis  Huntington  Snow  Entomo- 
logical Museum  of  the  University  of  Kansas;  seventeen  specimens 
taken  by  P.  A.  Glick  in  the  Superstition  Mountains,  Arizona. 

Comparative  Notes.  This  species  belongs  to  the  A',  mexicana  A. 
&  S.  group  and,  until  1925,  was  confused  with  N.  mexicana  A.  &  S.  It 
is  a  common  species  in  the  southwest  part  of  the  United  States. 

Data  on  Distribution: 

Arizona.  Superstition  Mts.,  November  11,  1922,  P.  A.  Glick;  Santa  Rita 
Mts.,  Alt.  8,000  ft.,  June  19,  F.  H.  Snow;  Santa  Rita  Mts.,  July  25,  1927,  and 
July  17,  1932,  R.  H.  Beamer;  Sabino  Canyon,  July  14,  1932,  R.  H.  Beamer; 
Grand  Canyon,  6  miles  below  North  Haxasn  Crk.,  September  16,  1923,  R.  C. 
Moore;  Huachuca  Mts.,  5,750  ft.,  March  16,  1919,  R.  D.  Camp  (Univ.  of 
Mich.) ;  Baboquivari  Mts.,  July  18,  1932,  R.  H.  Beamer;  Huachuca  Mts.,  July 
8,  1932,  R.  H.  Beamer;  Cochise  Co.,  July  29,  1927,  L.  D.  Anderson;  Pima  Co., 
July  27,  1927,  R.  H.  Beamer;  Gila  Co.,  August  5,  1927,  P.  A.  Readio;  Yuvapai 
Co.,  August  9,  1927,  L.  D.  Anderson;  Chiricahua  Mts.,  Pinerey  Canyon,  Hands 
Ranch,  5,000  ft.,  June  16,  1932,  D.  K.  Duncan;  S.  Catalina  Mts.,  Sabina  Canyon, 
3,000  ft.,  June  20,  1932,  D.  K.  Duncan;  Reservation  Springs,  12  miles  N.E.  of 
Globe,  September  17,  1932,  D.  K.  Duncan.  The  following  in  U.  S.  N.  M.:  Ariz. 
(Uhler  Coll.);  Sabina  Canyon,  Hubbard  Hot  Springs,  June  26,  H.  S.  Barber; 
Bright  Angel,  Colo.  Canyon  3,500  ft.,  H.  S.  Barber;  Senator  Mine,  near  Prescott, 
September  12,  1907,  H.  S.  Barber;  Catal,  Spr.  April  19,  H.  G.  Barber;  Huachuca 
Mts.;  Santa  Rita  Mts.,  July  26,  1925. 

New  Mexico.    San  Antonio,  July  15,  1927,  L.  D.  Anderson. 

Texas.  Valentine,  July  13.  1927,  L.  D.  Anderson ;  Presidio  Co..  July  15,  R. 
H.  Beamer.  The  following  in  U.S.N.M.:  Kerrville,  April  12,  1907,  F.  C. 
Pratt;  Brewster  Co.,  Chisos  Mts.,  June  10-12,  1908,  Mitchell  and  Cushman; 
Fort  Davis,  Jeff  Davis  Co.,  5,000  ft.,  Davis  Mts.,  Mrs.  O.  C.  Poling  (H.  M. 
Harris). 

Mexico.    Mex.  (Uhler  Coll.). 

Biological  Notes.  On  May  12,  1932,  Mr.  Douglas  K.  Duncan 
visited  a  small  pool  which  is  located  twelve  miles  northeast  from 
Globe,  Arizona,  on  the  Showlow  road  and  two  miles  due  east  from 
the  highway  down  a  canyon.  There  is  a  small  spring  forming  a 
small  pool  about  six  feet  by  six  feet  and  from  a  few  inches  to  eighteen 
inches  deep;  sandy  and  rock  bottom;  some  moss,  submerged  twigs, 
etc.  The  elevation  is  about  4,000  feet.  In  this  pool  he  took  all  the 
adults  he  could  find,  five  of  them,  and  noted  many  nymphs.  On 
May  16th  he  again  visited  the  pool  and  could  find  no  adults,  but 
took  more  than  fifty  nymphs  for  me,  leaving  about  twenty  others  to 
mature.    There  were  no  other  species  of  Notonecta  in  the  pool. 


72  The  University  Science  Bulletin 

Notonecta  hintoni  new  species 

(Text   figure   4) 
1934.    iV.  liintoni  Hungerford,  Jl.  Kans.  Ento.  Soc.  VII,  No.  3,  p.  97. 

Size.    Length,  14  mm.  to  15  mm.;  width  of  pronotum,  5.1mm. 

Color.  Typically  red  and  black.  Some  specimens  may  have  the 
red  replaced  by  tan.  All  have  the  head,  pronotum  and  limbs  yellow 
or  gray  and  scutellum  and  membrane  of  hemelytron  black.  The 
hemelytra  of. females  are  typically  red,  excepting  the  membrane 
and  a  spot  near  the  distal  angle  of  the  corium,  which  are  black.  The 
males  are  darker,  having  the  red  of  the  corium  and  clavus  more  or 
less  suffused  with  black.  Middle  and  hind  trochanters  and  femora 
with  longitudinal  median  streaks  of  brown  or  black. 


N   hintom 


Text  Figure  4.  Notonecta  hintoni  Hungerford, 
showing  male  genital  capsule.  Compare  with  drawings 
on  Plate  XI. 

Structural  Characteristics.  Head  large,  anterior  outline,  as 
viewed  from  above,  flattened;  vertex  of  male  slightly  longer  than 
its  anterior  width,  subequal  in  female;  anterior  margin  of  vertex 
less  convex  and  plainly  shorter  than  the  frontal  margin  of  the  eye; 
anterior  breadth  of  vertex  :  synthlipsis  : :  19:7.  Pronotum  not  more 
than  one  and  two-thirds  length  of  head;  lateral  margins  divergent, 
straight  to  faintly  concave;  anterior  angles  about  normal,  slightly 
more  produced  than  in  A^.  lobata  Hungerford;  lateral  ledge,  as  seen 
from  the  side  sinuate  and  oblique  and  shorter  than  rear  margin  of 
eye  below  it;  anterior  half  moderately  explanate.  Anterior  lobe  of 
membrane  larger  than  the  posterior  one.  Anterior  trochanter  of 
the  male  with  an  inconspicuous  hook.  Mesotrochanter  rounded. 
The  keel  of  the  fourth  abdominal  sternite  bare.  Last  abdominal 
sternite  of  female  slender,  sides  not  concave,  tip  incised.  Last 
abdominal  sternite   of  male  unusually   broad  and   rounded;   male 


Hungerford:    Genus  Notonecta  73 

genital  capsule  as  shown  in  figure  4.  The  clasper  strikingly  different 
from  any  other  member  of  the  subgenus  Erythronecta. 

Location  of  Types.  Holotype,  allotype  and  paratypes  in  the 
Francis  Huntington  Snow  Entomological  Museum.  Paratypes  will 
be  sent  to  United  States  National  Museum  and  to  British  Museum. 
Described  from  a  series  of  twenty-nine  specimens  taken  by  Mr. 
H.  E.  Hinton  in  Real  de  Arriba,  District  of  Temascaltepec.  Alti- 
tude, 1,960  meters,  Mexico. 

Comparative  Notes.  General  appearance  that  of  N.  lobata 
Hungerford.  The  males  are  distinguished  from  that  species  by  the 
very  different  shape  of  the  genital  capsules  and  claspers.  The 
females  by  having  only  an  inconspicuous  notch  at  tip  of  penultimate 
abdominal  sternite.  Lateral  margins  of  last  abdominal  sternite  of 
female  not  as  curved  as  in  A^.  lobata  Hungerford. 

Data  on  Distribution.    Known  only  from  the  type  locality. 

Notonecta  compacta  Hungerford  1925 

(Color  Plate  III,  fig.  6;   Plate  XI,  figs.  3  and' 8;   and  text  fig.   5) 

1925.    N.   compacta  Hungerford,  Can.  Ento..  Ivii,  p.   239. 

1933.    A^.  compacta  Hungerford,  Bull.  Brook.  Ento.  Soc,  xxviii,  p.   135  (allotype    ^  ). 

Referring  to  this  species,  also: 

1897.    N.  mexicana  Kirkaldy,  Trans.  Ento.  Soc.  London  (in  part). 

1901.  A^.  mexicana  Champion,  Biol.  Centr.  Amer.  Hem.  Het.  II,  p.  369  (Amula  in  Guer- 
reio). 

A  dark  male  from  Amula  studied  by  Champion  and  another  by 
Kirkaldy,  labeled  N.  mexicana  A.  and  S.  by  them.  There  is  another 
specimen  in  Oxford  Museum  labeled  by  Kirkaldy. 

Size.  Length,  12.5  mm.  to  14  mm.;  width  of  pronotum,  4.8  mm. 
to  5.7  mm. 

Shape.  A  comparatively  short,  plump-bodied  species.  The  lat- 
eral margins  of  the  prothorax  in  the  female  strongly  constricted  in 
the  middle. 

Color.  Females  are  usually  red  and  black  and  males  nearly  black. 
I  have  some  females  that  are  black  like  the  males.  Head,  pro- 
notum  and  limbs  dark  testaceous.  Femora  with  dark  longitudinal 
stripes  beneath  and  the  trochanters  each  with  a  dark  elongate  spot. 

Structural  Characteristics.  Head  large;  anterior  outline  of  head 
viewed  from  above,  flattened ;  vertex  slightly  longer  than  its  anterior 
width  in  the  male,  subcqual  in  the  female;  anterior  margin  of  vertex 
less  convex  and  plainly  shorter  than  the  frontal  margin  of  the  eye ; 
anterior  breadth  of  vertex  :  synthlipsis  ::  3  +  :1.  Pronotum  about 
twice  the  length  of  the  head  in  female.  Head  of  male  longer  than  half 


74 


The  University  Science  Bulletin 


of  the  pronotum;  lateral  margins  divergent,  moderately  constricted 
in  the  male  but  very  strongly  and  characteristically  constricted  in 
the  female.  Anterior  angles  embracing  the  eyes.  Lateral  ledge 
unlike  in  the  two  sexes,  sigmoid  and  oblique  in  both  sexes  as  seen 
from  the  side;  in  the  female  the  ledge  is  heavy  and  pronounced  on 
the  anterior  half  and  almost  obliterated  and  curved  on  the  posterior 
half;  in  the  male  the  anterior  portion  of  the  ledge  is  broader  than 
elsewhere  but  not  interrupted;  anterior  end  of  the  ledge  deflected  in 
both  sexes,  margin  of  prothorax  shorter  than  the  rear  margin  of  the 
eye  below  it.  Scutellum  a  little  longer  than  the  pronotum.  Pos- 
terior and  anterior  lobes  of  membrane  subequal,  the  posterior  some- 
times a  trifle  longer.  Anterior  trochanters  of  the  male  with  broad, 
blunt  hook.  Mesotrochanters  not  angulate.  The  carina  or  keel 
of  the  fourth  abdominal  sternite  bare;  this  segment  is  short  in  the 
male  and  long  in  the  female.  In  the  male  the  penultimate  ab- 
dominal sternite  is  enlarged.  The  terminal  abdominal  sternite  of 
the  female  is  as  broad  as  long  (exposed  part)  with  lateral  edges  con- 
cave and  tip  incised.  First  pair  of  gonapophyses  short.  Male  geni- 
tal capsule  as  shown  on  Plate  XI. 


N  compacta 


N.m. 


mexicana 


Text  Figure  5.     Notonecta  covipacta  Hungerford  and  Notonecta  m.  mexicana  A.   &  S. 
Underside  of  abdomen  of  females.     (Subgenus  Erythronecta.) 


Location  of  Type.    The  type  is  in  the  U.  S.  N.  M. 
Comparative  Notes.    This  species  belongs  to  the  N.  mexicana  A. 
and  S.  group.    The  last  abdominal  sternite  of  the  female  is  more 


Hungerford:    Genus  Notonecta  75 

nearly  quadrate  than  in  the  other  species.    No  other  species  has  the 
pronotum  so  conspicuously  constricted  laterally. 

Data  on  Distribution.  Colima,  Mexico,  coll.  by  L.  Conradt; 
Amula,  Guerrero,  6,000  ft.,  H.  H.  Smith;  Between  Cajones  and 
Rincon,  S.  of  Chilpancingo,  Guerrero,  Mex.,  July  1,  1932,  Hobart 
Smith  (18  specimens)  ;  Near  Taxco,  Guerrero,  Mex.,  July  4,  1932, 
Hobart  Smith  (2  specimens).  A  male  of  the  Hobart  Smith  collec- 
tion is  the  allotype. 

Notonecta  mexicana  mexicana  Amyot  and  Serville,  1843 

(Color  Plate  III,  fig.  8;  Plate  XI,  fig.  11;  text  figure  5.) 

1843.    .V.  mexicana  Amyot  &  Serville,  Hemipteres  p.   453,  PI.   viii,  fig.   7. 

1851.    -V.  mexicaiia  Amyot  &  Serville;   Fieber,  Rhynchotographieen,  p.   475. 

1851.    A',  mexicana  Amyot  &  Serville;   Fieber,  Rhynchotographien,  p.   51.      (Van  Duzee.) 

1S53.  A',  mexicana  Herrich-Schaeffer,  Wanzen.  Ins.  ix,  p.  43,  Tab.  ccxciv,  fig.  903  (in 
color). 

1884.    A',  mexicana  Uhler,  Standard  Nat.  Hist.,  ii,  p.  252  (in  part). 

1897.  .V.  mexicana  Amyot  &  Serville;  Kirkaldy,  Trans.  Ento.  Soc.  London  for  1897. 
p.   401  (in  part). 

1901.  A'',  mexicana  Amyot  &  Serville;  Champion,  Biol.  Centr.  Amer.  Hem.-Het.  ii,  p. 
368-369  (in  part,  but  not  figures).     (Mexico,  Salle;   Oaxaca,  Salle;    San  Luis  Potosi,  Palmer.) 

1904.  A',  mexicana  Amyot  &  Serville;  Kirkaldy,  Wien  Ento.  Zeit.,  xxiii,  pp.  94  and  132 
(in  part). 

1905.  N.  mexicana  Amyot  &  Serville;  Bueno,  Jl.  N.  Y.  Ento.  Soc.  xiii,  pp.  145,  150,  158. 
(Ref.  not  localities.) 

1909.  N.  mexicana  Amyot  &  Serville;  Kirkaldy  &  Bueno,  Proc.  Ento.  Soc.  Wash.,  x, 
p.   198  (in  part). 

1917.    A",   mexicana  Amyot  &  Serville;   Van  Duzee,  Catalogue  Hemip.,  p.  454  (in  part). 
1919.    N.  mexicana  Amyot  &  Serville;    Hungerford,   Kans.   Univ.    Sci.   Bull,   xi,   (in   part). 
1925.    .V.  mexicana  Amyot  &  Serville;   Hungerford,  Can.  Ento.  Ivii,  p.   238. 
1928.    A^.  mexicana  Amyot  &  Serville;    Hungerford,  Jl.  Kans.  Ento.   Soc.   v,  p.   53. 

Referring  to  this  species,  also : 

1851.    A^.  klugii  Fieber,  Rhynchotographieen  p.  475  (in  part). 

1851.    A'',  klugii  Fieber,  Rhynchotographien  p.   51   (Van  Duzee). 

1925.     -V.     macroccphala  Hungerford,   Can.   Ento.  Ivii,  p.   241,  PI.   0,  fig.   5.      (Syn.  new.) 

Size.  Length,  13  mm.  to  14  mm.;  width  of  pronotum,  4.6  mm. 
to  5.1  mm. 

Color.  Typically  a  red  and  black  species.  The  males  a  little 
darker  than  the  females  as  a  rule.  Probably  there  exist  black  and 
tan  females  and  black  males,  but  I  have  never  seen  them.  Anterior 
half  of  prothorax,  head  and  limbs  yellow,  the  middle  and  hind  tro- 
chanters and  femora  with  a  median  longitudinal  dark  stripe,  front 
femur  may  have  two  such  stripes  on  its  rear  margins,  posterior  half 
of  pronotum  may  be  darkened  by  the  black  mesonotum  beneath. 
Scutellum  black.*  Clavus  and  corium  orange  red  to  red.  Smoky 
or  black  areas  may  occur  along  hemelytral  suture,  beneath  corial- 

*  Footnote:  I  have  seen  a  specimen  or  two  with  yellowish  scutellum  that  were  otherwise 
fairly  well  colored. 


76  The  University  Science  Bulletin 

claval  suture,  on  the  embolium,  on  outer  half  of  corium  and  on  mem- 
brane, which  is  nearly  always  dark. 

Structural  Characteristics.  Head  large,  embraced  at  base  by  the 
anterior  angles  of  prothorax.  Anterior  outline  of  the  head,  viewed 
from  above,  flattened;  vertex  subequal  in  length  to  its  anterior 
width,  sometimes  in  the  male  a  little  longer,  anterior  margin  of  vertex 
less  convex  and  plainly  shorter  than  the  frontal  margin  of  the  eye; 
anterior  breadth  of  vertex  :  synthlipsis  : :  17:7.  Pronotum  not  quite 
twice  the  length  of  the  head,  lateral  margins  moderately  divergent, 
somewhat  concave  in  the  male,  more  so  in  the  female.  Anterior 
angles  acute  and  embracing  the  eyes,  more  pronounced  in  the  male 
than  in  the  female.  Lateral  ledge  moderately  broad  and  horizontal 
on  anterior  half  and  oblique  and  nearly  obliterated  on  posterior  half ; 
anterior  angle  slightly  deflected  in  both  sexes;  margin  of  prothorax 
shorter  than  the  rear  margin  of  the  eye  beneath.  Anterior  lobe  of 
membrane  of  hemelytra  a  little  longer  than  the  posterior.  Anterior 
trochanters  of  male  with  a  hook  of  moderate  size,  mesotrochanters 
rounded,  the  carina  of  the  fourth  abdominal  sternite  bare,  the  ter- 
minal abdominal  sternite  of  female  slender,  elongate,  but  slightly 
constricted  and  the  tip  incised.  First  pair  of  gonapophyses  rather 
slender.    Male  genital  capsule  as  shown  on  Plate  XL 

Location  of  Type.  I  have  been  unable  to  find  the  types  in  Paris 
or  elsewhere. 

Comparative  Notes.  The  description  I  have  given  above  applies 
to  the  species  that  fits  best  Amyot  and  Serville's  brief  description 
and  the  color  drawing.  The  size  (14  mm.),  the  parallel  sides  and 
the  distribution  point  to  this  species.  In  former  writings  I  have 
been  confused  by  Kirkaldy  and  Champion  and  believed  N.  mexicana 
A.  &  S.  to  be  something  else.  I  now  conclude  that  my  A',  macro- 
cephala  is  only  an  unusual  specimen  of  this  species  and,  therefore, 
suppress  it.  I  described  it  from  a  single  specimen  with  an  abnormally 
large  head,  but  the  genitalia  are  indistinguishable  from  the  species 
described  above.  The  slender,  tapering  last  abdominal  sternite  of 
the  female  distinguishes  this  species. 

Data  on  Distribution.  Described  from  "Mexico,"  and  unfortu- 
nately most  of  the  specimens  I  have  seen  bear  no  more  precise  label. 
However,  in  the  Museum  at  Vienna  there  is  a  specimen  labeled  "San 
Luis  Potosi,  Dr.  Palmer."  This  was  determined  by  Champion  as 
N.  mexicana.  There  is  another  specimen  from  Signoret's  collection 
labeled  "Bogota."  There  are  two  specimens  from  Mexico  by  Salle 
in  the  museum  at  Stockholm. 


Hungerford:    Genus  Notonecta  77 

Notonecta  mexicnna  v.  creaseri  Hungerford  1932 

(Color  Plate  II,  fig.  13;  Plate  XI,  figs.  1  and  9.) 
1932.    N.  mexicarta  v.  creaseri,  Hungerford,  Jl.  Kans.  Ent.  Sec,  v.,  p.  53. 

Size.    Length,  12  mm.;  width  of  pronotum,  4.3  mm. 

Color.  The  four  specimens  at  hand  are  tan  and  black  with  one 
specimen  showing  some  flecks  of  red  on  the  corium.  Head  and  legs 
testaceous.  The  femur  with  a  longitudinal  stripe  which  is  charac- 
teristic of  entire  group.  Scutellum  is  black  or  nearly  so,  three  of  the 
specimens  with  lateral  margins  tan.  The  clavus  and  corium  mostly- 
tan.  The  distal  third  of  corium  more  or  less  dark.  Membrane  nearly- 
black.  It  is  probable  that  this  variety,  like  other  species  in  the 
group,  may  have  red  or  nearly  black  in  place  of  the  tan  on  the 
hemelytra. 

Structural  Characteristics.  Anterior  outline  of  the  head,  viewed 
from  above,  flattened ;  vertex  longer  than  its  anterior  width ;  margin 
of  vertex  less  convex  and  shorter  than  frontal  margin  of  the  eye; 
anterior  breadth  of  vertex  :  synthlipsis  ::  11:4  (in  the  female).  In 
the  male  the  synthlipsis  is  a  little  narrower.  Pronotum  about  twice 
length  of  head;  lateral  margins  short,  divergent,  but  quite  concave 
in  the  females  and  plainly  so  in  the  males;  anterior  angles  embracing 
the  eyes ;  lateral  ledge  as  seen  from  the  side  with  anterior  three-fifths 
broad  and  nearly  horizontal  and  posterior  two-fifths  faint  and 
oblique;  the  anterior  angle  deflexed;  the  length  of  the  ledge  less 
than  the  rear  margin  of  the  eye  below  it.  Anterior  lobe  of  the 
membrane  of  the  hemelytron  distinctly  longer  than  the  posterior 
lobe.  The  anterior  trochanter  of  the  male  with  a  rather  narrow, 
pointed  hook.  Mesotrochanter  rounded.  The  genitalia  in  both 
sexes  like  A'',  mexicana  A.  &  S.  (See  Plate  XL) 

Location  of  Types.  Described  from  four  specimens  labeled, 
"Mexico,  Nueva  Leon,  Santiago,  April  19,  1930,  13.  Creaser-Gor- 
don."  Holotype  and  allotype  in  Museum  of  Zoology,  Michigan 
University.  Paratype  in  Entomological  Museum,  University  of 
Kansas. 

Comparative  Notes.  This  is  smaller  than  the  typical  A^.  mexicana 
A.  &  M.,  but  in  the  characters  of  the  genitalia  indistinguishable 
from  it. 


78  The  University  Science  Bulletin 

Notonecta  robusta  Hungerford  1932 

(Color  Plate  II,  fig.   14;    Plate  XI,  figs.   4  and   10) 
1932.    .V.   robusta  Hungerford.  Jl.  Kans,  Ento.  Soc,  v,  p.  54. 

Size.  Length  (of  male),  a  trifle  less  than  15  mm.;  the  width  of 
pronotum,  5.1  mm. 

Color.  The  holotype  has  the  head,  pronotum  and  limbs  dark 
testaceous.  The  trochanters  and  femora  marked  as  in  other  species 
of  the  N.  mexicana  A.  &  S.  group.  Scutellum  black  and  hemelytra 
nearly  black.  No  doubt  this  species  may  have  hemelytra  that  are 
tan  or  red  with  black  membrane. 

Structural  Characteristics.  Anterior  outline  of  the  head,  viewed 
from  above,  flattened;  vertex  about  as  long  as  the  anterior  width; 
margin  of  vertex  less  convex  and  shorter  than  the  frontal  margin  of 
an  eye;  anterior  breadth  of  vertex  :  synthlipsis  : :  10.5:4.  Pronotum 
about  twice  length  of  head;  lateral  margins  divergent  and  nearly 
straight  in  the  male ;  anterior  angles  not  embracing  the  eyes ;  lateral 
ledge,  as  seen  from  the  side,  slightly  undulate,  upturned  at  both  ends, 
oblique  and  shorter  than  the  rear  margin  of  the  eye  below  it. 
Scutellum  and  ridge  of  hemelytral  suture  equal  in  length.  Posterior 
lobe  of  membrane  a  little  longer  than  anterior  lobe.  The  anterior 
fouteri  margin  of  the  membrane  constricted  at  the  embolial  suture 
and  undulate.  Anterior  trochanter  of  male  with  short  hook.  Meso- 
trochanter  rounded.  Male  genital  capsule  much  like  A^.  mexicana 
A.  &  S.,  but  the  clasper  with  the  tip  broader  and  the  rear  margin 
more  concave.     (See  Plate  XI,  figs.  4  and  10.) 

Location  of  Type.  Described  from  a  single  male  specimen  which 
bears  no  locality  or  collector's  label.  Deposited  in  Entomological 
Museum,  University  of  Kansas. 

Com'parative  Notes.  Agrees  with  other  species  of  N.  mexicana 
A.  &  S.  group  in  having  the  keel  of  fourth  ventral  abdominal  segment 
bare,  the  hairs  confined  to  the  sides.  Head  broad,  eyes  large  and 
broad,  rear  margin  broader  than  the  length  of  the  lateral  margin  of 
prothorax.  Scutellum  conspicuously  broader  than  long.  It  differs 
from  N.  lobata  Hungerford  in  having  the  last  abdominal  sternite  of 
male  normal  in  size.  I  have  two  female  specimens,  red  and  black 
in  color,  from  San  Cristobal,  Chiapas,  Mexico,  A.  Dampf.,  that  are 
the  broadest,  most  robust  Notonecta  known  to  me.  The  lateral 
margins  of  the  pronotum  are  greatly  constricted  near  the  front  so 
that  the  anterior  angles  flare  out.  The  very  marked  undulation  of 
the  anterior  (outer)  margin  of  membrane  suggests  to  me  that  these 
may  be  females  of  A^  robusta  Hungerford. 


Hungerford:    Genus  Notonecta  79 

Notonecta  ceres  ceres  Kirkaldy  1897 

(Color  Plate  III,  figs.  2  and  3 ;   Plate  XI,  fig.  7;   text  figure  6) 

1897.    A',    mexicana  var.   ceres  Kirkaldy,   Ento.   Soc.   London,   p.   402,   1897. 

1904.  A',  mexicana  var.   ceres  Kirkaldj',  Wiener  Ento.  Zeit.,  xxiii,  p.   132. 

1905.  A',   mexican  var.   ceres  Kirkaldy;    Bueno.   Jl.  N.  Y.  Ento.  Soc,  xiii,  p.   l.'iO. 

1917.    A',   mexicana  var.   ceres  Kirkaldy;    Van  Duzee,  catalogue  of  Hemiptera    .    .    ji.    454. 
1925.    A',    mexicana    var.    ceres    Kirkaldy;     Hungerford,    Can.    Ento.,    Ivii,    p.    238.      (Says 
are  all  females,  the  variety  hades  all  males.) 

Referring  to  this  species,  also: 

1851.    A',    klugii  Fieber,  Rhynchotographieen,  p.   475  (in  part). 

1897.  A',  mexicana  var.  hades  Kirkaldy.  Ento.  Soc.  London,  p.  402,  1897.  This  is  only 
the  male  of  A'',  ceres  Kirkaldy.     See  list  of  references  above. 

1901.  A',  jnexicana  Champion.  Biol.  Centr.  Am.  Hem.-Het.,  11,  p.  369,  Tab.  22,  figs. 
6,  6a-d   (in  part). 

Size.  Length.  11mm.  to  12  mm.;  width  of  pronotum,  4.3  mm.  to 
4.8  mm. 

Color.  Red  and  black,  tan  and  black,  or,  in  case  of  males,  black. 
Anterior  half  of  pronotum,  head  and  limbs  yellow,  the  middle  and 
hind  trochanters  and  femora  with  the  dark  stripe  usual  for  the 
A'',  mexicana  A.  &  S.  group. 

Structural  Characteristics.  Head  large;  anterior  outline  of  head 
viewed  from  above,  flattened;  vertex  a  little  longer  than  its  anterior 
width;  anterior  margin  of  vertex  less  convex  and  plainly  shorter 
than  the  frontal  margin  of  the  eye;  anterior  breadth  of  vertex  : 
synthlipsis  ::  11:3.  Pronotum  nearly  twice  to  twice  the  length  of 
the  head;  lateral  margins  concave  and  moderately  divergent;  the 
margin  more  concave  in  females  than  in  males;  anterior  angles 
slightly  embracing  the  eyes,  but  rounded;  lateral  ledge  broader 
anteriorly  in  the  female  than  in  the  male,  as  seen  from  the  side, 
strongly  curved,  pronounced  on  anterior  half,  only  a  ridge  and 
curved  on  posterior  half;  margin  of  prothorax  shorter  than  the  rear 
margin  of  the  eye  below  it.  Anterior  lobe  of  membrane  of  hemelytra 
a  little  longer  than  the  posterior  one.  Anterior  trochanters  of  male 
with  moderate  hook.  Mesotrochanters  rounded.  The  carina  of 
fourth  abdominal  sternite  bare.  The  terminal  abdominal  sternite  of 
female  narrow,  broader  at  tip,  which  is  broadly  incised.  First  pair 
of  gonapophyscs  short.    Male  genital  capsule  as  shown  on  Plate  XI. 

Location  of  Types.  In  the  Kirkaldy  collection  at  U.  S.  N.  Museum 
at  Washington. 

Comparative  Notes.  Described  by  Kirkaldy  as  a  A'ariety  of  A^. 
mexicana  A.  &  S.,  but  really  a  distinct  species.  Kirkaldy  described 
the  red  and  black  females  as  N.  mexicana  var.  ceres  and  the  black 


80  The  University  Science  Bulletin 

males  as  N.  mexicana  var.  hades.  I  have  pointed  out  previously 
that  these  are  a  single  species.  This  species  is  smaller  than  N. 
mexicana  A.  &  S.  and  a  trifle  larger  than  N.  koffmanni  Hungerford. 
It  is  distinguished  from  the  former  by  the  less  acute  anterior  angles 
of  the  prothorax  and  from  the  latter  by  having  the  lateral  margins 
of  prothorax  concave. 


N.  ceres  stirtoni  N  ceres  ceres 


Text  Figure  6.     Notonecfa  ceres  stirtoni  Hungerford  and  A',  ceres  ceres  Kirkaldy.     Underside 
of  abdomen  of  females.      (Subgenus  Erythronocta.) 

Data  on  Distribution.  Costa  Rica:  San  Jose,  C.  R.  1160  M. 
August,  1905,  P.  Biolley.;  La  Palma,  1600  M.  P.  Biolley  (Det.  by 
Kirkaldy  as  var.  of  A^.  mexicana)  ;  Rio  Sarapiqui,  Alt.  2,000  m.  Hein- 
rich  Schmidt;  Rio.  Virilla,  December  26,  1931,  Heinrich  Schmidt; 
R.  Susio,  H.  Rogers. 

Notonecta  ceres  stirtoni  new  subsp. 

(Text  figure  6) 

1928.  N.  mexicana  var.  ceres,  Kirkaldy;  Hungerford,  Pan.  Pac.  Ento.,  iv,  No.  3,  p.  119. 
(Taken  mating  with   var.   hades.) 

In  this  subspecies  the  lateral  margins  of  the  prothorax  of  the  male 
are  almost  straight  as  seen  from  above.  In  the  female  the  penulti- 
mate sternite  projects  broadly  in  the  middle,  whereas  in  N.  ceres 
Kirk,  the  portion  of  the  sternite  surpassing  the  segment  is  narrow. 

Location  oj  Types.  Francis  H.  Snow  Coll.  From  El  Salvador,  C.  A. 


Hungerford:    Genus  Notonecta  81 

Notonecta  ceres  rogersi  Hungerford,  1932 

(Color  Plate  II,  fig.   12) 
1932.    ,V.  rogersi  Hungerford,  Jl.  Kansas  Ento.  Soc,  v.  p.  55. 

This  subspecies  is  larger  than  the  typical  form.  The  male  clasper 
is  almost  identical  with  N.  ceres  Kirk.  When  I  described  this  species 
I  was  confusing  A^.  ceres  stirtoni  with  N.  ceres  Kirkaldy.  I  have  one 
female  which  I  assign  to  this  subspecies.  In  this  red  and  black 
specimen  the  lateral  margins  of  pronotum  are  constricted  in  front 
of  the  middle  instead  of  near  the  middle  as  in  typical  N.  ceres 
Kirkaldy. 

Notonecta  impressa  Fieber,  1851 

1851.    .V.  impressa  Fieber,  Rhynchotographieen  Abh.  Bohm.  Gesel.  Wis.   (5),  vii,  p.  475. 

1897.  A',  impressa  Fieber;  Kirkaldy,  Trans.  Ento.  Soc.  London,  1897,  p.  403  (Made  it 
a  synonym  of  N.  insulata  Kirby  in  error). 

1901.  A',  impressa  Fieber;  Champion,  Biologia  Centr.  Americana,  Rhynchota,  ii,  p.  368 
(Places  doubtfully  under  N.  mexicana  A.  &  S.  in  error). 

1908.  A^.  impressa  Fieber;  Kirkaldy  and  Bueno,  Proc.  Ento.  Soc.  Washington,  x,  p.  198 
(Places  as  syn.  of  N.  insulata  Kirby  in  error). 

1917.    A',  impressa  Fieber;   Van  Duzee,  Cat.  of  Hemiptera  of  America,  p.  453. 

1925.  A'',  impressa  Fieber;  Hungerford,  Can.  Ento.,  Ivii,  p.  241  (Says  not  A'^.  insulata 
Kirby). 

Size.    Female  nearly  14  mm.  long. 

Shape.  Much  like  A^.  montezuma  Kirkaldy;  eyes  less  prominent; 
pronotum  transversely  crossed  by  a  strong  depression  that  does  not 
extend  to  the  margins. 

Color.  A  red  and  black  species.  The  head  and  limbs  of  usual 
color;  scutellum  black;  hemelytra  red  with  black  markings  such  as 
in  A^.  montezuma  Kirkaldy. 

Structural  Characteristics.  Head  not  prominent;  the  face  deeply 
depressed  with  surface  transversely  wrinkled;  a  longitudinal  ridge 
extending  from  the  depression  to  the  labrum;  anterior  margin  of  ver- 
tex :  synthlipsis  : :  6.7  :  2.7.  Pronotum  and  head  sloping  downward. 
Scutellum  large.  Hemelytral  suture  short.  Mesotrochanter  rounded. 
Ventral  abdominal  keel  without  hairs  on  the  ridge.  Last  abdominal 
sternite  of  female  broad;  short  and  depressed  on  the  sides. 

Location  of  Type.  In  the  Zoological  Museum  at  Berlin  I  found 
a  female  specimen  labeled,  "Notonecta  impressa"  "3623,  a  green 
label  "Mexico  Westw.,"  a  green  label  "Impressa  Fieb."  This  fits 
Fieber's  description  exactly!  The  head  is  not  prominent.  The 
synthlipsis  :  anterior  margin  of  vertex  : :  2.7  :  6.7,  as  seen  from  above 
with  the  body  horizontal.  The  pronotum  slopes  downward,  and  the 
head  follows  the  same  slope;  the  face  is  deeply  depressed,  the  sur- 
face of  the  depression  uneven  and  transversely  wrinkled.  Below 
the  depression  there  is  a  median  longitudinal  ridge  to  the  labrum; 


82  The  University  Science  Bulletin 

transverse  wrinkles  across  this  part  of  the  face;  labrum  cross- 
wrinkled.  The  pronotum  is  tranversely  crossed  just  in  front  of  the 
middle  by  a  depression  that  does  not  extend  to  the  margins  of  the 
pronotum,  which  are  straight.  The  anterolateral  angles  embrace  the 
eyes;  lateral  margins  are  moderately  ledged.  Mesotrochanter  is 
rounded;  ventral  abdominal  keel  is  without  hairs  on  the  ridge.  The 
comparative  longitudinal  measurements  from  above  are  as  follows: 
Head  :  Pronotum  :  Scutellum  :  Hemelytral  suture  (from  tip  of  scu- 
tellum)  :  tip  of  hemelytral  suture  to  wing  tip  ::  2.5  :  4.9  :  5.7  :  3.7  : 
10+. 

Coiwparative  Notes.  The  color  is  quite  like  N.  montezuma 
Kirkaldy,  but  it  is  a  little  smaller  than  the  female  N.  montezuma 
Kirkaldy  from  Valentine,  Texas,  with  which  I  compared  it.  The 
eyes  seem  smaller  and  the  pronotum  embraces  the  eyes  a  little  more. 
The  eyes  seem  flatter  on  top  and  the  transverse  pronotal  depression 
more  marked.  The  shape  of  the  last  ventral  abdominal  plates  is 
the  same  in  both  females.  While  I  am  retaining  N.  montezuma 
Kirkaldy  as  a  separate  species,  an  examination  of  a  long  series  may 
discover  specimens  like  Fieber's  female.  Since  A^.  montezuma 
Kirkaldy  in  the  Hope  Collection  came  from  the  Westwood  Collec- 
tion, according  to  information  given  by  Professor  Poulton  to  Mr. 
Champion,  instead  of  "West  Mexico"  as  Kirkaldy  recorded,  there 
is  additional  doubt  about  Kirkaldy's  species  being  specifically  dif- 
ferent from  Fieber's. 

N.  impressa  Fieber  is  not  N.  compacta  Hungerford,  which  also  has 
transversely  depressed  pronotum.  My  N.  compacta  has  sides  of 
pronotum  strongly  constricted,  N.  impressa  Fieber  does  not.  It  is 
not  N.  mexicana  A.  &  S.,  because  the  lateral  edge  of  the  prothorax 
of  A^.  impressa  Fieber  is  straight  and  the  ledge  is  even  and  longer, 
the  pronotum  longer,  head  smaller,  scutellum  very  large,  and  the 
hemelytral  suture  short.  A^  impressa  Fieber  is  not  A'',  kirbyi 
Hungerford,  which  has  a  much  wider  synthlipsis. 

Xotonecta  montezuma  Kirkaldy,  1897 

(Color  Plate  II,   fig.   1;    Plates  X,   fig.    2,   and  XV  fig.    4.) 

1897.    N.  montezuma  Kirkaldy,  Trans.  Ento.  See.  London,  1897,  p.   402. 
1901.    N.  montezuma  Kirkaldy;   Champion,  Biol.  Central!  Americana  Heteroptera,  vol.   II, 
p.  369,  pi.  22,  figs.  8  and  9. 

1904.  A'',  montezumn  Kirkaldy,  Wien.  Ento.  Zeit.,  xxiii,  pp.  94  and  132. 

1905.  N.  montezuma  Bueno,  Jl.  N.  Y.  Ento.  Soc,  xiii,  p.  162.  (Records  specimen  from 
California  in  error.*) 

*  The  specimen  labeled  "California"  in  the  American  Museum,  and  determined  by  Bueno 
as  A^.  montezuma  Kirk.,  is  an  oriental  species  with  erroneous  locality  label. 


Hungerford:    Genus  Notonecta  83 

1919.  N.  montezuma  Kirkaldy ;  Hungerford,  Kansas  Univ.  Sci.  Bull.,  xi,  pp.  108,  172 
and  331. 

1923.    A^.  montezuma  Kirkaldy;    Hungerford,  Kansas  Univ.   Sci.  Bull.,  xiv,  p.   42*;. 

1928.  A'^.  m^ntezumu  Kirkaldj';  Hungerford,  Annals  Ento.  9oc.  Am.,  xxi,  p.  1-12,  pi.  ix, 
fig.  8.     (Fig.  male  genital  capsule  and  records  species  from  Valentine,  Texas.) 

Size.    Length,  14  ram. -15  mm.;  width  of  pronotum,  4.8  ±  ram. 

Shape.    A  rather  elongate  species. 

Color.  A  red  and  black  species.  Head  and  legs  of  usual  color, 
often  marked  with  green.  Pronotum  often  having  a  median  dark 
spot  a  little  behind  the  anterior  margin.  Scutellum  black.  Hemelytra 
orange-red  with  black  markings;  the  clavus  may  be  marked  with 
dark  spots,  especially  at  the  tip  where  they  join  and  form  a  part  of 
an  irregular  black  band  that  traverses  the  corium  at  this  level; 
membrane  black.  Venter  dark  except  the  sides  of  prothorax  beneath 
the  ledge,  the  distal  end  of  the  metacoxal  shields  (extension  of 
mesothoracic  scutellum  according  to  Rich  1918),  the  connexivum 
and  median  abdominal  keel  which  are  usually  lighter  in  color. 

Structural  Characteristics.  Anterior  outline  of  the  head  viewed 
from  above,  rather  flattened;  face  transversely  depressed;  vertex 
longer  than  its  anterior  width ;  anterior  margin  of  vertex  less  convex 
and  shorter  than  the  anterior  margin  of  an  eye;  anterior  breadth  of 
vertex  :  synthlipsis  ::  8:3  (a  little  more  or  less) ;  head  a  little  more 
than  half  as  long  as  pronotum.  Lateral  margins  of  pronotum  only 
moderately  divergent  and  straight;  anterolateral  angles  somewhat 
embracing  the  eyes;  lateral  ledge,  as  seen  from  the  side,  nearly 
straight,  faintly  sigmoid  and  somewhat  oblique,  slightly  longer  than 
the  rear  margin  of  the  eye  below  it.  Scutellum  large,  longer  than 
the  pronotum.  Heraelytral  suture  short,  the  ridge  of  the  heraelytral 
suture  shorter  than  the  scutellura.  Anterior  lobe  of  membrane  longer 
than  the  posterior  lobe.  Anterior  trochanter  of  the  male  with  a 
black  pointed  tubercle  directed  only  slightly  forward  in  place  of 
the  hook.  Mesotrochanter  rounded.  Anteapical  tooth  of  middle 
femur  normal.  Ventral  abdominal  keel  smooth  and  thickened  down 
the  entire  length  of  the  fourth  and  fifth  segments  and  a  smooth 
space  down  the  middle  of  the  following  segment.  The  terminal 
abdominal  sternite  of  female  short,  broad  and  depressed  on  the  sides. 
First  pair  of  gonapophyses  of  female  intermediate  in  length.  Male 
genital  capsule  as  shown  in  Plate  XV,  fig.  4. 

Location  of  Types.  Male  and  female  in  Hope  Collection  at  Ox- 
ford, England.  I  find,  upon  examining  the  types,  that  Doctor 
Champion's  descriptive  notes  and  figures  are  excellent. 

Data  on  Distribution.    "W.  Mexico"  does  not  mean  West  Mexico, 


84  The  University  Science  Bulletin 

according  to  Professor  Poulton,  but  simply  that  specimens  are  from 
Westwood's  collection. 

In  1928  I  recorded  11  specimens  taken  by  the  University  of  Kan- 
sas party  as  follows:  10  specimens,  Valentine,  Texas,  July  13,  1927, 
R.  H.  Beamer.  One  specimen,  Presidio  County,  Texas,  July  16, 
1927,  R.  H.  Beamer.  Another,  Alpine,  June  5,  1927  (U.  S.  N.  M.). 
I  have  also  two  male  specimens  taken  by  Hobart  Smith  near  Santa 
Rosa,  Guanajuata,  Mexico,  August  14,  1932.  In  the  Stockholm 
Museum  there  is  a  male  labeled  "Mexico"  "Salle."  I  have  also  seen 
the  specimen  in  the  Paris  Museum  labeled  "Nord  de  ITnde  39-42" 
which  was  mentioned  by  Kirkaldy.    The  locality  is  indeed  erroneous. 

Notonecta  insulata  Kirby,  1837 

(Color  Plate  III,  fig.   1;   Plate  XV,  fig.  1.) 

1837.    TV.  insulata  Kirby,  in  Richardson's  Faun.  Bor.  Am.,  iv,  p.  285. 

1S78.    N.  insulata,  Kirby;   in  Bethune  reprint,  p.  140;   same  from  Can.  Ento.,  x,  p.  216. 

1851.    A'',   insulata  Kirby;  Fieber,  Rhynchotographieen,  p.  479. 

1851.    A',  insulata  Kirby;   Fieber,  Rhynchotographien,  p.  55   (Van  Duzee's  Catalogue.). 

187G.  A',  insulata  Uhler;  Bull.  U.S.  Geol.  Geog.  Surv.  II,  p.  73.  (Remarks  about  Balti- 
more, Md.,  specimens  refer  to  this  species.) 

1878.  N.  insulata  Kirby;  Uhler,  Proc.  Boston  Soc.  Nat.  Hist.,  xix,  p.  442,  "No.  17  Harris' 
Collection  July  1,  1823." 

1894.  A',  insulata  Kirby;  Van  Duzee,  Bull.  Buffalo  Soc.  Nat.  Sci.,  v,  p.  186.  (Not  un- 
common in  stagnant  pools  with  clayey  bottom.) 

1897.    -V.  insulata  Kirby;   Kirkaldy,  Trans.  Ento.  Soc.  London  for  1897,  p.   403  (in  part). 

1899.    A',   insulata  Kirby;   Smith,  Insects  of  N.  J.,  p.  144. 

1902.    N.  insulata  Kirby;  Bueno,  Jl.  N.  Y.  Ento.  Soc,  x,  pp.  230,  231,  232,  235. 

1904.  N.  insulata  Kirby;   Kirkaldy,  Wien.  Ento.  Zeit.  xxiii,  pp.  94  and  132. 

1905.  N.  insulata  Kirby;  Bueno,  Jl.  N.  Y.  Ento.  Soc,  xiii,  pp.  46,  150  and  162.  PI.  vii, 
fig.  9  (in  part). 

1908.  N.  insulata  Kirby;  Bueno,  M.  N.  Y.  Ento.  Soc,  xvi,  p.  237. 

1909.  N.  insulata  Kirby;  Kirkaldy  &  Bueno.  Proc.  Ento.  Soc.  Wash.,  x,  p.  198  (in  part). 

1910.  N.  insulata  Kirby;  Bueno,  Jl.  N.  Y.  Ento.  Soc,  xviii,  p.  33. 
1910.  N.  insulata  Kirby;   Smith,  catalogue  Ins.  N.  J.,  edn.  3,  p.  169. 

1913.  A^.  insulata  Kirby;   Browne,  Jl.  Exp.  Zool.,  xiv,  p.  61. 

1914.  A'',  insulata  KJrby;  Parshley,  Psyche,  xxi,  p.  140.  (Taken  on  April  27,  Orono, 
Maine.) 

1917.    A',  insulata  Kirby;   Parshley,  Occ.  papers  Boston  Soc.   Nat.  Hist.,  vii,  p.   113. 

1917.    A'',  insulata  Kirby;    Van  Duzee,  Catalogue  Hemiptera,  p.  453  (in  part). 

1917.    A',  insulata  Kirby;    Hungerford,  Ento.  News,  xxviii,  p.   175. 

1915.  N.  insulata  Kirby;  Hungerford,  Ento.  News,  xxix,  pp.  242-244,  PI.  xiv,  fig.  5  (ovi- 
position  of). 

1919.  AT.  insulata  Kirby;  Hungerford,  Kans.  Univ.  Sci.  Bull.,  xi,  pp.  168,  173,  181,  182, 
186,  187,  189.     PI.  viii,  figs.  3  and  4  (eggs)  PI.  xix,  tigs.  3,  4,  7.     (1st  instar  nymph.) 

1919.  N.  insulata  Kirby;  Hungerford,  Kans.  Univ.  Sci.  Bull.,  xi,  p.  332,  PI.  xxxi,  fig.  9 
(male  genital  capsule). 

1919.  A',  insulata  Kirby;  Parshley,  Occ  papers  Mus.  Zool.  Univ.  Mich.  No.  71.  (From 
Vancouver  Island,  probably  N.  kirbyi  Hungerford.) 

1923.    A'^.  insulata  Kirby;   Bueno  &  Hussey,  Bull.  Brookl.  Ento.  Soc,  xviii,  p.  107. 

1923.  N.  insulata  Kirby;  Bueno,  Conn.  St.  Geol.  &  Nat.  Hist.  Survey  Bull.  No.  34,  pp. 
405  and  407.     PI.  xvi,  fig.  8  (Mar.  &  Apr.). 

1926.  A',  insulata  Kirby;  Leonard,  Cornell  Univ.  Agri.  Exp.  Sta.  Memoir  101,  p.  139. 
(March  to  November.) 

1926.    A',    insulata  Kirby;    Blatchley,   Heteroptera     ...     p.    1052    (in    part). 


Hungerford:    Genus  Notoxecta  85 

1920.    -V.   insulata  Kirby ;   Hungerford,  Bull.  Brookl.  Ento.  Soc,  xxi,  p.  195. 
1929.    -V.   insulata  Kirby;    Hutchinson,  Annals  S.  Afr.   Mus.,  xxv,  pt,  3,  p.  364. 

Referring  to  this  species,  also : 

1851.  .V.  rufiosa  Fieber,  Rhynchotosraphit'en,  pp.  470-477  and  his  \ar\etiea  bicolor.plagiata 
and  cordigera  from  eastern  U.  S.  but  not  his  basalis  from  Brazil. 

1897.    jV.  impressa  Kirkaldy  (not  Fieber)  Trans.  Ento.  Soc.  London  for  1897,  p.  403. 

1897.  .V.  insulata  Kirkaldy,  Trans.  Ento.  Soc.  London  for  1897,  p.  404.  His  varieties 
odora  and  geala. 

1917.    -V.    insulata  var.  geala  Kirkaldy;   Van  Duzee,  Catalogue  Hemiptera,  p.  453. 

1917.    .V.   insulata  var.  odora  Kirkaldy;   Van  Duzee,  Catalogue  Hemiptera,  p.  453. 

Size.  Length,  14  mm.  to  15.5  mm.;  width  of  pronotiim,  4.5  mm. 
to  5  mm. 

Shape.    A  large  and  rather  elongate  species. 

Color.  Pattern  variable,  and  described  by  Kirkaldy  under  five 
varieties,  but  all  intergradations  occur.  Some  specimens  are  nearly 
immaculate;  gray  to  light  tan,  but  with  the  caudal  half  of  pronotum 
darkened  by  black  mesonotum  beneath;  scutellum  black  and  a 
streak  behind  hemelytral  suture  infuscated;  golden  hairs  present, 
especially  on  the  corium.  Other  specimens  are  strikingly  marked, 
the  hemelytra  being  gray,  tan  and  black  with  many  individuals 
showing  some  roseate  reflections,  this  last  especially  marked  on 
freshly  captured  specimens.  While  many  specimens  are  marked  like 
the  typical  A^.  glauca  L.,  the  following  may  be  given  as  the  typical 
color  pattern:  Head  and  anterior  half  of  pronotum  grayish  to 
testaceous,  often  marked  with  green.  Limbs  testaceous,  often  green- 
ish, with  dark  coxae;  hind  femora  longitudinally  streaked  beneath. 
Venter  dark  except  connexivum  which  is  more  or  less  testaceous, 
often  greenish.  Posterior  half  of  pronotum  darkened  by  the  black 
mesonotum  beneath  it.  Scutellum  black.  Hemelytra  with  clavus 
gray  or  tan  except  at  tip,  which  is  spotted  with  black;  corium  with 
black  or  brown  maculations  near  base  and  often  along  the  margin,  a 
transverse  black  band,  broadening  laterally,  starting  from  end  of 
hemelytral  suture,  behind  which  is  a  tan  or  roseate,  roughly  trian- 
gular spot,  that  is  quite  characteristic  for  the  species;  membrane 
and  distal  angle  of  corium  dark;  membrane  near  the  tip  may  be 
traversed  by  a  testaceous  or  tan  band;  corium  and  anterior  lobe  of 
membrane  especially,  clothed  with  golden  hairs.  Dorsum  of  ab- 
domen more  or  less  orange,  metaxyphus  typically  yellow. 

Structural  Characteristics.  Head  about  one-half  length  pronotum, 
transverse;  vertex  slightly  longer  than  its  anterior  width;  anterior 
margin  of  vertex  less  convex,  plainly  shorter  than  the  frontal  margin 
of  the  eye  and  projected  beyond  it;  anterior  breadth  of  vertex: 
synthlipsis  ::  2 — :1;  synthlipsis  usually  slightly  less  than  one-half 

7—3482 


86  The  University  Science  Bulletin 

width  of  the  eye.  Pronotum  about  twice  the  length  of  the  head; 
lateral  margins  only  moderately  divergent,  nearly  straight;  anterior 
angles  normal;  lateral  ledge  narrow,  nearly  straight  and  but  slightly 
oblique.  Anterior  lobe  of  membrane  slightly  longer.  Anterior  tro- 
chanters of  male  with  a  small  hook  beyond  the  middle.  Mesotro- 
chanter  not  angulate.  Carina  of  the  fourth  ventral  abdominal  seg- 
ment bare  on  keel.  The  tenninal  abdominal  sternite  of  female 
broad,  short  and  only  slightly  incised  at  tip.  Female  gonapophyses 
(first  pair)  of  intermediate  length.  Male  genital  capsule  as  shown 
on  Plate  XV. 

Location  of  Type.  This  species  was  described  from  a  single  speci- 
men.   I  have  failed  in  my  endeavor  to  locate  this  type. 

Comparative  Notes.  This  species  in  certain  particulars  appears 
to  be  a  connecting  link  between  the  subgenera  Notonecta  and  Para- 
necta,  as  pointed  out  by  Hutchinson  (1929).  It  is  one  of  our  largest 
American  species.  Its  American  relatives  are  N.  kirbiji  Hungerford 
long  confused  with  it,  and  A^.  montezuma  Kirkaldy.  While  this 
species  is  often  found  in  collections  with  A^.  undulata  Say,  it  is 
readily  distinguished  by  its  larger  size.  The  last  abdominal  sternite 
of  the  female  is  but  shallowly  incised  at  tip,  while  it  is  deepty  incised 
in  N.  undulata  Say.  The  male  genital  capsules  are  quite  different 
also. 

Data  on  Distribiition.  The  distribution  given  in  Van  Duzee's 
catalogue  embraced,  of  course,  A^.  kirbyi  Hungerford.  Its  true  range 
appears  to  be  the  states  north  of  the  fortieth  degree  of  latitude  and 
east  of  the  one  hundredth  meridian  and  the  region  of  Canada  ad- 
jacent thereto. 

The  records  based  upon  my  own  observations  are: 

United  States  of  America 

Maine:    Maine  (Ohio  State  Univ.) 

New  York:  Ithaca,  April  29,  1926;  July  17,  1917,  H.  B.  Hungerford;  Albany, 
April,  10,  1886  (Uhler  Coll.)  ;  White  Plains,  August  22,  1907  (U.  S.  N.  M.) ; 
Long  Island,  H.  Meeske  (U.  S.  N.  M.) ;  Staten  Island,  September  3,  1926 
(U.  S.  N.  M.);  West  Point,  September  9,  1917,  W.  Robinson;  also.  April  22, 
1926  (U.  S.  N.  M.) ;  Lake  Side  (N.  Y.?),  September  22,  October  18,  November 
16  (Uhler  Coll.) 

New  Jersey:  Madison  (Uhler  Coll.);  Palisades,  August  3,  1923  (U.  S.  N. 
M.). 

Connecticut:  Hartford,  October,  1892.  (U.  S.  N.  M.) ;  Milford,  March  21; 
New  Canaan,  April  3.  1919  (see  Conn.  Geol.  Sz  Nat.  Hist.  Surv.  Bull.  34). 

Mass.\chusetts :  Woods  Hole  (U.  S.  N.  M.)  ;  Wellesley,  October  10  (Uhlor 
Coll.). 


Hungerford:    Genus  Notoxecta  87 

Maryland:    Mt.  Zion  Valley,  Augvi.^t  16  (Uhler  Coll.). 

Ohio:    Cu5^ahoga  Co.,  June  1,  1912,  C.  J.  Drake. 

Michigan:  Cheboygan  Co.,  August  7,  1930.  July  20,  1930;  Douglas  Lake, 
Cheboygan  Co.,  July  29,  1928,  July  26,  1926 ;  Pellston  Rd.  Pool,  July  20,  1930. 
The  same  pool,  August  3,  1932;  Mackinac  Island  State  Park,  August  19,  1925. 
All  taken  by  H.  B.  Hungerford ;  Ann  Arbor,  March  7,  1894.  Wolcott  (Nebr.U.). 

Minnesota:  Grand  Marais,  August  13,  1922;  Pelican  Rapids,  August  22, 
1922.  Itasca  Park,  Green  Lake,  August  21,  1922.  St.  Louis  Co..  August  14, 
1922.    All  taken  by  H.  B.  Hungerford. 

Canada 

Newfoundland:    Kelligrews,  August  26,  1922,  F.  Johansen;  (Ottawa,  Can.). 
Ql-ebec:    Ivnowlton,  July  30,  1929,  L.  J.  Milne  (newly  emerged) ;   (Ottawa, 
Can.). 

Ont.^rio:    E.  Ottawa,  May  12,  1930,  G.  S.  Walley.    (Ottawa,  Can.) 

Other  Biological  Notes.  Long  ago  Doctor  Uhler  pointed  out  tliat 
this  insect  prefers  cool  waters.  Van  Duzee  has  reported  it  in  stag- 
nant pools  having  clayey  bottoms  near  Buffalo,  N.  Y.  I  found  it  a 
rather  rare  insect  around  Ithaca,  N.  Y.,  but  in  Michigan,  in  a  gravel- 
pit  pool,  I  took  seventy-six  specimens  in  a  couple  of  hours.  This 
.pool  contained  cold  water  about  two  feet  deep  at  the  west  end  and 
shallower  and  Avarmer  water  elsewhere.  Most  of  the  specimens 
were  taken  in  the  cold  water,  where  they  were  swimming  slowly  on 
an  even  keel  about  midway  between  the  bottom  and  top.  They  are 
able  to  remain  for  a  long  time  submerged,  their  backs  covered  with 
a  silvery  film  of  air.  At  Ithaca,  N.  Y.,  the  species  wintered  as  an 
adult;  eggs  were  attached  to  submerged  vegetation  during  the  last  of 
April  and  in  May.    The  incubation  period  was  twenty-four  days. 

Notes  on  Notonecta  rugosa  Fieber 

In  the  Zoological  Museum  at  Berlin  there  are  nine  specimens  in 
the  row  labeled  A^.  rugosa  Fieb.    They  are  labeled  as  follows: 

First:  "3628"  "Baltim.  Klug."  "Rugosa  Fieb."  The  last  looks  like  Fieber's 
writing. 

Second:  "Cat.  No.  3628"  "Nov.  Granada-Gond"  (I  cannot  be  sure  of  this 
written  label.)  "Notonecta  itigosa  Fieb.  and  below  it  another  "nigosa  Fieb. 
undulata  Say." 

Third:  "Cat.  No.  3628"  and  a  green  label  I  cannot  read  with  certainty. 
"Massachut.  Zim."?    Notonecta  rugosa  Fieb." 

Fourth:    Cat.  No.  3628"  "Massachus.  Zimmerm"  Notonecta  rugosa  Fieb. 

Fifth:    Labeled  as  above.    This  is  a  darker  specimen  than  the  four  above. 

Sixth  and  Seventh:    Labeled  as  above. 

Eighth:    "3629"  "Mexico  Deppe"  "Notonecta  rugosa  var." 

Ninth:    "3630"  "Baltim.  Klug."  "N.  nigosa  var."    This  is  a  white  specimen. 


88  The  Uxiversity  Science  Bulletin 

All  of  the  above  except  No.  8  are  N.  undulata  Say,  and  the  first 
specimen  looks  as  if  it  were  Fieber's  determination  label.  The 
eighth  one  is  a  dermestid-eaten  female.  It  is  not  N.  undulata  Say, 
but  I  do  not  know  what  to  call  it. 

Doctor  Fieber  described  his  A',  rugosa  as  "Llinge,  ^M^-ly2  linien.'" 
The  specimens  here  in  Berlin  are  only  51/2  English  lines.  Doctor 
Kirkaldy  places  N.  rugosa  Fieb.  as  a  synonym  of  N.  insulata  Kirby. 
no  doubt  being  influenced  by  Fieber's  name  and  by  his  statement  of 
length.  Fieber  named  four  varieties:  variety  bicolor  and  variety 
plagiata  froin  "Longisland,  Baltimore  (Mus.  Berol.  und  Hal.),"  var. 
cordigera  from  "Pennsylvanien  (Dr.  Germar),  Baltimore  (Mus. 
Berol.),"  and  var.  basalis  from  "Brazilien  (Mus.  Berol.)."  In 
Kirkaldy 's  Uber  Notonectiden.  p.  94,  he  says  that  Doctor  Breddin 
sent  him  from  the  Halle  Museum  a  specimen  labeled  N.  rugosa,  but 
that  it  was  a  variety  of  A^.  undulata  Say  and  too  small  to  be  Fieber's 
A'",  rugosa. 

I  have  seen  no  species  from  Brazil  (except  A^.  nigra  Fieb.)  that  is 
as  large  as  N.  insulata  Kirby.  Doctor  Fieber  recorded  his  var. 
basalis  as  from  Brazil.  I  did  not  find  it  in  the  Vienna  Museum. 
A^  rugosa  Fieber  is  either  a  synonym  of  A^.  insidata  Kirby  credited 
with  the  same  size  or  else  is  A^.  undulata  Say  with  some  strange  error 
in  Fieber's  measurement.  In  any  event,  the  localities  in  eastern 
ITnited  States  are  so  well  known  that  A^.  rugosa  must  be  one  or  the 
other  of  the  species  named  above. 

Notonecta  kirbyi  Hungerford,  1925 

(Color  Plate  III,  fig.  4;   Plate  XV,  fig.  2) 

1925.  A^  kirbyi  Hungerford,  Can.  Ento.  Ivii.  p.  241,  pi.  G,  fig.  2. 
1928.  A',  kirbyi  Hungerford,  Annals  Ent.  Sor.  Amer..  xxi,  p.  142. 
1928.    A',  kirbyi  Hungerford,  Ento.  New.s.   xxxix.  p.   l.'>fi. 

Referring  to  this  species,  also: 

1875.  A^.  insulata  Uhler;  Wheeler's  Rrpt.  upon  Geog.  &  Geol.  Expl.  and  Surveys  West 
of  One  Hundredth  Meridian,  Washington,  vol.  v.  eh.  xii,  p.  841.  (Owen's  valley,  Calif.,  by 
F.   Bischoff.) 

1876.  A',  insulata  Uhler,  Bull.  U.  S.  Geol.  Geog.  Surv.,  vol.  I.  Bull.  No.  .'.,  2d  ser.  p.  339 
(Mts.   of  Colorado). 

1876.  A^.  insulata  iJh'l^F,  Bull.  U.  S.  Geol.  Geog.  Surv.,  p.   73.     (Reprint  from  above.) 

1877.  N.  insulata  Uhler,  Wheeler's  Rept.  Chief  Engineer  for  1877,  p.  13.S2.     (In  Arizona.) 

1878.  N.  insulata  Uhler,  Bull.  U.  S.  Geol.  Geog.  Surv..  iv,  p.  509.  (Milk  river  region, 
Montana.) 

1897.    A',  insulata  var.  imprcssa,  Kirkiildy  (not  Fieber)  Tiaiis.   Ento.  Soc.   London  fc.r  1897, 

p.  404. 

1901.  .V.  insulata  Champion,  Biol.  Centr.  Amer.  Heni.-Het.,  If,   p.   .S69. 

1904.  -V.  insulata  Uhler.,  Proc.  U.  S.   Natl.   Mus.,  xxvii,  p.   364. 

1905.  A',  insulata  Bueno,  .11.   N.  Y.   Ento.  Soc,  xiii,  p.   46  (in  part). 


Huxgerford:    Gexus  Notoxecta  89 

1906.    N.  insulata  Snow,  Trans.  Kansas  Acad.  Sei.,  xx,  pt.  1,  p.   181. 

1909.    .V.  insulata  Kirkaldv  and  Bueno,  Proe.  Ento.  Soc.  Wash,  x,  p.  198  (in  part). 

1914.    X.  insulata  \'an  Duzee,  Trans.  San  Diego  Soc.  Nat.  Hist.  11,  p.  33. 

1917.    .V.  insulata  Van  Duzee,  Catalogue  Hemiptera,  p.   453  (in  part). 

1919.  N.  inmlata  Parshley,  Ocoas.  Papers  Mus.  Zool.  I'niv.  of  Mich.,  No.  71,  1919. 
(Probably.) 

1922.    A',  instdata  Hungerford,  Kansas  Univ.  Sci.  Bull.,  xiv,  pp.   417-419. 

1922.    N.  insulata  Parshley,  S.  Dakota  St.  College  Tedi.  Bull.   No.   II,  p.   22  (from  Capa, 

Jones  Co.,  S.  D.) 

Size.  Length,  13.8  mm.  to  15.5  mm.;  width  of  pronotum,  4.5  mm. 
to  4.8  mm. 

Shajje.    A  rather  elongate  species. 

Color.  Variable.  Some  specimens  nearly  like  A^.  insulata  Kirby, 
but  usual  pattern  considerably  darker;  limbs  usually  less  distinctly 
longitudinally  striped.  Typical  color  pattern  tan  and  black  or  red 
and  black.  The  clavus  and  basal  half  of  corium  tan,  orange  or  red 
mottled  more  or  less  with  black.  Distal  half  of  corium  and  basal 
three-fifths  of  membrane  typically  black,  the  black  field  of  the 
corium  broken  only  by  two  small  spots  colored  like  the  clavus. 
(These  spots  may  be  large  and  confluent  giving  a  triangular  patch 
like  that  of  N.  insulata  Kirby.)  Corium  and  anterior  lobe  of  mem- 
brane especially  clothed  with  golden  hairs.  Dorsum  of  abdomen 
in  the  red  forms  nearly  always  reddish ;  metaxyphus  typically  black. 
The  venter  dark  except  connexivum. 

Structural  Characteristics.  Head  less  than  one-half  length  of 
pronotum,  transverse ;  vertex  longer  than  its  anterior  width ;  anterior 
margin  of  vertex  less  convex,  as  long  as  the  frontal  margin  of  the 
eye  and  projected  beyond  it;  anterior  breadth  of  vertex  :  synthlip- 
sis  : :  2  —  :  1 ;  synthlipsis  usually  fully  one-half  to  two-thirds  width 
of  an  eye.  Pronotum  more  than  twice  as  long  as  the  head;  lateral 
margins  only  moderately  divergent,  nearly  straight;  anterior  angles 
normal  or  slightly  embracing  the  eye;  lateral  ledge  a  little  thicker 
than  in  N.  insulata  Kirby,  oblique  and  slightly  sigmoid.  In  side 
view  the  rear  portion  of  pronotum  more  elevated  than  in  N.  insulata 
Kirby.  Anterior  lobe  of  membrane  slightly  longer.  Anterior  tro- 
chanter of  male  with  a  small  hook  beyond  the  middle.  Alesotro- 
chanter  not  angulate.  Carnia  of  the  fourth  ventral  abdominal  seg- 
ment bare  on  keel.  The  terminal  abdominal  sternite  of  female 
broad,  short  and  only  slightly  incised  at  tip.  Female  gonapophyses 
(first  pair)  of  intemiediate  length.  Male  genital  capsule  as  shown 
on  Plate  XV. 

Location  of  Type.  In  the  Francis  Huntington  Snow  Entomological 
Museum  at  the  University  of  Kansas.  Holotype,  allotype  and  para- 
types  from  Emery  county,  Utah,  Aug.-Sept.,  1921,  Mrs.  Grace  A\'iley. 


90  The  University  Science  Bulletin 

Comparative  Notes.  This  species  has  long  been  confused  with  A^. 
insulata  Kirby,  from  which  it  can  be  distinguished  as  indicated  in 
the  key. 

Data  on  Distribution.  This  species  has  been  confused  heretofore 
with  A^  insulata  Kirby.  The  distribution  is  western  United  States 
and  Canada.    I  have  the  following  records: 

Canada 

British  Columbia:  Mt.  Cheam,  Mar.  9,  1924;  Bearfoot  Mt.,  Sept.  3,  1917 
(U.  S.  N.  M.);  Vernon,  Aug.  5,  1931,  L.  D.  Anderson;  Vernon,  Sept.  26,  1929 
(Ottawa,  Can.) ;  Peachland,  Oct.  7,  1921,  A.  N.  Gartrell;  Rolla,  Aug.  6,  1927,  P. 
N.  Vroom;  Revelstoke  Mt.,  6,000  ft.,  Aug.  12,  1923,  E.  R.  Buckell;  Creston, 
July  15,  1926,  A.  A.  Dennys;  Princeton,  May  6,  1923,  P.  N.  Vroom;  Brents  L. 
Summerland,  Oct.  29,  1931,  A.  N.  Gartrell;  Copper  Mts.,  Oct.  26,  1928,  G.  Stace 
Smith;  Oliver,  July  24,  1923,  E.  R.  Buckell;  Minnie  L.,  July  26,  1925,  N. 
Criddle;  Aspen  Grove,  May  26,  1922,  P.  N.  Vroom;  (above  in  Ottawa,  Can.). 

Alberta:  Tofield,  Aug.  25,  1923,  E.  H.  Strickland;  Cypress  Hills,  July  21, 
1930,  J.  H.  Pepper;  Waterton  Lakes,  July  2,  1930,  J.  H.  Pepper;  Medicine  Hat, 
Aug.  7,  1929,  J.  H.  Pepper. 

United  States  of  America 

S.  Dakota:   Capa,  Jones  Co.,  June  1,  1921,  H.  C.  Severin  (S.  D.  St.  College). 

Nebraska:  Monroe  Canyon,  Sioux  Co.,  August  15,  1908,  J.  T.  Zimmer  (Nebr. 
U.);  Pine  Ridge,  July  (Ohio  St.  Univ.). 

Colorado:  Pingree  Park,  Aug.  25,  1921,  Lawson  &  Beamer;  Northern  Colo., 
Aug.  25,  1920,  R.  C.  Moore;  Caisson,  July  1,  1931,  L.  D.  Anderson;  Colorado 
819  (Uhler  Coll.) ;  Colorado  Mts.,  July  to  Sept.  (Uhler  Coll.) ;  Colorado  2063 
Gillette  (Uhler  Coll.) ;  Denver,  B.  H.  Smith  (Uhler  Coll.) ;  Colorado,  C.  F. 
Baker  (U.  S.  N.  M.) ;  Hamilton,  July  3,  1928  (newly  emerged) ;  Ft.  Collins, 
Aug.  7,  1900,  Aug.  11,  1899,  Sept.  1,  1926,  Sept.  8,  1926,  Sept.  14,  1916,  Sept.  18, 

1928,  Sept.  19,  1931,  Oct.  2,  1917,  Oct.  5,  1931,  Oct.  30,  1930,  Nov.  1,  1926  (Ft. 
Collins  Exp.  Sta.). 

Tex.4s:  Valentine.  July  13,  1927,  R.  H.  Beamer;  Presidio,  July  16,  1927,  R. 
H.  Beamer. 

New  Mexico:    Torrance  Co.,  July  21,  1929,  P.  W.  Oman;  Estancia.  Sept.  6, 

1929,  C.  H.  Martin;  Cocorro  Co.,  Aug.  18,  1927,  R.  H.  Beamer;  Messilla  Park, 
July  18,  1927,  L.  D.  Anderson;  Las  Vegas,  Barber  &  Schwarz  (U.  S.  N.  M.). 

Arizona:  Yavapai  Co.,  July  1,  1929,  L.  D.  Anderson;  Cochise  Co.,  July  20, 
1927,  R.  H.  Beamer;  Gila  Co.,  Aug.  5,  1927,  R.  H.  Beamer;  Santa  Cruz,  Aug. 
7,  1927,  P.  A.  Readio;  Santa  Rita  Mts.,  July  25,  1927,  R.  H.  Beamer;  Navajo 
Co.,  Aug.  15,  1927,  R.  H.  Beamer;  Coconino  Co.,  Aug.  13,  1927,  P.  A.  Readio; 
Douglas,  San  Bernardino  Ranch,  3,700  ft.,  Aug.,  F.  H.  Snow;  Baboquivaria 
Mts.,  F.  H.  Snow;  Arizona  (Uhler  Coll.);  Yuvapai  Co.  (U.  S.  N.  M.); 
Senator  Mine,  near  Prescott,  Sept.  12,  1907;  Flagstaff,  Barber  &  Schwarz 
(U.  S.  N.  M.) ;  Lowell  (Uhler  Coll.) ;  Chiricahua  Mts.,  Pinerey  Canyon,  Boy 
Scout  Camp,  June  17,  1932,  D.  K.  Duncan;  Resen^ation  Springs,  12  miles 
N.  E.  of  Globe,  Sept.  17.  1932,  D.  K.  Duncan;  Peach  Springs,  Aug.  27  (Ohio 
State  Univ.). 


Hungerford:    Genus  Notoxecta  91 

Utah:  Emery  Co.,  Aug.  2,  1921,  Grace  0.  Wiley;  Emery  Co.,  July  22  and 
29,  1921,  Grace  O.  Wiley;  Emery  Co.,  Sept.  13,  1921,  Grace  O.  Wiley;  Far- 
west,  C.  J.  D.  Bro\vn;  Salt  Lake  Co.,  July,  1929;  Kane  Co.,  Summer,  1921, 
R.  C.  Moore;  Garfield  Co.,  Aug.  14,  1921,  R.  C.  Moore;  Antelope.  July  1, 
1931,  L.  D.  Anderson;  Ogden,  June  27,  1910,  F.  T.  Moore  (U.  S.  N.  M.);  Salt 
Lake  (U.  S.  N.  M.);  Lehi,  May  27,  1930,  G.  F.  Knowlton  (Knowlton  Coll.). 

Nevad.\:  S'unnyside,  C.  T.  Brues,  1930,  Hot  Sp.  Exp.,  1930,  Sp.  No.  97; 
Bottle  Mountain;  Fallon,  Aug.  9,  1929,  L.  D.  Anderson;  Carson  City,  Aug. 
9,  1929,  R.  H.  Beamer;  Meadow,  Nev.  Hot  Sp.  Expd.,  1930,  Sp.  No.  113, 
Gerlach  &  Soldier;  Reno,  Feb.  13,  1916,  H.  G.  Dyar  (U.  S.  N.  M.)  ;  Owens 
Valley  (Uhler  Coll.);  Vacity  (Uhler  Coll.). 

California:  Bautista  Canyon,  Apr.  8,  1931,  C.  H.  Martin;  Emery,  May  14, 
1921,  C.  T.  Dodds;  Oakland,  Feb.  22,  1921,  C.  T.  Dodds;  Laguna  Beach,  C.  T. 
Dodds;  Laguna  Mts.,  July  6,  1929,  P.  W.  Oman;  Alpine,  July  9,  1929,  L.  D. 
Anderson;  San  Diego  Co.,  July  28,  1929,  P.  W.  Oman;  Cuyamaca  Lake, 
July  6.  1929,  P.  W.  Oman;  Giant  Forest,  July  28,  1929,  P.  W.  Oman;  Indio, 
July  24,  1929,  L.  D.  Anderson;  Palo  Alto,  Mar.  5,  1892  (Cornell);  Giant 
Forest,  S'eq.  Nat.  Park,  July  21,  26,  1907,  6,000-7,000  ft.,  J.  C.  Bradley 
(Cornell);  Fish  Springs,  Hot.  Sp.  Expd.,  1930,  C.  T.  Brues;  San  Francisco 
(labeled  by  Uhler  N.  impressa  Fieb.)  (Uhler  Coll.) ;  San  Luis  Obispo  (U.  S. 
N.  M.);  Sonoma  Co.,  (U.  S.  N.  M.) ;  San  Jose,  Sept.  8,  1881  (U.  S'.  N.  M.); 
Menlo  Park,  Jan.  1905,  F.  Hornung;  Santa  Barbara  Co.,  Coquillet  (U.  S. 
N.  M.);  San  Diego  Co.,  Coquillet  (Uhler  Coll.);  California,  Aug.,  C.  V. 
Riley  (Uhler  Coll.) ;  Cahfornia,  L.  Lethierry  (labeled  N.  mexicana)  (U.  S. 
N.  M.) :  Santa  Clara  Co.,  Baker  (U.  S.  N.  M.) ;  Placer  Co.,  Aug.  (U.  S.  N.  M.) ; 
Palm  Springs,  H.  G.  Hubbard  (U.  S.  N.  M.) ;  Mt.  Diablo  (U.  S.  N.  M.) ; 
S.  California  (Uhler  Coll.);  San  Antonio  Canyon,  Ontario,  July  25,  1907; 
Campo.  Calif.,  Laguna  Mts.,  Aug.  25,  1932,  H.  W.  Capps. 

Oregon:  Gaston;  Corvallis,  Sept.  4  (C.  J.  Drake);  Dilley  (H.  G.  Barber); 
Hood  River,  July  17,  1931,  L.  D.  Anderson;  Boardman,  July  17,  1931,  L.  D. 
Anderson;  Union,  July  13,  1931,  L.  D.  Anderson;  Hot  Lake,  July  13,  1931, 
L.  D.  Anderson;  Oregon  (Uhler  Coll.);  Corvallis,  Aug.  8,  1925  (U.  S.  N.  M.) ; 
Mt.  Hood  Lodge,  Parkdale,  alt.  3,500  ft.,  July  30,  1931,  M.  C.  Lane  (U.  S. 
N.  M.);  Warner  Lake,  1877  (Uhler  Coll.). 

Wyoming:  Yellowstone  Nat.  Park,  Aug.  2,  1926,  George  Cady;  Grand 
Teton  Nat.  Park,  Aug.  8,  1931,  L.  D.  Anderson;  Ft.  Laramie,  Horn  (Uhler 
Coll.) ;  Grand  Teton  Nat.  Park,  Aug.  1931,  L.  D.  Anderson. 

Idaho:  Burley,  July  6,  1931,  L.  D.  Anderson;  Bliss,  July  7,  1931,  L.  D. 
Anderson;  Soldier,  (Nebr.  U.) ;  Council,  June  25,  1926,  alt.  3,059  ft.;  Moscow, 
Oct.  1928.  alt.  2,560  ft'. 

Washington:  Pullman;  L.  McElroy,  Paha,  1920;  Kalama.  July  21.  1931, 
L.  D.  Anderson;  Coulee  City,  Sept.  2,  1920,  M.  C.  Lane  (U.  S.  N.  M.) ;  Lind, 
May  13,  1921.  M.  C.  Lane  (U.  S.  N.  M.) ;  Stratford,  Sept.  3,  1920,  M.  C.  Lane 
(U.  S.  N.  M.) ;  Ritzville,  May  17,  1921,  M.  G.  Lane  (U.  S.  N.  M.) ;  L.  Mc- 
Elroy, Paha,  June,  1920,  M.  C.  Lane  (U.  S.  N.  M.) ;  Pullman,  C.  V.  Piper 
(U.  S.  N.  M.). 

Montana:  Glacier  Park,  Aug.  20,  1926,  George  Cady;  Anaconda,  Aug, 
12,  1931,  L,  D.  Anderson;  Whitehall,  Aug.  31,  1931,  L.  D.  Anderson;  Three 
Forks.  July  7.  1931,  L.  D.  .Anderfon. 


92  The  University  Science  Bulletin 

Notonecta  irrorata  Uhler,  1879 

(Color  Plate  I,  fig.   1;   Plates  X,  fig.   4,  and  XVI   fig.    1.) 

1879.  N.  irrorata  Uhler,  Proc.  Boston  Soc.   Nat.  His.,  xix,  p.   443. 

1888.  iV.  irrorata  Uhler;    Provancher,  Pet.   Faune  Ento.   Can.,  iii,  p.   200. 

1891.  -V.  irrorata  Uhler;   Summers,  Bull.  Agr.  Exp.  Sta.,  Univ.  of  Teiin.,  iv.  No.  3,  p.   82. 

1894.  iV.  irrorata  Uhler;  Van  Duzee,  Bull.  Buf.  Soc.  Nat.  Sci.,  v,  p.  186.  (Not  infre- 
quent in  stagnant,  muddy  iwoLs  about  Buffalo,  N.  Y.) 

1897.  N.  irrorata  Uhler;   Kirkaldy,  Trans.  Ento.   Soc.  Lond.   for  1897,  p.   418. 

1900.  N.  irrorata  Uhler;    Osborn,  Contr.  Dept.  Zool  &  Ent.  Ohio  St.   Univ,  No.   2  p.   79. 

1902.  N.  irrorata  Uhler:   Bueno,  Jl.  N.  Y.  Ento.  &oc.,  x,  pp.  230,  231,  235. 

1904.  .V.  irrorata  Uhler;   Kirkaldy,  Wien,  Ento.  Zeit.,  xxiii,  p.  132. 

1905.  iV.  irrorata  Uhler;   Bueno,  Jl.  N.  Y.  Ento.  Soc.  xiii,  pp.   46,  159;    pi.  vii,  fig.  6. 

1907.  N.  irrorata  Uhler;  Bueno  and  Brimley,  Ento.  News,  xviii,  p.  435.  (March  to 
November  four  newly  transformed  August  8,  N.  Car.) 

1908.  A'^.  irrorata  Uhler;   Bueno,  Jl.  N.  Y.  Ento.   Soc,  xvi,  p.   238. 

1909.  A'',  irrorata  Uhler;  Kirkaldy  and  Bueno,  Cat.  of  Am.  Aquatic  and  Semiaquatic 
Hemiptera,  Proc.  Ento.   Soc.   Wash,  x,  Nos.    3  and   4,  p.   198. 

1910.  N.  irrorata  Uhler,  Smith,  Cat.  Ins.  N.  J.,  edn.  3,  p.   169. 

1913.  A',  irrorata  Uhler;   Browne,  Jl.  Exp.  Zool.,  xiv,  p.  .61. 

1914.  A',  irrorata  Uhler;  Parshley,  Psyche,  xxi,  p.  140.  (May  22-September  19,  Orono 
&  Machias,  Maine.) 

1914.  A^.  irrorata  Uhler;  Barber,  Bull.  Am.  Mus.  Nat.  His.,  xxxiii,  p.  499.  (From 
Florida.) 

1917.    A^.  irrorata  Uhler;   Hungerford,  Ento.  News,  xxviii,  p.   271. 

1917.  A^.  irrorata  Uhler;  Parshley,  Occasional  papers  of  the  Boston  Soc.  of  Nat.  His., 
vii,  p.   112.     (Me.,  N.  H.,  Vt.,  Mass.,  R.   I.,  Conn.) 

1917.  N.  irrorata  Uhler;    Van  Duzee,  Cat.   of  Hemiptera,  pp.   450,   451. 

1918.  N.  irrorata  Uhler;  Hvuigerford,  Ento.  News,  xxix,  pp.  242-245;  pi.  xiv,  figs.  1-4; 
pi.  XV,  fig.  9.     (lUus.  ovipositor.) 

1919.  A^.  irrorata  Uhler;  Hungerford,  Kans.  Univ.  Sci.  Bull.,  xi,  pp.  32,  33,  166,  167,  171, 
177,  180-185,  187,  304,  319,  312,  339;   pi.  xix,  figs.  2,  6,  9  ;   pi.  xxii,  figs.  1-5:   pi.  xxiii,  fig.  9. 

1923.  A',  irrorata  Uhler;  Bueno,  Conn.  State  Geol.  and  Nat.  Hist.  Survey  Bull.,  xxxiv, 
pp.  405,  406,  figs.  44,  45;  pi.  xvi,  fig.  7.     (April  to  November.) 

1923.    A^.  irrorata  Uhler;   Bueno  and  Hussey,  Bull.   Brook.   Ento.   Soc,  xviii,  pp.    105,  106. 
1925.    A^.  irrorata  Uhler;   Hungerford,  Annals  Ento  Soc.  Am.,  xviii,  p.   417. 

1925.  A',  irrorata  Uhler;  Hungerford  and  Beamer,  Ento.  News,  xxx\i,  p.  297.  (Kansas 
Record  new.) 

1926.  N.  irrorata  Uhler;  Leonard,  Cornell  Univ.  Agri.  Exp.  Sta.  Memoir  101,  p.  138. 
(March  to  November.) 

1926.  A^  irrorata  Uhler;  Hungerford,  Bull.  Brook.  Ento.  Soc,  xxi,  p.  194. 
1926.  A^  irrorata  Uhler;  Blatchley,  Heteroptera  of  Eastern  N.  A.,  p.  1052. 
1928.    A',  irrorata  Uhler;    Hungerford,  Ento.  News,  xxxix,  p.   15G.     (Georgia.) 

1928.  A^.  irrorata  Uhler;    Hungerford,  Annals  Ento.  Soc.  Amer.,  xxi,  p.   142. 

1929.  A^.  irrorata  Uhler:    Hutchinson,  Annals  of  South   African   Mus.,   xxv,  pt.   3.  p.    363. 

Also,  referring  to  this  species: 

1878.  N.  irrorata  Say;  Packard,  Guide  to  the  Study  of  Insects,  Cth  Edn.  p.  537.  (With- 
out description,  a  manuscript  name  of  Say  used  in  1st  Edn.  1869.) 

Size.  Length,  12.9  mm.  to  15.5  mm.;  wi(ith  of  pronotum,  4.5  mm. 
to  5  mm. 

Shape.  The  rapid  divergence  of  the  lateral  margins  to  the  prono- 
tum  from  the  comparatively  small  head  give  the  appearance  of  a 
broad-shouldered  back  swimmer. 

Color.  General  color  is  irrorated  or  mottled  brown  and  blue-black 
or  black.     Head,  limbs  and  connexivum  of  usual  color,  or  a  little 


Hungerford:    Genus  Xotoxecta  93 

darker,  often  greenish.  Pronotinn  of  same  color,  but  darkened  on 
caudal  part  by  the  black  mesonotum  beneath  and  marked  by  a 
median  dark  brown  blotch  in  front.  Scutellum  black.  Hemelytra 
irrorated  with  brown  and  black  or  blue-black,  the  brown  usually  pre- 
dominating on  the  clavus  and  the  black  on  the  corium;  the  mem- 
brane is  black.  The  venter  is  black.  The  last  abdominal  sternite 
may  be  testaceous  or  partly  so,  and  the  metaxyphus  is  usually  mar- 
gined with  testaceous. 

Sti'uctural  Characteristics.  Anterior  outline  of  the  head,  as  seen 
from  above,  somewhat  flattened;  vertex  with  anterior  width  equal 
to  its  length;  anterior  margin  of  vertex  less  convex  and  a  little 
shorter  than  the  frontal  margin  of  an  eye ;  anterior  breadth  of  ver- 
tex :  synthlipsis  ::  3:1;  the  synthlipsis  narrow,  only  one-third  the 
width  of  the  eye;  the  inner  margins  of  the  eyes  diverging  widely. 
Pronotum  more  than  twice  the  length  of  the  head;  lateral  margins 
quite  divergent,  and  nearly  straight;  anterior  angles  not  embracing 
the  eyes;  lateral  ledge  slightly  sigmoid  and  oblique.  Scutellum 
slightly  wider  than  long.  Lobes  of  membrane  about  equal.  Anterior 
trochanter  of  male  without  hook  or  tubercle.  Mesotrochanters 
roughly  right  angulate.  The  terminal  abdominal  sternite  of  female 
large  and  constricted  near  the  tip,  which  is  entire ;  first  pair  of  gona- 
pophyses  long.    Male  genital  capsule  as  shown  on  Plate  XVI. 

Location  of  Type.  The  type  was  from  "Ipswich,  March,  Mr. 
Oakes;  Milton,  Mass.,  April  22,  1829."  I  have  seen  four  specimens 
in  the  Boston  Society  of  Natural  History  Museum  (Harris  Collec- 
tion) that  may  be  designated  the  "cotypes." 

Comparative  Notes.  This  species  is  related  to  the  Notonecta 
glauca  Linn,  group.  The  long  gonapophyses  and  the  large  terminal 
abdominal  sternite  of  the  female  and  the  digitate  process  on  the 
genital  capsule  of  the  male  place  it  clearly  with  the  European 
species.  The  only  related  species  of  North  America  is  N.  borealis 
Bueno  and  Hussey. 

Biological  Notes.  Bueno  tells  us  that  this  handsome  back  swim- 
mer likes  the  shadows  of  the  bank,  overhanging  limb,  or  of  aquatic 
vegetation.  I  think  it  does  prefer  such  places,  but  I  have  taken  it 
in  open  ponds.  It  winters  in  the  deeper  ponds  and  spring-fed,  open 
pools,  flying  from  these  quarters  to  shallower  waters  for  breeding. 
I  have  seen  them  swimming  slowly  under  the  ice  at  the  Field  Station 
at  Ithaca,  N.  Y.,  in  early  February.  Mating  has  been  observed  in 
April  in  a  pool  in  whicli  tiiey  overwintered  and  from  which  they 
soon  disappeared,  to  be  found  in  a  meadow  pool  a  quarter  of  a  mile 


94  The  University  Science  Bulletin 

away.  Unlike  all  other  North  American  species,  except  A^.  borealis 
Bueno  and  Hussey,  this  species  inserts  its  eggs  in  the  tissues  of 
plants.  One  female  contained  252  ova.  The  egg  stage  lasted  two 
or  three  weeks  in  late  May  and  first  half  of  June  at  Ithaca,  N.  Y., 
where  I  made  the  above  observations.  After  five  nymphal  instars 
the  adults  appeared  about  the  middle  of  July. 

Data  on  Distribution: 

Canada 

Quebec:  Knowlton,  Aug.  12,  1929,  G.  S.  Walley;  Aj-Imer,  April  20,  192r, 
G.  S.  Walley;  Kazubazua,  Aug.  18,  1931,  G.  S.  Walley;  Fairy  Lake,  Hull, 
Sept.  11,  1928,  G.  S.  Walley  (Ottawa,  Can.  Coll.);  Montreal,  Aug.  1903 
(U.  S.  N.  M.). 

Ontario:  Southampton,  Sept.  13,  1927,  G.  S.  Walley;  Black  Rapids, 
Ottawa,  Aug.  25,  1927,  G.  S.  Walley;  Arnprior,  Sept.  15,  1928,  G.  S.  Walley; 
Merivale,  May  9,  1930,  G.  S.  Walley;  Rockcliffe,  Ottawa,  July  2,  1928. 
G.  H.  Fiske;  Ventnor,  Sept.  2,  1928,  J.  A.  Adams;  Vineland  Sta.,  Sept.  11, 
1928,  W.  Putman.     (All  of  above  in  Ottawa,  Canada  Collection). 

Unitb^d  States  of  America 

Maine:    Orono,  Machias,  May  22  to  Sept.  19  (Parshley  1914). 

Massachusetts:  Mass.  (Uhler  Coll.);  Melrose  Heights,  Mar.  15,  D.  H. 
Clemons. 

Rhode  Isl.\nd:    R.  I.  (C.  F.  Baker  Coll.). 

Connecticut:  Conn.  (Uhler  Coll.).  April  24-Nov.  24.  (See  Conn.  Geol. 
and  Nat.  Hist.  Surv.  Bull.  34). 

New  York:  Ithaca,  every  month  of  year,  H.  B.  Hungerford;  New  York 
(Uhler  Coll.);  Long  Island  (Uhler  Coll.). 

New^  Jersey:  Pallisades,  Aug.  23,  1903  (U.  S.  N.  M.);  Ft.  Lee  Dist.,  Oct. 
10,  1903  (U.  S.  N.  M.) ;  Madison  (Uhler  Coll.) ;  N.  J.  (Uhler  Coll.). 

Pennsylv-ANIa :  Philadelphia,  1926,  F.  Anderson;  Penn.  (Uhler  Coll.); 
Philadelphia,  near  Narberth  Dec.  19,  1922,  W.  E.  Hoffmann  (Minn.  Coll.); 
Hummelstown,  April  1,  1917,  J.  N.  Knull. 

Ohio:  Wellington,  July  8,  1890  (Tenn.  Coll.);  Cincinnati,  Dec.  30,  1923, 
W.  E.  Hoffmann  (Minn.  Coll.) ;  Columbus.  Sept.  26,  1914,  C.  J.  Drake. 

Indiana:  Maxinkuckee;  Indiana  (Uhler  Coll.);  South  Bend  (U.  S.  N.  M.) ; 
Kosciusko  Co.,  July,  1932,  G.  E.  Gould. 

Illinois:  Toledo,  Aug.  1891,  Harvey;  111.  (Uhler  Coll.);  Champaign,  April, 
1907,  Hart  and  Hood;  Hilliary,  Mar.  23,  1907,  Hart  and  Hood;  Dubois,  June 
20,  1905;  Johnson  Co.,  July  20,  1877;  St.  Francesville,  Aug.  23,  1884;  Flora, 
Aug.  22,  1884  (above  in  111.  Nat.  Hist.  Surv.  Mus.). 

Michigan:  Ann  Arbor,  Sept.  15,  1927,  E.  Creaser;  Bryants  Bog,  Douglas 
L.,  July  17,  1923,  Smith's  Bog,  July  17,  1923,  Sedge  Pt.  Pool,  Aug.  16,  1923, 
Mackinac  Island,  Aug.  19,  1925,  Trout  L.,  Mich.  Aug.  25,  1925,  Nigger  Creek, 
Mullett  Lake,  Aug.  4,  1925,  Burt  Lake,  Fontinalis  Run,  July  7,  1923,  Bois 
Blanc  Island,  Aug.  14,  1932.    All  above  by  H.  B.  Hungerford. 

Minnesota:  Grand  Marais,  Aug.  13,  1922.  W.  E.  Hoffmann;  Ramsey  Co., 
Nov.  11,  1922,  W.  E.  Hoffmann  (Minn.  Coll.). 


Huxgerford:    Genus  Notonecta  95 

Marylaxd:  Crisfield.  Baltimore,  Frederick  (all  Uhler  Coll.);  Pliimmers 
Island.  April  25,  1913;  Laurel,  Aug.,  1909,  F.  B.  Marshall;  Annapoli.s,  Sept.  18, 
1932,  Paul  Oman. 

District  of  Columbia:  Rock  Creek,  Aug.  31,  1906,  D.  H.  demons  (U.  S. 
N.  M.). 

Virginia:  Chain  Bridge,  Sept.  3,  1906,  D.  H.  Clemons;  Great  Falls,  May  22, 
1906. 

Tennessee:  Monroe  Co.,  July  6,  1890,  H.  E.  Summers  (111.  Nat.  Hist.  Surv. 
Mus.). 

Kentucky:  East  Cairo,  June  7,  1881,  S.  A.  Forbes  (111.  Nat.  Hist.  Surv. 
Mus.);  Shakertown,  July  26,  1932,  H.  Carman  (Carman  Coll.). 

North  Carolina:   Southern  Pines. 

South  Carolina:  Clemson  College,  Mar.  19,  1927,  W.  E.  Jones;  Clemson 
College,  Nov.  28,  1927,  D.  Dunavan;  Jacassee,  June  19,  1927;  Salem,  June  30, 
1927.  D.  Dunavan  (all  above  in  S.  C.  Exp.  Sta.) ;  S.  C.  (Uhler  Coll.) ;  Hopkins. 
Sept.,  1920,  C.  S.  Washington  (U.  S.  N.  M.). 

Georgia:  Floj'd  Co.,  Sept.  1,  1929,  Creaser  and  Becker;  Bleckley  Co.,  Sept. 
5,  1929.  Creaser  and  Becker;  Wrens,  July  22,  1930,  P.  W.  Oman;  Prattsburg, 
July  25.  1930.  R.  H.  Beamer;  Baker  Co..  Oct.  23,  1927.  C.  H.  Martin. 

Florida:  Gainesville,  June  9,  1918,  Drake;  Hilliard,  Aug.  19,  1930,  R.  H. 
Beamer. 

Al.\bama:  Prattsville,  July  21,  1930.  P.  W.  Oman;  Mount  Meigs,  July  21. 
1930.  Leonard  Tuthill;  Ala.  (Uhler  Coll.). 

Mississippi:  Smithville,  July  17,  1930,  R.  H.  Beamer;  Hamilton,  July  15, 
1930,  P.  W.  Oman;  Scooba,  July  17,  1930,  P.  W.  Oman;  luka,  July  14,  1930, 
P.  W.  Oman;  Crowder,  Aug.  30,  1925,  H.  M.  Harris;  Charleston,  Sept.  7,  1925, 
H.  M.  Harris;  Agricultural  College,  Sept.  1,  1925. 

Louisiana:    New  Orleans;  Baton  Rouge,  Mar.  9,  1929,  R.  M.  DeCoursey. 

Texas:    Bowie  Co.,  Aug.  16,  1928,  R.  H.  Beamer;  Texas  (Uhler  Coll.). 

Ark.^nsas:    Imboden,  B.  C.  Marshall  (U.  S.  N.  M.). 

Kansas:  Douglas  Co.,  F.  H.  Snow;  Douglas  Co.,  Rock  Pool,  Apr.  1,  1933, 
Carl  Cummings;  Doniphan  Co.,  Aug.  25,  1924,  Linsdale;  Cherokee  Co.,  Aug., 
1920,  Hungerford  and  Beamer. 

Arizona:    Arizona  (Uhler  Coll.). 

Notonecta  borealis  Bueno  and  Hussey,  1923 

(Color  Plates  I,  fig.  4,  and  II,  figs.  5  and  10;   Plate  XVI,  fig.  2.) 

1923.  .V.  borealis  Bueno  and  Hussey,  Bull.  Brooklyn  Ento.  Soc,  xviii,  pp.  104-107,  fig.  1 
(fig.  of  female  gonapophyses). 

1924.  .V.  borealis  Bueno  and  Hussey;  Hungerford,  Kansas  Univ.  S"ci.  Bull.,  xiv,  p.  426 
(says  tig.  labeled  A',   lutea  on  plate  xxxi,  Ivans.    Univ.   Sci.   Bull,   xi,  is  -V.   borealis  B.   &   H.) 

1926.    -V.  borealis  Bueno  and  Hussey;   Blatchley,  Heteroptera.     ...     p.   1053. 
1926.    A',   borealis  Bueno  and  Hussey;    Hungerford,  Bull.  Brooklyn  Ento.  Soc,  xxi,  p.   194 
(Abundant  in  Michigan). 

1928.    A',   borealis  Bueno  and  Hussey;   Hungerford,  Annals  Ento.   Soc.   Amer.,  xxi,  p.   142. 

1928.  A',  borealis  Bueno  and  Hussey;  Hungerford,  Can.  Ento.,  Ix,  p.  76.  (Describes 
dark-colored  forms.) 

1929.  N.  borealis  Bueno  and  Hussey;  Hutchinson,  Annals  of  South  African  Museum,  xxv, 
pt.  3,  p.  363. 

1930.  A',   borealis  Bueno  and  Hussey;    Walley,  Can.   Ento.,  vol.   Ixii,   No.   4,  p.   77. 


96  The  University  Science  Bulletin 

Also  referring  to  this  species: 

1897.  N.  insutata  Kirkaldy,  Tians.  Ento.  Soc.  London  for  1897,  p.  405.  (Specimen  in 
Signoret's  Coll.  labeled  A'^.  rugosa.) 

1904.  N.   lutea  Bueno,  Ento.  News,  xv,  p.  220. 

1905.  iV.   lutea  Bueno,  Jl.  N.  Y.  Ento.  Soc,  xiii,  p.  160. 

190S.  A'^.  ?  lutea  Kirkaldy  and  Bueno,  Proc.  Ento.  Soc.  Wash.,  x,  p.  108.  (Says  .■\nierican 
species  is  not  Miiller's.) 

1917.  A'',   lutea  Van  Duzee,  Cat.  Hern.  Amer.  North  of  Mexico,  p.   450. 

1918.  A^.   lutea  Hungerford,  Ento.  News,  xxix,  p.  244,  pi.  xv,  fig.  1   (fig.  of  ovipositor). 

1919.  A'^.   lutea  Hungerford,  Kans.  Univ.  Sci.  Bull.,  xi,  p.  331. 

Size.  Length,  12.4  mm.  to  14.2  mm.;  width  of  pronotiim,  4.3  mm. 
to  5.1  mm. 

Shape.  Plump,  deep  bodied,  with  the  abdomen  broader  than  the 
pronotum  at  the  humeral  angles. 

Color.  General  color  luteous,  usually  with  a  dark  streak  follow- 
ing the  costal  margin  from  the  base  of  the  hemelytra;  membrane 
pale.  Scutellum  pale  yellow.  Pronotum,  head  and  limbs  ivory 
white  or  the  last  darker  or  streaked  with  brown;  sometimes  face 
and  limbs  more  or  less  greenish.  Venter  dark.  In  some  specimens 
the  brown  streak  nearly  covers  the  clavus,  and  in  rare  cases  speci- 
mens have  been  encountered  that  have  a  black  scutellum  and  heme- 
lytra as  dark  as  in  N.  irrorata  Uhler.  In  such  cases  of  melan- 
chroism,  however,  the  posterior  lobe  of  the  membrane  remains  pale. 

Structural  Characteristics.  Anterior  outline  of  head,  viewed  from 
above,  somewhat  flattened,  but  vertex  produced  beyond  the  eyes; 
vertex  at  least  a  fifth  broader  in  front  than  its  length;  anterior 
margin  of  vertex  less  convex  and  plainly  longer  than  the  frontal 
margin  of  the  eye;  anterior  breadth  of  vertex  :  synthlipsis  ::  2.3:1. 
Pronotum  slightly  more  than  twice  as  long  as  the  head ;  lateral  mar- 
gins divergent,  slightly  concave;  anterior  angles  not  embracing  the 
eyes;  lateral  ledge  sigmoid  and  oblique  and  shorter  than  the  length 
of  the  eye  below  it.  Scutellum  small,  slightly  shorter  than  the  prono- 
tum. Posterior  lobe  of  membrane  shorter  than  the  anterior  lobe. 
Anterior  trochanters  of  the  male  without  hook  or  tubercle ;  mesotro- 
chanters  rounded.  The  terminal  abdominal  sternite  of  the  female 
large  and  constricted  near  the  tip,  which  is  entire.  First  pair  of 
gonapophyses  long.    Male  genital  capsule  as  shown  on  Plate  XVI. 

Location  of  Type.  Type  and  paratypes  in  collection  of  J.  R.  de  la 
Torre-Buneo;  paratypes  in  collections  of  R.  F.  Hussey,  H.  B.  Hun- 
gerford, H.  M.  Parshley,  W.  E.  Hoffmann,  and  the  Museum  of  Zool- 
ogy, University  of  Michigan. 

Comparative  Notes.  This  species  is  related  to  N.  lutea  Miiller 
of  northern  Europe  from  which  it  differs  in  a  number  of  respects. 
Besides  the  differences  in  genitalia  in  both  sexes,  the  V.  lutea  Miiller 


Hungerford:    Genus  Notonecta  97 

has  a  hook  on  the  anterior  trochanter  of  the  male.  Both  the  head 
and  mesofemoral  spine  of  N.  borealis  B.  &  H.  are  relatively  smaller 
than  in  the  old  world  form. 

Biological  Notes.  This  species  deposits  its  eggs  in  the  tissues  of 
aquatic  plants.  The  eggs  may  be  completely  covered  or  be  partially 
exposed.  The  life  history  of  this  species  should  be  studied.  Miss 
Lucile  Rice  made  some  observations  (unpublished)  on  this  species 
at  the  Biological  Station  of  the  University  of  Michigan  on  Douglas 
Lake  during  the  summer  of  1927.  She  found  no  adults  before  June 
29th,  at  which  time  she  took  two,  and  observed  many  last  instar 
nymphs.  Seven  of  these  nymphs  transformed  in  the  laboratory 
within  the  next  twenty-four  hours.  By  July  6th  there  were  many 
adults  in  the  pool.  One  of  the  adults  brought  in  June  29th  deposited 
an  egg  on  July  9th.  Egg  laying  continued  till  August  11th.  How- 
ever, none  of  the  eggs,  either  from  laboratory  or  field,  showed  any 
signs  of  developing  during  the  summer.  Miss  Rice  therefore  took 
them  to  Tennes.see,  and  some  of  them  deposited  July  22d  hatched  in 
her  laboratory  on  December  15th. 

From  the  collecting  records  which  I  have  made  about  Douglas 
Lake,  during  a  period  of  ten  years,  and  from  Miss  Rice's  observa- 
tion, it  appears  that  this  northern  species  winters  in  the  egg  state. 

Data  on  Distribution: 

Canada 

Quebec:  East  Coast  James  Bay,  Sept.,  1920.  F.  Johansen;  Natashquan,  Que., 
Aug.  S,  10.  11.  1929,  W.  J.  Brown  (Ottawa,  Can.). 

British  Columbia:    Bearfoot  Mts.,  Sept.  17,  1903  (U.  S.  N.  M.). 

United  States  of  America 

Michigan:  Cheboygan  Co.,  Aug.  29,  1918;  Douglas  L.,  July  4,  1926,  C.  H. 
Martin;  Douglas  L.,  July  29,  1926,  H.  B.  Hungerford;  Douglas  L.  (Sedge  Pool), 
1925  and  July  6,  1923,  H.  B.  Hungerford;  Douglas  L.  (Biyants  Bog),  July  17, 
1925.  H.  B.  Hungerford;  Mackinac  Island,  Michigan,  Aug.  19,  1925,  and  Aug.  4. 
1929.  H.  B.  Hungerford. 

Minnesota:  Lake  Co.,  Stony  R.  Camp  No.  9,  Aug.  16,  1922,  H.  B.  Hunger- 
ford, H.  H.  Knight,  W.  E.  Hoffmann;  Hibbings,  Aug.  18,  1922,  H.  B.  Hungerford. 

Notonecta  melaena  Kirkaldy,  1897 

(Color  Plate  III,  fig.   7;   Plate  XIII,  fig.   11) 

1897.    K.   sliooterii  var.  melaena  Kirkaldy,  Trans.   Ento.   Soc.   London,   1897,   p.    406. 

1904.  K.  shoeterii  var.  melaena  Kirkaldy,  Wien.  Ento.  Zeit.,  XXIII,  p.   132. 

1905.  .V.  shootcrii  var.  melaena  Kirkaldy;   Bueno.  Jl.  N.  Y.  Ento.  Soc.  XIII,  p.   Kil. 

Referring  to  this  species,  also: 

1807.    A',  shooterii  var.  tearrn  Kirkaldy,  Trans.  Ento.  Soc.  London,   1897,  ]).  407. 
1925.    X.  distincta  Hungerford,  Can.  Ento.,  LVII,  p.  241,  pi.  VI,  fig.  4. 
192S.    .V.    diftinrtu   Hungerford,  .\nnals  Ento.  Soc.  .'\m.,  XXI,  pi.  IX,  fig.  S. 


98  The  University  Science  Bulletin 

S)ize.  Length,  10.8  mm.  to  11.7  mm.;  width  of  pronotiim,  4  mm. 
to  4.3  mm. 

Color.  A  bichromatic  species,  the  luteous  forms  having  a  tinge 
of  orange,  the  pigmented  forms  with  black  scutellum  and  hemelytra 
varying  from  light  brown  to  nearly  black ;  a  light-brown  streak  may 
be  present  along  costal  margin  and  above  the  claval  suture;  base  of 
membrane  black  and  tip  white.  Propleura  mostly  pale.  Pro-  and 
mesosternum  dark.     Abdominal  venter  with  black  maculations. 

Structural  Characteristics.  Head  less  than  one-half  length  of  pro- 
notum;  vertex  shorter  than  its  anterior  width  as  seen  from  above; 
anterior  margin  of  vertex  less  convex  than  frontal  margin  of  the 
eye;  anterior  breadth  of  vertex  :  synthlipsis  ::  11:7;  the  synthlipsis 
broad,  slightly  more  than  half  the  rear  margin  of  an  eye;  inner  mar- 
gins of  eyes  concave.  Pronotum  steeply  declivant;  lateral  margins 
of  pronotum  nearly  straight  and  diverging  at  an  angle  of  from  44 
to  50  degrees,  in  the  dark  form  slightly  convex  and  diverging  less 
markedly,  the  angle  being  about  42  degrees ;  anterolateral  angle  em- 
bracing the  eye,  more  in  the  luteous  forms  than  in  the  pigmented 
forms,  some  of  which  show  angles  but  slightly  acute;  lateral  ledge 
a  little  broader  in  the  luteous  forms  and  shorter  than  the  rear  mar- 
gin of  the  eye  beneath,  as  long  as  rear  margin  of  the  eye  in  the  pig- 
mented forms ;  lateral  ledge  faintly  sigmoid,  the  anterior  end  curved 
downward  and  posterior  end  turned  upward.  Posterior  lobe  of 
membrane  much  reduced  in  luteous  forms  and  slightly  shorter  in 
dark  forms.  Anterior  trochanter  of  male  with  basal  tubercle  and 
stout  hook  characteristic  of  the  A'',  shooteri  Uhler  group.  Meso- 
trochanters  rounded.  Penultimate  abdominal  sternite  of  female 
broadly  triangular,  broadly  notched  at  tip  and  overlapping  base 
of  following  sternite,  which  is  not  notched  at  tip.  Female  gona- 
pophvses  short.  ]Male  genital  capsule  as  shown  on  Plate  XIII, 
fig.  11. 

Location  of  Type.  Kirkaldy  named  this  as  a  variety  of  N. 
shooterii  Uhler,  from  Mexico  (Noualhier  coll.).  I  have  seen  this 
specimen,  a  male,  in  the  Paris  Museum.  My  N.  distincta  is  the  same 
species.  I  have  seen  the  pale  specimens  mentioned  by  Kirkaldy  as 
having  developed  posterior  lobes  to  the  membranes  and  for  which  he 
proposed  the  varietal  name  tearca.  He  says:  "This  form  I  have  seen 
only  from  Mexico  (in  Stockholm  Mus.  and  in  the  Paris  Mus.,  where 
it  was  labeled  "americana"  by  Fallou) ."  The  Paris  specimens  are 
labeled  "Mexique,  Coll.  G.  Fallou  259-95."  The  male  has  the  mem- 
brane lobes  subequal,  but  the  female  has  them  unequal  like  the 


Hungerford:    Genus  Notonecta  99 

Colombian  specimen  he  labeled  "ochrothoe."  As  far  as  I  can  de- 
termine the  liiteoiis  male  with  the  normal  posterior  lobe  of  the  mem- 
brane is  an  unpigmented  specimen  of  the  dark  form  of  N.  melaena 
Kirkaldy. 

Comparative  Notes.  The  pigmented  forms  have  been  confused 
until  recently  with  N.  distinctoidea  Hungerford,  being  about  the 
same  size  and  color.  The  presence  of  the  large  hook  on  the  anterior 
trochanter  of  the  male  and  the  penultimate  abdominal  sternite  of 
female  notched,  separate  this  species  from  N.  distinctoidea  Hunger- 
ford. 

Notes  on  Distribution.  Described  from  Mexico.  Paris  Museum 
has  a  series  of  dark  forms  labeled  ''Etat  Jalisco,  L.  Diguet  1903."  I 
have  specimens  labeled:  "10  miles  N.  E.  Leon  Quanajuato,  Mex. 
Aug.  17,  1932,  Hobart  Smith";  my  N.  distincta  type  is  labeled  ''Co- 
mondu,  Lower  Calif.  Chas.  D.  Haines,  March,  1889."  The  two 
specimens  I  recognize  as  the  luteous  form  of  this  species  are  labeled 
''Las  Parras,  Baja,  Calif.,  W.  M.  Mann." 

Notonecta  ochrothoe"^  Kirkaldy,  1897 

1897.     .V.  shooterii  var.  ochrothoe  Kirkaldy,  Trans.  Ento.  Soc.  London  for  1897,  i>.   407. 
1905.    TV.  shooterii  var.  ochrothoe  Kirkaldy;   Bueno,  Jl.  N.  Y.  Ento.  Soc,  XIII,  p.   161. 

Size.  Length,  10.8  mm.  to  11.4  mm.;  width  of  pronotum,  4  mm. 
to  4.4  mm. 

Color.  Only  the  luteous  form  known  to  me,  and  this  is  pale 
luteous. 

Structural  Characteristics.  Anterior  outline  of  head,  viewed  from 
above,  convex;  vertex  usually  shorter  than  its  anterior  width;  mar- 
gin of  vertex  less  convex  than  margin  of  the  eye;  anterior  breadth 
of  vertex  :  synthlipsis  ::  3:2.  Pronotum  not  markedly  declivant; 
pronotum  more  than  twice  length  of  head;  lateral  margins  slightly 
divergent  and  nearly  straight;  anterolateral  angles  acute  and  em- 
bracing the  eyes;  lateral  ledge,  as  seen  from  the  side,  nearly  straight, 
about  as  long  as  rear  margin  of  the  eye  beneath.  Scutellum  shorter 
than  pronotum.  Posterior  lobe  of  membrane  reduced.  Anterior 
trochanter  of  male  with  stout  hook  and  tuberculate  projection  on 
basal  angle.  Mesotrochanter  rounded.  The  penultimate  abdominal 
sternite  of  female  broadly  triangular,  tip  incised  and  projecting 
upon  the  base  of  the  last  sternite,  which  is  not  notched  at  tip. 

*  Kirkaldy.  1!MI4.  in  his  I'ber  Notnnectiden,  Wien.  Eiit.  Zict..  XXIII.  and  in  his  Cataloguo 
Proc.  Ent.  Soc.  Wash.,  X,  appears  to  have  dropped  this  variety.  Nevertheless  I  find  a  male 
in  the  Paris  Museum  from  "Colombie"  labeled  this  species  by  Kirkaldy,  and  since  it  is  not 
N.  fhooterii  Uhler,  I  make  it  the  type  and  raise  his  variety  to  specific  rank. 


100  The  University  Science  Bulletin 

Female  gonapophyses  short.  Male  genital  capsule  shaped  much 
like  N.  melaena  Kirk,  but  with  a  thickened  protuberance  on  keel. 

Location  of  Type.    In  Paris  Museum. 

Cojnparative  Notes.  Differs  from  luteous  form  of  N.  melaena 
Kirkaldy  by  less  declivant  pronotum  and  by  the  thickened  pro- 
tuberance on  the  keel  of  the  male  genital  capsule. 

Data  on  Distribution.  Colombia,  S.  A.,  and  the  following:  ''San 
Cristobel  M.  F.  3-7-1929";  San  Diego  Co.,  Calif.,  Doway  VaL,  4-9, 
1930,  C.  and  D.  Martin. 

Notonccta  rcpanda  new  species 

(Text   figure   7) 

Size.  Length,  11.7  mm.  to  13.8  mm. ;  width  of  pronotum,  4  mm.  to 
4.8  mm. 

Color.  Color  markings  as  in  A' .  shooteri  Uhler. 

Structural  Characteristics.  Anterior  outline  of  head,  viewed  from 
above,  less  convex  than  in  N.  shooteri  Uhler;  vertex  shorter  than 
its  anterior  width ;  margin  of  vertex  usually  less  convex  than  margin 
of  the  eye;  anterior  breadth  of  vertex  :  synthlipsis  : :  3:2.  Pronotum 
less  than  twice  the  length  of  the  head;  lateral  margins  slightly  di- 
vergent and  straight  to  slightly  convex,  more  divergent,  of  coui'se,  in 
the  dark  forms;  anterior  angles  acute  and  closely  embracing  the 
eyes;  lateral  ledge,  as  seen  from  the  side,  nearly  straight  throughout 
and  as  long  or  longer  than  the  rear  margin  of  the  eye  below  it; 
anterior  end  of  lateral  ledge  of  pronotum  broadly  flattened,  the 
ledge  plainly  wider  beneath  than  the  diameter  of  the  second  antennal 
segment.  Scutellum  short  as  in  N.  shooteri  Uhler.  The  color  forms 
as  in  A^.  shooteri  Uhler.  Trochanters  the  same.  The  penultimate 
abdominal  sternite  of  female  broadly  triangular,  the  apex  broader 
than  in  Uhler's  species,  but  projecting  upon  last  sternite,  which  is 
not  notched  at  tip.  Female  gonapophyses  short.  Male  capsule  not 
elongate  but  compact  much  as  in  N.  melaena. 

Location  of  Type.  In  the  Francis  Huntington  Snow  Entomolog- 
ical Museum  of  the  University  of  Kansas.  Described  from  35  speci- 
mens labeled:  "Cochise  Co.,  Arizona,  July  29,  1927,  P.  A.  Readio, 
R.  H.  Beamer,  L.  D.  Anderson." 

Comparative  Notes.  This  species  has  the  appearance  of  N. 
shooteri  Uhler,  but  the  lateral  ledge  of  the  pronotum  is  more  ex- 
planate,  the  genital  capsules  of  males  different  and  the  penultimate 
abdominal  sternite  of  female  broader  in  the  portion  that  overlaps 
the  last  segment. 


Hungerford:    Genus  Notonecta 


101 


Data  on  Distribution: 

Akizona:  Cochise  Co.,  July  29,  1927  (type  series);  Baboquivari  Mts.,  July 
18,  1932,  R.  H.  Beamer;  Douglas,  Aug.,  San  Bernardino  Ranch,  3,750  feet, 
F.  H.  Snow. 

Mexico:  D.  F.  Texoco  Sea,  Jan.  31,  1926,  A.  Dampf.;  Mexico  D.  F.,  Apr. 
22,  1910;  D.  Fedrl.  L.  Conradt  (U.  S.  N.  M.) ;  Xochimilco  Lake,  near  Mexico 
City,  June  2.  1926,  A.  Dampf. 


N.  repanda 


N   shooten 


Text  Figure  7.     Notonecta  repanda  Hungerford  and  Notonecta  shooteri  Uhler. 
Underside  of  abdomen  of  females.     (Subgenus  Bichromonecta.) 


Notonecta  shooteri  Uhler,  1894 

(Color  Plate  I,  fig.  6;  Plate  XIII,  fig.  8,  Text  fig.  7) 

1894.  .v.    shooterii  Tlhler;   Pioo.  Calif.  Acad.  Sci.  (2),  iv,  p.  292. 

1897.  N.  shooterii  Uhler;   Kirkaldy,  Trans.  Ento.  Soc.  London  for  1897,  p.  406  (in  part). 

1901.  N.  shooterii  Uhler;  Champion,  Biol.  Centr.  Amer.,  Hem.-Het.  II,  p.  370  (in  part). 

1904.  .V.   shooterii  Uhler;    Kirkaldy,  Wien.  Ento.  Zeit.,  xxiii,  pp.  94  and  132. 

1905.  A',  shooterii  Uhler;   Bueno,  Jl.  N.  Y.  Ento.  Soc,  xiii,  p.   161. 

1909.    A^.  shooterii  Uhler;  Kirkaldy  &  Bueno,  Proo.  Ent.  Soc.  Wash.,  x,  p.  199. 

1916.  A',   shooteri  Uhler;    Browne,  Jl.   Morph.,  xxvii,  p.   129   (Cytology). 

1917.  A',  shooterii  Uhler;    Van  Duzee,  Catalog  Hemipt.,  p.   453   (in  part). 

1918.  A'.   .'/(oo^crM  I'hler;    Hungerford,  Ento.  News,  xxix,  p.  245,  pt.  xv,  fig.  8.     (Ovipositor.) 

1919.  A^.  shooterii  Uhler;    Hungerford,  Kans.  Univ.  Sci.  Bull.,  xi,  pp.   168,  172,  181,  312, 
331;   pi.  xxiii,  fig.   8,  pi.  xxxi,  fig.   5. 

1922.  A^  shooterii  Uhler;   Hungerford,  Kans.  Univ.  Sci.  Bull.,  xiv,  p.  426. 

1923.  N.  shooterii  Uhler;   Bueno  &  Hussey,  Bull.  Brookl.  Ent.  Soc,  xviii,  p.   106. 

1924.  N.  shooterii  Uhler;    Hale,  Proc.  Lin.  Soc.   N.   S.  Wales,  xlix,  pt.  4,  p.   462. 

1928.    A",   shooterii  Ihler;    Hungerford,  Annals  Ento.  Soc.  Auier.,  xxi,  p.  143;   pi.  ix.  fig.  5. 
(Anterior  trochanter  of  male.) 

1928.  A',   shooterii  Uhler;   Hungerford,  Can.  Ento.,  Ix,  p.   76. 

1929.  A',  shooterii  Uhler;   Hutchinson,  Annals  S.  Afr.  Mus.,  xxv,  pt.  iii,  p.   364. 

Size.    Length,  11.7  mm.  to  13.5  mm. ;  wicith  of  pronotum,  4.4  mm. 
to  4.8  mm. 

Color.     A  bichromatic  species.     One  form  pale  luteous  with  or 
.S— 3482 


102  The  University  Science  Bulletin 

without  a  submarginal  stripe  and  one  or  two  elongate  triangular 
brown  spots  on  corium.  The  other  form  dark,  with  black  scutellum 
and  brown  hemelytra,  which  are  streaked  and  spotted  with  pale 
luteous,  the  pale  markings  being  a  broad  streak  on  clavus,  a  trans- 
verse irregular  spot  at  end  of  basal  third  of  corium,  a  spot  at  distal 
end  of  corium  and  another  on  anterior  lobe  of  membrane.  The 
posterior  lobe  of  membrane  may  be  pale  to  nearly  black.  The 
abdominal  venter  luteous  with  brown  maculations  in  both  color 
phases.  Face,  limbs  and  connexivum  may  be  yellowish  or  suffused 
with  green. 

Structural  Characteristics.  Anterior  outline  of  head,  viewed  from 
above,  convex;  vertex  usually  longer  or  subequal  to  its  anterior 
width;  margin  of  vertex  more  convex  than  margin  of  the  eye;  an- 
terior breadth  of  vertex  :  synthlipsis  ::  9.3:5.5.  Inner  margins  of 
eyes  slightly  concave.  Pronotum  usually  less  than  twice  the  length 
of  the  head;  lateral  margins  slightly  divergent  and  straight  to 
slightly  convex;  anterolateral  angles  acute  and  closely  embracing 
the  eyes;  lateral  ledge,  as  seen  from  the  side,  straight  in  front,  un- 
dulate behind,  shorter  than  the  rear  margin  of  the  eye  below  it,  and 
nearly  horizontal.  Scutellum  short,  about  as  long  as  pronotum  and 
shorter  than  ridges  of  hemelytral  commissure.  Anterior  lobe  of 
membrane  much  longer  and  more  developed  than  the  posterior  lobe 
in  luteous  phase  and  plainly  longer  in  the  dark  phase.  The  luteous 
forms,  besides  the  reduced  posterior  lobe  of  hemelytra,  have  the 
flight  wings  reduced,  and  this  is  accompanied  by  a  reduction  in  the 
prominence  of  the  humeri  and  in  the  size  of  the  scutellum.  The 
reduced  humeri  cause  the  lateral  margins  of  the  pronotum  to  appear 
more  nearly  parallel.  Anterior  trochanters  of  the  males  with  broad, 
stout  hook  in  usual  position  and  in  addition  a  tuberculate  projection 
on  basal  angle  (see  plate  IX,  fig.  4) .  Mesotrochanter  rounded.  The 
penultimate  abdominal  sternite  of  female  elongate  triangular,  the 
sides  converging  caudally,  tip  incised  and  projecting  upon  the  base 
of  last  sternite,  which  is  not  notched  at  tip.  Female  gonapophyses 
(first  pair)  short.     Male  genital  capsule  as  shown  on  Plate  XIII. 

Location  of  Type.  In  the  Uhler  collection  at  the  U.  S.  N.  M., 
Washington,  D.  C.  The  type  series  was  found  near  San  Diego, 
California,  on  October  19,  by  Mr.  Shooter,  for  whom  Doctor  Uhler 
named  the  species.  The  spelling  of  the  name  might  have  been 
N.  shooteri. 

Comparative  Notes.  This  species  is  typical  of  a  group  that  have 
the  large,  stout  hook  on  anterior  trochanters  of  the  males  and  fe- 


Hungerford:    Genus  Notonecta  103 

males,  with  the  penultimate  sternite  elongate  triangular,  thus  in- 
termediate between  the  A^.  mexicana  A.  &  S.  series  and  the  N.  un- 
dulata  Say  group.  The  males  readilj^  recognized  by  the  elongate 
portion  of  the  capsule  behind  the  keel  protuberence. 

Data  on  Distribution.  The  group  as  a  whole  ranges  from  Cali- 
fornia to  Colombia,  S.  A.,  but  the  species  N.  shooteri  Uhler  appears 
to  be  restricted  to  California.    I  have  the  following  records: 

California:  Marin  Co.,  Aug.  3,  1929,  L.  D.  Anderson;  Winters,  Aug.  6,  1929, 
L.  D.  Anderson;  Campo,  Laguna  Mts.,  Aug.  25,  1932,  H.  W.  Capps;  Fresno, 
June  20,  1926,  C.  J.  Drake;  Laguna  Beach,  C.  T.  Dodds;  Los  Angeles  (Uhler 
Coll.);  San  Diego,  Oct.  19,  Mr.  Shooter  (Uliler  Coll.). 

"Viti-Ins":     Distant  coll.  1911-383  (British  Mus.). 

Notonecta  uhleri  Kirkaldy,  1897 

(Color  Plate  III,    fig.   9;    Plate  XIII,   fig.    10) 

1897.  N.  uhleri  Kirkaldy,  Annals  &  Mag.  Nat.  Hist.,  ser.  6,  xx,  p.  58. 

1897.  N.' uhleri  Kirkaldy,  Trans.  Ento  Soc.  London,  1897,  p.  415. 

1902.  N.  uhleri  Kirkaldy;   Bueno,  Jl.  N.  Y.  Ento.  Soc,  x,  pp.   231,  235. 

1904.  N.  uhleri  Kirkaldy,  Wien.   Ento.  Zeit,  xxiii,  p.   132. 

1905.  N.  uhleri  Kirkaldy;  Bueno,  Jl.  N.  Y.  Ento.  Soc,  xiii,  pp.  46  and  157,  pi.  vii, 
fig.  4. 

1906.  N.  uhleri  Kirkaldy;   Bueno  and  Brimley,  Ento.  News,  xviii,  p.  436. 

1909.  N.  uhleri  Kirkaldy  and  Bueno,  Proc.  Ento.  Soc  Washington,  x,  No.  3-4,  p.  199. 
(Cat.  of  Am.  Aquatic  &  Semiaquatic  Hemiptera.) 

1910.  N.   uhleri  Kirkaldy;    Smith,  Cat.  Ins.  N.  J.,  edn.   3,  p.   169. 

1914.    N.  uhleri  Kirkaldy;  Barber,  Bull.  Am.  Mus.  Nat.  Hist.,  xxxiii,  p.  499. 
1917.    N.  uhleri  Kirkaldy;   Parshley,  Occasional  Papers  of  the  Boston  Society  of  Natural 
History,  vii,  p.  112. 

1917.  N.   uhleri  Kirkaldy;    Van   Duzee,    Catalog   of   Hemiptera,    p.    451. 

1918.  N.   uhleri  Kirkaldy;    Hungerford,  Ento.  News,  xxix,   p.  245,  PI.  xv,  fig.   10. 

1919.  N.  uhleri  Kirkaldy;  Hungerford,  Kans.  Univ.  Sci.  Bull.,  xi,  pp.  167,  170,  180, 
181,  312,  331;   PI.  xxiii,  fig.  10. 

1923.  N.  uhleri  Kirkaldy;  Bueno,  Conn.  State  Geol.  &  Nat.  Hist.  Survey  Bull.  No. 
34,  p.  405. 

1923.  N.  uhleri  Kirkaldy;  Hungerford,  Kansas  Univ.  Sci.  Bull.,  xiv,  p.  426.  (Mention 
only). 

1926.    N.  uhleri  Kirkaldy;    Blatchley,   Heteroptera  of  Eastern  North  America,   p.    1055. 

1926.  N.  uhleri  Kirkaldy;  Leonard,  Cornell  Univ.  Agri.  Exp.  Sta.  Memoir  101,  p.  138, 
(Apr.  to  Nov.) 

1928.    N.  uhleri  Kirkaldy:    Hungerford,  Ento.   News,  xxxix,  p.   156. 

Size.  Length,  10  mm.  to  12  mm. ;  width  across  pronotum,  3.3  mm. 
to  4  mm.    The  males  usually  smaller  than  the  females. 

Shape.    The  males  at  least  are  slender. 

Color.  Orange  to  brick  red  and  black.  Anterior  half  of  pronotum, 
head,  limbs  and  underside  of  connexivum  of  usual  color  or  a  little 
darker  and  sometimes  greenish;  a  dark  linear  longitudinal  band 
usually  present  on  pronotum  behind  the  caudolateral  margin  of  the 
eye.  Scutellum  black.  Hemelytra  orange  to  brick  red;  corium 
crossed  by  broad,  irregular  black  band  extending  from  near  tip  of 


104  The  University  Science  Bulletin 

clavus  to  costal  margin.  Posterior  lobe,  at  least  of  membrane,  dark, 
and  both  lobes  crossed  by  black  or  nearly  black  band  that  often 
covers  the  lobes  to  their  tips  and  sometimes  darkens  entire  mem- 
brane. The  hemelytra  usually  clothed  v^ith  silvery  hairs  that  are 
more  pronounced  on  the  corium;  a  few  such  hairs  may  also  be  seen 
on  scutellum.    The  venter  is  dark. 

Structural  Characteristics.  Anterior  outline  of  the  head,  as  seen 
from  above,  slightly  convex;  vertex  longer  than  its  anterior  width 
and  "shaped  like  an  inverted  wine-decanter";  anterior  margin 
of  vertex  shorter  than  the  frontal  margin  of  an  eye;  the  anterior 
breadth  of  vertex  :  synthlipsis  : :  6  or  8:1;  the  eyes  at  synthlipsis 
often  nearly  contiguous  and  their  inner  margins  curved.  Head 
shorter  than  length  of  pronotum.  Lateral  margins  of  pronotum 
strongly  divergent  and  moderately  concave ;  anterior  angles  normal ; 
lateral  ledge,  as  seen  from  the  side,  curved  (convex  above),  oblique 
and  plainly  shorter  than  the  rear  margin  of  the  eye  below  it.  The 
scutellum  plainly  longer  than  the  pronotum  and  also  longer  than 
the  ridge  of  the  hemelytral  suture.  Lobes  of  membrane  subequal, 
the  anterior  lobe  sometimes  slightly  longer.  Anterior  trochanter 
of  the  male  with  a  slight  tubercle  in  place  of  a  hook.  Mesotro- 
chanter  right-angulate  or  slightly  produced.  Terminal  abdominal 
sternite  of  female  carinate  and  slightly  notched  at  tip ;  the  first  pair 
of  gonapophyses  short.  The  male  genital  capsule  as  shown  on  Plate 
XIII,  fig.  10. 

Location  of  Type.    In  the  P.  R.  Uhler  collection  at  the  U.  S.  N. 
Museum. 

Comparative  Notes.    The  small  size,  the  reddish  and  black  color 
and  the  very  narrow  synthlipsis  readily  distinguish  this  species. 

Data  on  Distribution.  A  Uhler  specimen  is  labeled  "Stony  Run, 
Pa."  "Among  grass  in  water."  In  the  Paris  Museum  I  found  a 
specimen  compared  with  type  by  Kirkaldy  labeled  "Mus.  Paris, 
Neue  Orleans,  Coll.  Guerin  Menev.  ex.  coll.  A.  Salle,  1897."  The 
British  Museum  has  specimens  from  Mass.  The  Uhler  collection 
from  Florida  and  Massachusetts.  Bueno  reports  it  from  New  York, 
Washington,  D.  C,  New  Jersey  and  Louisiana.  Doctor  Drake  took 
it  at  Gainesville,  Florida.  Doctor  Blatchley  adds:  Maryland,  North 
Carolina  and  southern  Illinois.  There  are  specimens  in  the  Uni- 
versity of  Kansas  collection  labeled  as  follows:  "Boardman,  N.  C, 
Sept.  21,  1915,  R.  W.  Leiby";  "Raleigh,  N.  C,  Oct.  16,  1905,  C.  S. 
Brimley";  "Florence,  S.  C,  July  2,  1930,  D.  Dunavan  (Clemson 
College  Coll.);"  "Putnam  Co.,  N.  Y.";  "Leland,  Miss.,  9-14-21"; 


Hungerford:    Genus  Notonecta  105 

"Baker  Co.,  Ga.,  Oct.  23,  1927,  0.  H.  Martin";  "Prattsbiirg,  Ga., 
7-25-1930,  R.  H.  Beamer";  "Hilliard,  Fla.,  8-31-30,  Paul  W.  Oman"; 
"Norfolk,  Va.,  8-15-28,  G.  E.  Gould;"  also  "Oct.  1,  1932,  L.  D.  An- 
derson; Annapolis,  Md.,  Sept.  18,  1932,  Paul  Oman." 

Thus  N.  uhleri  Kirkaldy  has  been  taken  in  the  following  states: 
Mass.,  New  Jersey,  Pa.,  N.  Y.,  Md.,  Va.,  Dist.  of  Columbia,  111., 
N.  Carolina,  La.,  Ga.,  Fla.,  Miss.  The  range,  so  far  as  we  know  it, 
extends,  therefore,  from  Massachusetts  south  to  Florida  and  west- 
ward to  the  ^Mississippi  river. 

Notonecta  raleighi  Bueno,  1907 

(Color  Plate  I,  fig.  7;   Plate  XIII,  fig.  6) 

1907.    .V.  raleighi  Bueno,  Can.  Ento.  xxxix,  p.  225. 

1907.  .V.  raleighi  Bueno  and  Brimley,  Ento.  New=,  x\Tii,  p.  436.  (Feb.  to  Nov.  in  N. 
Carolina.) 

1909.  N.  raleighi  Bueno;  Kirkaldy  &  Bueno,  Proc.  Ento.  Soc.  Wash.,  x,  p.  199. 

1910.  A^.  raleighi  Bueno;   Smith,  Catalog  Ins.  N.  J.,  edn.  3,  p.   169. 

1917.  -V.  raleighi  Bueno;    Van  Duzee,  Catalog  Hemip.,  p.   452. 

1918.  -V.  raleighi  Bueno;  Hungerford,  Ento.  News,  xxix,  p.  245,  pi.  xv,  fig.  2.  (Fig. 
of  ovipositor.) 

1919.  A',  raleighi  Bueno;  Hungerford,  Kans.  Univ.  Sci.  Bull.,  xi,  pp.  167,  169,  181,  239, 
331;  pi.  xxiii,  fig.  2. 

1923.  A^.  raleighi  Bueno;  Conn.  St.  Geol.  &  Nat.  Hist.  Survey  Bull.  No.  xxxiv,  pp. 
404-405. 

1925.  A'',  raleighi  Bueno;   Hungerford  &  Beamer,  Ento.  News,  xxxvi,  p.   297. 

1926.  N.  raleighi  Bueno;  Leonard,  Cornell  Univ.  Agri.  Exp.  Sta.  Memoir  101,  p.  139. 
(March,  White  Plains,  N.  Y.  det.  Bueno.) 

1926.    A'',  raleighi  Bueno;    Blatchley,   Heteroptera     ...     p.   1055. 

1930.    N.  raleighi  Bueno;   Hungerford,  Bull.  Brookl.  Ento.  Soc,  xxv,  p.  140. 

Referring  to  this  species,  also: 

Probably  some  specimens  prior  to  1907  were  included  under 
A^.  variabilis  Fieber  in  error. 

Size.  Length,  8  mm.  to  8.9  mm.;  width  of  pronotum,  2.5  mm.  to 
2.9  mm. 

Color.  A  pale  species  that  may  be  pale  yellowish  throughout  or 
with  scutellum  black  save  for  lateral  margins  and  hemelytra  more 
or  less  smudged  with  black.  Bueno  says:  "Scutellum  ranges  from 
pure  light  yellowish  to  black,  disk  margined  with  smoky  orange- 
yellow  on  hemelytral  margins.  Hemelytra  ranging  from  white  with 
vague  beginnings  of  the  corial  fasciae  and  black  humeri  with  white 
membrane,  through  all  intergrades  to  a  form  with  a  blackish  stripe 
along  the  anterior  margin  of  the  corium;  black  margins  to  the  clavus 
along  the  scutellar  edges;  a  black  streak  along  the  corium  near  to 
and  parallel  to  the  claval  suture;  black  corial  fasciae  merging  into 
the  black  membrane  which  shade  off  into  smoky  and  then  white 
at  apex.    One  of  the  types  is  the  most  pronouncedly  melanic  speci- 


106  The  University  Science  Bulletin 

men  of  the  species  in  a  series  of  sixty  or  so  specimens.  In  this 
the  extreme  of  scutellar  darkness  with  orange-red  edges  obtains. 
The  external  edges  of  the  clavus  are  broadly  black,  shading  into 
smoky  to  the  corial  suture ;  the  dark  band  on  the  corium  parallel  to 
this  suture  is  broad;  the  humerus  has  a  black  streak  running  into 
the  corium,  which  is  dark  luteous,  except  for  the  black  fascise  which 
coalesce  with  the  black  membrane,  which  in  turn  lightens  to  smoky 
at  the  apex." 

Structural  Characteristics.  Our  smallest  North  American  species. 
Anterior  outline  of  head  somewhat  rounded,  vertex  and  eyes  in 
nearly  an  even  curve :  vertex  slightly  longer  than  its  anterior  width ; 
anterior  margin  of  vertex  about  equal  to  the  frontal  margin  of  the 
eye.  Anterior  breadth  of  vertex  :  synthlipsis  : :  5:1,  often  less.  Syn- 
thlipsis  usually  less  than  one-fourth  the  width  of  the  eye.  Pronotum 
two-  to  four-tenths  longer  than  the  head,  lateral  margins  straight, 
divergent,  anterior  angles  normal  to  slightly  embracing  the  eyes, 
lateral  ledge  a  little  bowed  and  oblique  as  seen  from  the  side.  Mem- 
brane lobes  subequal.  Anterior  trochanters  of  male  with  minute 
hook;  mesotrochanter  right  angulate  to  acutely  angulate  (one  speci- 
men with  rounded  trochanter).  Terminal  abdominal  sternite  of 
female  incised  at  tip,  male  genital  capsule  as  shown  on  Plate  XIII. 

Location  of  Types.  Described  from  sixteen  specimens  from 
Raleigh,  N.  C.  Types:  Collections  U.  S.  National  Museum,  Ameri- 
can Museum  of  Natural  History,  C.  S.  Brimley  and  Bueno. 

Comparative  Notes.  Although  this  species  was  described  as  hav- 
ing "vertex  more  than  six  times  as  wide  as  synthlipsis,"  I  have  seen 
no  specimens  with  such  a  narrow  synthlipsis.  Many  specimens  from 
the  type  locality,  some  deteraiined  by  Bueno,  have  come  under  my 
observations.  In  my  "Biology  of  Aquatic  and  Semiaquatic  Hemip- 
tera"  I  separated  this  species  from  N.  variabilis  (now  N.  lunata)  as 
follows: 

"Vertex  3  times  S3'nthlipsis N.  variabilis. 

Vertex  6  times  synthlipsis N.  raleighi." 

I  took  these  statements  from  the  literature,  and  Doctor  Blatchley, 
in  his  "Heteroptera,"  ^  used  the  same  statements.  The  problem  of 
separating  these  species  is  by  no  means  so  simple.  After  studying 
much  material  involving  an  examination  of  all  specific  characters 
known  to  me  I  must  confess  my  failure  to  find  definite  distinctive 
characters.  All  gradations  appear  to  occur.  The  male  genitalia 
are  not  clearly  distinguishable.  Typical  specimens  of  N.  raleighi 
Bueno  are  smaller  and  synthlipsis  is  narrower  than  in  the  form  we 


Hungerfoed:    Genus  Notonecta  107 

know  as  N.  lunata  Hungerford.  The  latter  appear  to  be  the  common 
form  in  New  York  state  and  in  JNIichigan,  while  the  former  is  more 
southern  in  distribution. 

Notonecta  lunata  Hungerford,  1926 

(Color  Plate  I,  fig.  5 ;   Plate  XIII,  fig.  5) 

192G.  -V.  lunata  Hungerford,  Psyche,  xx.xiii,  p.  12.  (new  name  for  A'',  variabilis  maculata 
Fieb.). 

1926.  jV.  lunata  Hungerford,  Bull.   Brooklyn  Ento.   Soc,  xxi,  p.   195. 

1928.  N.  lunata  Hungerford,  Annals  Ento.   Soc.  Amer.,  xxi,  p.   143. 

1930.  N.  lunata  Hungerford,  Bull.  Brooklyn  Ento.   Soc.,  .xxv,  p.   140. 

Referring  to  this  species,  also: 

1851.  A\  variabilis  var.   maculata  Fieber,  Rhynchotographieen,   p.   477. 

1851.  .V.  variabilis  var.  maculata  Fieber,  Rhynchotographien,  p.   53   (Van  Duzee). 

1878.  .V.  undulata  Uhler.  Proc.  Boston  Soc.  Nat.  Hist.,  xix,  p.   442  (var.  2). 

1885.  N.  undulata  Uhler,  Stand.   Nat.   Hist.,   II,   p.   252    (in  part). 

1897.  iV.  variabilis  Kirkaldy,  Trans.  Ento.  Soc.  London  for  1897,  p.  414  (the  N.  A. 
forms). 

1899.  N.  am^icana  Ashmead,  Smiths  Ins.  N.  J.,  p.  144  (Bueno). 

1901.  N.  variabilis  CJiampion,  Biol.   Centr.   Amer.   Hem.-Het.,  II,  p.   370. 

1902.  N.  variabilis,  Bueno,  Jl.  N.  Y.  Ento.  Soc,  x,  pp.  231  and  234. 

1904.  A',  variabilis  Kirkaldy,  Wien.   Ento.  Zeit.,  xxiii,  p.   95  (in  part). 

1905.  A',  variabilis  Bueno,  Jl.  N.  Y.  Ento.  Soc,  xiii,  pp.  45  and  155. 

1908.  N.  variabilis  Bueno,  Jl.  N.  Y.  Ento.  Soc,  xvi,  p.  238. 

1909.  N.  variabilis  Kirkaldy  and  Bueno,  Proc.  Ento.  Soc.  Wash.,  x,  p.  199  (in  part). 

1910.  N.  variabilis  Smith,  Catalog  Ins.  N.  J.,  edn.  3,  p.  169. 
1910.  A'',  variabilis  Bueno  and  Brimley,  Ento.  News,  xviii,  p.   435. 
1912.  N.  variabilis  Bueno,  Can.  Ento.,  xliv,  p.   213. 

1914.    A'',  variabilis  Parshley,  Psyche,  xxi,  p.   140.     (Apr.  to  Sept.  in  Maine.) 
1914.    A',  variabilis  Barber,  Bull.  Am.   Mus.   Nat.  Hist.,  x.xxiii,  p.   499  (Florida). 
1917.    N.  variabilis  Hungerford,  Ento.  News,  xx\-iii,  p.  267. 
1917.    A',   variabilia,  Parshley,   Occ.   Papers  of  Boston  Soc.   Nat.   Hist.,  vii,   p.    112. 

1917.  A'',  variabilis  Van  Duzee,  Catalog  Hem.,  p.   451. 

1918.  A',  variabilis  Hungerford,  Ento.  News,  xxix,  pp.  242,  243,  245;  PI.  xv,  fig.  4 
(Ovipositor). 

1919.  N.  variabilis  Hungerford.  Kans.  Univ.  Sci.  Bull.,  xi,  pp.  22,  167,  169,  170,  180, 
181,  186,  187,  193;    PI.  viii  fig.   4;    PI.  xix,  figs.   1,  5,  8,   11,   13;    PI.  xxiii,  fig.   4. 

1919.  N.  variabilis  Hungerford,  Kans.  Univ.  Sci.  Bull.,  xi,  pp.  331,  332;  PI.  xxxi,  fig.  II 
(male  genital  capsule). 

1923.  N.  variabilis  Bueno,  Conn.  St.  Geol.  and  Nat.  Hist.  Survey  Bull.,  No.  34,  pp.  404, 
405;  PI.  xvi,  fig.  5.     (May  to  Oct.) 

1925    N.  variabilis,  Hungerford,  and  Beamer,  Ento.  News,  xxxvi,  pp.  264  and  297. 

1926.  .V.  variabilis  Leonard,  Cornell  Univ.  Agri.  Exp.  Sta.,  Memoir  101,  p.  139.  (Mar. 
to  Nov.) 

Size.  Length,  8.5  mm.,  to  10.7  mm.;  width  of  pronotum,  2.8  mm. 
to  3.5  mm. 

Color.  A  pale  species  from  nearly  white  to  yellowish  with  scu- 
tellum  nearly  always  yellowish.  Head,  pronotum  and  limbs  often 
greenish.  Hemelytra  only  occasionally  with  a  smoky  band  in  corium 
and  a  spot  near  tip  of  corium. 

Structural  Characteristics.  Difficult  to  distinguish  from  N.  raleighi 
Bueno.    Vertex  slightly  shorter  than  anterior  width;  anterior  margin 


108  The  University  Science  Bulletin 

of  vertex  slightly  longer  than  frontal  margin  of  the  eye;  anterior 
breadth  of  vertex  :  synthlipsis  ::  3:1;  synthlipsis  usually  about  a 
third  the  width  of  an  eye.  Pronotum  four-  to  six-tenths  longer  than 
the  head;  lateral  margins  nearly  straight,  divergent;  anterior  angles 
normal,  lateral  ledge  nearly  straight  and  oblique  as  seen  from  the 
side.  Membrane  lobes  about  equal,  anterior  lobe  may  be  trifle 
longer.  Anterior  trochanters  of  male  with  the  usual  small  hook, 
mesotrachanters  acutely  angulate,  terminal  abdominal  sternite  of 
female  incised  at  tip.  Male  genital  capsule  as  in  N.  raleighi  Bueno. 
(See  plate  XIII,  fig.  5.) 

Location  of  Type.  N.  lunata  was  a  name  proposed  to  replace 
the  name  N.  variabilis  var.  maculata  Fieber,  a  name  previously 
used.  I  have  not  been  able  to  find  any  specimens  from  Baltimore 
in  the  Berlin  Museum  that  might  have  been  studied  by  Fieber. 
Whether  Fieber  had  N.  raleighi  Bueno  or  N.  lunata  Hungerford  is 
impossible  to  determine.  By  general  consent  both  were  called  N. 
variabilis  until  Bueno  described  A'',  raleighi,  leaving  the  larger 
forms  to  be  called  N.  variabilis.  As  I  have  pointed  out,  N.  variabilis 
Fieb.  must  be  a  South  American  form. 

I  have  labeled  a  series  of  insects  from  Ithaca,  N.  Y..  as  typical 
of  N.  lunata  Hungerford.  They  are  in  the  Francis  Huntington  Snow 
collection  at  the  University  of  Kansas. 

Data  on  Distribution.  The  following  records  have  been  furnished 
by  G.  S.  Walley,  who  has  kindly  determined  the  specimens  in  the 
Ottawa  collection  and  given  me  the  data: 

Canada 

New  Burnswick:   Ludlow,  July  7,  1928;  W.  J.  Brown. 

Quebec:  Fairy  Lake,  Hull,  May  3-17,  1927,  G.  S.  Walley;  May  16.  1928, 
W.  J.  Brown;  Sept.  11,  1928,  G.  S.  Walley.  Knowlton,  Aug.  12.  1929,  G.  S. 
Walley.  Missisquoi  Bay,  July  11,  1927,  G.  S.  Walley.  Brome  Lake,  July  8, 
1927,  G.  S.  Walley. 

Ontario:  Black  Rapids,  Ottawa,  Apr.  27,  May  11.  1927,  G.  S.  Walley.  Jock 
River,  Ottawa,  Apr.  20,  May  21,  Aug.  25,  1927,  G.  S.  Walley.  Britannia,  Aug. 
21,  1927,  W.  J.  Brown.  Ventnor,  Aug.  5,  Sept.  1,  1928,  J.  A.  Adams.  Chesley 
Lake,  Aug.  23,  1928,  G.  S.  Walley.  Arran  Lake,  June  20,  1921,  G.  S.  Walley. 
Southampton,  Sept.  13,  1927,  G.  S.  Walley.  Dundas,  May  31,  1931,  G.  S. 
Walley.  Pt.  Pelee,  June  3,  6,  1929,  G.  S.  Walley  and  L.  J.  Milne.  Leamington, 
July  6,  1931,  G.  S.  Walley. 


Hungerford:    Genus  Notoxecta  109 

Notonecta  unifasciata  unifasciata  Guerin,  1857 

(Color  Plate  I,  Sg,  12;   Plate  XIV,  figs.  1,  2,  3.) 

IS.JT.  iV.  unifasciata  Guerin-Meneville,  Le  Moniteur  Universel,  26  Nov.,  1857,  Numero 
330,  p.  1298. 

1857.  A'',  unijascinta  Guerin-Meneville,  Revue  et  Magasin  de  Zoologie,  2d  ser.,  Tom.  ix, 
pp.  522-527. 

1858.  X.  vnifasciata  Guerin-Meneville,   L'lllustration,   Tom.    xxxii,   p.    47,  Juillet   17,1858. 
1858.    -V.   unifasciata  Guerin-Meneville,   Comptes  Rendiis,  Nov.   23,   1857,   p.   865.      (From 

Annals  &  Mag.  ref.  below.) 

1858.  N.  unifasciata  Guerin-Meneville,  Annals  and  Mag.  of  Nat.  Hist.  (3),  I,  p.  79. 
(Eng.  review.) 

1858.    N.  unifasciata  Guerin-Meneville,  Bull.  Soc.  Zool.  Acclim.  iv,  p.  581,  1857. 

1858.    iV.   unifariata   Guerin-Meneville,   Bull.  Soc.  Ento.  Fr.  (3)  v,  pp.  cxlviii  -  cli. 

1897.  N.  unifasciata  Guerin;  Kirkaldy,  Trans.  Ento.  Soc.  London  for  1897,  p.  426.  (Says 
unknown  to  him.) 

1898.  N.  unifasciata  Guerin;  Kjrkaldy,  Ento.  Mon.  Mag.  (2)  ix  (Old  ser.,  vol.  xxxiv), 
p.  173.     (Says:    "doubtless  N.  a-mericana  Fabr.) 

1899.  N.   unifasciata  Guerin;    Kirkaldy,  Ento.,  xxxii,  (Says  =  A'',  undulata  Say.) 
1917.    iV.  unifasciata  Guerin;    Hungerford,  Ento.   News,  xxviii,  p.   181. 

1929.  A'',  unifasciata  Guerin;  Hungerford,  Pan  Pacific  Ento.,  vi,  No.  2,  pp.  73-77.  (Re- 
port on  the  tj'pe.) 

1930.  N.  unifasciata  Guerin;   Hungerford,  Can.  Ento.,  Ixii,  p.  218. 

1930.    A'^.  unifasciata  Guerin;   Hungerford,  Bull.  Brooklyn  Ento.  Soc,  xxv,  p.  139. 
1933.     A',   unifasciata   Guerin;    Ancona,  Anales  del   Inst,   de  Biol.   IV,  p.   68.      (Mexico.) 

Referring  to  this  species,  also: 

1897.  N.  americana  Kirkaldy,  Trans.  Ento.  Soc.  London  for  1897,  p.  409.  (In  part: 
Spec.  Det.  by  him  in  Vienna  and  Paris.) 

1899.  A^.  americana  Kirkaldy,  Revue  d'Entomologie,  xviii,  p.   94  (Desc.  of  egg). 

1900.  A'^.  indica  Kirkaldy,  Ento.,  xxxiii,  p.   10. 

1901.  A'',   undulata  Champion,  Biol.  Centr.  Amer.,  Hem.-Het.,  ii,  p.  370  (in  part). 

1904.  A'^.  indica  Kirkaldy,  Wien,  Ento.  Zeit.,  xxiii,  p.  95. 

1905.  N.  indica  Bueno,  Jl.  N.  Y.  Ento.  Soc,  xiii,  pp.  149,  151 ;  pi.  vii,  fig.  1. 

1906.  A'',  indica  Snow,  Trans.  Kans.  Acad  Sci.,  xx,  pt.  1,  p.  181. 
1909.  A'',  indica  Kirkaldy  &  Bueno,  Proc.  Ento.  Soc.  Wash.,  x,  p.  198. 

1916.  N.  indica  Browne,  .71.  Morph.,  xxvii,  pp.  119-162.     (Cytology.) 

1917.  A'',   indica  Van  Duzee,  Catalogue  Hemipt.,  p.  452. 

1917.  A'',  indica  Hungerford,  Ento.  News,  xxviii,  p.  175. 

1918.  A'^.   indica  Hungerford,  Ento.  News,  xxix,  pp.  243,  245  ;   pi.  xv,  fig.  11. 

1919.  N.  indica  Hungerford,  Univ.  Kans.  Sci.  Bull.,  xi,  pp.  167,  168,  181,  332;  pi.  xxiii, 
fig.  11.     (Ovipositor)  pi.  xxxi,  fig.  6. 

1928.    N.  indica  Hungerford,  Ento.  News,  xxxix,  p.  156. 

Size.  Length.  9.3  mm.  to  10.2  mm.;  width  of  pronotum,  3.3  mm. 
to  3.6  mm. 

Color.  Typically  black  and  white.  Scutellum  black  with  lateral 
margins  fiavous.  Hemelytra  white  with  black  markings;  clavus 
often  black  along  inner  margin  and  apex;  distal  end  of  corium  and 
basal  two-thirds  of  membrane  crossed  by  solid  black  band. 

Structural  Characteristics.  Anterior  outline  of  head,  as  seen  from 
above,  somewhat  flattened;  vertex  shorter  than  its  anterior  width; 
anterior  margin  of  vertex  less  convex  and  longer  than  frontal  margin 
of  an  eye;  anterior  breadth  of  vertex  :  synthlipsis  : :  5:3.  Pronotum 
slightly  more  than  twice  as  long  as  the  head;  lateral  margins  nearly 
straight  and  moderately  divergent;  anterior  angles  somewhat  acute 


110  The  University  Science  Bulletin 

and  embracing  the  eyes;  lateral  ledge  oblique.  Anterior  lobe  of 
membrane  longer  than  the  posterior.  Anterior  trochanter  of  male 
with  medium-sized  hook.  Mesotrochanters  angulate.  Terminal 
abdominal  sternite  of  female  relatively  broad  and  not  plainly 
notched  at  tip.  Female  gonapophyses  short.  Male  genital  capsule 
as  shown  on  Plate  XIV. 

Location  of  Types.  Museum  of  Paris.  Kirkaldy  in  his  ''Revision" 
listed  the  species  as  an  "Unrecognized  species,"  but  I  found  the  types 
in  the  Paris  Museum  in  a  box  prepared  by  Guerin  labeled  "Hautle." 
For  a  full  report  of  this  discovery  see  my  paper  in  the  Pan-Pacific 
Entomologist,  volume  VI,  p.  73,  1929. 

Comparative  Notes.  In  general  facies  this  is  a  characteristic 
species.  The  head  is  quite  short,  the  black  scutellum  is  margined 
with  fiavous  and  the  mesotrochanter  is  angulate.  There  is  some 
variation  in  the  shape  of  the  head,  in  the  angle  of  the  mesotrochanter 
and  in  the  male  claspers.  The  material  from  the  type  locality  (near 
the  city  of  Mexico),  from  Guadalupe,  D.  F.  Mex.,  Socorro,  N.  M., 
and  some  specimens  from  Southern  California,  while  showing  some 
variation  in  the  male  claspers,  appear  to  be  N.  unifasciata  Guerin. 
The  variations  mentioned  above  make  it  necessary  to  designate  sub- 
species as  follows: 

Notonecta  unifasciata  cochisiana  new  subsp. 

This  subspecies  conforms  to  the  general  description  of  the  species, 
but  has  a  very  characteristic  male  clasper  that  separates  it  from  the 
typical  N.  unifasciata  Guerin,  see  Plate  XIV,  fig.  5.  Sixty  specimens 
from  Cochise  Co.,  Arizona,  represent  the  type  series  of  this  sub- 
species. 

Notonecta  unifasciata  andersoni  new  subsp. 

In  this  subspecies  the  anterior  margin  of  vertex  is  more  rounded 
and  the  clasper  longer  than  in  typical  A^.  unifasciata  Guerin.  See 
Plate  XIV,  fig.  6.  This  is  a  smaller  species  than  N.  spinosa  Hunger- 
ford,  and  although  the  shape  of  the  head  is  somewhat  like  it  the 
mesotrochanter  does  not  have  the  long  spine-like  projection  of  the 
mesotrochanter  so  characteristic  of  N.  spinosa.  The  type  series  was 
taken  at  Oliver,  B.  C.  Material  collected  in  Navajo,  Ariz.;  Carson 
City,  Nevada;  Lehi  and  Am.  Forks,  Utah;  and  Hot  Lake,  Oregon, 
shows  a  gradation  away  from  the  typical  N.  unifasciata  Guerin. 


Hungerford:    Genus  Notonecta  111 

Notonecta  unifasdata  angulata  new  subsp. 

This  subspecies  conforms  in  general  shape  and  appearance  with 
the  typical  species,  but  the  clasper  of  the  male  is  stouter  than  any 
of  the  others.  (See  PI.  XIV,  fig.  4.)  The  vertex  of  the  female  is 
more  rounded.  The  mesotrochanter  is  angulate  but  not  as  produced. 
This  subspecies  is  described  from  one  male  and  one  female  in  the 
Uhler  Collection.  The  specimens  bear  the  label  "Kirkaldy,  S. 
Amer." 

Biological  Notes.  This  species  was  described  from  a  measure  of 
dried  insects  offered  for  sale  in  Mexico  as  food.  It  formed  part  of 
a  mixture  of  water  bugs  consisting  largely  of  Corixids  of  two  or 
three  species,  one  of  which  Guerin  described  as  new  at  the  time  he 
described  this  Notonectid.  Collections  of  eggs  of  Corixids  gathered 
and  offered  for  food  nearly  always  contain  some  eggs  of  this  back- 
swimmer.  C.  T.  Brues  took  this  species  and  its  nymphs  from  water 
having  a  temperature  of  97.3°  F.  He  took  it  from  a  number  of  hot 
springs  with  temperatures  of  85°  F.  and  above. 

Data  on  Distribution.  This  species  (broad  sense)  is  widely  dis- 
tributed from  Mexico,  across  Western  United  States  to  Western 
Canada.  Two  specimens  labeled  "S.  Amer."  in  the  Uhler  collection 
appear  to  be  a  subspecies  of  N.  unifasdata  Guerin. 

Mexico 
D.  F.  Texcoco  Lake,  Oct.  21,  1923,  Jan.  31,  1926,  Alf.  Dampf ;  Ahuatle  from 
Lake  Texcoco,  L.  Ancona  H.,  9-1-1932;  Guadalupe,  D.  F.,  Aug.  31,  1903,  W.L. 
Tower;  D.  F.  Lago  de  Texcoco,  May  14,  1930,  Creaser-Gordon.     (Mich.  U.) 

United  States 

Texa.<:  Presidio  Co.,  Julj^  16,  1927,  P.  A.  Readio;  Valentine,  July  13,  1927, 
R.  H.  Beamer. 

New  Mexico:  Torrance  Co.,  Sept.  10,  1925,  C.  H.  Martin;  Estancia,  Sept. 
6,  1925,  C.  H.  Martin;  Socorro  Co.,  Aug.  18,  1927,  R.  H.  Beamer. 

Arizona:  Douglas,  Aug.,  F.  H.  Snow;  Navajo  Co.,  Aug.  15,  1927,  R.  H. 
Beamer;  Apache  Co.,  Aug.  16,  1927,  R.  H.  Beamer;   Maricopa  Co.,  Aug.  7, 

1929,  L.  D.  Anderson;  Cochise  Co.,  July  20,  1927,  R.  H.  Beamer. 
California:    L.  Elsinore,  Aug.  2,  1911;  Los  Angeles  Co.,  Coquillett  (U.S. 

N.  M.) ;  San  Diego  Co.,  Poway  Val,  Apr.  9,  1930,  C.  &  D.  Martin;  Niles  Can- 
yon, Alameda  Co.;  San  Antonio  Canyon,  Ontario,  July  25,  1907;  Kelly  Springs, 
Camby,  C.  T.  Brues;   Cedarville,  1930,  C.  T.  Brues;   Bassetts  Hot  Springs, 

1930,  C.  T.  Brues. 

'Nevada:  Carson  City,  Aug.  9,  1929,  Oman,  Anderson  and  Beamer;  Soda 
Lake.  July  14,  1911;  Fallon,  Aug.  9,  1929,  L.  D.  Anderson;  37  mi.  s.  of  Battle 
Mts.,  Tem.30°C.,  C.  T.  Brues;  Goldfield,  Tem.  35.4°C.,  C.  T.  Brues;  29  mi.  s. 
of  Winnemucca,  Tem.  36.3°C.,  C.  T.  Brues  (nymphs  also) ;  Sunnyside,  1930, 
C.  T.  Bnies;  Austin,  Aug.  9,  1929,  P.  W.  Oman. 


112  The  University  Science  Bulletin 

Utah:  Emery  Co.,  Aug.  26,  1921,  Grace  Wiley;  Garfield,  July  9,  1911,  Loco- 
motive Spgs.,  April  10,  1930,  Grantsville,  April  2,  1930,  Pleasant  Grove,  April 
4,  1930,  Lehi,  April  4,  1930,  American  Fork,  April  8,  1930,  all  by  G.  F.  Knowl- 
ton;  Farwest,  Sept.  12,  1929,  C.  J.  D.  Brown;  Tremonton,  July  5,  1931,  L.  D. 
Anderson. 

Oregon:  Hot  Lake,  Temp.  80°  F.,  July  13,  1931,  L.  D.  Anderson;  Board- 
man,  July  15,  1931,  L.  D.  Anderson;  Hood  River,  July  17,  1931,  L.  D.  Ander- 
son. 

Canada 

Oliver,  B.  C,  Aug.  6,  1931,  L.  D.  Anderson;  Peachland,  Oct.  7.  1931,  A.  N. 
Gartrell;  Brent's  Lake,  Summerland,  Oct.  29,  1931,  A.  N.  Gartrell ;  Vernon, 
June  21,  1921,  R.  Hopping;  Penticton,  Oct.  14,  1931,  A.  N.  Gartrell;  Westbank, 
April  21,  1932,  A.  N.  Gartrell.     (All  but  first  in  Ottawa  Museum.) 

Notonecta  spinosa  Hiingerford,  1930 

(Color  Plate  11,  fig.   7;    Plate  XIV,   fig.   7) 
1930.    A'^.  spinosa  Hungerford,  Can.  Ento.,  Ixii,  pp.  217,  218. 

Size.  Length,  about  12  mm.,  the  females  may  be  a  little  longer 
and  the  males  a  little  shorter  than  this.  Width  of  pronotum,  3.3 
mm.  to  3.6  mm. 

Color.  General  color  of  N.  undulata  Say  except  that  the  black 
scutellum  is  broadly  margined  with  orange-yellow.  It  is  therefore 
a  black  and  white  species  with  venter  dark  to  black;  the  head,  limbs 
and  venter  often  suffused  with  green;  dorsal  view  of  head  and  an- 
terior part  of  pronotum  pale  horn  in  color,  posterior  part  of  pro- 
notum dark,  due  to  black  mesonotum  beneath.  Scutellum  black, 
broadly  margined  with  orange-yellow,  the  ends  of  this  V-shaped 
figure  slightly  broader,  not  attaining  the  base  of  the  scutellum. 
Hemelytra  white  with  sooty  black  markings  as  follows:  Small,  dark, 
irregular  blotch  at  base  that  may  extend  as  narrow  line  to  near  the 
middle,  one  as  a  median  dark  line  on  the  membrane;  a  zigzag  M- 
shaped  figure,  which  is  thin,  transversing  hemelytra  just  in  front  of 
the  membrane. 

Structural  Characteristics.  Anterior  outline  of  head,  as  seen  from 
above,  roundly  curved;  eyes  not  prominent;  vertex  with  anterior 
width  slightly  greater  than  its  length;  anterior  margin  of  vertex 
broadly  rounded  and  wider  than,  the  frontal  margin  of  an  eye; 
anterior  breadth  of  vertex  :  synthlipsis  ::  2+ :1.  Length  of  pro- 
notum :  length  of  head  : :  3+:2;  lateral  margins  straight  and  diver- 
gent; anterior  angles  normal;  lateral  ledge  oblique  and  shorter  than 
the  rear  margin  of  the  eye  beneath  it.  Lobes  of  membrane  about 
equal.  Anterior  trochanter  of  male  with  a  hook.  Mesotrochanter 
with  angle  produced  into  a  long,  sharp  spine.    Terminal  abdominal 


Huxgerford:    Genus  Notonecta  113 

sternite  of  female  with  broad,  but  very  shallow,  notch  at  tip.  Fe- 
male gonapophyses  (first  pair)  short.  Male  genital  capsule  as 
shown  on  Plate  XIV. 

Location  of  Types.  Holotype,  allotype  and  some  paratypes  in 
Francis  Huntington  Snow  Entomological  Museum  in  the  University 
of  Kansas.  Other  paratypes  in  U.  S.  National  Museum;  Canadian 
National  Collection,  Ottawa,  Canada;  Mus.  of  California  Academy 
of  Science  and  Knowlton's  collection.  Others  have  since  been  sent 
to  Mr.  Torre-Bueno,  Mr.  Esaki,  the  Paris  Museum  and  the  British 
Museum. 

Comparative  Notes.  Size  and  color  as  in  A'',  undulata  Say  except 
that  the  scutellum  is  always  margined  with  orange-yellow.  The 
more  produced  front  margin  of  the  vertex,  the  spinose  mesotro- 
chanter  and  the  genitalia  of  both  sexes  separate  it  from  Say's  species. 
It  is  larger  than  N.  unifasciata  Guer.,  has  a  narrower  synthlipsis 
and  a  long  spinose  mesotrochanter  instead  of  angulate  or  slightly 
produced  process  as  in  Guerin's  species. 

Data  on  Distribution : 

Canada 

British  Columbia:  Vernon,  Sept.  26,  1919,  W.  Downes  (Ottawa,  Canada) ; 
Oliver,  Aug.  6,  1931,  L.  D.  Anderson. 

United  States  of  America 

Utah:  Lehi,  April  4,  5,  8  and  May  3-27  1930,  G.  F.  Knowlton;  Logan, 
Oct.  5,  1930,  M.  J.  Jones;  Hyde  Park,  April  19,  1930,  G.  F.  Knowlton;  Brigham, 
April  26,  1930,  G.  F.  Knowlton;  American  Fork,  May  3,  1930,  G.  F.  Knowlton. 

Nevada:   Carson  City,  Aug.  9,  1929,  R.  H.  Beamer. 

Yellowstone  National  Park:    Fry  Pan  L.,  Aug.  15,  1931,  L.  D.  Anderson. 

Montana:  Whitehall,  Aug.  13,  1931,  L.  D.  Anderson;  Three  Forks,  July  22, 
1931,  L.  D.  Anderson. 

Oregon:    Hot  Lake,  July  13,  1931,  L.  D.  Anderson. 

Notonecta  indica  Linne,  1771 

(Color  Plate  I,  fig.  2;   Plate  XIII,  fig.  3) 

1771.    A^.  indica  Linnsus,  Mantissa  Plantarum,  ii,  p.  534. 

1806.    N.  indica  Linnaeus;    Turton  in  Linn.   Syst.   Nat.,   edn.    13,   Eng.   Transl.,  ii,  p.   605. 

1930.    iV.  indica  Linnaeus;   Hungerford,  Bull.  Brook.  Ento.   Soc,  .\xv,  p.   139. 

Referring  to  this  species,  also: 

1775.    -V.  americana  Fabricius,  Syst.  Ento.,  p.  690. 

1811.  A',  americana  Olivier,  Encyclopedic  Methodique  Histoire  Naturelle  Insectes,  viii, 
pp.  387  and  389,  "Saint  Domingue." 

1832.  A',  undulata  Say,  N.  Harm.,  p.  39;  Fitch  reprint,  p.  812;  Compl.  Writ.  I,  p.  368 
(in  part). 

1851.  A',  variabilis  var.  acutellaris  Fieber,  Rhynchotographieen,  p.  477.  (Those  from 
Portorico.) 


114  The  University  Science  Bulletin 

1856.    N.  variabilis  Gueriu,  in  Ramon  de  la  Sagra's  Cuba,  Hist.  Nat.,  vii,  p.   176. 

1881.  N.  pallipes  Lethierry,  Ann.  Soc.  Ento.  Belg.,  xxv,  p.  13  (three  specimens  from 
St.  Barthelemy). 

1884.    N.  undulata  Uhler,  Standard  Nat.  Hist.  II,  p.  252   (in  part.) 

1891.  N.  undulata  Summers,  Bull.  Agri.  Exp.  Sta.,  Univ.  Tenn.,  iv.  No.  3,  p.  82  (in 
part). 

1894.  N.  americana  Uhler,  Proc.  Zool.  Soc.  London  for  1894,  p.  223  (a  single  nymph 
taken  Aug.  20). 

1897.    N.  undulata  Kirkaldy,  Trans.  Ento.  Soc.  London,  p.  410  (in  part). 

1897.    N.  undulata  van  charon  Kirkaldy,  Trans.  Ento.  Soc.  London,  p.   411   (in  part). 

1899.    A^.  undulata  Kirkaldy,  The  Ento.,  xxxii,  p.  29  (Flight  to  lights  in  Jamaica). 

1901.  A^.  undulata  Champion,  Biol.  Centr.  Amer.  Hem.  Het.,  ii,  p.  370,  Tab.  xxii, 
fig.  10. 

1905.    N.  howardii  Bueno,  Jl.  N.  Y.  Ento.  Soc.,  xiii,  p.  151. 

1905.    A',  undulata  Bueno,  Jl.  N.  Y.  Ento.  Soc,  xiii,  p.  153  (in  part). 

1909.    N.  undulata  Delcourt,  Bull.   Sci.   Fr.   et  Belg,  Tom  xliii   (7),  vol.    1,   pi.   iv,   line   8. 

1909.    N.  undulata  Kirkaldy  &  Bueno,  Proc.  Ento.  Soc.  Wash.,  x,  p.   199   (in  part). 

1909.    N.  howardii  Bueno;  Kirkaldy  &  Bueno,  Proc.  Ento.  Soc.  Wash.,  x,  p.  198  (in  part). 

1914.    N.  undulata  Barber,  Bull.  Am.   Mus.  Nat.   Hist.,  xxxiii,  p.   499. 

1917.    A^  undulata  Van  Duzee,  Catalogue  Hemiptera,  p.   451  (in  part). 

1917.    N.  howardii  Bueno;   Van  Duzee,  Catalogue  Hemiptera,  p.  452. 

1919.    N.  howardii  Bueno;    Hungerford,  Kans.  Univ.  Sci.  Bull.,  xi,  p.   168. 

1919.    A^.  undulata  Hungerford,  Kans.  Univ.  Sci.  Bull.,  xi,  p.  169  (in  part). 

1922.  A",  howardii  Bueno;  Hungerford,  Kans.  Univ.  Sci.  Bull.,  xiv,  p.  426.  (Says  fig.  8, 
pi.  xxxi  of  Sci.  Bull,  xi  is  this  species). 

1922.    N.  howardii  Bueno;   Drake,  Ohio  Jl.  Sci.,  xxii,  p.   116. 

1926.    A'',  howardii  Bueno;    Blatchley,  Heteroptera     ...     p.   1053. 

1926.    AT.  undulata  Blatchley;    Blatchley,  Heteroptera     ...     p.   1054   (in  part). 

1928.    A^.  howardii  Bueno;    Hungerford,  Ento.  News,  xxxix,  p.  156  (Georgia). 

Size.  Length,  10  mm.  to  11  mm.;  width  of  pronotum,  3.36  mm.  to 
3.75  mm. 

Color.  Color  variable,  from  luteous  to  nearly  black,  the  hemely- 
tra  having  only  the  base  of  the  clavus  tan.  The  typical  color  is 
black  and  white;  the  scutellum  and  a  broad  band  covering  the 
distal  end  of  corium  and  the  basal  two-thirds  of  the  membrane 
black;  the  scutellum  often  pale  at  tip  and  marked  near  each  basal 
angle  by  a  light  spot. 

Structural  Characteristics.  Head  more  than  one-half  length  of 
pronotum ;  vertex  as  long  or  slightly  longer  than  its  anterior  width 
as  seen  from  above;  anterior  margin  of  vertex  less  convex  and 
shorter  than  the  frontal  margin  of  the  eye;  anterior  breadth  of 
vertex  :  synthlipsis  : :  5:2;  synthlipsis  about  two-fifths  the  width  of 
the  eye.  Pronotum  not  twice  length  of  head,  lateral  margins  but 
slightly  divergent  on  anterior  two-thirds,  then  divergent,  making  the 
lateral  margin  appear  concave  and  yet  giving  the  appearance  of  a 
somewhat  parallel  sided  pronotum;  anterior  angles  embracing  the 
eye ;  lateral  ledge  but  slightly  oblique  and  straight.  Lobes  of  mem- 
brane equal.  Anterior  trochanter  of  male  with  a  stout  conspicuous 
hook.  Mesotrochanter  rounded.  Terminal  abdominal  sternite  of 
female  with  a  shallow  notch  or  none  at  tip.  Female  gonapophyses 
short.    Male  genital  capsule  as  shown  on  Plate  XIII. 


Hungerford:    Genus  Notonecta  115 

Location  of  Type.  I  did  not  find  any  specimens  in  Upsala, 
Sweden,  but  in  the  Collection  of  the  Linnean  Society  at  London  there 
is  a  male  specimen.  It  lacks  one  middle  leg  entirely  and  the  tibia 
and  tarsus  of  the  other.  It  still  has  both  front  legs  and  one  hind 
leg  (1928).  The  specimen  has  a  black  scutellum  with  flavous  marks, 
which  are  irregular,  as  in  many  specimens  of  what  we  have  hereto- 
fore known  in  America  as  N.  howardii  Bueno.  The  mesotrochanter 
is  rounded  as  in  Bueno's  species,  not  angulate  as  in  N.  unifasciata 
Guer,  which  we  called  N.  indica  for  many  years.  The  label  "indica" 
appears  not  to  be  in  Linnaeus  handwriting.  It  is,  however,  altogether 
probable  that  this  specimen  was  in  the  Linnaean  collection  purchased 
by  Sir  James  E.  Smith,  who  was  for  forty  years  president  of  the 
Linnean  Society  of  London.  I  therefore  designate  it  the  type  of 
N.  indica  Linn. 

In  the  Museum  at  Copenhagen,  under  the  name  Notonecta  ameri- 
cana  Fabr.,  I  found  several  specimens.  The  first  one  bears  the 
labels:  "Musemn  Seh.  and  T.  Lund,"  'W.  americana"  and  is  N. 
indica  Linn.  It  is  a  female  specimen  and  the  scutellum  has  the 
pale  spot  at  each  basal  corner  as  described  by  Fabricius  (Fabricius, 
however,  said  his  specimen  was  from  "Mus.  Tottianum").  The 
second  specimen  bears  one  label  I  cannot  read  and  another  "Mus. 
Seh.  and  T.  Lund."  It  is  N.  undulata  Say.  The  third  and  fourth 
specimens  are  females  labeled  "Mus.  Westerm"  'W.  americana"  and 
are  N.  indica  Linn.  There  is  no  doubt  that  the  first  two  specimens 
at  least  were  seen  by  Fabricius  and  the  first  one  fits  his  description 
exactly. 

Comparative  Notes.  Due  to  its  size  and  color  variations  it  has 
been  included  under  N.  undulata  Say  in  most  collections.  The  broad 
black  band  across  the  hemelytra  of  typical  specimens  separate  it 
from  typical  specimens  of  N.  undulata  Say.  The  head  is  more 
truncate  as  seen  from  above.  The  shape  of  the  pronotum  appears 
more  parallel  sided  than  in  Say's  species.  The  last  abdominal  ster- 
nite  of  the  female  is  shallowly  notched,  if  notched,  and  the  anterior 
trochanters  of  the  male  are  armed  with  larger  hooks  than  in  .V. 
undulata  Say. 

Data  on  Distribution.  This  is  one  of  the  few  species  common  to 
both  North  and  South  America.  Its  range  extends  across  the 
southern  United  States  south  of  37°  latitude.  I  have  seen  but  one 
collection  north  of  this  line.  This  is  from  Maryland,  which  indicates 
that  east  of  the  Allegheny  Mountains  it  extends  northward  to  in- 
clude Maryland.  Southward  I  have  it  from  Mexico,  Yucatan,  Guate- 


116  The  University  Science  Bulletin 

mala,  Colombia,  South  America  (Bogota),  Brazil*  ?,  and  from  the 
Islands  of  Cuba,  Jamaica,  St.  Croix,  St.  Thomas,  and  Porto  Rico. 
The  following  collections  are  before  me: 

United  States  of  America 

Maryl.'^nd:  Crisfield,  Sept.  26,  U.  S.  N.  M.  (Uhler  called  it  iV.  punctata 
Fieb.). 

North  Carolina:    Southern  Pines,  Sept.  14,  1916. 

South  Carouna:    Clemson  CoUege,  April  13,  1929,  Oct.  30,  1927,  Nov.  25, 

1929,  D.  Dunavan;  Walhalla,  Mar.  24,  1929,  D.  Dunavan. 

Kentucky:  Pond  near  Brooklyn  Bridge,  Jan.  3,  1933,  H.  Garmen,  (taken 
along  with  A'^.  undulata  Say.) 

Tennessee:  Murfreesboro,  Aug.  29,  1929,  Creaser  &  Becker  (Michigan); 
Kno.xville,  July  13,  1890;  Neubert  Springs,  Dec.  1,  1932,  L.  Chester  Marsten 
Jr.  (34  adults  and  3  nymphs). 

Georgl\:  Baker  Co.,  Aug.  24,  1927,  C.  H.  Martin;  Ga.  (Uhler  Coll.  U.  S. 
N.  M.). 

Florida:  Inverness,  Aug.  1,  1930,  J.  O.  Nottingham;  Cocoanut  Co.,  Aug. 
9,  1930,  R.  H.  Beamer;  Archer,  July  31,  1930,  Paul  Oman;  Sanford,  Aug.  4, 

1930,  Paul  Oman;  Wildwood,  Aug.  2,  1930,  J.  O.  Nottingham;  Plant  City, 
Jan.  4,  1927,  C.  O.  Bare;  Dunedin,  W.  S.  Blatchley,  Feb.  9,  1919  (Purdue 
Univ.). 

Mississippi:  Fayette,  July  23,  1921;  Miss.  Agri.  Coll.,  Aug.  20,  1913,  J.  G. 
Hester;  Natchez,  Sept.  8,  1924,  H.  M.  Harris. 

Louisiana:  Baton  Rouge,  Mar.  9,  1929,  R.  M.  DeCoursey;  Mound,  Mar. 
9,  1929;  Opelousas,  C.  R.  Pilate  (Baker  Coll.  U.  S.  N.  M.) ;  New  Orleans,  July, 
1916,  H.  E.  Hubert  (U.  S.  N.  M.). 

Arkansas:  Arkansas  Co.,  Apr.  9,  1930;  Fayette ville,  Feb.  27,  1930;  Law- 
rence Co.,  B.  C.  Marshall;  Imboden,  Mar.  19,  1925,  B.  C.  Marshall. 

Tex.\s:  Hidalgo  Co.,  July  30,  1928,  R.  H.  Beamer;  Sutton  Co.,  July  16 
and  Aug.  20,  1928,  R.  H.  Beamer;  Victoria  Co.,  Aug.  9,  1928,  R.  H.  Beamer; 
Victoria,  Dec.  27,  1910,  J.  D.  MitcheU  (U.  S.  N.  M.) ;  Colorado  Co.,  Apr.  25, 
1922,  Grace  Wiley;  Cameron  Co.,  Aug.  13,  1928,  A.  M.  James;  Brooks  Co.,- 
July  25,  1928,  R.  H.  Beamer;  Jim  Wells  Co.,  July  24,  1928,  A.  M.  James; 
Valentine,  July  13,  1927,  R.  H.  Beamer;  Kerrville,  Apr.  11,  1907,  F.  C.  Pratt; 
Calvert,  July  22,  1907,  C.  R.  Jones  (U.  S.  N.  M.);  Alfred,  July  24,  1932, 
R.  H.  Beamer. 

New  Me.xico:  Socorro  Co.,  Aug.  18,  1927,  L.  D.  Anderson;  Eddy  Co.,  July 
9,  1927,  P.  A.  Readio;  San  Antonio,  July  15,  1927,  L.  D.  Anderson;  Mesilla 
Park,  July  18,  1927,  P.  A.  Readio. 

Arizona:  Douglas,  F.  H.  Snow;  Gila  Co.,  Aug.  5,  1927,  R.  H.  Beamer; 
Cochise  Co.,  July  29,  1927,  L.  D.  Anderson;  S.  W.  edge  Tuscon,  July  20, 
1932,  R.  H.  Beamer. 

Caufornia:    Holtville,  July  2,  1929,  R.  H.  Beamer;  Calipatria,  July  14,  1923. 


See  p.  141  where  I  have  called  these  Fieber's  .V.  variabilis. 


Huxgj:rford:    Genus  Xotonecta  117 

Mexico 
Tamaulipas,  San  Carlos  Mts.,  July,  1930,  Dice  &  Bartlett  (Michigan) ;  Los 
Mochis,  Senaloa,  June  13,  1922,  C.  T.  Dodds;  Cuernavaca,  Nov.  4,  1922,  E.  G. 
Smyth  (U.  S.  N.  M.) ;  Guadalajara,  Etat  de  Jalisco,  M.  Diguet,  1901  (labeled 
N.  undulata  var.  charon  Am.  M.  N.  H.) ;  Yucatan,  Tabi,  F.  D.  G.  (labeled 
"B.  C.  A.  Rhyn.  II.    .V.  undulata  Say"  U.  S.  N.  M.). 

GU.4TEM.\LA 

S.  Geronimo,  Champion  B.  C.  A.  Rhyn  II  (labclcMl  "N .  xinduluta  by  Cham- 
pion) ;  Guatemala  City,  Ainslei  (C.  J.  Drake). 

COLOMBU 

Bogota,  Coll.  Signoret  (Vienna);  Colombie,  Steinhcil  (det.  by  Kirkaldy  as 
A',  wuhdata  Paris  Mus.). 

West  Indies 

Cuba  :  Havana,  Jan.  25.  1932,  P.  J.  Bermudez  ;  Havana,  Baker  (U.  S.  N.  M)  ; 
Soledad.  Apr.  4,  1925,  J.  G.  Myers;  Littletowm,  Jan.  1,  1878. 

Jam.uca:  Claremont  Baron  Hill,  Trelawny,  Mar.  4,  1928,  L.  G.  Perkins; 
Montego  Bay,  Mar.  15,  1911  (Am.  Mus.  Nat.  Hist.);  Port  Antonio,  A.  E. 
Wight  (Mus.  Comp.  Zool.,  Cambridge,  Mass.) ;  Yardley  Chase,  Santa  Cruz 
Mts.,  St.  Elizabeth,  taken  at  light  Oct.  31,  1899. 

PoRTO  Rico:    Desengano,  June,  1924  (Cornell  Univ.  lot  719). 

St.  Thom.vs:    Charlotte  Amalie;  V.  Ids.,  June  2,  1917,  Harold  Morrison. 

St.  Croix:   "F.  5022,"  Mar.  4,  1925  (Am.  Mus.  Nat.  Hist.). 

Notonecta  undulata  Say,  1832 

(Color  Plate  I,  fig.   3;    Plates  IX,   fig.   3 ;    X,  fig.   3 ;    XIII,  fig.    1) 

1832.  .V.  undulata  Say,  Heter.  New  Harm.,  p.  39;  Fitch's  reprint,  p.  812:  Complete 
Writ.  I,  p.   3CS. 

1851.  -V.  undulata  Say;  Fieber,  Rhynchotographieen  p.  479  (not  known  to  him).  Rhyn- 
chotographien,  p.  55  (Van  Duzee). 

1853.    -V.  undulata  Say;   Herrick-Schaffer,  Wanzenartigen  Insecten,  i.x,  p.   137. 

1874.  A',  undulata  Say;  Packard,  Half  hours  w.  Insects,  pt.  C,  pp.  139-141,  fig.  103 
(Egg  desc.   p.    158). 

1876.  A\  undulata  Say;  Uhler,  Bull.  U.  S.  Geol.  Geog.  Surv.,  vol.  I,  Bull.  V,  2d  ser., 
p.  339,  pi.  21,  fig.  33. 

1876.  N.  undulata  Say;  Uhler,  Bull.  U.  S.  Geol.  Geog.  Surv.  Reprint  from  above,  p.  73. 
(Inhabits  foulest  pools.) 

1877.  A",   undulata  Say;    Uhler,  Bull.  U.   S.  Geol.   Geog.   Surv.,  Ill,  p.   453. 

1877.  A',  undulata  Say;   Uhler,  Wheeler's  Rept.  Chief  Eng.  for  1877,  p.  1332.     (N.  Mex.) 

1878.  A',  undulata  Say;   Uhler,  Bull.  U.  S.  Geol.  Geog.  Surv.,  iv,  p.  509. 

1878.  A',   undulata  Say;   Uhler,  Proe.  Bost.  Soc.  Nat.  Hist.,  xix,  p.  442.     (Except  Var.  -.) 

1878.  A',  undulata  Say;   Packard's  Guide,  6th  Ed.,  p.   537,  fig.   542. 

1884.  A'',  undulata  Say;   Uhler,  Stand.  Nat.  Hist.,  II,  p.  252  (in  part.) 

1884.  A'',   undulata  Say;   Popenoe,  Trans.  Kans.  Acad.  Sci.,  ix,  p.  62. 

1888.  A',  undulata  Say;    Comstock,  Introduction,   p.    186,  fig.    157. 

1888.  A',  undulata  Say;    Provancher,  Pet.  Fauna  Ent.  Can.,  iii,  p.   201. 

1889.  A^.  undulata  Say;  Weed,  Bull.  Ohio.  Agri.  Sta.  Tech.,  .'^er.  I,  p.  12;  pi.  ii,  fig.  3 
(Feeding  and  defense). 

1880.    A^  undulata  Say;   Carman,  Bull.  111.  Lab.  X.  H.,  .Art.  ix,  vol.  iii,  p.  174. 

1890.  A',  undulata,  Say;   Hyatt  and  Arms,  Insecta,  p.   121,  fig.   70. 

1891.  y.  undulata  Say;    Summers,  Bull.  Agr.  Exp.  U.  of  Tenn.,  iv,  No.   3,  p.   82. 
1894.    A',  undidata  Say;    Uhler,  Proc.  Calif.  Acad.  Sci.   (2)  iv,  p.  292. 

1894.    A',  undulata  Say;   Van  Duzee,  Bull.  Buffalo  Acad.  Nat.  Sci.,  v.  p.  186. 

9—3482  i 


118 


The  University  Science  Bulletin 


N.  undulata  Say;   Packard,  Ento.  for  Beginners,  p.  83. 
N.  undulata  Say;  Comstock,  Manual,  p.  130,  fig.  49. 

Osbom,   Contr.    Dept.    Zocil.    and    Ento 


N.   undulata  Say ; 
N.   undulata  Say; 


Ohio 


1895.    A',    undulata   Say;    Gillette   &   Baker,   Bull,    xxxi,    Colo.    Asr.    Exp.    .-^ta.    Tech.    ser. 
p.  63. 

1897.    -V.  undulata  Say;   Kirkaldy,  Trans.   Ento.   Soc.  London   for  1897,  p.    410. 

1897.    -V.  undulata  i^*ay ;    Smith,  Ins.  N.  J.,  p.   144. 

1899. 

1899. 

1900. 
,  P-  'i'9. 

1900. 
.   15. 

1901. 

1901. 

1902. 

1904. 

1905. 

1907. 

1908. 

1909. 

1909. 


A'',  undulata  Say 
A'',  undulata  S'ay 
A',  itndulata  Say 
.V.  undulata  Say 
A"",  undulata  Say 
.V.   imdulata   Sav 


State   Univ.,    No. 
Lugger,   Bull.   69,   Ento.   Div.   U.  of  Minn.   Agr.   Coll.    Exp.   Sta, 
Hem.-Het.,   ii,   p.    370    (in   part). 


Ctiampion,   Biol.   Centr.   Amer, 

Howard,  Insect  Book.,  p.   275. 

Bueno,  Jl.   N.   Y.   Ento.   Soc,   x,   pp.    231   and   233. 

Kirkaldy,  Wien.  Ento.  Zeit.,  xxiii,  pp.  94,  95  and   132   (in  part) 

Bueno,  Jl.  N.  Y.  Ento.  Soc,  xiii,  p.  45   (in  part). 

Bueno  and  Brimley,  Ento.   News,  xviii,  p.   435. 
N.  undulata  Say;    Bueno,  Jl.  N.  Y.  Ento.  Soc,  xvi,  p.  238. 
A^   undulata  Say;   Kirkaldy  &  Bueno,  Proc  Ento.  Soc  Wash.,  x.  p.   100   fin  part) 

Delcourt,  Bull.  Sci.  Fr.  et  Belg.,  xliii,  (7)  I,  pi.  iv.  line  C,  No.  5; 


A'',  undulata  Say 
line  7,  Nos  1,  2,  3,  4. 

1910.    A',  undulata  Say 
ming  under  thin  ice.) 

1910.    A',  undulata  Say 
1910.    A',  undulata  Say 


Bueno,  Jl.  N.  Y.  Ento.  Soc,  xviii,  p.  33.     (Dec.  and  Feb.  swim- 


Severin,  Can.  Ento.,  xlii,   p.   340.      (Food  habits.) 
Smith,  Catalog  Ins.  N.  J.,  edn.  3,  p.  169. 

1912.  -Y.  undulata  Say;   Bueno,  Can.  Ento.,  xliv,  p.   213. 

1913.  A',  undulata  Say;   Browne,  Jl.  Exp.  Zool.,  xiv,  p.   61.     (Male  germ  cells.) 
A'',   undulata  Say;    Barber,   Bull.   Am.    Mus.   Nat.    Hist.,   xxxiii,   p.    499.      (Cited   in 


1914 

error.) 
1914 
1017 
1917 


X.  undulata  Say;  Parshley,  Psyche,  xxi,  p.  140.  (May  8,  Oct. 
.V.  undulata  Say;  Hungerford,  Ento.  News,  xxviii,  pp.  175-18 
Hungerford,   Ento.   News,   xxviii,   pp.   267-21 


A',   undulata  Say 
(Life  history.) 

1917.    A',  undulata  Say 

1917.  N.  undulata  Say 

1918.  N.  undulata  Say 
(ovipositor). 

1919.  N.  undulata  Say 
1919.    N.  undulata  Say 


17.  Orono,   Maine.) 
!:    pi.   xiii,   fig.    7. 
7 :   pis.  xix  and  xx. 


Parshley,  Occ.  Papers  Boston  Soc.   Nat.  Hist.,  vii,  p.   113. 
Van  Duzee,  Catalog.  Hemipt.,  p.   451,   452  (in  part). 
Hungerford,  Ento.   News,  xxix,  pp.   242,  243,   245  ;    pi.   xv,  fig 


Parshley,  Occ  Papers  Mus.  Zool.  Univ.  Mich.,  No.   71,  1919. 

Hungerford,  Kans.  Univ.  Sci.  Bull.,  xi,  pp.  21,'  96,  166-169,  177, 
180,  181,  184,  186,  187,  190,  193,  195,  196,  258,  259,  263,  268.  Color  Plate  II,  fig.  1  and 
pis.  i,  iv,  viii,  xix,  xx,  xxi,  xxiii,  xxiv.     (Biology  of.) 

1910.    A',   undulata  Say:    Hungerford,   Kans.   Univ.   Sci.   Bull.,   xi,   pp.    320,   332;    pi.   xx.xi, 

Parshley,  Proc  British  Columbia  Ento.  Soc,  No.  IS  (Syst.  Ser.), 

Parshley,  S.   Dakota  St.   College  Tech.   Bull.   II,  p.   22. 
Bueno  &  Hussey,  Bull.  Brooklyn  Ento.  Soc.  xviii,  p.   107. 
Bueno,  Conn.  St.  Geol.  &  Nat.  His.  Surv.  Bull.  No.   34,  pp.   404 

Hale,  Proc.  Lin.  Soc.  N.  S.  Wales,  xlix,  pt.  4.  p.   462. 
Hungerford,  Annals  Ento.   Soc   Amer.,  xxviii,  )i.   417. 
Comstock,  Introduction  to  Ento.,  p.  362,  fig.   410. 
Clark,  Bull.  Brooklyn  Ento.  Soc,  xx,  pp.   186  and  187. 
Hungerford  &  Beamer.     Ento  News,  xxxvi,  pp.  263,  264,  297. 
]926.    A',  undulata  Say;    Leonard,    Cornell    Univ.    Agri.    Exp.    Sta.    Memoir   101,    p.    139. 
(March  to  Nov.) 

Bare,  Annals  Ento.   Soc.  Amer.,  xix,  p.   93.     (Life  history.) 
Hungerford,  Bull.   Brooklyn  Ento.   Soc,  xxi,  p.   195. 
Blatchley,  Heteroptera     ...     p.   1053. 
Hungerford,  Annals  Ento.  Soc.  Amer.,  xxi,  pi.   ix,   fig.    4. 

1928.  A",   undulata  Say;    Hungerford,   Ento.   News,  xxxix,  p.   156.      (From   N.    M.) 

1929.  N.  undulata  Say;    Hutchinson,  Annals  S.  Afr.  Mus.  xxv,  pt.  3,  p.   304. 

1930.  A",    undulaia  Say;   Hungerford,  Bull.   Brooklyn  Ento.  Soc,  xxv,  p.   138. 


fig.  10. 

1921. 

A-. 

undulata  Say; 

p.  24. 

1922. 

A^. 

undulata  Say; 

1923. 

A^. 

undulata  Say; 

1923. 

N. 

undulata  Say ; 

and  405. 

(April  to  Oct.) 

1924. 

N. 

undulata  Say; 

1925. 

X. 

undidata  Say; 

1925. 

N. 

undulata  Say; 

1925. 

S. 

undidata  Say ; 

1925. 

\. 

undulata  Say  ; 

1926.  A^.  undulata  Say; 

1926.  A',  undulata  Say; 

1926.  A^   undulata  Say; 

1928.  A',  undulata  Say; 


Hungerford:    Genus  Notoxecta  119 

Referring  to  this  species,  also: 

1847.  jV.  glauca  Shepherd,*  Sillimans  Amer.  Jl.   (2)  iv,  pp.  423,  424   (migration). 

1848.  A^.  glauca  Shepherd,  Ann.  &  Mag.  Nat.  Hist.  (2)  I,  p.  158. 

1848.  A^.  glauca  Simpson,  Quoted  by  Spence  in  Trans.  Ento.  Soc.  London,  v,  proc.  p.  30. 

1851.  A',  piiiirtafri  FieVjer,  Rhynchotographieen,  p.  47().  Rliync  liotographien,  p.  52.  (Van 
Duzee.) 

1853.     A',   aincriraitn    Herrifk-Schaeffer,    Wanzen.    Ins.,   ix,    p.    44,    fig.    902.      (Color  illus. 

not  typical  for  any  species.) 

1897.  A',  undulata  var.  cliaron  Kirkaldy,  Trans.  Ento.  Soc.  London  for  1S97,  p.  411  (in 
part). 

Size.  Length,  10.5  mm.  to  12.6  mm.;  width  of  pronotum,  3.6  mm. 
to  4  mm. 

Color.  Typically  black  and  white.  Head  and  legs  of  usual  color, 
often  greenish ;  abdominal  venter  usually  marked  with  black  areas ; 
connexivum  like  the  legs;  thoracic  venter  dark;  posterior  part  of 
pronotum  darkened  by  the  black  mesonotum  beneath.  Scutellum 
black.  Hemelytra  white  with  an  undulate  transverse  black  band 
across  apex  of  corium  and  base  of  membrane.  The  species  may  vary 
from  pale  luteous  forms  with  pale  scutellum  to  forms  darker  than 
typical,  having,  beside  the  usual  black  marks,  distal  end  of  clavus, 
a  band  on  middle  of  corium  and  a  streak  on  embolium  nearly  black. 
All  gradations  occur. 

Structural  Characteristics.  xVnterior  outline  of  head,  as  seen  from 
above,  somewhat  rounded ;  vertex  about  as  long  as  its  anterior  width ; 
anterior  margin  of  vertex  slightly  less  convex  and  longer  than  frontal 
margin  of  an  eye;  anterior  breadth  of  vertex  :  synthlipsis  : :  2  +  :1. 
Pronotum  less  than  twice  as  long  as  the  head;  lateral  margins 
straight  and  moderately  divergent;  anterior  angles  slightly  em- 
bracing the  eyes;  lateral  ledge  oblique.  Lobes  of  membrane  about 
equal.  Anterior  trochanter  of  male  with  a  medium-sized  hook. 
Mesotrochanter  not  angulate.  Terminal  abdominal  sternite  of 
female  notched  at  tip,  the  notch  deeper  than  wide.  Female  gona- 
pophyses  (first  pair)  short.  Male  genital  capsule  as  shown  on 
Plate  XIII. 

Location  of  Type.  In  the  T.  AV.  Harris  collection  at  the  Boston 
Society  of  Natural  History  are  the  siiecimens  reported  by  Doctor 
Uhler  in  the  Proceedings  of  the  Boston  Society  of  Natural  History 
XIX,  pt.  4,  Apr.,  1878,  p.  442.  In  this  paper,  entitled  "Notices  of  the 
Hemiptera  Heteroptera  in  the  Collection  of  the  late  T.  W.  Harris," 
Doctor  Uhler  writes  concerning  N.  undulata  Say:  "No.  17  (3  Harris 
collection  "Rivers  and  Ponds.  Sept.  20, 1821.  May  15, 1828  j  ,  5  ." 
Determined  by  Mr.  Say.     Four  varieties  remain  in  the  collection, 

*  This  and  two  following  relate  to  a  flight  of  backswimmers  in  the  upper  Mississippi 
valley,  probably  N.  undulata  Say,  but  cannot  be  certain. 


120  The  University  Science  Bulletin 

noted  ^.  y.  8  2  .  •  •"  The  remaining  notes  by  Uhler  are  of 
little  interest  now  except  to  show  that  his  own  idea  of  the  species 
included  N.  lunata  Hungerford  and  N.  indica  Linn.  I  have  examined 
the  specimens  mentioned  above.  /?.  is  typical,  y.  is  less  colored,  the 
scutellum  is  yellow  margined,  and  there  is  a  small  black  spot  on 
outer  tip  of  emboHum  and  corium.  8  has  lost  its  head  and  pronotum, 
the  wings  are  pale  and  there  is  a  central  spot  of  black  on  scutellum. 
2  is  N.  lunata  Hungerford. 

There  can  be  no  doubt  as  to  the  identity  of  Say's  N.  undulata 
which  he  described  from  "Missouri  and  Indiana."  The  species  I 
have  described  is  not  only  the  most  common  species  in  the  regions 
he  names,  but  the  most  common  and  widespread  back-swimmer  in 
the  United  States.  It  is  the  only  one  with  the  typical  undulate 
band  and  has  been  recognized  by  all  American  students.  I  have 
specimens  from  Missouri  that  fit  Say's  original  description.  There 
can  be  no  objection  in  designating  y8.  which  was  determined  by  Say 
as  the  type  of  N.  undulata  Say. 

Comparative  Notes.  Kirkaldy  included  several  distinct  species 
under  this  name  which  lead  him  to  give  a  distribution  extending  all 
the  way  down  to  Chile,  S.  A.  There  are  in  North  America  several 
black  and  white  species  of  approximately  the  same  size,  and  w^hen 
pale  specimens  are  encountered  they  have  often  been  confused  with 
N.  lunata-  Hungerford.  The  rounded  mesotrochanter  separates  N. 
undulata  Say  from  A^.  hmata  Hungerford,  N.  unijasciata  Guer.  and 
N.  spinosa  Hungerford,  which  have  angulate  to  spinose  mesotro- 
chanters.  The  straight,  divergent  lateral  margins  of  the  pronotum, 
the  moderate  size  of  the  hook  on  the  anterior  trochanter  of  the  male, 
and  the  deep,  narrow  notch  in  the  last  abdominal  sternite  of  the 
female  separate  it  from  A^  iridica  Linn.,  which  has  lateral  margins 
of  the  pronotum  curved,  conspicuous  hook  on  anterior  trochanter  of 
male  and  a  shallow  notch,  or  none,  in  the  last  abdominal  sternite  of 
the  female.     (See,  also,  the  key  to  species  on  page  68.) 

Biological  Notes.  This  common  species  has  been  the  subject  of 
several  studies  in  behavior  and  life  history.  In  1917  Hungerford 
described  and  figured  the  elongate  white  egg,  which  is  attached  to 
submerged  objects,  and  the  five  nymphal  instars.  In  the  earlier 
instars  the  food  consists  largely  of  ostracods  and  similar  organisms. 
Winter  is  passed  in  the  adult  stage,  and  in  Kansas  there  are  two 
main  broods  in  a  season.  Bare  (1926)  obtained  the  emergence  of  an 
adult  in  the  laboratory  on  July  30,  and  13  days  later  it  began  lay- ' 


Hungerford:    Genus  Notonecta  121 

ing  eggs.     He  also  reared  the  species  to  maturity  in  from  forty  to 
fifty-two  days,  using  mosquito  larvae  as  food. 

Data  on  Distribution.  This  North  American  species  ranges  from 
coast  to  coast  and  from  Canada  to  the  Gulf.  It  is  the  most  common 
species  of  the  back  swimmers  east  of  the  Rocky  Mountains  and  north 
of  the  37°  of  latitude.  South  of  this  line  it  is  more  and  more  re- 
placed in  abundance  by  N.  indica  Linn,  as  one  goes  southward. 
My  data  include  the  following: 

Canada 

Nova  Scotia:    Truro,  Sept.  23,  1926,  F.  Johansen. 

Quebec:  Fairy  Lake,  May  14,  1927,  and  Sept.  11,  1928,  G.  S.  Walley;  Kazu- 
bazua,  Aug.  18,  1927,  and  Aug.  28,  1928,  W.  J.  Brown,  and  Aug.  18,  1931,  G.  S. 
Walley;  Otter  Lake,  Aug.  6,  1931,  G.  S.  Walley;  Knowlton,  June  21,  1927, 
G.  W.  Walley. 

Ontario:  Rockcliffe,  Aug.  2,  1928,  G.  H.  Fisk;  Mer  Bleue,  May  28,  1927, 
G.  S.  Walley;  Westboro,  Sept.  10,  1928,  J.  A.  Adams;  Toronto,  Sept.  18,  1930, 
L.  J.  Milne;  Rondeau  Park,  June  13,  1929,  G.  S.  Walley;  Pt.  Pelee,  June  6, 
1929,  G.  S.  Walley;  Ventnor,  Aug.  1,  1928,  J.  A.  Adams. 

Manitoba:  Aweme,  July  6,  1922,  N.  Griddle;  Treesbank,  Aug.  15,  1922, 
N.  Griddle. 

Alberta:    Lethbridge,  May  8-13,  1920,  J.  H.  Pepper. 

British  Columbia:  Peachland,  Oct.  7,  1931,  A.  N.  Gartell;  Wellington; 
Victoria,  June  6,  1923,  K.  F.  Auden;  Chilliwack,  Sept.  20,  1925  (Univ.  of 
Kans.) ;  Mt.  Cheam,  Sept.  13,  1924  (Univ.  of  Kansas) ;  Van  Couver  Island, 
Saanich  Dist.,  Sept.  1,  1917,  W.  D.  (Kept,  by  Parshley,  1919)  Oliver;  Aug.  6, 
1931,  L.  D.  Anderson  (Univ.  of  Kansas). 

All  above  Canadian  records  from  material  in  Ottawa,  Canada,  unless  other- 
wise stated. 

United  States 

M.-une:    Orono,  May  8  to  Oct.  17  (see  H.  M.  Parshley). 

Massachusetts:    Amherst,  May  16,  1922. 

Rhode  Island:    Providence  (Bueno  Coll.). 

Connecticut:  Apr.  9  to  Oct.  24  (see  Conn.  Geol.  and  Nat.  Hist.  Surv. 
Bull.  34). 

New  York:    Long  Island,  May;  Ithaca,  Nov.  19,  1921,  P.  W.  Claassen. 

Ne-w  Jersey:    Rancocas,  Aug.  29,  1927,  E.  M.  Becton. 

Pennsylvania:    Rockville,  Apr.  1,  1917,  J.  G.  Sanders  (C.  J.  Eh-ake). 

Virginia:    Wan-enton,  June  7,  1928,  L.  Woodruff. 

Kentucky:  Pond  near  Brooklyn  Bridge,  Jan.  3,  1933,  H.  Garman  (taken 
along  with  V.  indica  Linn.). 

Ohio:  Columbus,  Oct.  2,  1914,  C.  J.  Drake;  Zane.^ville,  Aug.  10.  1915, 
C.  J.  Drake. 

Michigan:  Cheboygan  Co.,  July  12  and  31,  H.  B.  Hungerford;  Mackinac 
Island,  Aug.  19,  1925,  H.  B.  Hungerford;  Nigger  Creek,  MuUett  L.,  Aug.  4, 
1925,  H.  B.  Hungerford;  Bois  Blanc  Island,  Aug.  14,  1932,  H.  B.  Hungerford. 


]22  The  University  Science  Bulletin 

Many  other  collections  near  Douglas  L.,  Cheboygan  Co.,  by  H.  B.  Hungerford, 
covering  ten  summers. 

Indiana:    Kosciusko  Co.,  July,  1932,  G.  E.  Gould  (Purdue  Univ.). 

Illinois:  Normal,  June  22,  1883;  Towanda  Pond,  April,  1882;  Havana. 
July  7,  1910.  and  many  other  records  in  Illinois  Nat.  Hist.  Surv.  Mus. 

Wisconsin  :    Lone  Rock,  Aug.  14,  1906. 

Minnesota:  Bengal,  Aug.  18,  1922,  H.  B.  Hungerford;  Becker  Co.,  Aug.  22, 
1922,  H.  B.  Himgerford;  Beaver  Dam,  Minn.,  Aug.  12.  1922,  H.  B.  Hungerford; 
Ita.sca  Park,  Aug:  21,  1922,  H.  B.  Hungerford. 

Iowa:    Ames,  Oct.  11,  1924,  H.  M.  Harris. 

Missouri:  Kansas  City,  F.  Rogers;  St.  Louis,  April,  1900,  J.  Abbott; 
Columbia,  Nov.  24,  1911,  J.  F.  Abbott. 

Arkansas:    Arkansas  Co.,  Oct.  6,  1930.  D.  Isely;  Fayettcville.  Nov.  14,  1929, 

D.  Isely. 

Texas:  Sutton  Co.,  Aug.  20,  1928,  A.  M.  James;  Eastland  Co.,  Jan.  2,  1921, 
Grace  Wiley;  Randall  Co.,  July  7,  1927,  R.  H.  Beanier;  Valentine,  Aug.  13, 
1927.  R.  H.  Beamer;  Presidio  Co.,  July  16,  1927,  R.  H.  Beamer. 

Oklahoma:  Ardmore,  April  14,  1923,  H.  B.  Hungerford;  Grant  Co.,  Feb. 
20.  1921,  W.  E.  Hoffmann. 

Kansas:  Doniphan  Co.,  Aug.  27,  1921,  W.  J.  Brown;  Leavenworth  Co., 
Feb.  6,  1921,  W.  E.  Hoffmann;  Douglas  Co.  (every  month  of  the  year);  Riley 
Co.,  Oct.  1920.  H.  B.  Hungerford;  Allen  Co.,  Apr.  24.  1921,  W.  E.  Hoffmann; 
Woodson  Co.,  Feb.  24.  1921,  W.  E.  Hoffmann;  Cowley  Co.,  Mar.  16,  1921,  W. 

E.  Hoffmann;  Lyons  Co.,  June  18,  1923,  Herbert  Darby;  Linn  Co.,  1915,  R. 
H.  Beamer;  Comanche  Co.,  June  19,  1927,  H.  B.  Hungerford;  Meade  Co., 
(no  data);  Morton  Co.,  Aug.  4,  1924,  C.  O.  Bare;  Scott  Co.,  June  22,  1929, 
Howard  Deay;  Cheyenne  Co.,  summer,  F.  X.  Williams. 

South  Dakota:  Sand  Hills,  Martin,  Sept.  15,  1930,  H.  C.  Severin;  Brook- 
ings, June  4.  1928.  H.  C.  Severin;  Springfield,  June  15,  1928,  L.  Hendricks, 
July  12,  1922,  H.  C.  Severin  (S.  D.  St.  College.) 

Mont.\na:     Three  Forks,  July  22,  1931,  L.  D.  Anderson. 

Id.\ho:     Caldwell,  May  9,  1926,  C.  J.  Drake. 

CoLOR\Do:  Estes  Park,  Aug.  22,  1919.  H.  B.  Hungerfortl;  Pingree  Park, 
Aug.  21,  1926.  R.  H.  Beamer;  Las  Animas  Co.,  Aug.  22,  1927,  P.  A.  Readio; 
Lamar,  Aug.  22,  1927,  P.  A.  Readio;  La  Junta,  Aug.  22.  1927,  L.  D.  Anderson; 
Boulder,  June  28,  1931.  L.  D.  Anderson;  Caisson,  July  1.  1931,  L.  D.  Anderson; 
Ft.  Collins,  Aug.  19.  1898. 

Utah:  Farwest,  Sept.  12,  1924,  C.  J.  D.  Brown;  Emery  Co.,  Aug.  26,  1921, 
Grace  Wilej-;  Antelope,  Jul.y  1,  1931,  L.  D.  Anderson. 

New  Mexico:  Socarro  Co.,  Aug.  18,  1927,  L.  D.  Anderson;  Torrance  Co., 
Sept.  10,  1925,  C.  H.  Martin;  Santa  Cruz  Co.,  Aug.  20,  1927,  R.  H.  Beamer; 
Estancia,  Sept.  6,  1925,  C.  H.  Martin. 

Arizona:  Gila  Co.,  Aug.  6.  1927,  P.  A.  Readio;  Navajo  Co.,  Aug.  15,  1927, 
P.  A.  Readio;  Cochise  Co.,  July  29,  1927,  R.  H.  Beamer;  Coconini  Co.,  Aug. 
13,  1927,  P.  A.  Readio. 

California:     Calipatria,  July  14,  1923. 

Nevada:     Carson  City,  Aug.  9,  1929,  R.  H.  Beamer. 

Oregon:  Hot  Lake,  July  13,  1931,  L.  D.  Anderson;  Corvallis,  June  26,  1926, 
C.  J.  Drake. 


Hungerford:    Genus  Notoxecta  123 

Washington:  Kalania  R.,  July  21,  1931,  L.  D.  Anderson;  Takoma  (Ham- 
burg, GermanjO. 

I  lack  material  from  the  following  states:  Maryland,  District  of  Columbia, 
West  Virginia,  Tennessee,  North  Carolina,  South  Carolina,  Georgia,  Florida, 
Alabama,  Mississippi,  Louisiana,  Nebraska,  North  Dakota,  Wyoming. 

Mexico 
San  Antonio,  Juh^  15,  1927,  L.  D.  Anderson. 

Notes  on  Notonecta  punctata  Fieber,  1851 

Doctor  Fieber  described  this  from  Baltimore,  Nortli  America. 
Without  doubt  it  is  A^.  undulata  Say  and  Kirkaldy  was  correct  in 
so  considering  it.  Fieber  does  not  mention  the  museum  where  the 
insects  belong.  There  are  in  the  Zoological  jMuseum  in  Berlin  4 
specimens  under  the  name  A^.  ^punctata  Fieber.  They  are  labeled 
as  follows: 

First.  ''3(324"  "Mexico  Deppe"  "punctata  Fieb."  (This  is  near 
N.  shootcri  Uhl.    Fieber's  type  was  from  Baltimore.) 

Second.    Label  like  the  first  but  lacking  determination  label. 

Third.  -Cat.  N.  3624"  'Alexico,  Deppe."  This  is  not  A",  shooteri, 
a  smaller  species  with  rounded  trochanter. 

Fourth.    "7195"  "Mexico."    This  is  near  N.  shooteri  Uhler. 

All  of  the  above  coming  from  Mexico  cannot  be  Fieber's  types. 

Notonecta  indicoidea  Hungerford,  1927 

(Color  Plate  II,  fig.  2  ;   Plate  XIV,  fig.  8) 
1927.    -V.  indicoidea  Hungerford,  Bull.  Brookl.  Ento.  Soc.  XXII,  p.  250. 

Referring  to  this  species,  also: 

1897.  -V.  undulata  var.  charon  Kirkaldv.  Trans.  Ento.  Soc.  London,  p.  411  (in  part). 
In  Stockholm  Mus.  lab.  by  Kirkaldy. 

1897.  A',  shooterii  Kirk.  Trans.  Ento.  Soc.  London,  p.  408  ("Two  handsome  Notonectae 
from  Guanajuata  in  Mex.  Duges  in  Mus.  Paris"). 

8ize.    Length,  10.8  mm.  to  11  mm.;  width  of  pronotum.  3.9  mm. 

Color.  General  facies  dark.  Head,  anterior  half  of  pronotum 
and  limbs  yellow.  Rear  half  of  pronotum  darkened  by  the  black 
mesonotum  beneath.  Scutellum  black.  Hemelytra  black,  save  two 
oblique  tan  streaks  near  base  of  corium  and  clavus  and  tip  of  mem- 
brane, which  is  pale.  Venter  black  except  connexivum  and  a  spot 
on  median  line  of  fourth  and  fifth  abdominal  sternites. 

Structural  Characteristics.  Head  and  eyes  not  prominent;  anterior 
outline  of  head  rather  flattened ;  vertex  subequal  to  shorter  than  its 
anterior  width;  anterior  margin  of  vertex  less  convex  than  margin 
of  the  eye ;  anterior  margin  of  vertex  longer  than  frontal  margin  of 


124  The  University  Science  Bulletin 

an  eye.  Anterior  breadth  of  vertex  :  synthlipsis  : :  12:5.  Pronotum 
a  trifle  less  than  twice  the  length  of  the  head ;  lateral  margins  faintly 
convex  and  explanate;  anterolateral  angles  slightly  embracing  the 
eyes;  lateral  ledge,  as  seen  from  the  side,  straight,  slightly  turned 
down  in  front  and  not  as  long  as  the  rear  margin  of  the  eye  beneath. 
Scutellum  longer  than  either  pronotum  or  ridge  of  hemelytral  com- 
missure. Posterior  lobe  of  membrane  slightly  shorter  than  anterior 
lobe.  Anterior  trochanter  of  male  with  minute  hook.  Mesotro- 
chanter  rounded.    Male  genital  capsule  as  shown  on  Plate  XIV. 

Location  of  Type.  In  Francis  Huntington  Snow  Entomological 
Museum,  University  of  Kansas. 

Comparative  Notes.  Distinguished  from  A^.  indica  Linn,  by  wider 
synthlipsis  and  from  N.  unifasciata  Guer.  by  the  black  scutellum. 

Data  on  Distribution.    Known  to  me  only  from  Mexico,  D.  F. 

Notonecta  distinctoidea  Hungerford,  1930 

(Color  Plate  II,  fig.   6;   Plate  XIII,  fig.  9) 
1930.    N.   distinctoidea  Hungerford.     Bull.  Brook.  Ento.  Soc.  XXV,  p.  141. 

Size.  Length,  10.8  mm.  to  12  mm.;  width  of  pronotum,  4  mm.  to 
4.3  mm. 

Color.  General  facies  dark.  Head,  anterior  part  of  pronotum 
and  legs  pale  yellow  to  horn.  Face,  legs^  and  lateral  margin  of 
abdominal  venter  more  or  less  tinged  with  green.  Scutellum  black. 
Hemelytra  purplish-black  save  two  short,  oblique  tan  streaks  near 
base  of  corium  and  clavus,  and  tip  of  membrane,  which  is  pale. 

Structural  Characteristics.  Anterior  outline  of  head  viewed  from 
above  convex;  vertex  shorter  than  its  anterior  width;  margin  of 
vertex  less  convex  than  the  margin  of  an  eye;  anterior  breadth  of 
vertex  :  synthlipsis  : :  13:6.  Inner  margins  of  the  eyes  straight  and 
divergent.  Pronotum  declivant,  not  quite  twice  length  of  head; 
lateral  margins  straight  and  divergent;  anterolateral  angles  slightly 
embracing  the  eyes;  lateral  ledge,  as  seen  from  the  side,  straight  and 
not  as  long  as  rear  margin  of  the  eye  beneath,  obhque.  Scutellum 
longer  than  the  pronotum.  Hemelytral  commissure  shorter  than 
scutellum.  Posterior  lobe  of  membrane  slightly  shorter  than  anterior 
lobe.  Anterior  trochanter  of  male  with  a  very  small  hook.  Meso- 
trochanters  rounded.  Penultimate  abdominal  sternite  of  female 
very  slightly  notched  and  last  one  not  notched  at  tip.  First  pair  of 
gonapophyses  of  female  short.  Male  genital  capsule  as  shown  on 
Plate  XIIL 

Location  oi  Type.    Holotype  in  Paris  Museum.    This  species  was 


Hungerford:    Genus  Notonecta  125 

described  from  three  males  labeled  as  follows:  "Mexique,"  Coll. 
Noualhier  1898";  Mexique,  Etat  de  Jalisco,  L.  Diguet  1900"; 
Mexico,  W.  H.  Ashm.  determined  N.  americana  Fab.  var.  by 
Kirkaldy  '97." 

Comparative  Notes.  See  notes  under  A^.  melaena  Kirk.  In  com- 
paring this  species  with  N.  indicoidea  Hungerford  it  may  be  noted 
that  the  species  is  plumper  of  body  (more  like  A^.  melaena  Kirk.) 
and  differs  in  slmpe  of  male  genitalia. 

Data  on  Distnhution.    Known  only  from  type  localities. 

Key  to  Notonecta  of  South  America 

1.  Keel  of  fourth  abdominal  sternite  bare,  the  hairs  confined  to  the  sides.  .N.  mexicana* 

(Colombia,  p.   75.) 
Keel  of  fourth  abdominal  sternite  not  bare (2) 

2.  Last   abdominal    sternite    of    female   large    and    strongly    constricted    just    before    the 

tip,  male  with  digitate  prolongation  on  genital  capsule K .  nigra. 

(Brazil?,  p.  126.) 
•    Last   abdominal    sternite    of    female   normal,    male    without    digitate    prolongation    on 

genital    capsule    (3) 

3.  Pronotuni  broad  in  front  embracing  the  eyes,  which  are  flattened,  and  receding  from 

anterior  margin  of  vertex.  Synthlipsis  broad,  one-half  anterior  margin  of  vertex 
as  seen  from  above.  Scutellum  plainly  broader  than  long.  Male  with  stout 
tubercle  at  angle  of  front  trochanter  and  a  very  stout,  broad  hook  as  shown  on 

Plate    IX,    fig.    i (4) 

Pronotum  and  eyes  not  as  above.  Scutellum  not  plainly  transverse.  Males  without 
stout  tubercle  at  angle  of  front  trochanter  and  with  small  hook  on  its  anterior 
surface     (5) 

4.  Length  usually  under   12   mm , N.   ochrothoe. 

(Colombia,  p.  99.) 

Length  uj^ually  more  than  12  mm -V.   colombiana. 

(Colombia,  p.  128.) 

5.  Mesotrochanter  more   or   less   angulate (6) 

Mesotrochanter  rounded    (8) 

6.  Mesotrochanter   distinctly   angulate N.   unifasciata   angulata. 

("S.  Amer.,"  p.  111.) 
Mesotrochanter    feebly    angulate (7) 

7.  Length   less   than   10   mm A',    bifasciata. 

(Argentina,  p.   129.) 

Length  more  than  10  mm A',  confusa. 

("S.  Amer.,"  p.  130.) 

8.  Length  usually  less  than  10  mm (9) 

Length  usually  more  than  10  mm (17) 

9.  Typical   facies  dark (10) 

Typical  facies  distinctly  black  and  white (14) 

10.  Facies  nearly  black.     Spots  on  end  of  corium  not  conspicuous (11) 

Facies  dark  but  with  spots  on  end  of  corium  distinct  or  clavus  pale (12) 

11.  .Synthlipsis  less  than  one-third  width  of  an  eye.     Size  small. 

(Max.  length,  9  mm.) ^'.  pulchra. 

(Paraguay,  p.  131.) 
Synthlipsis  more  than  one-third  width  of  an  eye.     Size  larger. 

(Min.  length,  9  mm.) -V-  polystolisma. 

(Brazil,  p.  132.) 

12.  Synthlipsis  :  width   of   eye  behind  :  :  5.3  :  9 A'',   peruviana. 

(Peru,  p.  134.) 


*  Other  members  of  this  group  are  keyed  out  in  Key  to  North  and  Central  America,  which 
see  if  description  does  not  fit. 


126  The  University  Science  Bulletin 

SynthlipsiS  :  width   of   eye   behind  :  :  4  :  9    (about) (13) 

13.  SynthlipsiS  :  width  of  eye  behind  :  :  4.1  :  9.     (See  drawing  of  male  clasper, 

page  155)    ^-  virescens. 

(Chile,  p.  135.) 
.SynthlipsiS  :  width    of   eye   behind  :  :  3.9  :  9.      (See    drawing    of    male   clasper,    Plate 

XII    fig.   3 -^  •   bicircoidea. 

(Argentina,  p.  136.) 

14.  Hemelytra,    viewed    from    above    with    two    large    nearly    circular    pale    spots    in   the 

transverse  black  band.     Vertex  to   synthlipsis  :   :  5  :  2 • .V.    bircirca. 

(Chile,  p.  137.) 
Hemelytra,   viewed   from  above,   with   small   or  transvers^e  tan   or  pale   spots   in  the 

transverse  black  band.     Vertex  to  synthlipsis  at  least  as  3  :  1 (15) 

15.  Vertex  to  synthlipsis  near  3:1 (!*') 

Vertex  to  synthlipsis  near  4:1 ^'-   minuta. 

(Bolivia,  p.  138.) 

16.  Clasper  of  male  genital  capsule  broail  and  bifurcate.     (See  Plate  XII, 

fig    4)   iV.  sellata. 

(Argentina,  p.  139.) 
Clasper  of  male  genital  capsule  not  as  above.     (See  Plate  XII,  iig.  7).  .-V.  disturbata. 

(Brazil  and  Paraguay,  p.  140.) 

17.  Color  black  and  white ^-  variabilis* 

(Brazil,  p.  141.) 
And  N.  indica. 
(Colombia,  p.  113.) 
Color  not  black  and  white (18) 

18.  Synthlipsis  less  than  three-fifths  of  width  of  eye  behind A',  fazi. 

(Chile,  p.  142.) 

Synthlipsis  more  than  three-fifths  of  width  of  eye  behind N.   vereertbruggheni. 

(Argentina,  p.  144.) 

Notonecta  nigra  Fieber,  1851 

(Color  Plate  I,  fig.  15;   Plate  XVI,  fig.  3) 

1851.  N.  nigra  Fieber.  Rhynchotogiaphieen,   p.    473. 

1897.  N.  nigra  Fieber.  Kirkaldy,  Trans.  Ento.  Soc.  London  for  1897,  p.   424. 

1904.  N.   nigra  Fieber.  Kirkaldy,  "VVien,  Ento.   Zeit.   xxiii,  p.   132. 

1909.  A',   nigra  Fieber.  Kirkaldy  and  Bueno,  Proc.  Ento.   Soc.  Wash,  x,  p.   199. 

Size.  Length,  16  mm. ;t  width  of  thorax  5.5  mm.  (female).  The 
male  is  smaller,  being  4.8  mm.  across  thorax. 

Color.  Chocolate  brown  to  nearly  black.  Pronotmn,  head  and 
limbs  shining  and  also  chocolate  brown  to  nearly  black.  Scutellum 
and  venter  black. 

Structural  Characteristics.    As  in  A^.  glauca  L.     (See  p.  48.) 

Location  of  Types.  Three  specimens,  two  females  and  one  male, 
in  Vienna  Museum,  and  one  specimen,  a  female,  in  Paris  Museum. 

*  See  discussion  under  these  two  species. 

t  Notes  ox  Fibber's  Measurements:  Doctor  Fieber  gave  the  length  of  his  Notonecta 
species  in  "linien."  Since  several  different  units  called  lines  have  been  employed,  I  wrote  to 
my  friend.  Doctor  Walther  Horn,  concerning  the  problem.  With  his  usual  kindness  he 
promptly  informed  me  that:  "We  have  had  in  the  past  time  in  Europe  almost  as  many 
different  kinds  of  'linien'  as  small  states.  Sometimes  a  iinie'  was  one-tenth,  sometimes  one- 
twelfth  of  an  inch.  In  the  following  I  give  you  three  kinds :  Paris  'Iinie,'  2.2558  rnm. ; 
Rheinprovinz  Iinie,  2.179  mm.;  English  line,  2.il6  mm.  There  have  been  many  other  kinds. 
For  example,  there  is  before  me  a  measuring  instrument  where  12  "linien"  =  27  cm."  For- 
tunately, I  have  studied  Fieber's  A',  nigra  types  in  the  Vienna  museum.  They  measure  16 
mm.  in  length.  Since  Doctor  Fieber  gives  "TVs  linien"  as  the  length  of  this  species,  it  is  easy 
to  figure  that  he  employed  the  Rheinprovinz  Iinie,  which  equals   2.179  mm. 


Huxgerford:    Genus  Notonecta  1127 

One  of  the  Vienna  females  lacks  distal  half  of  abdomen,  and  the 
male  has  lost  the  hind  legs  and  the  tibia  of  one  front  leg.  The 
Vienna  specimens  are  labeled  "Brasilien"  and  the  Paris  specimen 
"Bresil."  I  have  compared  the  labels  side  by  side;  the  paper  is  the 
same  in  texture  and  of  the  same  width.  The  "Brazilien"  label  is 
longer.  The  handwriting  and  the  ink  look  identically  the  same  to 
me.  The  pins  are,  however,  quite  different,  but  both  specimens  show 
that  they  have  been  repinned.  The  Paris  specimen  bears  also  the 
label  '']Museum  Paris,  Coll.  Noualhier  1898."  It  is  even  nearer 
black  than  the  Vienna  specimens  and  is  labeled  ''type."  I  would 
give  it  cotype  standing,  and  have  labeled  the  Vienna  specimens 
with  red  cotype  labels.  Fieber  wrote  "Aus  Brasilien  (kais.  kon 
Hofnat.  Cabinet  in  Wien;  gesammelt  von  Schott). 

Comparative  Notes.  This  species  is  a  puzzle  to  me.  I  can  find 
no  structural  characteristics  to  distinguish  it  from  A^  glauca  Linn, 
of  Europe.  The  genital  capsules  of  the  males  are  alike  and  the 
female  genitalia  are  alike.  The  specimens  of  N.  nigra  Fieb.  are 
larger  than  the  average  N  glauca  Linn,  and  have  a  little  different 
shape.  The  unusual  color  of  N.  nigra  Fieb.  certainly  gives  it  a  dif- 
ferent facies.  At  times  I  have  been  convinced  that  these  four  speci- 
mens might  be  only  stained  specimens  of  N.  glauca  Linn.,  labeled 
"Bresil"  and  "Brasilien"  by  some  unscrupulous  dealer.  I  have  seen 
Notonecta  collected  from  acrid  coffee-colored  waters  of  some  bog- 
land  pool  that  were  nearly  as  dark  as  N.  nigra  Fieb.  For  example, 
in  the  ^luseum  of  Paris  there  is  a  series  of  specimens  labeled 
"Takersan."  "Museum  Paris,  Algerie,  P.  Lesne  6-97,"  determined 
as  N.  glauca  by  Kirkaldy  and  more  recently  N.  maculata."  One  of 
these  has  the  venter  covered  with  bits  of  wood  fiber,  as  if  taken  in 
a  pool  of  coffee-colored  water,  and  is  nearly  as  black  as  the  cotypes 
in  Vienna.  There  are  other  specimens  that  have  the  black  legs  and 
are  but  slightly  lighter  in  color,  and  still  others  that  are  of  typical 
A',  maculata  color.  Even  the  darkest  specimens  of  this  series,  how- 
ever, have  the  orange  band  on  the  abdominal  dorsum.  I  recall  also 
a  female  .specimen  labeled  "Soeul,  Korea."  in  the  Hungarian  Museum 
at  Budapest.  This  specimen  is  light  chocolate  in  color.  The  venter 
is  covered  with  bits  of  wood,  as  if  the  specimen  had  been  collected 
in  amber-colored  bog  waters.  The  scutellum  is  black;  the  anterior 
margin  of  hemelytra  shows  faintly  the  darker  maculations  that  are 
so  characteristic  of  N.  glauca  Linn.  Compared  with  N.  canariensis 
Kirkaldy,  -V.  nigra  Fieb.  is  a  more  slender  species;  the  head  is 
smaller  and  the  limbs  very  plainly  more  slender.     Compared  with 


128  The  University  Science  Bulletin 

N.  V.  viridis  Delcourt,  it  does  not  have  the  anterior  hiteral  angles 
of  the  pronotum  embracing  the  eyes,  and  has  longer  and  more 
slender  front  and  middle  legs. 

There  will  continue  to  be  some  doubt  in  my  mind  about  the  au- 
thenticity of  the  label  "Brazil"  until  some  further  evidence  is  avail- 
able. I  have  many  South  American  Notonecta  from  various  places 
in  Brazil,  but  no  species  even  akin  to  this  A",  nigra  Fieber  have  been 
taken  in  South  America  in  modern  times. 

In  the  museum  at  Oxford,  England,  there  are  two  insects  (a  male 
and  a  female)  that  look  like  N.  obliqua  Gallen  and  bear  a  tiny  green 
square  and  a  label,  "Miers  coll.;  Presented  1880  by  J.  "W.  Miers." 
Professor  Poulton  says  most,  if  not  all,  of  Miers'  insects  came  from 
South  America. 

While  there  is  still  much  uncertainty  about  N.  nigra  Fieber,  I  do 
not  wish  to  take  the  responsibility  of  denying  the  existence  of  this 
species  in  South  America,  and  am  sure  that  if  further  material  be- 
comes available  it  will  be  possible  to  find  in  a  longer  series  the 
proper  position  of  this  species.  Every  South  American  collector 
should  be  on  the  alert  for  this  species. 

Notonecta  colomhiana  ne^  species. 

Size.    Length,  12  mm.  to  12.6  mm.;  width  of  pronotum,  4.5  mm. 

Color.  No  doubt  a  bichromatic  species,  but  only  the  dark  form 
known  to  me.  The  scutellum  is  black.  The  hemelytra  are  brown 
and  black  with  a  luteous  streak  along  the  claval  suture  and  a  luteous 
transverse  spot  at  end  of  basal  third  of  corium.  Lower  half  of 
propleurse,  presternum,  mesoternum  and  spots  on  abdominal  venter 
black.  Face,  limbs  and  connexivum  may  be  yellowish  or  suffused 
with  green. 

Structural  Characteristics.  Anterior  outline  of  head  viewed  from 
above  quite  convex ;  vertex  longer  or  subequal  to  its  anterior  width ; 
margin  of  vertex  more  convex  than  margin  of  the  eye;  anterior 
breadth  of  vertex  :  synthlipsis  : :  17:12.  Pronotum  steeply  declivant 
as  viewed  from  the  side.  Pronotum  twice  length  of  the  head; 
lateral  margins  somewhat  divergent  and  straight;  anterior  angles 
acute  and  embracing  the  eyes;  lateral  ledge  as  seen  from  the  side 
nearly  straight,  as  long  as  the  rear  margin  of  the  eye  below  it,  and 
nearly  horizontal.  Scutellum  about  as  long  as  pronotum.  Anterior 
lobe  of  membrane  a  little  longer  than  posterior  lobe.  Anterior  tro- 
chanter of  male  with  the  stout  hook  and  protuberance  characteristic 
of  the  N.  shooteri  group.  Male  genital  capsule  much  as  in  N. 
melaena  Kirk. 


Hungerford:    Genus  Notoxecta  129 

Location}  of  Type.  Holotype,  allotype  and  two  paratypes  in  the 
Zoological  ^Museum  at  Berlin;  two  paratypes  in  Francis  Huntington 
Snow  Entomological  Museum  of  University  of  Kansas.  The  above 
labeled  "Bogata  8000?,  Kalte  Region,  Bobisch  L."  Hamburg 
Museum  has  two  paratypes  labeled  "Bogata,  Dr.  0.  Burger  leg.  Ill, 
1897,  Vend.  1, 1,  1898."  In  the  Vienna  Museum  there  is  one  labeled 
"Bogata,  Coll.  Signoret/'  and  is  marked  "N.  rugosa  Fieb." 

Comparative  Notes.  Mistaken  for  N.  shooteri  Uliler,  from  which 
they  differ  by  the  shape  of  male  genital  capsule,  by  the  straighter 
margins  of  the  pronotum  and  more  declivant  pronotum.  It  is  a 
larger  species  than  N.  melaena  Kirkaldy,  with  more  rounded  head 
and  luteous  stripe  on  claval  commissure. 

Data  on  Distribution:  Colombia,  South  America:  Bogata  8000', 
Bobisch  L.  (Berlin  Mus.  I  ;  Bogata,  Dr.  0.  Burger,  leg.  Ill,  1897. 
Vend.  1,  I,  1898,  Hamburg  Mus.;  Bogata  coll.  Signoret  (labeled 
rugosa  Fieb.). 

Notonecta  bifasciata  Guerin,  18^14 

(Plate  XII,  fig.   1) 

1844.    A',   bifasciata   Guerin-Meneville.     Icon,   du  Regne  Anim.  Ins.,  Ill,  p.  354. 

ISol.  -V.  bifasciata  Guerin.  Fieber,  Rhynchotgiaphieen  (Abh.  Bohm.  Ges.  Wissensch. 
Prag  V,  7,  p.   480).     (Known  to  Fieber  only  by  desc.) 

1853.  -V.  bifasciata  Guerin.  Herrich-Schaffer,  Die  Wanzenartigen  Insecten  IX,  p.  130 
(in  Index). 

1879.  .V.  bifasciata  Guerin.  Berg,  Hemiptera  Argentina  .  .  .,  p.  197  of  reprint. 
(Under  A',  variabilis  Fieb.  in  error.) 

1897.  A",  bifasciata  Guerin.  Kirkaldy,  Trans.  Ento.  Soc.  London,  1897,  p.  413. 
(Kirkaldy  saw  the  type.) 

1899.  A',  bifasciata  Guerin.  Kirkaldy,  Boll.  Mus.  Torino,  xiv.  No.  347,  p.  2.  (Only 
in  part.) 

1904.  A',  bifasciata  Guerin.  Kirkaldy,  Wien,  Ento.  Zeit.  xxiii,  pp.  95-132.  (Kirkaldy 
thought  only  a  ^•ar.  of  A\  undulata  Say,  but  was  in  error.) 

1909.  A',  bifasciata  Guerin.  Kirkaldy  and  Bueno.  Catalogue  .  .  .Proc.  Ento.  Soc. 
Washington,   x,  p.   198.      (Only  in   part.) 

192G.  A',  bifasciata  Guerin.  Hungerford,  Psyche  xxxiii,  pjj.  12  and  15.  (My  error; 
not  this  species.) 

1930.  A',  bifasciata  Guerin.  Hungerford,  Bull.  Brooklyn  Ento.  Soc,  xxv,  p.  140. 
(Corrects  error  of  1926.) 

Size.  Given  by  Guerin  as  8.5  mm.  long  and  3  mm.  wide.  Accord- 
ing to  my  measure,  the  type  is  9.3  mm.  long. 

Color.  I  have  seen  the  type,  which  is  a  male,  and  give  the  follow- 
ing descriptive  information:  The  general  facies  dark,  the  scutellum 
black;  hemelytra  dark  except  a  pale  stripe  on  basal  two-thirds  of 
clavus,  the  inner  margin  against  the  scutellum  being  brown  through- 
out; against  the  clavocorial  suture  there  is  a  triangular  pale  patch 
on  the  corium ;  at  distal  end  of  embolium  (on  the  suture)  there  is  a 
transverse  orange  spot  and  above  it  a  larger  one  irregular  in  shape; 


130  The  University  Sciexce  Bulletin 

tip  of  membrane  pale,  hemelytra  (the  dark  part  particularly) 
covered  with  silvery,  narrowly  spatulate  hairs. 

Structural  Characteristics.  Anterior  margin  of  vertex  :  synthlip- 
sis  ::  11:4;  the  width  of  an  eye  at  its  base  equals  the  width  of  the 
anterior  margin  of  the  vertex.  The  anterior  trochanter  of  the  male 
has  the  little  hook,  as  in  other  small  South  American  species,  such 
as  N.  bicirca  Hungerford,  for  example.  The  mesotrochanter  is  not 
angulate,  but  not  rounded  as  in  some  other  species.  The  male 
genital  capsule  is  shown  on  Plate  XII,  figure  1. 

Location  of  Type.  There  is  a  specimen  in  the  Paris  ]\Iuseum 
labeled  "type,"  and  from  the  top  down  the  labels  read:  1st.  "Type"; 
2d,  "Museum  Paris,  Maldonado  Coll.  Guer.-Menev.  ex  coll.  A. 
Salle  1897"  (printed);  3d,  "Notonecta  bifasciata  Guer.  Ic.  R.  A. 
Platta  (type)  Maldonade"  (written)  ;  4th,  "Notonecta  polystolisma 
Fieb.,  p.  53,  Maldonado  bifasciata  Guer.  Ic.  R.  A.  (type)";  5th, 
''Notonecta  bifasciata  Guer.  det  Kirkaldy  1897."  Kirkaldy  quotes 
these  on  page  413  of  his  "Revision."  The  type  is  a  male  and  appears 
to  have  been  broken  into  two  or  three  parts  and  glued  together.  One 
hemelytron  is  in  two  pieces  and  glued  to  the  side  of  the  body,  and 
the  pronotum  sometime  has  been  parted  from  the  after  body. 

Comparative  Notes.  I  was  surprised  to  find  that  this  is  not  the 
species  I  named  N.  bifasciata  in  my  paper  of  1926,  where  I  figured 
the  genital  capsule  of  what  I  thought  to  be  Guerin's  species.  The 
type  has  a  male  clasper  more  like  my  N.  pulchra,  but  it  is  not  the 
same  species.  N.  pulchra  Hungerford  is  a  smaller  species  with 
longer  head  and  narrower  synthlipsis.  On  the  other  hand,  it  is  a 
smaller  species  than  my  iV".  bicirca  or  N.  bicircoidea.  Of  course,  it 
is  not  what  I  named  N.  polystolisma  var.  spatulata.  While  the  shape 
of  the  male  genital  capsule  resembles  somewhat  my  N.  disturbata,  a 
comparison  of  the  species  side  by  side  shows  them  to  be  distinct. 
iV.  disturbata  Hungerford  has  a  longer  head  and  a  narrower  syn- 
thlipsis.   Indeed,  it  proved  to  be  a  species  I  had  never  seen. 

Data  on  Distribution.  Known  to  me  only  by  the  type.  I  cannot 
accept  the  distribution  records  of  former  workers,  who  most  ob- 
viously have  confused  the  species.    Maldonado  is  in  Uruguay,  S.  A. 

Notonecta  confusa  Hungerford,  1930 

(Color  Plate  II,  fig.    4  ;    Plate  XIII,   fig.    7) 
1930.    .Y.  confusa  Hungerford.     Bull.  Brooklyn  Ento.   See,  xxv,  p.   140. 

Referring  to  this  species,  also : 

1926.    N.  species?  Hungerford.     Psyche,  xxxiii,  pi.  2,  fig.   6. 

Size.    Length,  12  mm.;  width  of  thorax,  4  mm. 


Huxgerford:    Genus  Notonecta  131 

Color.  The  color  of  the  solitary  type  specimen  is  pale  yellow 
throughout.  The  typical  color  is  probably  black  and  white  as  in 
N.  undulata  Say. 

Structural  Characteristics.  Size  and  general  appearance  might 
confuse  this  species  with  N.  undulata  Say,  from  which  it  differs  in 
having  the  mesotrochanter  feebly  angulate  and  the  male  genital 
clasper  with  broader  branches,  as  shown  on  Plate  XIII. 

Location  of  Type.  Described  from  a  single  male  specimen  from 
''S.  Amer."  type  in  the  P.  R.  Uhler  collection  in  the  U.  S.  National 
Museum,  Washington,  D.  C. 

Comparative  Notes.  Until  a  series  of  specimens,  including  both 
sexes,  is  found,  it  is  impossible  to  offer  any  further  comparisons. 

Data  on  Distribution.  Beside  the  type,  I  have  a  specimen  labeled 
"Cuba"  which  belongs  to  this  species. 

Notonecta  pulchra  Hungerford,  1926 

(Color  Plate  I,  fig.  10;   Plate  XII,  fig.   8) 
1926.    N.  pulchra  Hungerford.     Psyche  XXXIII,  p.   14,  PI.   II,  fig.   3. 

Size.  Length,  7.5  mm.  to  9  mm.;  width  of  pronotum,  2.18  mm.  to 
3.1  mm. 

Color.  General  facies  dark.  Scutellum  entirely  black  or  marked 
with  a  tan  spot  on  each  lateral  margin.  Hemelytra  black  save  two 
pale  spots  at  base  of  hemelytra  and  an  orange  irregular  transverse 
band  at  end  of  corium.  Silvery  hairs  on  hemelytra.  When  typically 
colored,  a  beautiful  little  species. 

Structural  Characteristics.  Anterior  outline  of  head  viewed  from 
above  nearly  flattened;  vertex  as  long  as  anterior  width;  margin  of 
vertex  less  convex  than  anterior  margin  of  an  eye ;  anterior  breadth 
of  vertex  :  synthlipsis  ::  11:3;  synthhpsis  less  than  one-third  the 
rear  margin  of  an  eye.  Pronotum  less  than  one  and  one-half  times 
as  long  as  head;  lateral  margins  straight  and  quite  divergent;  antero- 
lateral angles  not  embracing  the  eyes;  lateral  ledge,  as  seen  from 
the  side,  curved,  oblique  and  shorter  than  rear  margin  of  eye  be- 
neath. Scutellum  a  little  longer  than  pronotum.  Membranal  lobes 
about  equal.  Anterior  trochanter  of  male  with  hook.  Mesotro- 
chanter rounded.  Penultimate  abdominal  sternite  of  female  notched. 
Last  one  not  elongate,  notched  at  tip.  Male  genital  capsule  as 
shown  on  Plate  XII. 

Location  of  Types.  Holotype,  allotype  and  some  paratypes  in 
Francis  Huntington  Snow  Entomological  Museum,  others  in  U.  S. 
N.  M.  at  Washington,  D.  C. 


132  The  University  Science  Bulletin 

Comparative  Notes.  The  typical  color  of  this  species  sets  it  apart 
from  the  others  of  its  size.  The  plump  capsule  of  the  male  with  its 
broader  clasper  separates  it  from  A^.  disturbata  Hungerford  and  N. 
hijasciata  Guerin. 

Data  on  Distnbution.  Paraguay:  Villarica,  Dec.  16,  1927,  F. 
Schade;  Villarica,  Estero  Granda,  Nov.  1,  1924,  F.  Schade;  Vil- 
larica, Sept.  16,  1923,  F.  Schade;  Villarica,  March  26,  1924,  F. 
Schade;  Caraveni,  L.  Montis,  Oct.  30,  1924,  F.  Schade;  Villarica, 
Jan.  19,  1926,  F.  Schade. 

Notonecta  polystolisma  Fieber,  1851 

(Color  Plate  I,  fig.  11;  PL  XII,  fig.  2) 

1851.  -V.  polystolisma  Fieber.  Rhynehotographieen,  pp.  477-478. 
192C.  .V.  polystolisma  Fieber.  Hungerford,  Psyche  xxxiii,  p.  12. 
1928.    X.  polystolisma  Fieber.     Jaczewski,  Annales  Musei  Zoologici  Poloniei  vii,  p.   121. 

Referring  to  this  species,  also: 

1851.  -V.   polystolisma  var.   guttata  Fieber.      Rhynehotographieen,  p.   478. 

1851.  .V.  polystolisma  var.   bipunctata   Fieber.      Rhynehotographieen,   p.    478. 

1851.  -V.  polystolisma  var.  fasciata  Fieber.     Rhynehotographieen,   p.    478. 

1851.  -V.  polystolisma  var.   ivimaculata   Fieber.      Rhynehotographieen,    p.    478. 

1897.  .V.  bifasciata  Kirkaldy.  Trans.  Ent.  Soc.  London  for  1897,  p.  413.  (As  syn.  in 
error.) 

1904.  -V.   bifasciata  Kirkaldy.     Wien.  Ent.   Zeit.,  xxiii,  p.   132.     (As  syn.  in  error.) 

1909.  -V.  bifusciata  Kirkaldy  &  Bueno.  Proc.  Ent.  Soc.  Wash,  x,  p.  198.  (As  syn.  in 
error. ) 

192G.  .V.  polystolisma  var.   spatulata  Hungerford.      Psyche   xxxiii,   p.    12. 

Size.  Length,  9  mm.  to  9.6  mm.;  width  of  pronotum,  3.1  mm.  to 
3.45  mm. 

Color.  General  facies  dark.  Anterior  half  of  pronotum,  propleura, 
head  and  legs  from  horn  to  brownish  yellow  often  tinged  with  green. 
Venter  brown  to  black  except  metaxyphus  and  connexivum,  which 
are  usually  colored  like  the  legs.  Scutellum  black  except  in  teneral 
specimens.  Hemelytra  black  or  blue  black  with  oblique  bifurcate 
tan  streak  along  basal  half  of  claval  commissure,  and  either  two 
spots  or  transverse  band  across  corium  at  tip  of  clavus.  Tip  of 
membrane  usually  pale. 

Structural  Characteristics.  Anterior  outline  of  head,  viewed  from 
above,  only  slightly  convex;  vertex  slightly  shorter  than  anterior 
width  and  less  convex  than  anterior  margin  of  the  eye;  anterior 
breadth  of  vertex  :  synthlipsis  ::  3:1  (sometimes  10:3);  the  syn- 
thlipsis  much  less  than  half  the  rear  margin  of  the  eye.  Pronotum 
less  than  one  and  one-half  times  as  long  as  the  head;  lateral  margins 
straight  and  divergent;  anterolateral  angles  not  embracing  the  eyes; 
lateral  ledge  as  seen  from  the  side  faintly  sigmoid  and  decidedly 


Hungerford:    Genus  Notonecta  133 

oblique,  shorter  than  the  rear  margin  of  the  eye  beneath.  Scutellum 
longer  than  the  pronotum.  Membranal  lobes  subequal.  Anterior 
trochanters  of  male  with  hook.  Mesotrochanters  rounded.  Penulti- 
mate abdominal  sternite  of  female  notched,  last  one  rather  elongate 
and  roundly  notched  at  tip.  Female  gonapophyses  of  moderate 
length.  Last  abdominal  sternite  of  male  rather  large  and  con- 
stricted before  the  tip.  Male  genital  capsule  large  and  as  shown  on 
Plate  XII. 

Location  of  Type.  Fieber  described  five  varieties  under  this 
species,  one  of  which  is  A'',  sellata  and  a  very  distinct  species.  The 
other  four  are  from  Brazil  and  appear  to  be  stages  of  pigmentation 
of  a  single  species.  I  named  a  variety  ''spahdata,"  but  now  suppress 
it,  as  I  consider  the  dark  form  as  really  the  usual  color  of  the  species. 
There  are  no  specimens  in  the  Berlin  Museum  that  I  can  mark  with 
certainty  as  types.  Under  the  name  Notonecta  polystolis?na  Fieb. 
there  is  a  mixed  lot  of  five  specimens,  three  are  N.  indica  Linn,  and 
other  two  labeled  "Cat.  No.  3631,  Brazilien,  Sellow"  are  what  I  con- 
sider correctly  named.  There  are  some  unnamed  Notonecta  in 
Berlin  INIuseum  that  belong  under  this  name:  One  pale  specimen 
labeled  "3633  Brasil,  Sellow,"  and  another  "3634  Brasil,  Sellow." 
The  latter  is  a  male  with  pale  scutellum  and  hemelytra.  The 
female  "3633"  is  partly  pigmented. '  It  has  a  dirty  brownish  to  black 
median  area  on  scutellum  with  an  uneven  paler  stripe  along  lateral 
margins,  the  clavus  and  upper  part  of  corium  are  pale,  a  thin  brown 
streak  on  clavus  bordering  a  brown  streak  between  corium  and  em- 
bolium  from  base  to  middle  of  corium  where  it  widens  into  a  patch 
of  brown  covering  distal  half  of  embolium  and  corium  but  not  reach- 
ing up  to  claval  margin.  Some  brown  on  base  of  membrane.  There 
are  some  dark  forms  labeled  as  follows:  "Lages.  Brasilien,  St.  Cath- 
arina,  Fruhstorfer";  Brasilien,  Osten  Sacken;  Allegrette  3635;  8014 
labeled  "pallipes,  Fabr.,"  "Brasil  Coll.  Germ." 

Comparative  Notes.  A  little  larger  than  most  of  the  other  small 
typically  South  America  species.  The  male  genital  capsule  is  large 
and  distinctive. 

Biological  Notes.  Doctor  Jaczewski,  who  collected  this  species 
in  the  state  of  Parana,  Brazil,  S.  A.,  found  it  in  ponds  and  pools  in 
open  country  and  in  forests.  Some  of  the  pools  were  formed  in  the 
course  of  small  streams.  Adults  were  taken  from  January  to  June 
and  nymphs  in  March,  April  and  June. 

Data  on  Distribution.    In  addition  to  the  data  published  by  Doc- 

10—3482 


134  The  University  Science  Bulletin 

tor  Jaczewski  in  Notonectidse  from  the  state  of  Parana,  1928.  I  can 
add  the  following  from  the  collection  before  me: 

Brazil:  Ypirango,  S.  Paulo,  R.  Spitz;  Itapuaqvieaetuba,  Via  Poa,  EFCB 
Sao  Paulo,  E.  D.  Townsend;  Sao  Paulo.  Aug.  7.  1927,  E.  D.  Townsend  (and 
nymphs) ;  Estado  de  Sao  Paulo,  Nov..  1928,  E.  D.  Townsend. 

Notonccta  peruviana  new  species 

Size.    Length,  9.3  mm.  to  10.3  mm.;  width  of  prothorax.  3.45  mm. 

Color.  General  facies  dark  when  fully  pigmented.  Scutellum 
black.  Hemelytra  dark  with  broad  oblique  streak,  tan  or  horn 
along  base  of  clavocorial  line;  oblique  orange  line  on  caudal  margin 
of  embolium  and  an  obscure  orange  spot  on  corium  above ;  membrane 
mostly  brown. 

Structural  Characteristics.  Anterior  outline  of  head  somewhat 
rounded,  slightly  more  convex  in  female  than  in  male;  vertex 
shorter  than  its  anterior  width;  margin  of  vertex  less  convex  than 
frontal  margin  of  the  eye;  anterior  breadth  of  vertex  :  synthlipsis 
::  15:8.  Vertex  a  little  wider  in  the  females.  Synthlipsis  :  width 
of  the  eye  behind  ::  5.3:9.  Pronotum  trifle  less  than  twice  length 
of  the  head;  lateral  margins  straight,  moderately  divergent;  antero- 
lateral angles  slightly  embracing  the  eyes;  lateral  ledge  as  seen  from 
the  side  curved  and  oblique,  shorter  than  rear  margin  of  the  eye 
beneath.  Scutellum  and  pronotum  subequal  in  length.  Ridge  of 
hemelytral  commissure  as  long  as  scutellum.  Lobes  of  membrane 
of  hemelytra  equal  in  length.  Anterior  trochanter  of  male  rough 
but  hook  obscure.  Mesotrochanters  rounded.  Penultimate  ab- 
dominal sternite  of  female  with  deep  but  narrow  notch,  last  one 
faintly  or  not  notched.  First  pair  of  female  gonapophyses  short. 
Male  genital  caj^sule  as  shown  on  page  155. 

Location  of  Type.  Holotype  and  allotype  in  Hungarian  ^Museum 
at  Budapest.  Paratype  in  Francis  Huntington  Snow  Entomological 
Museum,  University  of  Kansas. 

Comparative  Notes.  Separated  from  N.  virescens  Blancliard  and 
A'',  bioircoidea  Hungerford  by  its  wider  synthlipsis  and  by  shape  of 
male  genital  claspers. 

Data  on  Distribution.  Known  only  by  the  male  and  two  females 
labeled  "Peru,  Lauramarca." 


Huxgerford:    Genus  Notonecta  135 

Notonccta  virescens  Blanrhard,  1852 

(Color  Plate  II,   fig.    11) 

1S52.  A',  virescens  Blanchaid,  in  Gay's  Cliile,  Zool.,  viii,  p.  2HS.  Atlas  Zoijl.  Ins.  Heniipt., 
PI.    II,   p.    10. 

1863.  N.  virescens  Blanchard?  Signoret,  Revision  des  Heniiptt'ies  du  Chile.  (Seance  du 
22  .\viil   18G3.)     (Doubtful — gives  length   10  mm.) 

1926.    A^    virescens  Blanchard.     Hungerford,  Psyche  xxxiii,  p.   11. 

1930.    A',    virescens   Blanchard.      Hungerford,    Bull.   Brooklyn   Ent.   Soc,   xxv,   p.    139. 

Referring  to  this  species,  also: 

1897.    .V.    iindulata  var.  virescens  Kirkaldy.     Trans.  Ent.  Soc.   London  for  1897,  p.   411. 
1904.    .V.    Iindulata  var.   virescens  Kirkaldy.     Wien.   Ent.   Zeit.,  xxiii,   pp.   95  and   132. 
1909.    .V.    Iindulata  var.  virescens  Kirkaldy  and  Bueno.     Proc.  Ent.  Soc.  Wash.,  x,  p.   199. 

Size.  Length,  8.7  mm.  to  9.3  mm.;  width  of  pronotum,  3  mm.  to 
3.4  mm. 

Color.  The  type  is  luteous  with  hcmelytra  incompletely  marked 
with  brown.  The  upper  pale  spot  of  corium  large  and  not  closed, 
silver,  narrowly  spatulate  hairs  on  hemelytra.  The  second  specimen 
is  entirely  luteous.  I  have  before  me  three  pigmented  specimens 
that  must  be  this  species.  These  have  black  scutellum  with  brown 
stripe  on  either  margin  and  another  before  the  tip.  Clavus  is  horn 
except  at  hemelytra  1  commissure  where  it  is  smoky  brown,  two 
pale  stripes  near  base  of  corium  separated  by  smoky,  brown  streak. 
Distal  half  of  corium  and  basal  half  of  membrane  black  eml)racing 
a  small  tan  spot  on  embolial  suture  and  a  large  irregular  one  above 
it.  Colored  more  like  .V.  bicircoidea  Hungerford  than  like  .V. 
bircirca  Hungerford. 

Structural  Characteristics.  Anterior  outline  of  head,  viewed  from 
above,  somewhat  convex;  vertex  shorter  than  its  anterior  width; 
anterior  breadth  of  vertex  :  synthlipsis  ::  5:2;  anterior  breadth  of 
vertex  a  little  greater  than  the  rear  margin  of  an  eye;  the  synthlipsis 
a  little  more  than  two-fifths  the  rear  margin  of  an  eye.  Pronotum 
about  one  and  five-sevenths  as  long  as  the  head;  lateral  margins  in 
the  type  less  divergent  than  in  A'',  bicirca  Hungerford;  lateral  mar- 
gins straight;  anterolateral  angles  slightly  embracing  the  eyes; 
lateral  ledge,  as  seen  from  the  side,  curved  and  shorter  than  rear  mar- 
gin of  the  eye  beneath.  Scutellum  a  little  longer  than  the  pronotum. 
Alembranal  lobes  subequal.  Anterior  trochanter  of  male  with  hook. 
Mesotrochanter  rounded.  Penultimate  abdominal  segment  notched 
and  last  one  shallowly  and  roundly  notched.  Male  genital  capsule 
as  shown  in  Text  Figure  on  jiage  155. 

Location  of  Type.  In  Paris  Museum.  The  type  is  a  dermestid 
eaten  female,  labeled  "Museum  Paris.     Chili,  Cav  15  =  43."     The 


136  The  University  Science  Bulletin 

head  and  prothorax  are  gone.  Only  one  leg  left  and  this  is  the  right 
middle  leg.  The  hemelytra  are  incompletely  marked  with  brown 
as  stated  above.  A  second  specimen  is  entirely  pale.  There  are  two 
specimens  labeled  "Museum  Paris,  Chili,  Coll.  G.  Fallou  259-95," 
but  this  has  an  angulate  trochanter,  and  the  label  must  be  wrong,  for 
this  is  A^.  lunata  Hungerford.  There  are  two  other  specimens  labeled 
"Museum  Paris,  Chili,  Coll.  G.  Fallou  259-95"  det.  N.  inrescens  by 
Fallou  and  A^.  undulata  var.  virescens  by  Kirkaldy.     These  are 

males. 

Comparative  Notes.  I  have  compared  my  own  types  with  this 
species  and  find  it  to  be  different.  The  male  clasper  is  more  pointed 
behind  than  in  either  N.  bicirca  or  N.  bicircoidea. 

Data  on  Distribution.  From  Chile,  S.  A.  There  are  therefore  two 
very  closely  related  little  species  from  Chile  and  another  across  the 
divide  in  Argentina. 

Notonecta  bircircoidea  Hungerford,  1928 

(Color  Plate  II,  fig.   8;    Plate  XII,  fig.   3.) 
1928.     A',    bircircoidea   Hungeiforcl.      Annals  Ent.   Soo.   Am.   xxi.  p.   120. 

Size.    Length.  8.8  mm.  to  9.6.  mm.;  width  of  pronotum,  3.15  mm. 

to  3.3  mm. 

Color.  General  facies  quite  like  N.  bicirca  Hungerford  but  darker. 
The  elongate  yellow  spot  on  lateral  margins  of  scutellum,  which  is 
present  in  N.  bicirca  is  nearly  always  absent  or  tan  in  this  species. 
Hemelytral  commissure  and  tip  of  clavus  black. 

Structural  Characteristics.  The  head  slightly  less  prominent  than 
in  N.  bicirca  and  the  anterior  margin  of  vertex  a  little  more 
rounded;  vertex  shorter  than  its  anterior  width;  anterior  breadth 
of  vertex  :  synthlipsis  ::  5:1.9.  Anterior  breadth  of  vertex  greater 
than  rear  margin  of  an  eye.  Synthlipsis  less  than  half  the  rear 
margin  of  an  eye.  Pronotum  less  than  twice  as  long  as  the  head; 
lateral  margins  straight,  moderately  divergent ;  anterolateral  angles 
nearly  normal,  slightly  embracing  the  eyes;  lateral  ledge,  as  seen 
from  the  side,  curved,  oblique  and  shorter  than  rear  margin  of  the 
eye  beneath.  Scutellum  a  little  longer  than  the  pronotum.  Mem- 
branal  lobes  subequal.  Anterior  trochanters  of  male  with  hook. 
Mesotrochanters  rounded.  Penultimate  abdominal  sternite  of  fe- 
male notched,  last  one  broadly  and  shallowly  notched.  Male  genital 
capsule  as  shown  on  Plate  XH.  It  is  relatively  larger  than  in  N. 
bicirca  Hungerford. 

Location  of  Types.    In  Francis  Huntington  Snow  Entomological 


Hungerford:    Genus  Xotonecta  137 

Museum,  University  of  Kansas.  Described  from  fourteen  specimens, 
four  males  and  ten  females,  taken  by  Mr.  Vereertbrugghen  in  Rio 
Negro  Territory,  Argentina. 

Comparative  Notes.  Head  slightly  less  prominent  than  in  -V. 
hir circa,  color  pattern  difference  as  shown  on  Plates  I  and  II.  Male 
capsule  relatively  larger  and  clasper  broader,  with  the  front  and  rear 
curves  differently  located. 

Data  on  Distribution.    Known  to  me  only  by  the  types. 

Xotonecta  bicirca  Hungerford 

(Color  Plate  I,  fig.   16;   Plate  XII,  fig.   5.) 

1926.    .V.  bicirca  Hungerford.     Psyche  xxxiii,  p.  12,  PI.  2,  fig.  1. 
192S.    .V.  bicirca  Hungerford.     Ann.  Ento.  Soc.  Amen,  xxi,  p.  120. 

Size.  Length,  8.4  mm.  to  9.3  mm.;  width  of  pronotum,  3  mm.  to 
3.3  mm. 

Color.  General  facies  spotted,  white  with  scutellum  and  two  circles 
on  hemelytra  black.  Closer  inspection  reveals  posterior  half  of 
pronotum  darkened  by  black  thorax  beneath,  an  elongate  yellow 
spot  on  lateral  margins  of  scutellum;  hemelytra  yellowish  white 
covered  with  silvery  hairs  and  with  the  following  typical  macula- 
tions:  Submarginal  band  of  black  on  base  of  corium;  distal  end  of 
clavus  black;  broad  transverse  black  band  traversing  distal  third 
of  corium  and  base  of  membrane  and  embracing  a  small  orange- 
yellow  spot  at  end  of  embolivmi  and  entirely  surrounding  a  much 
larger  nearly  circular  spot  above.  The  general  effect  produced  is 
that  of  two  black  circles  surrounding  orange-yellow  spots  upon  the 
insects,  the  lateral  orange-yellow  spots  upon  the  margins  of  hemely- 
tra and  opening  upon  the  embolial  sutures  being  unnoticed. 

Structural  Characteristics.  Anterior  outline  of  head  slightly  con- 
vex, more  so  in  male  than  in  female;  vertex  longer  than  its  anterior 
width  in  male,  subequal  in  female ;  margin  of  vertex  less  convex  than 
frontal  margin  of  the  eye;  anterior  breadth  of  vertex  :  synthlipsis 
::  5:2;  anterior  breadth  of  vertex  greater  than  rear  margin  of  an 
eye;  synthlipsis  slightly  less  than  twice  the  rear  margin  of  an  eye. 
Pronotum  less  than  twice  as  long  as  the  head;  lateral  margins  nearly 
straight  and  moderately  divergent;  anterolateral  angles  about 
normal;  lateral  ledge  as  seen  from  the  side,  curved,  oblique  and 
shorter  than  the  rear  margin  of  the  eye  beneath.  Scutellum  a  little 
longer  than  the  pronotum.  ]Membranal  lobes  subequal.  Anterior 
trochanter  of  male  with  hook.  ]Mesotrochanters  rounded.  Penulti- 
mate abdominal  sternite  of  female  notched  and  last  one  broadly 


138  The  University  Science  Bulletin 

and  shallowly  notched.  Female  gonapophyses  short.  IMale  genital 
capsule  as  shown  on  Plate  XII. 

Location  of  Types.  In  Francis  Huntington  Snow  Entomological 
Museum,  University  of  Kansas.  Some  paratypes  in  U.  S.  N.  Mu- 
seum, Washington,  D.  C.  Described  from  50  specimens  from  Chile, 
S.  A.  Some  from  Santiago  and  others  from  Termas  Cauquences, 
Dec.  15,  1922,  by  Alfredo  Faz. 

Comparative  Notes.  The  characteristically  spotted  appearance  of 
this  species  is  distinctive.  AVhile  now  and  then  a  N.  sellata  Fieb. 
specimen  may  have  same  general  color  appearance,  Fieber's  species 
has  narrower  synthlipsis  and  entirely  different  male  clasper.  The 
closest  relatives  are  A',  virescens  Blanehard  and  N.  bircircoidea 
Hungerford.  It  recjuires  an  examination  of  the  claspers  of  the  male 
to  distinguish  precisely  these  three  species  when  the  typical  color- 
ing is  lacking.  Both  of  the  latter  species  are  usually  more  deejily 
pigmented. 

Data  on  Distribution.  Besides  the  type  series  I  have  the  follow- 
ing from  Chile:    Limache,  Nov.  12,  14,  15,  1923,  A.  Faz. 

Notonecta  mimita  Hungerford,  1926 

(Color  Plate  I,  fig.  9;   Plate  XII,  fig.   6) 
192(i.     .v.    inituita   HungPifoicl.     Psyche  xxxiii,  p.   14,  pi.   II,  fig.   7. 

S^ize.    Length,  7.2  mm.;  width  of  pronotum,  2  mm. 

Color.  Head,  pronotum,  legs  and  basal  third  of  hemelytra  yel- 
low. Scutellum  and  distal  two-thirds  of  hemelytra  black  with  tips 
of  membrane  pale.  In  the  broad,  black  area  across  the  hemelytron 
are  two  small,  irregular  orange  spots  of  nearly  equal  size.  On  one 
of  the  three  specimens  there  are  ill-defined  yellowish  spots  near  the 
base  and  at  the  tip  of  the  scutellum. 

Structural  Charactenstics.  Anterior  outline  of  head,  viewed  from 
above,  but  slightly  convex;  vertex  longer  than  its  anterior  width; 
margin  of  vertex  less  convex  than  anterior  margin  of  the  eye;  an- 
terior breadth  of  vertex  :  synthlipsis  ::  9:2.1;  width  of  eye  at  base 
e(]ual  to  width  of  vertex;  synthlipsis  less  than  a  third  the  rear  mar- 
gin of  an  eye.  Pronotum  but  little. longer  than  the  head;  lateral 
margins  divergent  and  straight ;  anterolateral  angles  acute,  slightly 
embracing  the  eyes;  lateral  ledge  nearly  straight,  shorter  than  the 
rear  margin  of  the  eye  beneath.  Lobes  of  membrane  subequal.  An- 
terior trochanter  of  male  with  hook.  Mesotrochanters  rounded. 
Male  genital  capsule  as  shown  on  Plate  XII. 

Location  of  Types.     Described  from  specimens  in  the  Carnegie 


Hungerford:    Genus  Notoxecta  139 

iNIuseiini  bearing  the  label  "Prov.  del  Sara  Bolivia  450  m.  J.  Stein- 
bach."  One  paratype  in  Francis  Huntington  Snow  Entomological 
Museum,  University  of  Kansas. 

Comparative  Notes.    The  smallest  Notonecta  known  to  me. 

Data  on  Distribution.    Known  only  from  the  types. 

Notonecta  sellata  Fieber,  1851 

(Color  PUite  I,  fig.   14;    Plate  XII,  fig.   4.) 

1851.  -V.  Polystolisma  \av.  .•icUnta  Fit'hor.  Rli>nchotosrapliieen,  Bcihm  Gesell.  der  Wis- 
sensch.  .Al>li.  \,  7,  p.   478. 

193(1.     .V.   scllota  FielMT.      Huiifieifoid,  Bull.   Brooklyn  Ent(j.   Soc,  xxv,  p.   140. 

Referring  to  this  species,  also: 

1861.  .V.  variabilis  Biinnei.ster.  Reise  duicli  die  Plata-Staaten  I,  p.  492  (1861)  (prob- 
ably in  part).     Says  he  found  it  in  Montevideo,  but  not  close  to  Parana. 

1879.  .V.  polystolisina  Bvvii.  Hemiptera  Argentina,  p.  198  (reprint  from  Ann.  S.  Cient. 
Arjrent.,  viii,  yi.  7.')). 

1897.     .V.   variabilis  Kirkaldy.     Tran.«.   Ento.  Soc.   London,  1897,  p.   414   (in  part). 

1904.  -V./  variabilis  Kirkaldy.  Wien.  Ento.  Zeit,  xxiii,  p.  95.  (Which  he  considered  a 
var.  of  A^   undidata.  .%y,  in  error). 

1909.    A',   undulata  Kirkaldy  and  Bueno.    Proc.  Ento.  Soc.  Washington,  x,  p.  199  (in  part). 

192G.     A',   bifasciata   Hungerford.      Psyche,  xxxiii.  p.   1'2,  plate  II,  fig.   5. 

Size.  Length,  8.7  mm.  to  9.6  mm.;  width  of  pronotum,  2.8  mm. 
to  3.2  mm. 

Color.  Typical  color  white,  with  transverse  black  band  across 
clavus,  in  which  are  two  tan  spots,  and  with  black  scutellum  mar- 
gined laterally  by  short  pale  stripe.  The  species  may  vary  from 
pale  to  strongly  pigmented,  as  in  A',  undulata  Say,  for  example. 
Even  the  darkest  forms  do  not  have  the  scutellum  solid  black, 
and  all  but  the  teneral  forms  have  abdominal  dorsum  dark  and 
metathorax  black. 

Structural  Characteristics.  Anterior  margin  of  head,  viewed  from 
above,  only  slightly  convex;  vertex  longer  than  anterior  width  and 
less  convex  than  anterior  margin  of  an  eye.  Anterior  breadth  of 
vertex  :  synthlipsis  ::  3:1;  the  synthlipsis  much  less  than  half  the 
rear  margin  of  the  eye.  Pronotum  about  one  and  one-half  times  as 
long  as  head:  lateral  margins  straight  in  females,  slightly  concave 
in  males  and  divergent;  anterolateral  angles  not  embracing  the 
eyes;  lateral  ledge  as  seen  from  the  side  slightly  curved,  oblique, 
and  shorter  than  rear  margin  of  eye  beneath.  Scutellum  longer 
than  pronotum.  Membranal  lobes  equal.  Anterior  trochanters  of 
male  with  hook.  Mesotrochanter  rounded.  Penultimate  abdominal 
sternite  of  female  notched,  last  one  not  elongate,  notched  at  tip. 
Female  gonapophyses  (first  pair)  short.    Last  abdominal  sternite  of 


140  The  University  Science  Bulletin 

male  normal  in  size  and  constricted  before  the  tip.     Male  genital 
capsule  as  shown  on  Plate  XII. 

Location  of  Type.  There  is  a  specimen  in  Berlin  Museum  labeled 
"3637  La  Platta,  Besoke"  "Sellata,  Germ."  This  fits  description  of 
N.  polystolisma  var.  sellata  Fieber  exactly,  but  bears  another  label, 
N.  variabilis  var.  sellata  Germ.  If  it  is  not  Fieber's  type  it  is  cer- 
tainly Fieber's  "sellata.''  Fieber's  type  was  designated  "Notonecta 
sellata  Germ.  Coll.  Buenos  Ayres.  (Mus.  Berol.  Hal. I  The  com- 
monest species  about  "Buenos  Ayres"  is  this  species. 

Comparative  Notes.  The  characteristic  clasper  of  the  male  geni- 
tal capsule  readily  separates  this  species  from  all  other  species. 

Data  on  Distribution.    I  have  before  me  the  following: 

Argentina:  Buenos  Aires,  Mercides,  IV,  12.  1923  R.  Hosford;  Buenos  Aires, 
Mercides,  XI,  19,  1923,  R.  Hosford;  Buenos  Aires,  Jan.  18,  1904  (U.  S.  X.  M.) ; 
Lopez,  Mendoza,  S.  A.,  C.  S.  Reed;  Alrededores  de  La  Rioja,  M.  S.  Penning- 
ton; Buenos  Aires,  Republique  Argentine,  J.  Kiinchel  1899,  XI,  1897.  (Paris 
Museum);  Rep.  Argentine  Rosario,  J.  Claine,  1898  (Paris  Mus.)  Entre  Rios. 
(Paris  Mus.);  Montevideo,  Coll.  Noualhier,  1898  (Paris  Mus.);  Cosquin,  Arg. 
S.  A.,  March  8,  1921,  J.  C.  Bradley  (Cornell  Univ.);  Argentina.  S.  A.  1925; 
Argentina,  La  Granja,  Altagracia,  Cordoba,  1923,  A.  Bruch. 

Paraguay:  Villarica,  Loma,  Aug.  S,  1924.  F.  Schade;  Molinasque,  .June, 
1928,  F.  Schade. 

Brazil:  Alto  Parana,  Aug.  29,  1929,  F.  Schade;  Manacapuru,  Manaos, 
Amaz.,  March,  1928,  S.  M.  Klages. 

Bolivia:  Dr.  A.  Barelli  (bears  two  det.  labels,  1st  "N.  bifasciata  Guer.  det. 
Kirkaldy,  97";  2d  '^V.  bifasciata  =  polystolisma,  det.  Kirkaldy). 

Notonecta  disturbata  Hungerford,  1926 

(Color  Plate  I,   fig.   13;    Plate  XII,   fig.    7.) 
1926.    .V.   disturbata  Hungerford.     P.^^yche,  xxxiii,   p.   13,   PI.   II,  fig.   2. 

Size.  Length,  8  mm.  to  8.4  mm.;  width  of  pvonotum.  2.8  mm.  to 
2.9  mm. 

Color.  From  luteous  to  black  and  white.  The  pale  ones  entirely 
light,  the  pigmented  ones  having  a  black  scutellum  with  brown 
stripe  on  each  lateral  margin.  The  hemelytra  white  with  distal  end 
of  clavus  brownish  black  and  distal  half  of  corium  and  basal  two- 
thirds  of  membrane  black. 

Structural  Characteristics.  Anterior  outline  of  head  somewhat 
convex;  vertex  longer  than  its  anterior  width;  vertex  less  convex 
than  margin  of  the  eye;  anterior  breadth  of  vertex  :  synthlipsis  :: 
10:2.7;  synthlipsis  less  than  one-third  rear  margin  of  an  eye,  which 
is  equal  to  or  less  than  width  of  anterior  margin  of  vertex.  Pro- 
notum  not  more  than  a  third  longer  than  the  head ;  lateral  margins 


Hungerford:    Genus  Xotonecta  141 

straight  and  divergent,  anterolateral  angles  normal;  lateral  ledge, 
as  seen  from  the  side,  curved,  oblique  and  shorter  than  rear  margin 
of  an  eye  beneath.  Scutellum  a  little  longer  than  the  pronotum. 
Membranal  lobes  equal.  Anterior  trochanter  of  male  with  hook. 
Mesotrochanters  rounded.  Penultimate  abdominal  sternite  with 
narrow,  deep  notch  and  last  one  shallowly  notched.  Male  genital 
capsule  as  shown  on  Plate  XII. 

Location  of  Type.  Sao  Paulo  Museum,  paratype  in  Francis 
Huntington  Snow  Entomological  Museum.  Three  specimens  from 
Campo,  Bello  Staat,  Rio  de  Janeiro,  IV,  1906. 

Comparative  Notes.  The  unusual  shape  of  the  male  genital 
capsule  distinguishes  this  species. 

Data  on  Distribution.  I  have  before  me  the  following  series  of 
this  species.  Villarica,  Paraguay,  Dec.  6,  1923,  Dec.  15,  1922,  Dec. 
5,  1923,  F.  Schade;  Natterer,  Brasilien  (Det.  by  Kirkaldy  as  .V. 
variabilis  Fieb.). 

Notonecta  variabilis  Fieber,  1851 

ISol.    iV.   variabilis  Fieber.     Rhynchotographieen  Abh.   bohm  Ges.  Wiss.,  v.   7,  p.   477. 

Referring  to  this  species,  also:  Fieber  described  four  varieties 
of  this  species.  Three  from  Brazil  and  one  from  Baltimore.  His 
South  American  varieties  were  named  scutellaris,  dorsalis  and  uni- 
color  and  were  not  worthy  of  varietal  names.  His  maculata  from 
Baltimore  is  another  species,  now  N.  lunata  Hungerford. 

Size.  Fieber  gives  the  size  as  4  4/6  linien,  which  would  be  10.17 
mm. 

Color.  Variable.  Some  specimens  are  yellowish-white  with  face, 
limbs  and  connexi\aim  more  or  less  greenish;  scutellum  in  such 
cases  uniformly  yellowish-white  or  more  or  less  suffused  with  brown. 
Other  specimens  may  have  a  broad,  black  band  covering  all  but  the 
tip  of  the  membrane,  across  the  caudal  third  of  the  hemelytra;  in 
such  specimens  the  scutellum  may  be  entirely  black,  black  with 
lateral  margins  flavous,  or  black  wdth  from  one  to  three  flavous 
spots — one  at  the  tip  and  one  at  the  base  on  either  side — or  in  the 
case  of  a  single  spot,  only  the  tip  pale.  It  is  probable  that  still 
darker  fonns  occur  as  in  Notonecta  indica  Linn.,  to  which  this 
species  is  very  closely  related,  if  not  identical  with  it. 

Structural  Characteristics.  Almost  identical  with  those  of  Noto- 
necta indica  Linn,  even  to  the  clasper  of  the  male  genital  capsule. 
Fieber  says  that  the  front  half  of  the  pronotum  is  finely  cross- 


142  The  Uxiversity  Sciexce  Bi  lletix 

wrinkled,  which  character  does  distinguish  these  specimens  from  N. 
polystoliHm  Fieber. 

Location  of  Type.  Doctor  Kirkaldy  could  not  locate  tlie  type. 
I  have  examined  the  collections  at  Berlin.  Vienna  and  Halle,  where 
Fieber  said  his  specimens  were  located,  and  did  not  find  types.  In 
the  Berlin  Musemn  there  is  a  pair  of  specimens  labeled  "Cat.  No. 
3632"  "Brasilien,  Sallow."  One  of  these  is  a  male  and  resembles 
A',  indica  Linn.  (=  N.  hoivardii  Buenol  in  the  shape  of  the  clasper 
of  the  male,  in  size  and  color,  rounded  mesotrochanter,  etc.  It  has 
the  broad,  black  vitta  across  the  wings.  The  scutellum  is  yellow, 
except  tiny  brown  spots  at  base  and  in  the  middle.  There  is  a 
brown  line  along  margin  of  clavus  where  it  meets  the  scutellum,  a 
brown  streak  along  front  wing  margin  extending  from  base  of  wing 
to  the  large  black  transverse  vitta  and  an  indefinite  sooty  patch 
on  corium.  The  female  has  a  black  scutellum  except  a  yellow  spot 
on  each  anterior  corner  and  at  the  tip.  It  has  the  broad,  black 
vitta  across  the  hemelytra  as  in  the  male.  It  might  well  have  served 
for  Fieber's  description  of  A^.  variabilis  var.  scutellaris.  Since  Doctor 
Fieber  said  this  variety  came  "Aus  Brasilien,  Portorico  (AIus.  Yienn. 
et  Berol.l"  and  since  the  above  specimens  are  exceedingly  close  to, 
if  not  identical  with,  the  species  common  in  the  islands,  I  hereby 
propose  to  consider  these  two  specimens  as  plesiotypes  of  A^.  vari- 
abilis Fieber. 

Data  on  Distribution.  I  have  chosen  to  retain  A",  variabilis  Fieb. 
as  a  name  for  the  specimens  that  come  from  Brazil,  in  spite  of  their 
very  close  resemblance  to  the  specimens  found  in  Porto  Rico  which 
belong  to  A",  indica  Linn.  This  I  believe  to  be  the  safest  policy  until 
we  find  more  Brasilian  specimens  to  study  or  other  si)ecimens  from 
intervening  territory. 

Notonecta  fazi  Hungerford,  1930 

(Color  Plate  I,   fig.   17;    Plate  XIII,   tig.    2) 
]'J8(>.    .V.   fazi  Hungerford.     Bull.  Brook.  Ento.  Soc,  x.\v,  pp.   141-142,   fig.   1. 

Referring  to  this  species,  also: 

1S97.  A',  americana  Kirkaldy.  Trans.  Ento.  Soc.  London,  p.  409,  1807.  ("Chili, 
Valdivia   Kami).   Mus."). 

1899.    .V.   undulata  Kirkaldy.     Above  specimen  redetermined  by  Kirkaldy   (not  published). 

1901.  .Y.  virescens  Reed.  Revista  Chilena  de  Historia  Natiual.  \.  Reprint  "Sinopsis 
de   lo.s    Hemiptcros   de   Chile,   i>.    TOG. 

Size.  Length,  12  mm.;  width  of  pronotum,  4  mm.  Alales  a  little 
smaller. 

Color.     General  facies  dark.     Head,  anterior  part  of  pronotum 


Huxgkrford:    Gents  Xotonkcta  143 

and  legs  pale  yellow.  Posterior  half  of  pronotum  darkened  by  the 
black  mesothorax  beneath,  scutelluni  black.  Hemelytra  reddish 
brown  to  black  marked  with  tan;  the  tan  markings  are  variable, 
typically  as  follows:  on  base  of  clavus  and  extending  as  indefinite 
streak  near  the  outer  margin  to  near  its  tip;  a  small  area  near  base 
of  corium  sometimes  lacking,  another  of  variable  form  in  outer  half 
of  corium,  sometimes,  but  not  usually,  reaching  membrane;  mem- 
brane dark,  sometimes  with  a  pale  spot  on  distal  half,  sometimes 
with  the  tip  pale  instead.  Venter  dark,  except  connexivum,  metaxy- 
phus  and  parts  of  abdominal  venter  (especially  fourth  sternite) 
which  are  yellowish  brown. 

Structural  Characteristics.  Anterior  outline  of  head,  as  viewed 
from  above,  truncate  but  slightly  rounded;  vertex  slightly  shorter 
than  its  anterior  width  in  female  but  longer  in  tlie  male;  anterior 
margin  of  vertex  less  convex  and  shorter  than  the  eye;  anterior 
breadth  of  vertex  :  synthlipsis  : :  3  +  :  2;  synthlipsis  less  than  three- 
fifths  as  wide  as  an  eye.  Pronotum  about  twice  as  long  as  the  head 
in  the  female,  but  sometimes  less  in  the  male,  due  to  its  relatively 
longer  head;  lateral  margins  not  very  divergent,  straight;  anterior 
angles  embracing  the  eyes;  lateral  ledge  somewhat  oblique  and 
shorter  than  the  eye  below  it.  Lobes  of  membrane  about  equal. 
Anterior  trochanter  of  male  with  a  hook.  Mesotrochanters  rounded. 
Last  abdominal  sternite  of  female  notched  at  tip.  Female  gona- 
pophyses  (first  pair)  short.  Male  genital  capsule  as  shown  on  Plate 
XIIL 

Location  of  Type.  Holotype.  allotype  and  many  paratypes  in  the 
Francis  Huntington  Snow  Entomological  Museum  at  the  University 
of  Kansas.  Some  paratypes  in  the  LT.  S.  N.  M.  and  in  the  collection 
of  J.  R.  de  la  Torre-Bueno.  The  types  were  collected  for  me  bj' 
Doctor  Alfredo  Faz  from  Limache,  Santiago  and  Termas  Cauquenes, 
Chile. 

Comparative  Notes.  This  common  Chilean  species  was  known  to 
Doctor  Kirkaldy,  who  called  it  A',  americana  Fabr.  in  his  1897  paper 
and  later  labeled  it  A',  undulata  Say.  Its  short  truncate  head,  gen- 
eral color  pattern  and  male  genitalia  readily  separate  it  from  .V. 
undulata  Say.  Its  closest  relative  is  A",  vereertbruggheni  Hungcrford, 
from  which  it  differs  in  its  narrower  vertex,  less  declivant  head, 
more  oblique  lateral  prothoracic  ledge  and  shape  of  the  male  genital 
claspers. 

Data  on  Distribution.  Appears  to  be  widespread  in  Chile,  S.  A., 
and  extenfls  north  into  Peru,  S.  A. 


144  The  University  Science  Bulletin 

South  America 

Chile:  Besides  those  from  type  localities,  I  have  seen:  Valparaiso,  R. 
Martin  1922  (Paris  Museum);  "Fundort?"  (Berlin  Museum);  Valdivia.  Cham- 
elcha  "Coll.  Breddin"   (Deutsches  Entomologisches  Museum,  Berlin-Dahlem). 

Peru:  "Coll.  Breddin"  (Deutsches  Entomologische  Museum,  Berlin-Dahlem). 

Notonecta  vereertbruggheni  Hungerford,  1928 

(Color  Plate  II,  fig.   3;    Plate  XIII,  fig.   4) 

1928.    N.  vereertbruggheni  Hungerford.     Annals  Ento.   Soc.  Amer.,  xxi,   pp.    119,   120. 
1930.    N.   vereertbruggheni   Hungerford.      Bull.    Brooklyn   Ento.    Soc,    xxv,    p.    142,    fig.    2 
(Male  genital  capsule). 

Size.    Length,  11  mm.  to  12  mm.;  width  of  pronotum,  4.2  mm. 

Shape.  Normal  as  viewed  from  above.  From  the  side  the  thorax 
appears  more  elevated  than  usual. 

Color.  General  faeies  dark,  typical  coloring  as  follows:  Head, 
anterior  half  of  pronotum  and  limbs  yellow,  the  last  named  often 
somewhat  clouded.  Posterior  half  of  pronotum  darkened  by  the 
black  mesothorax  beneath,  scutellum  black.  Hemelytra  black 
streaked  with  tan;  clavus  with  longitudinal  streak  of  tan;  corium 
with  a  longitudinal  pale  stripe  that  is  often  obliterated  in  the  middle, 
leaving  tan  spots  on  base  and  distal  field;  membrane  black.  Venter 
dark,  except  connexivum,  metaxyphus  and  parts  of  abdominal  ven- 
ter (especially  fourth  sternite)  which  are  yellowish  brown. 

Structural  Characteristics.  Head  depressed;  anterior  outline,  as 
viewed  from  above,  rounded;  vertex  shorter  than  its  anterior  width; 
anterior  margin  of  vertex  nearly  as  convex  as  and  longer  than  the 
eye;  anterior  breadth  of  vertex  :  synthlipsis  : :  3:2;  synthlipsis  more 
than  three-fifths  as  wide  as  an  eye.  Pronotum  about  twice  as  long 
as  the  head;  lateral  margins  not  very  divergent,  straight;  anterior 
angles  embracing  the  eyes;  lateral  ledge  nearly  horizontal  in  front 
two-thirds,  somewhat  deflected  behind,  shorter  than  rear  margin  of 
the  eye  below  it.  Lobes  of  membrane  about  equal.  Anterior  tro- 
chanter of  male  with  a  hook.  Mesotrochanter  rounded.  Last  ab- 
dominal sternite  of  female  notched  at  tip.  Female  gonapophyses 
(first  pair)  short.     Male  genital  capsule  as  shown  on  Plate  XIH. 

Location  of  Type.  Francis  Huntington  Snow  Entomological  Mu- 
seum at  the  University  of  Kansas.  Some  paratypes  in  the  U.  S.  N. 
M.  Described  from  15  males  and  11  females  taken  by  Mr.  Vereert- 
brugghen,  in  whose  honor  the  species  is  named.  This  series  includes 
the  holotype,  allotype  and  paratypes,  which  were  captured  in  Rio 
Negro  territory  of  Argentina,  South  America. 

Comparative  Notes.     This  species  has  the  general  faeies  of  iV. 


Htjngerford:    Genus  Notonecta  145 

fazi  Hungerford  from  which  it  differs  in  the  wider  interocular  space 
and  in  the  shape  of  genital  claspers  of  the  males. 

Data  on  Distribution.    Known  only  from  the  type  locality. 

CHECK  LIST  OF  NOTONECTA— Linn jev& 

Subgenus  Paranecta  Hutchinson 

1.  .V.  lactitans  Kirkaldy.     (Africa.) 

2.  -V.  viinuta  Hungerford.    (Bolivia,  S.  A.) 

3.  N.  -peruviana  Hungerford.     (Peru,  S.  A.) 

4.  A',  disturbata  Hungerford.     (Brazil  and  Paraguay,  S.  A.) 

5.  A',  pulchra  Hungerford.     (Paraguay,  S.  A.) 

6.  A',  bifasciata  Guerin.    (Uruguay,  S.  A.) 

7.  A',  virescens  Blanchard.    (Chile,  S.  A.) 

8.  A',  bicirca  Hungerford.     (Chile,  S.  A.) 

9.  A^.  bicircoidea  Hungerford.    (Southern  Argentina,  S.  A.) 

10.  N.  sellata  Fieber.    (Argentina,  Brazil,  Paraguay  and  Bolivia,  S.  A.) 

11.  A",  polystolisma  Fieber.     (Brazil,  S.  A.) 

12.  A',  vereertbruggheni  Hungerford.    (Southern  Argentina,  S.  A.)  . 

13.  A',  fazi  Hungerford.    (Chile  and  Peru,  S.  A.) 

14.  A',  variabilis  Fieber.     (Brazil,  S.  A.) 

15.  A^.  indica  Linnreus.     (South,  Central  and  North  America.) 

16.  A',  conjusa  Hungerford.     (South  America.) 

17.  A'",  undulata  Say.     (North  America.) 

IS.  X.  distinctoidea  Hungerford.     (Mexico.) 

19.  A',  indicoidea  Hungerford.     (Mexico.) 

20.  A',  raleighi  Bueno.     (Southern  U.  S.,  America.) 

21.  A'.  lunata  Hungerford.     (Northern  U.  S.,  America,  and  Canada.) 

22.  A',  unifasciata  unifasciata  Guerin.     (Mexico.) 

23.  A',  u.  cochisiana  Hungerford.     (Arizona,  U.  S.  A.) 

24.  A',  u.  angulala  Hungerford.     (South  America.) 

25.  N.  u.  andersoni  Hungerford.    (Western  U.  S.  A.  and  B.  C.  Canada.) 

26.  y.  spinosa  Hungerford.   (Northwestern  U.  S.  A.  and  B.  C.  Canada.) 

27.  A',  kiangsis  Kirkaldy.     (China  and  Eastern  Siberia.) 

28.  N.  uhleri  Kirkaldy.     (Southeastern  U.  S.  A.) 

29.  A',  kirkaldyi  Martin.     (China.) 

30.  X.  saramoa  Esaki.    (Formosa.) 

31.  X.  chinensis  Fallou.     (China.) 

32.  A',  immediata  Kiritshenko.     (Eastern  Siberia.) 

33.  A",  triguttata  Motschulsky.    (Japan.) 

34.  A',  insulata  Kirb3^     (Eastern  U.  S.  A.  and  Canada.) 

35.  X.  kirbyi  Hungerford.     (Western  U.  S.  A.  and  Canada.) 

36.  A',  impressa  Fieber.    (Mexico.) 

37.  A',  viontezuma  Kirkaldy.     (Mexico  and  Te.xas,  U.  S.  A.) 

Subgenus  Bichromonecta  Hungerford 

38.  X .  melaena  Kirkaldy.     (Mexico.) 

39.  X.  ochrothoe  Kirkaldy.     (Southern  California  to  Colombia,  S.  A.) 


146  The  University  Science  Bulletin 

40.  N.  colomhiaita  Hungerford.     (Colombia,  S.  A.) 

41.  N.  re-panda  Hungerford.     (Arizona  to  Mexico.) 

42.  N.  shooteri  Uhler.     (California.) 

Subgenus  Ehytheonecta  Hungerford 

43.  N.  hojfmanni  Hungerford.     (California,  Arizona  and  Lower 

California.) 

44.  N.  ceres  ceres  Kirkaldy.    (Costa  Rica.) 

45.  A^.  c.  slirtoni  Hungerford.     (Mexico.) 

46.  N.  c.  rogersi  Hungerford.     (Costa  Rica.) 

47.  A^  compacta  Hungerford.     (Mexico.) 

48.  A^.  robusta  Hungerford.     (Mexico.) 

49.  A^.  mexicana  Amyot  and  Serville.     (Mexico.) 

50.  N.  VI.  creaseri  Hungerford.     (Mexico.) 

51.  N.  hintoni  Hungerford.     (Mexico.) 

52.  N.  lobata  Hungerford.     (New  Mexico,  Texas,  Mexico.) 

Subgenus  Notoxect.-v  Linnseus 

53.  X.  maculata  Fabricius.     (Europe,  North  Africa  east  to  India.) 

54.  N.  canariensis  Kirkaldy.     (Canary  Islands.) 

55.  A'^.  pallidula  Poisson.    (Algeria  and  Morocco.) 

56.  A^.  meinertzhageni  Poisson.     (Hoggar  Mts.,  Sahara,  Africa.) 

57.  A^.  arabiensis  Hungerford.     (Arabia.) 

58.  A",  lutea  Miiller.     (Northern  Europe.  Siberia.) 

59.  A^.  borcalis  Bueno  and  Hussey.    (Northern  North  America.) 

60.  N.  reuteri  Hungerford.     (Northern  Europe.) 

61.  A',  viridis  viridis  Delcourt.     (France  and  England.' 

62.  N .  V.  mediterranea  Hutchinson.     (About  Mediterranian  and 

eastward  to  India.) 

63.  A".  V.  kashmiriana  Hungerford.     (Kashmir.) 

64.  N .  gUmca  gla^ica  Linnaeus.     (Europe.) 

65.  A^.  g.  poissoni  Himgerford.     (Asia  Minor.) 

66.  A",  g.  hybrida  Poisson.     (Southern  France  to  North  Africa.) 

67.  A-",  g.  rujesccns  Poisson.     (Guernsey  Islands.) 

68.  N.  g.  fidva  Fuente.     (Iberian  Peninsula.) 

69.  A'^.  g.  kirvillei  Poisson.    (Asia  Minor.) 

70.  A^  nigra  Fieber.    (Brazil.) 

71.  N.  obliqua  obliqua  Gallen.     (Northern  and  Middle  Palearctic.) 

72.  A^.  o.  meridionalis  Poisson.     (About  the  Mediterranian.) 

73.  A^  o.  delcourti  Poisson.     (Belgium  and  Northwest  France.) 

74.  N .  amplifica  Kiritshenko.     (Southeastern  Siberia.) 

75.  A^.  violacea  Kirkaldy.     (Burma.) 

76.  A',  montandoni  Kirkaldy.     (China.) 

77.  N .  irrorata  Uhler.     (Eastern  North  America.)  » 

Subgenus  Enitharonecta  Hungerford 

78.  A',  handlirschi  Kirkaldy.     (Australia.) 


Hungerford:    Gexus  Notoxecta  147 

ACKNOWLEDGMENTS 

Through  the  years  during  which  I  have  been  interested  in  the 
study  of  the  Notonecta  I  have  received  assistance  from  many 
sources  and  in  various  ways.  Without  the  splendid  cooperation 
of  many  friends  the  completion  of  this  task  would  have  been  im- 
possible.    My  gratitude  is  therefore  expressed: 

To  the  graduate  Research  Conmiittee  of  the  University  of  Kansas 
for  financial  assistance. 

To  Dr.  Kathleen  Docring,  who  made  the  color  illustrations  and 
other  drawings  for  this  paper. 

To  Doctors  L.  0.  Howard  and  J.  M.  Aldrich,  who  made  it  possible 
for  me  to  spend  several  weeks  at  the  U.  S.  National  Museum  in 
1922,  at  which  time  I  studied  the  Notonecta  there. 

To  the  following  men  of  my  own  country  M-ho  have  given  me  the 
privilege  of  studying  the  collections  in  their  charge:  Wm.  J.  Baerg, 
Fayetteville,  Ark. ;  Nathan  Banks,  Museum  of  Comparative  Zoology, 
Harvard;  H.  G.  Barber,  hemipterist  at  U.  S.  National  ]\luseum; 
J.  C.  Bradley,  Cornell  University;  T.  H.  Frison,  Illinois  State 
Natural  History^  Survey;  F.  M.  Ciaige,  University  of  Michigan; 
Wm.  J.  Gerhardt,  Field  Museum,  Chicago;  Hugo  Kahl,  Carnegie 
]\Iuseum,  Pittsburg;  George  F.  Knowlton,  Utah  State  Agricultural 
College;  Frank  Lutz,  American  Museum  of  Natural  History,  New 
York  City;  H.  C.  Severin,  Brookings,  S.  Dak.;  Franklin  Sherman, 
Clemson  College,  South  Carolina;  G.  S.  Walley,  Ottawa,  Canada. 

To  J.  R.  de  la  Torre-Bueno,  who  has  the  largest  private  collection 
of  Notonecta  in  this  country,  who  has  published  on  Notonecta,  and 
whose  splendid  hospitality  has  made  me  welcome  in  his  home  on 
numerous  occasions. 

To  the  following  who  have  helped  me  with  the  loan  of  material 
from  their  private  collections:  Doctors  Herbert  Osborn,  C.  J.  Drake. 
H.  M.  Harris  and  C.  T.  Dodds. 

To  the  following  colleagues,  who  have  given  me  specimens  in  ex- 
change for  determination  of  material  used  in  studies  they  were 
making:  Doctors  J.  G.  Needham  and  C.  T.  Brues.  Also  the  fol- 
lowing who  made  collections  at  my  request:  H.  Garman,  G.  M. 
Bentley,  R.  M.  DeCoursey  and  D.  K.  Duncan. 

To  my  colleagues  in  this  university  who  have  added  specimens 
for  my  studies:  Doctors  R.  H.  Beamcr,  P.  B.  Lawson,  P.  A.  Readio 
and  R.  C.  Moore. 

To  the  following  former  students  who  have  remembered  my  in- 
terest in  Notonecta  and  who  have  sent  me  specimens  from  manv 


148  The  University  Science  Bulletin 

different  places:  L.  D.  Anderson.  C.  0.  Bare,  E.  M.  Becton,  E.  P. 
Breakey,  C.  J.  D.  Brown,  W.  J.  Brown,  George  Cady,  H.  W.  Capps, 
Carl  Cummings,  P.  A.  Glick,  G.  E.  Gould,  W.  E.  Hoffmann,  Clarence 
Hoffmann,  Dwight  Isely,  A.  M.  James,  Jean  Lindsdale,  C.  H. 
Martin,  J.  0.  Nottingham,  Paul  Oman,  Leonard  Tuthill,  Grace 
Wiley,  Laurence  Woodruff. 

To  the  following,  who  by  their  kindness  in  sending  me  specimens 
from  foreign  countries  have  aided  me  immeasurably  in  these  studies: 
Ernest  de  Bergevin,  P.  J.  BeiTnudez,  A.  Bruck,  E.  A.  Butler,  P.  W. 
Claassen,  Ed  Creaser,  A.  Dampf,  C.  Deane,  J.  B.  Edwards,  T. 
Esaki,  A.  Faz,  A.  C.  Jensen-Haarup,  H.  Hacker,  Herbert  M.  Hale, 
H.  E.  Hinton,  Ruby  Hosford,  R.  W.  Jack,  A.  Kiritshenko,  H. 
Leuderwaldt,  H.  Lindberg.  0.  Lundblad,  Harold  Morrison,  J.  G. 
IVIyers,  Stuart  Pennington,  Lilly  G.  Perkins,  F.  W\  Petty,  C.  S. 
Reed,  F.  Schade,  Heinrich  Schmidt,  Hobart  Smith,  R.  Spitz,  R. 
Stirton,  E.  Suenson,  E.  D.  Townsend,  B.  Vereertbrugghen,  Oscar 
Whittaker,  Chenfu  F.  Wu. 

To  Dr.  R.  Poisson,  of  Rennes,  Franc^,  who  has  been  a  most  helpful 
correspondent  and  who  is  much  interested  in  the  Notonecta  of 
Europe. 

To  Prof.  Teiso  Esaki,  of  Fukuoka,  Japan,  who  so  generously 
placed  at  my  disposal  before  its  publication  his  manuscript  on  the 
Notonecta  of  Europe,  and  whose  researches  on  the  synonomy  of 
this  group  I  have  so  largely  followed. 

To  Dr.  Hem  Singh  Pruthi  for  the  opportunity  of  studying  the 
undetermined  Notonecta  of  the  Indian  Biological  Survey. 

To  the  European  entomologists,  who  received  me  so  kindly  upon 
my  visit  to  their  museums,  who  placed  every  facility  of  their  labora- 
tories at  my  disposal  and  made  possible  the  examination  of  the 
types  and  other  historical  material  in  their  charge.  Major  E.  E. 
Austen,  custodian  of  the  insect  collections  at  the  British  Museum 
of  Natural  History,  and  Mr.  W.  E.  China,  directly  in  charge  of  the 
extensive  collections  of  Hemiptera,  extended  every  courtesy.  No 
man  could  have  been  more  generous  with  his  time  or  more  interested 
in  helping  me  with  the  studies  in  which  I  was  engaged  than  Mr. 
China.  This  fact  can  be  appreciated  by  students  of  the  Hemiptera 
and  Homoptera  the  world  over  who  have  had  occasion  to  enlist  his 
help  concerning  types  in  his  charge. 

Dr.  E.  P.  Poulton,  at  Oxford,  most  obligingly  made  it  possible  for 
me  to  complete  my  studies  there  in  the  very  brief  time  at  my  dis- 
posal. 


Huxgerford:    Genus  Notonecta  149 

Dr.  E.  L.  Bouvier,  at  that  time  in  charge  of  the  insect  collections 
at  the  Paris  Museum,  was  most  cordial,  and  the  help  given  me  by 
Doctor  Berland  and  Dr.  E.  Seguy  during  the  two  montiis  I  worked 
there  will  always  be  appreciated. 

Dr.  A.  Handlirsch,  of  the  Museum  in  Vienna,  and  Dr.  F.  Maidl 
most  cordially  made  arrangements  that  placed  the  collections  as 
much  at  my  disposal  as  those  in  my  own  department — a  privilege 
most  gratefully  acknowledged. 

Dr.  Geza  Horvath,  at  the  National  INIuseum  of  Hungary,  at 
Budapest,  with  whom  I  have  enjoyed  many  years  of  correspondence, 
made  my  studies  in  his  laboratory  most  profitable.  Because  of 
Doctor  Horvath's  assiduous  studies  in  the  Hemiptera  during  a 
verv^  long  and  productive  career  h.is  museum  must  be  considered 
one  of  the  most  important  centers  in  the  world  for  the  hemipterist. 
I  feel  under  unusual  obligation  to  him  and  to  Mr.  Josef  Ujhelyi,  his 
assistant,  who  spent  many,  many  hours  overtime,  at  the  expense  of 
another  task,  that  I  might  have  more  opportunity  to  study  than 
the  official  hours  of  the  museum  would  have  permitted. 

Dr.  Walther  Horn,  another  friend  of  many  years'  standing,  made 
my  visits  to  his  institute  in  Berlin-Dahlem  very  pleasant  ones.  His 
knowledge  of  early-day  entomologists,  the  location  of  their  collec- 
tions and  the  character  of  pins  and  labels  has  been  a  great  help 
to  me  in  checking  the  authenticity  of  types.  I  am  indebted  to  Di. 
H.  Bischoff.  custodian  of  the  entomological  collections  at  the  Berlin 
Museum,  for  the  pri^'ilege  of  studying  the  collections  there.  Dr. 
Willy  Ramme,  in  charge  of  the  Hemiptera,  and  Dr.  H.  Hedicke  con- 
tributed greatly  to  my  convenience  and  comfort  while  working  in 
their  museum. 

Dr.  Ludwig  Reh  and  Mr.  Eduard  Wagner,  at  the  Hamburg  Mu- 
seum, were  very  cordial  and  astonished  me  with  the  large  collections 
of  undetermined  material  in  their  charge. 

Dr.  0.  ^chroeder,  at  the  Zoological  Museum  of  Kiel  University, 
and  Dr.  L.  Bruel  and  Dr.  Veit  Ziehen,  at  Halle,  made  my  brief 
studies  at  their  institutions  profitable. 

At  the  University  Zoological  Museum  in  Copenhagen  I  found 
much  historical  material  of  interest  to  me,  and  am  pleased  to  ac- 
knowledge the  kind  assistance  of  Doctors  Wm.  Lundbeck  and  K. 
Stephensen. 

Dr.  S.  Bengtsson  and  Mr.  Ossian  Larsen,  at  Lund  University,  gave 
me  the  opportunity  of  examining  the  collections  in  their  charge, 
and  the  latter  most  kindly  showed  mc  his  experimental  work  on 

11—3482 


150  The  University  Science  Bulletin 

some  aquatic  Hemiptera — a  line  of  investigation  that  has  proved 
most  fruitful  since  that  time. 

Dr.  Y.  Sjostedt,  in  charge  of  the  entomological  section  of  the 
Royal  ]\Iuseum  at  Stockholm,  made  it  possible  for  me  to  spend  long 
hours  with  the  valuable  collections  studied  by  Stal  and  others,  and 
which  are  now  located  in  magnificient  quarters.  Dr.  A.  Roman 
was  also  very  cordial.  Dr.  0.  Lundblad,  whose  splendid  studies 
on  the  aquatic  Hemiptera  have  been  enriching  our  literature  for 
many  years,  was  exceedingly  kind,  and  our  many  hours  together  will 
always  remain  a  pleasant  memory.  The  hospitality  of  his  family 
and  his  own  sincere  desire  to  assist  me  made  possible  a  deligthful 
trip  to  Upsala,  his  own  alma  mater,  where  I  saw  not  only  the  trays 
of  insects  studied  by  the  great  Linnaeus,  but  the  gardens  and  the 
country  home  of  the  great  savant  as  well. 

xA.t  Brussels  Dr.  G.  Severin  welcomed  me  and  Dr.  Antoine  Ball 
not  only  placed  at  my  disposal  the  collections  of  the  Royal  Museum 
of  Natural  History,  but  took  me  to  Tervueren  to  see  the  Museum 
of  the  Belgian  Congo,  the  insect  collections  of  which  are  under  the 
direction  of  Dr.  H.  Schouteden. 

It  will  always  be  a  matter  of  sincere  regret  that  I  could  not  accept 
the  cordial  invitations  of  Doctor  Lindberg,  of  Helsingfors,  and 
Doctor  Jaczewski,  of  Warsaw,  to  study  the  rich  collections  in  their 
charge. 


HrNGERFORD:    Genus  Notoxecta  151 

BIBLIOGRAPHY  ON  NOTONECTA 

(Other  references  cited   under  species) 

1.  Alverdes,  F.     1925.     Notonecta  nach  einseitger  Blendung  in:    Zs.   Wiss. 

Zool.  Leipzig,  CXXIV,  pp.  382-405. 

2.  AsHTON,  R.  I.     1837.    On  the  Wings  of  Hemiptera  m :    Trans.  Ento.  Soc. 

London,  III,  p.  95  (Union  of  wings  of  Notonecta). 

3.  B.'VRE,  Clarence  O.    1926.    Life  Histories  of  Some  Kansas  Backswimmers 

in:  Annals  Ento.  Soc.  Amer.,  XIX,  pp.  93-101. 

4.  Bare,  Clarence  O.     1928.     Haemoglobin  Cells  and  Other  Studies  of  the 

Genus  Buenoa  in:  Univ.  of  Kansas  Science  Bulletin,  XVIII,  pp.  265-349 

5.  Berlese,  a.    1909.    Gli  Insetti,  I,  p.  265,  fig.  304   (female  abdomen  of  A\ 

glauca  L.) ;  p.  299,  fig.  342  (female  genitalia  of  .V.  glauca  L.) ;  p.  322, 
fig.  384  (male  genitalia  of  A',  glauca  L.). 

6.  Blatchley,  W.  S.    1926.    Heteroptera   or  True   Bugs   of  Eastern   North 

America     .     .     .    Nature  Pub.  Co.,  Indianapolis,  1116  pages,  with  illus. 

7.  Bollweg,  Wilhelm.    1915.    Verhandlungen  des  Naturhistorischen  Vereins 

der  Prussischen  Rheinlande  und   Westfalens,   LXXI  Jahrgang.     1914. 
(Appeared  1915.) 

8.  Brocher,  Frank.    1909.    Sur  Forgaue  imlsatile  observe  dans  les  pattes  des 

Hemipteres  aquatique  in :  Annales  de  Biologie  Lacustre,  Bruxelles,  IV, 
pp.  33-41  (Notonecta).    (For  natural  history  of  Notonecta  see  pp.  9-32.) 

9.  Brocher,  Frank.     1913.    Researches  sur  la  respiration  des  insectes  aqua- 

tiques  adultes.    La  Notonecte  in:  Zool.  Jahrb.  Jena.  Abt.  f.  allg.  Zool., 
XXXIII,  pp.  225-234.     (See,  also,  Annales  Biol.  Lacustre  IV.) 

10.  Bueno,  J.  R.  DB  LA  Torre-.    1902.    Notonectidse  of  New  York  in:  Jl.  N.  Y. 

Ento.  Soc,  X,  pp.  230-236. 

11.  Bveno,  J.  R.  de  LA  Torre-.    1902.    Some  Preliminary  Notes  on  the  Early 

Stages  of  Notonecta  in:  Proceedings  of  New  York  Ento.  Soc,  X,  p.  250. 

12.  Bueno,  J.  R.  de  l.^  Torre-.     1905.     The   Genus  Notonecta  in   America 

North  of  Mexico  in:  Jl.  N.  Y.  Ento.  Soc,  XIII,  pp.  143-167. 

13.  Bueno,  J.  R.  de  la  Torre-.    1908.    Concerning  the  Notonectidae  and  Some 

Recent  Writings  on  Hemipterology  in:   Canadian  Entomologist,  XL. 
p.  211. 

14.  Bueno,  J.  R.  de  la  Torre-.     1923.    Family  Notonectidae  in:    Conn.  State 

Geol.  and  Nat.  Hist.  Survey  Bull.,  No.  34. 

15.  Browne,  E.  N.    1913.    Study  of  Male  Germ  Cells  in:  Jl.  of  Exp.  Zoology 

XIV,  pp.  61-122.    (Philadelphia.) 

16.  Broavne,  Ethel  Nicholson.    1916.    A  Comparative  Study  of  the  Chromo- 

somes of  Six  Species  of  Notonecta  in:   Jl.  of   Morpholog.y,   XXVII, 
pp.  119^162,  7  plates,  115  figures. 

17.  Champlain,  a.  B.     1923.     Fish-killing  Insects   in:    Nature   Mag.  II,  pp. 

119-120. 

18.  Delcourt,  a.    1907.    Quelques  ob.?ervations  sur  la  variabilite  de  Notonecta 

glauca  L.,  in:  Comptes  rendus  des  seances  de  la  societe  de  Biologie, 
LXIL  p.  11. 

19.  Delcourt,  A.    1907.    De  I'influence  de  la  temperature  sur  de  development 

des  Notonectes  in:  Compt.  R.  Assoc  franc  avanc  Sc,  XXX\'T,  p.  244. 
(Rheims.) 


152  The  University  Science  Bulletin 

20.  Delcourt,  a.     1907.     De  la  necessite  d'line  revision  des  Xotonectes  de 

France  in:  Feuille  des  jeunes  Naturalistes.  l^r  aout.  1907. 

21.  Delcovkt,    a.      1909.      Amixie   regionale    chez    Notonecta    glauca    L..    in: 

Comptes  rendus  de  la  societe  de  Biologie,  LXVI,  p.  589. 

22.  Delcourt,  A.    1909.    Recherches  sur  la  variabilite  dii  Genre  "Notonecta," 

Bulletin  Scientifique  Fr.  et  Belg.  (7)  XLIII,  pp.  373-461. 

23.  DeichmI'-ller,  J.  V.   1886.  Eichstatt  in  Bayern.    Lithogr.  Kalk.  Malm,  in : 

Ins.  lith.  Sch.  Dresd.,  p.  64,  t.  5,  f.  7. 

24.  Despax,  R.   ■  1922.     Notonecta  maculata  F.  et  Notonecta  glauca  L.,  in : 

Bull.  Soc.  Hist.  Nat.  Toulouse,  pp.  97-112. 

25.  Edwards,  James.     1918.     On  the  British  Species  of  Notonecta  in:    Ento. 

Mo.  Mag.,  LIV,  pp.  56-59. 

26.  Embody,  G.  C.     1912.     Preliminary  Study  of  the  Distribution,  Food  and 

Reproductive   Capacity  of  Some   Freshwater  Amphipods  in:    Interna- 
tionale Revue  der  gesamten  Hydrobiologie. 

27.  Enock,  Fred.     1899.     Oviposition   of   Nepa   cinerea   and    Notonecta    in: 

Proc.  Ento.  Soc.  London,  p.  XIV.     (In  Trans.) 

28.  ESAKI,  T.,  and  Chin.\,  W.  E.     1928.     A  Monograph  of  the  Helotrephidte, 

Subfamily  Helotrephina;  in:  Eos,  Madrid,  IV,  pp.  165-168. 

29.  Essenberg,  Christine.     1915.    Habits  and  Natural  History  of  the  Back- 

swimmers  in:  Jl.  Animal  Behavior.    Camb.  V,  p.  381. 

30.  Fieber,  Franz  Xav.  1851.  Rhynchotographieen  Abh.  K.  Bohm.  Ges.  Wiss., 

V.  Pt.  7.     (Die  Gruppe  der  Notonectse,  pp.  469-486.) 

31.  Findler,    W.      1922.      Experimentelle    Erzegung    von    Pigmentierung    und 

Zeichnung  der  Fliigeldecken  am  Riickenschwimmer  (Notonecta  glauca), 
in :    Anz.  Ak.  Wiss.  Wien.  LIX,  p.  208. 

32.  Hale,  Herbert  M.     1923.    Studies  in  Australian  Aquatic  Hemiptera,  No. 

II,  In:  Rec.  S.  Aust.  Mus.,  Vol.  II,  No.  3. 

33.  Hale,  Herbert  M.    1924.    Two  New  Hemiptera  from  New  South  Wales  in: 

Proc.  Lin.  Soc.  New  South  Wales,  Vol.  XLIX,  Pt.  4. 

34.  Handlirsch,  Anton.     1906-1908.     Die   Fossilen   Insekten   und   die    Phy- 

logenie  der  Rezenten  Formen.    Leipzig. 

35.  Holmes,  S.  J.     1904.    The  Reactions  of  Ranatra  to  Light  in  :   Jl.  Comp. 

Neurology  and  Psychology,  XV,  p.  232. 

36.  HOPPE,   Julian.      1912.      Die    Atmung    von    Notonecta    glauca,    in:    Zool. 

Jahrb.  (Jena)  Abt.  f.  allg.  Zool.,  XXXI.  pp.  189-244. 

37.  HrNGERFORD,  H.  B.     1918.     Concerning  the  Oviposition  of  Notonecta  in: 

Ento.  News,  XXIX,  pp.  241-245. 

38.  HuNGERFORD,   H.  B.     1919.     The   Biology   and   Ecology   of   Aquatic   and 

Semiaquatic  Hemiptera  in:  Kansas  ITniv.  Science  Bull.,  XI,  pp.  166-173 
and  179. 

39.  HrNGERFORD,  H.  B.     1919.    The  Male  Genitalia  as  Characters  of  Specific 

Value  in  Certain  Cryptocerata  in:  Kansas  Univ.  Science  Bulletin,  Vol. 
XI.  pp.  329-331.  2  pi. 

40.  HrNGERFORD,  H.  B.    1928.    Some  Recent  Studies  in  Aquatic  Hemiptera  in  : 

Annals  Ento.  Soc.  America,  XXI.  pp.  139-146. 

41.  Hutchinson,  G.  E.    1929.    A  Revision  of  the  Notonectidse  and  Corixidse 

of  South  Africa  in:  Annals  of  the  South  African  Museum,  Vol.  XXV, 
Pt.  3,  p.  359. 


Huxgerford:    Genus  Xotoxecta  153 

42.  Jaczewski,  Tadf.usz.     1926.     Notes   on   Some   West-African   Heteroptera 

in :    Annalibus   Zoologicis    Musei    Polonici    Historise    Naturalis,    T.    Y, 
zesz.  2,  IX,  1926,  pp.  62-106. 

43.  J.ACZEWSKi,  Tadeusz.     1928.     Xotonectidae   from   the  State   of  Parana  in: 

Annales  Musei  Zoologici  Polonici,  VII,  pp.  121-136. 

44.  Kerkis,  J.     1926.     Zur  Kenntnis  de.s   inneren   Geschlechtsapparates   der 

wasserbewohnender  Hemiptera-Heteroptera   in:   Revue  Russe  d'Ento., 
XX,  p.  300. 

45.  KiRK.ALDY,  G.  W.     1897.     Revision  of  the  Xotonectidae,  Pt.  I.  in:   Trans. 

Ento.  Soc.  London,  pp.  391-426. 

46.  KiRKALDY,   G.   W.     1898.     An   Economic  Use   for   Water  Bugs  in :    Ento. 

Monthly  Mag.  2d  Ser.,  IX,  p.  173. 

47.  KiRKALDY,  G.  W.     1904.    Uber  Notonectiden  in:  Wiener  Entomologische 

Zeitung,  XXIII,  pp.  93-135. 

48.  KiRK.'VLDY,  G.  W..  and  Bueno,  J.  R.  de  l.\  Torre-.    1909.    A  Catalogue  of 

American  Aquatic  and  Semiaquatic  Hemiptera  in :  Proceedings  of  the 
Entomological  Society  of  Washington,  X,  pp.  173-215. 

49.  L.^RSEN,  OssiAN.     1930.    Biologische  Beobachtungen  an  sch\Vedischen  Xo- 

tonecta-Arten  in:  Entomologisk  Tidskrift,  p.  219. 

50.  Larsen,  Ossian.     1930.    W^elche  Bedeutung  hat  ein  A'crlust  des  Flugver- 

mogens  f iir  die  lokale  Verbreitung  der  Wasserhemipteren  in :  Zeitschrift 
fiir  wissenschaftliche  Insektenbiologie,  XXV,  p.  173. 

51.  Le.\ch,  W.  E.    1817.    Classification  of  Notonectides  in:  Trans.  Linn.  Soc. 

London,  XII,  pp.  11,  13. 

52.  Lehmann,  Hans.    1923.    Biologische  Beobachtungen  an  Notonecta  glauca 

L.  in:  Zool.  Jahrb.  Abt.  Syst.,  XLVI,  pp.  121-158,  figs.  A-C,  Pi.  5  and  6. 

53.  LrxuBLAD,   O.     1916.     Orthoptera,   Odonata   och    Hemiptera    Hetero'i)tera 

in:  S,ion  Takerns  Fauna  och  Flora,  II,  p.  11. 

54.  M.\iN,  James.    1837.    Habits  of  the  Boatfly  (Xotonccin  giauca).  in:  Mag. 

Nat.  Hist.,  new  ser.,  I,  pp.  557-558. 

55.  Xeedham,  James  G.,  1903.     Aquatic  Insects  in  New  York  State  //( :  New 

York    State    Mus.    Bull.,    LXVIII.      (Notonecta    eaten    by    dragon-fly 
naiad.) 

56.  OsHANix,  B.  F..   1906-1909.     ^'erzcichnis  der  Pakcarktischen   Hemipteren 

.     .     .    Svo,  3  vols.,  St.  Petersburg. 

57.  OsHANiN,  B.  F.     1912.     Katalog  der  paliiarktischen  Hemipteren     .     .     . 

in :  8vo,  Berlin. 

58.  Pantel.  J.,  and  Sinety,  R.     1902.     Sur  revolution  de  la  spermatide  chez 

le  Notonecta  glauca,  in:  Compt.  R.  Acad.  Sc  Paris,  Tome  135.  pp.  997- 
1000. 

59.  P.-iNTEL,  J.,  and  Sinety,  R.     1902.     Sur  revolution  de  Tacrosome  dans  la 

spermatidae  du  Notonecta  in:   Compt.  R.  Acad.  Sc.  Paris,  Tome   135, 
pp.  1124-1126. 

60.  Pantel,  J.,  and   Sinety,   R.     1902.     Sur  I'origine   du    Nebenkern   ct  les 

mouvements  nucleiniens  dans  la  spermatide  du  Xotonecta  glauca,  in: 
Compt.  R.  Acad.  Sc.  Paris,  Tome  135.  pp.  1359-1362. 

61.  Pantel,  J.,  and  Sinety,  R.     1906.    Les  cellules  de  la  lignee  male  chez  le 

Notonecta   glauca   L.   avec   des   details   plus    etendus    sur    la    periode 


154  The  University  Science  Bulletin 

d'accroissement  et  sur  celle  de  transformation  m :  La  Cellule,  XXIII, 
pp.  89-303. 

62.  Pettit,  a.,  and  Krohn,  A.     1905.    Sur  la  structure  de  la  glande  salivaire 

du  Notonecta  in:  Arch.  Anat.  Micr.,  VII,  pp.  351-368. 

63.  Pettit,  a.,  and  Krohn,  A.    1905.    Sur  revolution  des  cellules  des  glandes 

salivaires  du  Notonecta  glauca  L.,  in:  Bn.  Mus.  Paris,  pp.  113-114. 

64.  PoissoN,  R.    1923.    Les  Notonectes  de  la  region  Normande  in:  Bull.  Soc. 

Ento.  France,  Paris,  pp.  299-301. 

65.  PoissoN,  R.  ■  1924.    Contribution  a  I'Etude  des  Hemipteres  Aquaticiues  in : 

Bull.  Biol,  de  la  France  et  de  la  Belgique,  LVIII,  Fa>c.  1. 

66.  PoissoN,  R.     1925.     Hemipteres  aquatiques  d'Asie  Mineure  recueillis  par 

M.  H.  Gadeau  de  Kerville  en  1912,  Remarques  sur  les  Notonectes,  in: 
Bull,  de  la  Societe  entomologique  de  France,  pp.  328-330. 

67.  PoissoN,    R.     1925.     Quelques   Hemipteres    aquatiques    des    en\irons    de 

Banj'uls.  in:  Bull,  de  la  Soc.  Ento.  France,  Annee  1925,  No.  16. 

68.  PoissoN,  R.     1926.     L'Anisops  producta  Fieb.  (Hemiptere  Notonectidae) , 

Observations  sur  son  anatomie  et  sa  biologie,  in:  Archives  de  Zool. 
Exp.  et  Gen.,  Tome  65,  Fasc.  4,  pp.  182-208. 

69.  PoissoN,  Raymond.    1927.    Recherches  sur  Quelques  Processus  Spermato- 

genetique  .  .  .  de  Notonecta  maculata  Fabr.  in:  Archives  de  Zool. 
Exp.  et  Gen.,  LXVI,  p.  23. 

70.  PuTox,  A.     1899-1900.     Catalogue  des  Hemip.  fauna  paleartic  in:   Revue 

d'Entomologie,  XVIII,  4th  Ed.,  p.  80. 

71.  Redfern,  p.     1859.    On  the  method  of  production  of  Sound  by  a  species 

of  Notonecta  in:  Rept.  of  the  29th  meeting  of  British  Assn.  for  Adv. 
Sci.  Trans.,  p.  173. 

72.  REGiMB.'iRT,  M.     1875.     Observations  sur  la  ponte  du  Dytiscus  marginalis 

et  de  Quelques  autres  Insectes  Aquatique  in:  Ann.  Soc.  Ento.  Fr.  (5), 
V,  p.  204. 

73.  Rich,  S.  G.     1918.     The  Respiratory  Organs  of  a  Notonectid  in:   South 

African  JI.  Sci.,  Cape  Town. 

74.  ScHLECHTEKDAL,  D.  VON.     1892.    Ueber  das  Vorkommon  fossilen  "Riicken- 

schwimmer"  in:  Zeit.  fur  Naturwissenschaften,  LXV,  pp.  141-143. 

75.  ScHLECHTENDAL,  D.  VON.   1894.  Rott  im  Siebengebirge,  Rheinlande.  Oberes 

Oligocan.  in:  Abh.  Halle.,  XX.  N.  Harnacki,  pp.  25-26,  Taf.  14,  fig.  1. 
N.  DeichmiXUeri,  pp.  27-30,  Taf.  14,  figs.  4-6.  A^.  juhata,  pp.  30-31,  Taf. 
13,  fig.  12;  Taf.  14,  figs.  7-10.  A^.  comata,  pp.  31-32,  Taf.  13,  figs.  10,  11; 
Taf.  14,  fig.  11. 

76.  ScuDDER,  S.  H.     1890.     Florissant,   Colorado.   Nordamerika.   Miocan.  in: 

Tert.  Ins.  346,  t.  22,  fig.  11. 

77.  Shepherd,  Forest.    1847.    On  the  Habits  of  N.  glauca,  in :  Sillimans  Amer. 

Jl.  of  Sci.  and  Arts  (2),  IV,  pp.  423-424.  (Remarkable  flight  of  an 
American  species  in  1846.)  (See,  also,  Ann.  &  Mag.  Nat.  Hist.  (2)  1, 
pp.  158-159,  1848.) 

78.  Simpson,  S.  Geo.     1848.     Quoted  by  Dr.  Spence  in:   Trans.  Ento.  Soc. 

London,  V.  Proc.  p.  36.  (A  flight  of  Notonecta,  near  headwaters  of 
Mississippi  River,  25  miles  long.) 


Hungerford:    Genus  Notonecta  155 

79.  Spinola,  Marchese  Maximilian.     1837.     Essais  sur  les  Genres  d'Insectes 

appartenants  a  I'Ordre  des  Hemipteres  Linn,  ou  Rhyngotes  Fabr.  a  la 
'  section  des  Heteropteres  Dufour.    S  vo.    Genes. 

80.  SpinoLlA,   Marchess   Maximilian.     1840.     Essai    sur   les   Insects   Hemip- 

teres RJiyngotes  ou  Heteropteres.    8  vo.    Paris. 

81.  SwiNTON,  A.  H.    1877.    On  the  Stridulation  in  the  Hemiptera  Heteroptera 

m:  Ento.  Monthly  Mag.,  XIV,  pp.  29-31. 

82.  Van  Duzee,  Edward  P.     1917.     Catalogue  of  the  Hemiptera  of  America 

Xorth  of  Mexico     .     .     .    in:   Univ.  of  California  Publ.  Tech.  Bull., 
Vol.  II,  pp.  I-XIV,  1-902. 

83.  Wefelscheid,  Heinreich.     1912.     Uber  die  Biologie  und  Anatomie  von 

Plea  in:  Zool.  Jahrb.  (Jena),  XXXII,  pp.  387-474. 

84.  Wesbnberg-Lund,   C.     1915.     Insektlivet   I    Ferske   Vande,    pp.    118-123. 

(Biology  of  A'',  glavca  and  A'',  lutea  Miiller.) 

ADDENDA 

(Incomplete  references  not  seen  by  me) 

Malek.    Biol.  Zbl.  L.,  p.  182.     (A  study  of  the  orientation  of  N.  gJauca  L.) 

1930? 
Kertesz  Rovart.     Lap.  XXIV,  p.   163.     (Experiments  on  feeding  habits   of 

A^.  glaiica  L.)     1930? 


N.  viresccns.  N.  peruviana. 

Text  Figure  8.     Male  genital  capsules  of  N.  virescens  Blanchard  and  A',  jyeruviar^a, 
new  species.     Drawn  by  Philip  Levereault. 


INDEX 

PAGE 

americana  Fabr.  1875  ( =  N.  indica  Linn.  1771) 113 

americana  Ashm.  1899  ( =  N.  lunata  Hung.  1926) 107 

americana  Kirk.  1897  ( =  .V.  uiiifasciata  Guer.  1857  in  part) 109 

americana  H.  S.  1853  ( =  .V.  undidata  Say  1832) 119 

americana  Kirk.  1897  (  =  A' .  fazi  Hung.  1930  in  part) 142 

amplifica  Kirit.  1930 55 

andersoni  Hung,  (new  subsp.  of  N.  unifasciata  Guer.) 110 

angulata  Hung,  (new  subsp.  of  N.  rmifascinta  Guer.) Ill 

arabiensis  Hung.  1926 44 

basalis  Fieb.  1851  (as  var.  of  A",  riigosa  Fieb.  1851) 85 

bergrothi  Esaki  1925  ( =  N.  kiangsis  Kirk.  1897) 33 

BiCHROMONECTA  Huug.  ucw  subg 24 

bircirca  Hung.  1926 137 

bicircoidea  Hung.  1928 136 

bicolor  Fieb.  1851  (as  var.  of  N.  rugosa  Fieb.  1851) 85 

bifasciata  Guer.  1844 129 

bifasciata  Kirk.  1897  (see  under  N.  polystoUfiina  Fieb.) 132 

bifasciata  Hung.  1926  (  =  A',  sellata  Fieb.  1851) 139 

bipunctata  Fieb.  1851  (as  var.  of  A",  polystolisma  Fieb.  1851) 132 

bivittata  Mats.  1905  ( =  A^  montandoni  Kirk.  1897) 37 

borealis  B.  &  H.  1923 95 

canariensis  Kirk.  1897 40 

ceres  Kirk.  1897 79 

charon  Kirk.  1897  (as  var.  of  N.  undulatn  Say) 119 

chinensis  Fallou  1887 31 

cochisiana  Hung,  (new  subsp.  of  A',  unifasciata  Guer.) 110 

colombiana  Hung,  (new  species) 128 

compacta  Hung.  1925 73 

confusa  Hung.  1930 130 

cordigera  Fieb.  1851  (as  var.  of  A",  rugosa  Fieb.  1851) 85 

creaseri  Hung,  (new  subsp.  of  A'^.  mexicana  A.  &  S.) 77 

delcourti  Poisson  1933  (as  var.  of  A',  glauca  Linn.) 54 

distincta  Hung.  1925  ( =  N.  melaena  Kirk.  1897) 97 

distinctoidea  Hung.  1930 124 

disturbata  Hung.  1926 140 

dorsalis  Fieb.  1851  (desc.  as  var.  A",  variabilis  Fieb.  1851) 141 

Enitharonecta  Hung,  (subgenus) 25 

Erythronecta  Hung,  (new  subgenus) 24 

fabricii  Fieb.  1851  (  =  A' .  ohliqua  Gallen  1787) 52 

fabricii  Fieb.  1851  (see  imder  A",  glauca  Linn.) 48 

fasciata  Fieb.  1851  (desc.  as  var.  of  A',  pouistolisma  Fieb.) 132 

fasciata  Fieb.  1861  (desc.  as  var.  of  A.  fabricii  Fieb.  1851) 52 

fazi  Hung.  1930 1-12 

fulva  Fuente  1897  (desc.  as  var.  of  N.  glauca  Linn.) 50 

furcata  Fabr.  1794  ( =  N.  ohliqua  Gallen  1787) 52 

fuscata  Spin.  1840  ( =  N.  ohliqua  Gallen  1787) 52 

(157) 


158  The  University  Science  Bulletin 

PAGE 

geala  Kirk.  1897  (desc.  as  var.  insulatn  Kirby  1837) 85 

glauca  Linn.  1758 46 

glauca  Shep.  1847  (probably  .V.  undulata  Say  1832) 119 

guttata  Fieb.  1851  (desc.  as  var.  N.  polysfolisma  Fieb.) 132 

hades  Kirk.  1897  ( =  N.  ceres  Kirk.  1897) 79 

halophila  Edwards  1918  ( =  A' .  viridis  Dele.  1909) 61 

handlirschi  Kirk.  1897 27 

hybrida  Poisson  1933  (desc.  as  subsp.  of  A^  glauca  Linn.) 50 

hintoni  Hung.  1934 72 

hoffmanni  Hung.  1925 68 

horvathi  Esaki  1927  ( =  A^.  pallidula  Poisson  1926) 39 

howardii  Bueno  1905  ( =  A^.  indica  Linn.  1771) 114 

immaculata  Fieb.  1851  (desc.  as  var.  A^.  polysfolisma  Fieb.) 132 

immediata  Kirit.  1930 35 

impressa  Fieb.  1851 81 

indica  Linn.  1771 113 

indica  Kirk.  1900  ( =  A^.  unifasciata  Guer.  1857) 109 

indicoidea  Hung.  1927 123 

insidata  Kirby  1837 84 

insulata  Uhler  1875  ( =  A^.  kirbyi  Hung.  1925  in  part) 88 

inversa  Xoualhier  (Ms.sp.?  see  Delcourt,^^  p.  406) 

irrorata  Uhler  1878 92 

kashniiriana  Hung.  (n.  subsp.  of  N'.  viridis  Dele.  1909) 65 

kiangsis  Kirk.  1897 33 

kirbyi  Hung.  1925 88 

kirkaldyi  Martin  1902 30 

kirvillei  Poisson  1934  (desc.  as  subsp.  of  N.  glauca  Linn.) 26 

klugii  Fieb.  1851  ( =  N.  mexicana  A.  &  S.  1843) 75 

klugii  Fieb.  1851  (included  A^.  ceres  Kirk.  1897  in  part) 79 

lactitans  Kirk.  1897 28 

latona  Kirk,  (listed  by  Van  Duzee  as  var.  A^.  undulata  Say) 117 

lobata  Hung.  1925 70 

lunata  Hung.  1926  (new  name  for  A'',  variabilis  of  American  authors) 107 

lutea  Muller  1776 56 

lutea  Bueno  1904  (=  N.  borealis  B.  &  H.  1923) 96 

macrocephala  Hung.  1925  ( =  A'',  mexicana  A.  &  S.  .1843) 75 

marginata  Aliiller  1776  (Esaki  says  probably  a  Corixid) 53 

marmorata  Spin.  1840  ( =  N.  mediterranea  Hutch.  1928?) 63 

marmorea  Fabr.  1803  ( =  A^.  maculata  Fabr.  1794) 42 

maculata  Fabr.  1794 41 

maculata  Fieb.  1851  (desc.  as  var.  of  A'',  variabilis  Fieb.  1851) 141 

mediterranea  Hutch.  1927  (desc.  as  var.  N.  viridis  Dele.) 63 

meinertzhageni  Poisson  1934 44 

melaena  Kirk.  1897 97 

melanota  L.  &  R.  1826  ( =  A",  obliqua  Gallen  1787) 52 

meridionalis  Poisson  1926  (desc.  as  subsp.  of  A^.  obliqua  Gallen) 54 

meridionalis  Hutch.  1927  (desc.  as  var.  of  A',  viridis  Dele.) 64 


Hungerford:    Gexus  Notoxecta  159 

PAGE 

mexicana  A.  &  S.  1843 75 

mexicana  Uhler  1884  ( =  .V.  lohata  Hung.  1925) 70 

mexicana  Kirk.  1897  (  =  .V.  compada  Hung.  1925  in  part) 73 

mexicana  Champ.  1901  ( =  A",  ceres  Kirk.  1897  in  part) 79 

minuta  Hung.  1926 138 

montandoni  Kirk.  1897 3^ 

montezunia  Kirk.  1897 82 

nigra  Heb.  1851 126 

NoTONECTA  as  subgenus 25 

obliqua  Gallen  1787 51 

ochrothoe  Kirk.  1897 99 

octopunctata  Goeze  1778  (uncertain) 50 

see  note  under  N.  giauca  hybrida  Poisson 50 

odara  Kirk.  1897  (desc.  as  var.  A' .  insulata  Kirby) 85 

pallida  Amyot  1848?  nom.  invalid 61 

pallidula  Poisson  1927 38 

pallipes  Leth.  1881  (see  under  A',  indica  Linn.) 114 

Paranecta  Hutch,  subgenus 24 

peruviana  Hung,  (new  species) • 134 

plagiata  Fieb.  1851  (desc.  as  var.  A',  rugosa  Fieb.) . .  .• 88 

poissoni  Hung,  (new  subsp.  of  A^.  giauca  Linn.) 49 

polystolisma  Fieb.  1851 132 

polystolisma  Berg.  1879  (see  under  A^.  sellata  Fieb.) 139 

prima  Schaffer  1766  nom.  invahd.  et  nudum,  (see  A",  giauca  Linn.) 48 

pulchra  Hung.  1926 131 

punctata  Fieb.  1851  ( =  N.  undulaia  Say  1832) 123 

raleighi  Bueno  1907 105 

repanda  Hung,  (new  species) 100 

reuteri  Hung.  1928  (new  name  for  A",  scutellaris  Renter) 59 

robusta  Hung.  1932 "8 

rufescens  Poisson  1933  (desc.  as  var.  of  A^  gLauca  Linn.) 51 

rugosa  Fieb.  1851  ( =  N.  insulata  Kirby  1837) 87 

saramoa  Esaki  1933 32 

schumacheri  Schirm.  1911  (desc.  as  var.  of  N.  lutea  Mtiller) 58 

scutellaris  Fieb.  1851  (desc.  as  var.  of  A'',  variabilis  Fieb.) 141 

scutellaris  Fieb.  1851  (see  under  A^.  indica  Linn.  1771) 113 

scutellaris  Renter  1886  ( =  A' .  reuteri  Hung.  1928) 59 

secunda  Schaffer  1766  (nom.  invalid  et  nud.) 50 

(see  note  under  A'',  octopunctata  Goeze) 50 

selene  Kirk.  1897  (desc.  as  var.  A",  undulata  Say) 117 

sellata  Fieb.  1851 139 

shooteri  Uhler  1894 101 

shooteri  Kirk.  1897  (see  under  N.  indicoidea  Hung.  1927) 123 

sinica  Walker  1873  ( =  A",  chijiensis  Fallou  1887) 31 

spatulata  Hung.  1926  (desc.  as  var.  A^.  polystolisma  Fieb.) 132 

spinosa  Hung.  1930 112 

stygica  Kirk.  1897  ( =  A' .  lactitans  Kirk.  1897) 29 

suensoni  Hung.  1925  ( =  A',  kiangsis  Kirk.  1897) 33 


160  The  University  Science  Bulletin 

PAGE 

teurca  Kirk.  1897  (,  =  A',  mdatna  Kirk.  1897) 97 

trigvittata  Motsch.  18G1 36 

tiimida  Gerinar  (fide  Fieber  1851)  (=  A',  hdea  MuUer  1776) 57 

uhleri  Kirk.  1897 103 

iiinbrina  Fieb.  1851  ( =  N.  maculata  Fabr.  1794) 42 

unicolor  H.  Sch.  1835  ( =  A^  lutea  Miiller  1776) 57 

unicolor  Fieb.  1851  (desc.  as  var.  A^.  variaJniis  Fieb.  1851) 141 

unifasciata  Guerin  1857 109 

imdulata  Say  1832 117 

undulata  Say  1832  (see  also  under  A^  indica  Linn.  1771) 113 

undiilata  Kirk,  and  B.  1909  (see  under  A' .  sellata  Fieb.  1851) 139 

variabilis  Fieb.  1851 141 

variabilis  Fieb.  1851  (see  under  A^.  indica  Linn.  1771) 114 

variabilis  Burm.  1861  (see  under  A'',  sellata  Fieb.  1851) 139 

variegata  L.  &  R.  1826  ( =  A^.  maculata  Fabr.  1794) 42 

vereertbruggheni  Hung.  1928 144 

violacea  Kirk.  1897 •45 

viridis  Dele.  1909 60 

virescens  Blanch.  1852 135 

virescens  Reed  1901  (see  .V.  Jazi  Hung.  1930) 142 


162  The  University  Science  Bulletin 

COLOR  PLATE  I 

(Magnification  approximately   X  3) 

.,  Western  Hemisphere  Notonecta 

i  IGURE 

1.  Notonecta  irrorata  Uhler.     Range:     Eastern  Canada  and  eastern  half  of 

United  States,  from  Canada  to  the  Gulf  and  westward  to  eastern  Kan- 
sas.    (See  page  92.) 

2.  Notonecta  indica  Linn.    Range :     Across  southern  United  States  south  of 

37°  latitude  and  southward  through  insular  America  (Greater  Antilles) 
and  thi-ough  Mexico  to  Colombia,  S.  A.     (See  page  113.) 

3.  Notonecta  nndulata  Say.    Range :    Across  southern  Canada  and  throughout 

United  States  from  coast  to  coast  and  south  to  northern  Mexico.    The 
most  common  species  in  the  United  States.     (See  page  117.) 

4.  Notonecta  borealis  Bueno  and  Hussey.     Range:     Canada  and  northern- 

most states  of  the  United  States.     Distribution  inadequately  known, 
but  a  Boreal  species..    (See  page  95.) 

5.  Notonecta   lunata   Hungerford.      Range:      Eastern    Canada    and    eastern 

United  States.    (See  page  107.) 

6.  Notonecta  shooteri  Uhler.     Range :     The  species  (lat.  sens.)  ranges  from 

California  through  Arizona  to  Mexico.     The  specimen  drawn  is  from 
Mexico,  D.  F.     (See  page  101.) 

7.  Notonecta  raleighi  Bueno.    Range:     Southeastern  states  of  United  States. 

(See  page  105.) 

8.  Notonecta  colombiana  Hungerford.     Range:     Described  from  Colombia, 

S.  A.,  previously  confused  with  N.  shooteri  Uhler.     (See  page  128.) 

9.  Notonecta  minuta  Hungerford.    Range:     Described  from  Proy.  del  Sara, 

Bolivia,  S.  A.     (See  page  138.) 

10.  Notonecta  pulchra  Hungerford.    Range:     This  is  a  very  common  species 

in  Paraguay,  S.  A.     (See  page  131.) 

11.  Noto?iecta  polystolisma  Fieber.     Range:     Common  in  the  region  of  Sao 

Paulo,  Brazil.     (See  page  132.) 

12.  Notonecta  unifasciata  Guerin.    Range:     (Lat.  sens.)     From  Mexico  across 

western  United  States  to  western  Canada.     (See  page  109.) 

13.  Notonecta  disturbata  Hungerford.     Range:     Known  only  from  "Brazil,"' 

but  doubtless  region  around  Sao  Paulo.     (See  page  140.) 

14.  Notonecta  sellata   Fieber.     Range :      Argentina,   Paraguay   and  southern 

Brazil.     (See  page  139.) 

15.  Notonecta  nigra  Fieher.    Range:    Described  from  Brazil,  but  doubtful.  See 

discussion  under  name.     (See  page  126.) 

16.  Notonecta  bicirca  Hungerford.    Range:     Chile,  S.  A.     A  common  species 

easily  confused  with  N.  bicircoidea  Hungerford  and  with  A',  vircscens 
Blanchard.     (See  page  137.) 

17.  Notonecta  fazi  Hungerford.     Range :      Very   common   in   Chile,   ranging 

northward  into  Peru.     (See  page  142.) 


HUNGERFORD — Notonecta — PLATE  I. 


0"ai 


N.  undulata   ? 


N.  ihootecii  vac.   rf 


N.  lunata   9 


N .  minuta  'S 


N.  iellata   cf 


14 


N.  raleighi   ? 


N.  bocealis    :f 


N .  colombiana 


a  ^ 


rW 


'2 

N.  unifasciata   rf 


A^.  polystoltsma  d' 


N.  nigra 


N.  fazi   r'    17 


Holland  Engraving  Co. 
Capper  Printing  Co. 


Katlileen  Doering 
Delineator 


164  The  University  Science  Bulletin 

COLOR  PLATE  II 

(Magnification  approximately   X  3) 

Western  Hemisphere  Notonecta 

IlGUItK 

1.  Xotonccla  montezuina  Kirkaldy.     Range:      Mexico  into  Texas,  U.  S.  A. 

(See  page  82.) 

2.  Notonecta  indicoidea  Hungerford.    Range:     Described  from  Mexco,  D.  F. 

(See  page  123.) 

3.  Notonecta   vereerthruggheni   Hungerford.     Range:      Described   from   Rio 

Negro,  territory  of  Argentina.     Compare  with  N.  fazi  Hungerford  on 
Plate  I.     (See  page  144.) 

4.  Notonecta  confma  Hungerford.    Range:     Known  only  by  the  type  which 

was  labeled  "S.  Amer."     (See  page  130.) 

5.  Notonecta  borealis  Bueno  and  Hussey.    Range :    See  Plate  I.  figure  4,  and 

this  plate,  figure  10.     A  stud^-  in  melanchroism.     (See  discussion  under 
this  species,  page  95.) 

6.  Notonecta  distinctoidea  Hungerford.    Range:     Known  only  from  Mexico. 

(See  page  124.) 

7.  Notonecta  spinosa  Hungerford.     Range:     Described  from  Utah  and  now 

known  from  Nevada,  Montana.  Oregon,  and  British  Columbia,  Canada. 
(See  page  112.) 

8.  Notonecta  bicircoidea  Hungerford.     Range:     Described  from  Rio  Negro, 

territory  of  Argentina.    Compare  with  N.  bicirca  Hungerford  on  Plate  I. 
(See  page  136.) 

9.  Notonecta  indica  Linn.    Range:     (See  Plate  I.)     This  specimen  has  color 

of  the  type  specimen,  and  until  carefully  studied  led  to  confusion  with 
Notonecta  unijasciata  Guerin.     (See  Plate  I,  figure  12,  and  page  113.) 

10.  Notonecta  Borealis  Bueno  and  Hussey.     (See  figure  5  and  page  95.) 

11.  Notonecta  virescens  Blanchard.     Specimen  from  Paris  Museum. 

12.  Notonecta  rogemi  Hungerford.    Now  considered  by  me  to  be  a  subspecies 

of  Notonecta  ceres  Kirkaldy.     (See  page  81.) 

13.  Notonecta  jnexicana  creaseri  Hungerford.    Range:    Known  only  from  type 

locality  in  Mexico.     (See  page  77.) 

14.  Notonecta  robusta  Hungerford.     Range:    Type  bears  no  label,  but  I  be- 

lieve it  comes  from  Chia;i)as.  Mexico.     (See  page  78.) 

15.  Notonecta   robusta?     Hungerford.     Range:     Female   specimen   fi'om   San 

Cristobal,  Chiapas,  Mexico,  taken  by  A  Dampf,  and  I  believe  it  to  be 
female  of  N.  robusta.     (See  page  78.) 


HUNGERFORD — Notonecta — PLATE    II. 


<njt 


C'Ti 


#^ 


I    N.  montezuma  ? 


2   N.  indicoidea  d     3'^    vereertbruggheni  cf  4  W.  confusa  d" 


#--^ 


7  W-  ipinosa  9 


5    N    boiealii  d 


8  N.  bicircoidea  9 


6  N.  distinctoidea  d 


9    W    mdica   9 


It    N    oitetcent    9 


12  N.  ro9««i  d  13   N.  m.  createti  d 


|4   N.  robusta  d 


15    N-  robusta f  9 


Holland    Engraving    Co. 
Grimcs-Joycc   Ptg.   Co. 


Kathleen    Doering 
Delineator 


12—3482 


166  The  University  Science  Bulletin 


COLOR  PLATE  III 

(Magnification  approximately   X  3) 

Pj^^jjg  Western  Hemisphere  Notonecta 

1.  Xolonecia  insulata  Kirby.     Range:     Eastern  Canada  and  United  States 

north  of  fortieth  degree  of  latitude  and  east  of  one  hundredth  meridian. 
(See  page  84.) 

2.  Notonecta  ceres  Kirkaldy.     Range:     A  common  species  in  Costa  Rica. 

The  female  shown  in  figure  3  was  named  as  A^.  mexicana  var.  ceres  and 
the  male  as  A^  mexicana  var.  hades  by  Kirkaldy.     (See  page  79.) 

3.  Notonecta  ceres  Kirkald.y.     (See  note  above.) 

4.  Notonecta   kirbyi   Hungerford.     Range:      Western   Canada    and   Western 

United  States.  Hemelytra  more  often  I'ed  and  black  ihan  like  the 
illustration.  (See  page  88.) 
.5.  Notonecta  lobata  Himgerford.  Range:  Foimd  in  Texas,  New  Mexico  and 
Arizona.  An  abundant  species  in  Arizona.  Have  seen  specimens 
labeled  "Mexico."  In  this  species  the  females  are  colored  as  in  figures 
5  and  10.  The  males  are  nearly  always  darker  (see  figures  11  and  13.) 
(See  page  70.) 

6.  Notonecta    compacta    Hungerford.      Range:    Known    from    the    states   of 

CiueiTero  and  Colima,  Mexico.     The  males  are  nearly  black  as  in  N. 
ceres  Kirkaldy  and  N.  lobata  Hungerford.     (See  page  73.) 

7.  Notonecta  melaena  Kirkaldy.    Range:     States  of  Jalisco,  Guanajuato  and 

Lower  California,  Mexico.    The  typical  color  is  darker  than  the  speci- 
men showTQ.     (See  page  97.) 

8.  Notonecta   mexicana  Amyot  and  Serville.     Range:     Mexico,  but  appears 

to  be  by  no  means  as  common  as  other  species  that  have  been  mistaken 
for  it,  N.  lobata  Hungerford,  for  example.     (See  page  75.) 

9.  Notonecta  uhlcri  Kirkaldy.    Range:     In  United  States  from  Massachusetts 

south  to  Mississippi.    Its  western  limit  appears  to  be  Illinois  and  south- 
ward, east  of  Mississippi  River.     (See  page  103.) 

10.  Notonecta  lobata  Hungerford.     (See  figure  5.)      A  typical   female.     (See 

page  70.) 

11.  Notonecta  lobata  Hungerford.     A   male,  showing  that  the  red  males  are 

darker  than  the  females.     (See  page  70.) 

12.  Notonecta  hoffmanni  Hungerford.    Range:     California  and  Arizona  in  the 

United  States  and  Lower  California  in  Mexico.     (See  page  68.) 

13.  Notonecta  lobata  Hungerford.     Range:     (See  figure  5.)     The  asual  color 

of  the  male,  and  showing  its  size  to  be  a  little  larger  than  the  female 
(.see  figure  10).     (See  page  70.) 


HUNGERFORD Notonecta PLATE   III. 


€>'ii 


eo 


N.  insulata   $ 


€f^ 


N.  lobata  ?         5 


N.  compacta    + 


N.  melaena  cf 


N.  uhleti  d" 


N.  kitbyi  9 


mexicana 


^    .0 

II 

N.  hoffrr 

ami 

■'   12 

N.  lobata    r'    13 

N.  lobata   ? 

N. 

lobata  d" 

IloUiind  F.iigruviiiif  Co. 

Kathleen  Doeiii 

Capper  Printing  Co. 

Delineator 

168  Thf.  University  Sciexce  Bulletin 


COLOR  PLATE  IV 

(Magnification  approximately   X  3) 

T,-,  .,,„^  Eastern  Hemispheke  Notonecta 

1.  XutoiKcta  inoiiUniiUiiii  Kirkaldy.     Range:     Tibet,  south  and  eastward  in 

China.     (See  page  37.) 

2.  Notonecta  handlirschi  Kirkaldy.     Range:     Known  only  from  West   Aus- 

tralia. Only  member  of  the  genus-  known  from  Australia.  (See  page 
27.) 

3.  Notonecta  lactitans  Kirkaldy.    Range:     South  Africa,  and  to  date  the  only 

member  of  the  genus  known  south  of  the  Sahaia.  (See  Plate-  \ ,  figure 
12,  for  the  male.)     (See  page  28.) 

4.  Noto)Hcta  ti-ig)iltala  Motschulsky.     Range:    Japan;  may  occur  in  eastern 

China,  but  not  confirmed  by  me.     (See  page  36.) 

5.  Notonecta  nmrmorca  marmoreu.     This  is  not  N.  m.  marmorea  Fabr.     It 

comes  from  Morocco  and  may  be  a  very  pale  specimen  of  A',  viridi^ 
mcditerraiica  Hutchinson,  but  I  really  believe  that  it  is  A'.  I'iridis  viiidis 
Delcourt.  A',  viridis  meditcrnnu  a  Hutchin.-on  typically  lia.s  more  nearly 
the  color  pattern  of  my  A'.  I'iridis  kashmiriana.     (Plate  V.) 

6.  Notonecta  obliqua  obliqna  Gallen.     Range:     Northern  and  middle  puhf- 

arctic.    A  well-known  Ein-opean  species.     (See  page  51.) 

7.  A"olo)iecta  lutea  Miiller.     Range:  Northern  Europe.     (See  page  56.) 

8.  Notonecta  canariensis  Kirkakhr    Range:    Canary  Islands.     (See  page  40.) 

9.  Notonecta  reiiteri  Huniierford.    Range:    Northern  Europe.     (See  page  59.) 

10.  Notonecta   glauca    Linnanis.      Range:      Widely    di.stributed   over   Europe. 

(See  page  46.) 

11.  Notonecta  violacrn  Kiik:il(iy.     Range:    Burma.     (See  page  45.) 


HUNGERFORD — Notonecta — PLATE   IV. 


N.  montandoni  cT 


N.  handlirschi  d' 


N.  lactttans    ? 


N.  tciguttata  ? 


€5^ 


N.  m,  marmorea  cf 


N.  o    obiiqua 


N.  lutea  d" 


N.  teuteci  d     9 


N.  glauca   d     10 


N .  canariensis    '^    ** 


lliiliiiiKl  Engraving  Co. 
(.'apper  Printing  Co. 


Katlileen  Doei'ing 
Delineator 


170  The  University  Science  Billetin 


COLOR  PLATE  V 

(Magnification  approximately   X  3) 

Eastern  Hemisphere  Notonecta 
FuiUKi: 

1.  Xolohecta  arabiensis  Himgerford.    Range:     Known  only  from  type  speci- 

mens from  Arabia.     (See  page  44.) 

2.  Notonecta   pollidula   Pois.son.     Range:      Algeria    and   Morocco   of   North 

Africa.     (See  page  38.) 

3.  Notonecta  glauca  -pohsoni  Himgerford.    This  is  one  of  the  specimens  men- 

tioned by  Kirkaldy  on  page  421  of  his  re^-ision.  It  comes  from  Mali — 
Nousky,  Erzerum.  There  are  eight  specimens  of  this  in  Vienna.  (Sec 
page  49.) 

4.  Xotomcta    amph'ficn   Kiritshenko.     Range:      Southeastern   Siberia.      (See 

page  55.) 

5.  Notonecta  obliqua    meridional  is  Poisson.     Range:      Around  the   Mediter- 

ranean.    (See  page  54.) 

6.  Notonecta  immediata   Kiritshenko.     Range:      Eastern  Siberia.     I  cannot 

distinguish  this  structurally  from  N.  tnguttata  Motsch.     (See  page  35.) 

7.  Notonecta  kiangsis  Kirkaldy.     Range:     Central  and  Northern  China  and 

Eastern  Siberia.    (See  page  33.) 

8.  Notonecta  chinensis  Fallou.    Range:     A  common  species  in  China.     (See 

page  31.) 

9.  Notonecta  kirkaldyi  Martin.     Range:    Described  from  Yun-Nan-Nansen. 

China.    (See  page  30.) 

10.  Notonecta  maculata  maculata  Fabricius.    Range:     Western  Europe,  south 

from  Denmark  to  North  Africa  and  eastward  to  Baluchistan. 

11.  Notonecta  glauca  julva  Fuente.     Range:    Not  well  established,  but  de- 

scribed from  Iberian  Peninsula.  The  specimen  shown  is  from  Guernsey 
Island.  This  form  needs  further  study  as  to  identity  and  relationships 
(See  page  50.) 

12.  Notonecta  lactitans  Kirkaldy.     Range:      (See  Plate  IV,  figure  3.)     This 

male  was  called  variety  stygica  by  Kirkaldy,  but  is  only  the  male  of  the 
above  species.     (See  page  28.) 

13.  Notonecta  viridis  kashmiriana  Hungevfovd.    Range:     Kashmir.     (Seepage 

65.) 

14.  Notonecta  m.  viridis  Delcourt  according  to  Esaki.    I  believe  should  be  N. 

viridi'i  jnridis  Delcoin-t.  Range:  England  and  France.  (See  page  60.) 
The  specimen  here  shown  is  an  unusually  highly  colored  specimen  from 
Gravesend,  England. 


HUNGERFORD Notonecta PLATE    V. 


2     N.  pallidula  d  .    .,      ,  o 

3    N.  glauca  vat.   V 


I    N.  arabiensit   9 


^yp    /.^^gi^ 


w 


4     N.  ampliUca  d 


6      N.  immediata  9 


7     W.  kiangsis   9 


8     N.  chinemis  d  9    ^r.  kickaldyi  d 


N.  o.  meridionati$  d 


12    N.  /actitans  cT 


10  II     N.  m.  fuloa  d 

N.  m.  maculata  9 


13  N.  kashmtriana  d     |4   W.  m.  viridis  d 


Holland    Engraving    Co. 
Crimes-Joyce  Ptg.  Co. 


Kathleen   Doering 
Delineator 


172  Thk  University  Science  Bulletin 


PLATE  VI 
This  plate  illustrates  the  genera  of  the  family  Xotonectidse. 

Figure 

1.  Notonecta.     Anterior  half  of  (hjrMua.     V^  vertex;  S  ^  synthlipsis;  Sc.  = 

scutellum. 

2.  Enithares.    Anterior  half  of  dorsum.     Note  that  the  anterolateral  margins 

of  prothorax  are  fo\eate,  indicated  in  drawing  by  F. 

3.  N.ychia.    Shows  the  foveate  prothorax. 

4.  Martarega.    Compare  with  figure  3.    The  genus  Martarega  is  from  Western 

Hemisphere,  while  Nychia  is  found  in  Africa. 

5.  Notonecta.     Middle  femur,  showing  the  anteapical  tooth  oi-  jiointed  pro- 

tuberance, characteristic  of  the  tribe  Notonectini. 

6.  Paranisops.     On  the  left  is  the  form  with  large  scutellum  anil  developed 

wings;  on  the  right  is  the  form  with  small  scutellum  and  reduced  meta- 
thoracic  wings.     Note  the  claval  orifice  just  behind  the  .>^cutellum. 

7.  Buenoa.    Anterior  half  of  dorsum  to  show  the  claval  orifice  (0)  just  behind 

the  scutellum.    In  the  key  this  is  called  the  hemelytral  commis.sure  pit. 


HuxtiERFORD:  (  texls  Xotonecta 
PLATE  VI 


17:5 


F^- 


1.    Notonecta. 


2.   Enithares. 


3      Nychia.  4.    Martarcga. 


5.    Notonecta    (middle  femur). 


6.     Paranisops. 


174  The  University  Science  Bulletin 


PLATE  VII 

Figure 

1.  Nychia.    Genital  capsule  of  male  viewed  from  the  left  side.    Note  that  the 

capsule  is  cleft  behind  and  that  the  claspers  or  parameres  are  unlike  on 
the  two  sides.  '  (After  Hutchinson.) 

2.  Martarega.    Side  view  of  the  head,  showing  the  four-segmented  antenna; 

the  terminal  segment  is  small.    Nychia  has  lost  the  terminal  segment. 

3.  Martarega.    Genital  capsule  of  the  male  viewed  from  the  left  side.    Note 

that  the  capsule  is  cleft  behind  and  that  the  claspers  are  unlike.    (After 
Jaczewski.) 

4.  Buenoa.    Genital  capsule  of  the  male  viewed  from  left  side.    The  capsule 

is  not  cleft  behind;  the  claspers  are  unlike. 

5.  Anisops.     Left   anterior  leg  of  male   (inside  or  anterior  view)   shows  the 

one-segmented  tarsus  and  stridular  area  (Str.)  on  tibia. 

6.  Buenoa.    Right  anterior  leg  of  male.    Shows  the  two-segmented  tarsus  and 

stridular  areas  (Str.)  on  femur  and  tibia. 


Hungerford:  Genus  Notonecta 


175 


PLATE  VH 


1.  Nychia 


ight  clasper 


2.  M 


artarega. 


5.  Anisops   (anterior  leg   $).        6.      Buenoa    (anterior  leg    $). 


176  The  I'xiversity  Science  Bulletin 


PLATE  Vni 

Figure 

1.  Eiiithares  bmsiUcims  Spinola.     Genital  capsule  of  nude  viewed  from  left 

side. 

2.  Eiiithares  brasiUeu.sLs  Spinola.     Genital  cap.sule  of  male  viewed  from  right 

side.    Note  the  differences  in  claspers  or  parameres. 

3.  Paranisops  incomtans  var.  liitea  Hale.     Male  genital  capsule  viewed  from 

left  side.    The  parameres  (P)  are  alike. 

4.  Xntonecta  handlirschi  Kirkaldy  (subg.  Enitharonecta.)     Male  genital  cap- 

sule \iewed  from  the  left  side.    The  parameres  ai'e  alike. 

5.  Xotonecta.    Head  and  pronotum  to  show  the  anterior  margin  of  vertex  for 

comparison  with  figure  8. 

6.  Buenoa.     Genital  capsule  viewed  from  right  side.     R.  C.  =  right   clasper 

L.  C.  =  left  clasper. 

7.  Buenoa.     Genital  capsule  viewed  from  the  loft  side.     Note  the  differences 

in  the  parameres  or  claspers.    L.  C.  =  left  clasper.   R.  C.  =  right  clasper. 

8.  Notonecta.     Head  and  pronotum — to  compare  shape  with  figure  5.     (Sec, 

also,  Plate  XVH,  figure  4.) 

9.  Xotonecta  lactitans  Kirkaldy   (subg.  Paranecta.)     Genital  capsule  viewed 

from  the  left  side. 


HuNCiERFORi):  (Iexus  Xotonecta 
PLATE  VI TT 


177 


1.  Enitbares  brasilicnsis  (left  side).  2.  Enithares  brasiliensis  (right  side). 


4.  N.  handlirschi  (subg.  Enitharonecta). 


3.  Paranisopsinconstansvar.  lutea 


6.  Buenoa---(nght  side).        7.  Bucno:i---(lcft  sid:). 


5.  Notonecta---head  and  pronotum. 


8.  Notonecta— head  and  pronotum.  9.  N.  lacf  itans  (subg.  Paranecta). 


178  The  University  Science  Bulletin 


PLATE  IX 

Figure 

1.  Notonecta    Dorsal  view: 

^^  =  vertex. 

S  ^  synthlipsis. 

A.  angle  ^  anterior  lateral  angle  of  the  pronotum. 

L.  angle  ^  lateral  angle  of  pi'onotuni. 

P.  angle  =  posterior  angle  of  pronotum. 

O.  Z.  =z  opaque  zone  of  membrane. 

C.  Z.  =  clear  zone  of  membrane. 

A.  L.  =  anterior  lobe  of  membrane. 

P.  L.  =  i)Osterior  lobe  of  membrane. 

2.  Xotonecta.     Male,   ventral   view,   showing   antenna    (A) ;    beak,   showing 

clypeus  (C);  the  three  pairs  of  legs,  showing  the  hook  on  the  anterior 
trochanters;    metaxyijhus  and  fourth  abdominal  sternite    (4.) 

3.  N.  undulata  Say.    Trochanter  of  male,  to  show  the  hook.    This  hook  may 

be  lacking  in  some  species,  represented  by  small  jiroturbance  or  greatly 
enlarged  as  shown  in  figure  4. 

4.  Notonecta  shooteri  Uhler.     Anterior  trochanter  of  male,  to  show  greatly 

enlai'ged  hook  and  basal  proturbance. 

5.  Middle  leg  of  a  Notonecta,  with  rounded  trochanter. 

6.  Trochanter   and   basal    part   of   femur   of   a    Notonecta.   to   show   spinose 

mesotrochanter. 

7.  Notonecta  handlirschi  Kirkal<I>-.     Middle  leg.  without  coxa. 

8.  Notonecta  lactitans  Kirkaldy.    Middle  leg. 

9.  Middle  leg  of  a  Notonecta  to  show  the  mesotrochanter  with  angle  pro- 

duced. 


HUN'GERFORD :    GeXUS    XoTOXECTA 

PLATE  IX 


179 


Tibia—  - 

Tarsus-  -! 

M.  femur 

Mesotrochanter^ 
Metaxyphus. — 


2.  Notonecta  (venter). 


Hook -+  — 


3.  N.  undulata  (male). 


4.  N.  shooterii  (male). 


c  __Mesotrochanter__L  _ 


6.  Mesotrochanter  spinose. 


5.  Mesotrochanter  rounded. 


8.  N.  lactitans. 


7.  N.  handlirschi. 


9.  Mesotrochanter  angle  produced. 


180  The  University  Science  Bulletin 


PLATE  X 

Figure 

1.  Xotuvecta  lobata  Hungerfonl.    Ventral  view  of  abdomen  of  female.    Note 

that  the  carina  of  the  fourtli  alidominal  sternite  (4th  St.)  is  bare,  that 
the  fifth  sternite  is  narrow  and  notched  at  tiji  and  that  the  last  ab- 
dominal sternite  (L.  St.) — tyjiical  for  X.  //u.r/ro»a  group — is  character- 
isticalh'  different  from  the  others. 

2.  Nutonecta   monlezuma  Kirkaldy.     Ventral   view   of  abdomen   of   female. 

Note  that  the  carina  is  bare  on  fourth,  fifth  and  most  of  sixth  sternites. 
The  last  sternite  is  intermediate  between  the  Notonecta  undulata  Say 
group  and  the  Noto)iecta  irrorata  Uhler  group. 

3.  Xotonecta  wididata  Say.    Ventral  view  of  abdomen  of  female.     Note  the 

notch  in  the  distal  end  of  the  fifth  sternite  and  the  notch  in  the  tip  of 
the  last  abdominal  sternite. 

4.  Xotoitecta  irrorata  Uhler.    Ventral  view  of  abdomen  of  female.    Note  that 

the  last,  abdominal  sternite  is  large  and  con.stricted  before  the  tip,  wliich 
is  not  notched. 


Hungerford:  Genus  Notonecta 
PLATE  X 


181 


N.  lobata  (abdominal  venter,  female). 


2.  N.  montezuma  (abdominal  venter,  female) 


.    •        /  i_j  ,1  .,„^.«,  fo™^l<.^       4.  N.  irrorata  (abdommal  venter,  female) 

N   undulata  (abdommal  venter,  tcmalej.  ^  ' 

13-3482 


182  The  University  Science  Bulletin 


PLATE  XI 

The  Notonecta  mcxicana  group.  Male  genital  capsules 
viewed  from  the  left  side.  Figures  4,  5,  8,  and  9  are  sketches 
of  the  tip  of  the  "internal  stay,"  which  in  this  group  is  large 
and  strongly  sclerotized. 


Hungerford:  Genl'S  Notonecta 


183 


PLATE  XI 


11.  N.m.  mexicana. 


10.  N.  robusta 


184  The  University  Science  Bulletin 


PLATE  XII 

The  male  genital  capsules  of  some  South  American  Noto- 
necta.    Views  from  the  left  side. 


Hungerford:  Genus  Notonecta 


185 


PLATE  XII 


7.  N.disturbata. 


8.  N.  pulchra. 


!186  'J'he  University  Science  Bulletin 


PLATE  XIII 

The  male  genital  capsules  of  some  American  Notonecta  be- 
longing to  the  subgenus  Paranecta. 


Hungerford:  Genls  Xotonecta 


187 


PLATE  XIII 


9.  N.  distinctoidci  10.  N.  uhleri 


11.  N.  mclaena. 


188  The  University  Science  Bulletin 


PLATE  XIV 

The  male  genital  capsules  of  Nolonecia  unijasciata  Guerin, 
its  saibspecies  and  close  relatives. 


Huxgerford:  Genus  Notonecta 
PLATE  XIV 


189 


1.  N.  unifasciata  (paratype). 


2.  N.  unifasciata. 


3.  N.  unifasciata. 


4.  N.  u.  angulata. 


5.  N.  u.  cochisiana. 


6.  N.  u.  andersoni. 


7.  N.  spinosa. 


8.  N.  indicoidea. 


190  The  University  Science  Bulletin 


PLATE  XV 

The  male  genital  capsules  of  some  Notonecta. 
Figures  1,  2  and  4  are   North  American  species  and   tlit; 
others  are  oriental. 

Figures  3,  5  and  8  are  from  China. 
Figure  6  is  from  Japan. 
Figure  7  is  from  Formosa. 


Hungerford:  Genus  Notonecta 
PLATE  XV 


191 


1 .  M.  saramoa. 


8.  N.  chinensis. 


192  The  University  Science  Bulletin 


PLATE  XVI 

The  male  genitalia  of  some  Notonecta  of  the  subgenus 
NoTONECTA.  Figures  1  and  2  represent  American  species,  the 
others  are  Old  World  species.  In  figure  7  the  clasper  has  been 
drawn  out  to  show  its  shape. 


Hungerford:  Genus  Notonecta 
PLATE  XVI 


193 


2.  N.  borealis. 


4  N.  amplifica. 


6.  N.m.  fulva. 


7.  N.m.  maculata. 


194  The  University  Science  Bulletin 


PLATE  XVn 

Male  genital  capsules  of  the  subgenus  Notonecta.     (Eastern  Hemisphere.) 

Figures  1  and  2  are  of  species  related  to  N.  borealis  Bueno  and  Hussey,  on 
Plate  XVI,  figure  2. 

Figure  4  shows  the  acute  anterolateral  angles  of  the  pronotum.  Compare 
with  Plate  VIII,  figure  8. 

Figures  6  and  11  are  oriental  species. 

Figure  8.  The  left  parameres  or  claspers  of  -V.  pallidula  Poisson  (left)  and 
of  A",  macidata  Fabr.  (right).     (After  Poisson.) 

Figure  13.    Head  and  pronotum  of  A'^.  macidata  Fabr.     (After  Poisson.) 

Figure  14.    Head  and  pronotum  of  A",  pallidula  Poisson.     (After  Poisson.) 


Hungerford:  Genus  Notonecta 


195 


PLATE  XVH 


13.  N.  m.  maculataj 

AR  12.  N.  V. 


10.  N.  canariensis.         \3       ^^  j^  pallidula. 


viridis. 


THE  UNIVERSITY  OF  KANSAS 

SCIENCE  BULLETIN 

Vol.  XXI.  1  March,  1933.  [No.  2. 


The  Giant  Water  Bugs 

( BelostomatidsG — Hemiptera) 

CARL   CUMMINGS,   Lawrence.   Kansas  *  ■ 


Abstract  :  This  paper  gives  the  key  to  the  eleven  genera  of  Belostomatida; 
and  keys  to  the  genera  Lethocerus  and  Diplonychus,  together  with  descrip- 
tions of  three  new  species  of  Lethocerus. 


THE  giant  ^vater  bugs  have  attracted  more  attention  than  any  of 
the  other  aquatic  Hemiptera.  They  have  been  called  "fish 
killers"  and  are  quite  widely  known  in  this  country  under  the  name 
"electric-light  bugs."  These  large  insects  measure  from  40  to  110 
mm.  in  length.  They  live  and  develop  in  ponds  and  other  water 
bodies,  where  they  feed  upon  small  fish  and  any  other  small  animals, 
including  insects  which  they  can  capture.  At  night  the  adults  take 
wing  and  are  attracted  to  street  lights,  where  because  of  their  large 
size  they  command  attention. 

The  family  Belostomatidse  comprise,  for  the  most  part,  large,  fiat, 
brown  bugs,  with  hind  legs  ciliated  and  flattened  for  swimming. 
They  bear  short,  flat,  straplike  appendages  at  the  tip  of  the  ab- 
domen; these  are  retractile.  The  antennae  are  hidden  in  pockets 
beneath  the  eyes,  and  the  front  legs  are  raptorial.  The  family  can 
be  divided  roughly  into  the  "giant  water  bugs"  and  the  "toebiters." 


*  Contribution   from   the   Department    of   Entomology,   University   of   Kansas. 

t  It  is  with  deep  regret  that  I  must  report  the  untimely  death  of  Carl  Oumniings  two 
days  after  he  submitted  this  paper  in  partial  fulfillment  for  the  requirement  of  the  master 
of  arts  degree.  Mr.  Cummings,  in  company  with  two  other  young  men,  was  on  a  collecting 
expedition  up  the  Kaw  river,  when  rough  water  capsized  the  canoe,  and  Mr.  Cunnnings, 
although  a  strong  swimmer,  could  not  overcome  the  handicap  of  the  heavy  clothing  he  was 
wearing  and  went  down  in  thirty  feet  of  water.  The  death  of  this  promising  young  man  is  a 
real  lo.ss  to  science,  becau.se  he  has  demonstrated  in  this  paper  an  unusual  ability  to  fix  upon 
structural  characters  of  specific  value  that  have  been  overlooked  by  former  students  of  this 
diiiicult  family.  To  my  mind,  Mr.  Cummings  has  given  us  the  first  satisfactory  key  to  the 
genus  Lethocerus.  It  was  his  intention  to  undertake  the  study  of  the  genus  Belostoma  as  hia 
next  assignment. — H.   B.   Hunoerford. 

14—3482 

(197) 


198  The  University  Science  Bulletin 

The  former  comprise  the  genera  Lethocerus,  Benacus,  Kirkaldyia 
and  Diplonychus ;  the  latter  include  the  remaining  genera.  Diplony- 
chus  is  included,  for  convenience,  in  the  first  group  because  it  con- 
tains a  species  or  two  of  large  size.  The  habit  of  depositing  the  eggs 
on  the  back  of  the  males,  as  in  Belostoma  and  Sphserodema,  however, 
really  relate  it  to  the  second  group. 

Since  there  are  only  one  or  two  species  in  Benacus  and  Kirkaldyia, 
it  is  necessary  only  to  provide  a  key  for  Lethocerus  and  Diplonychus. 
Key  to  Genera  of  Belostomatid^ 

(Based  upon  Bueno,  Can.  Ento.,  1907,  p.   333,  with  the  addition  of  Horvathinia  Montandon 
and  Kirkaldyia  Montandon) 

A.    Anterior  femora  not  sulcate Benacus  Stal. 

(See  p.   200) 
AA.    Anterior  femora  sulcate. 

B.    Anterior  tarsi  without  claws Horvathinia  Montandon. 

BB.    Anterior  tarsi  with  claws. 

C.    Anterior  tarsi  with  two  claws. 

D.    Claws  of  anterior  tarsi  of  equal  length,  minute. 

E.    Anterior  femora  more  or  less  incrassate,  much  larger  than  tibiae. 

F.    Species  with  two  sulci  between  the  eyes.  . .  .Nectocoris  Mayr. 

FF.    Species  without  such  sulci Sphcerodema  Laporte. 

EE.    Anterior    femora    scarcely    incrassate,    but    little    larger    than 

tibia    Limnogeton   Mayr. 

DD.    Claws  of  anterior  tarsi  of  equal  or  imequal  length  but  elongate. 

Diplonychus  Laporte. 
(See  p.  214) 
CC.    Anterior  tarsi  with  one  claw. 

D.    Head  conically  produced,   rostrum  long,   thin. 

E.    Membrane  of  hemelytra  large Belostoma  Latreille. 

EE.    Membrane  of  hemelytra  much  reduced Abedus  Stal. 

DD.    Head  not  conically  produced,  rostrWn  short,  stout. 

E.    Front  femur  with  two  unequal  sulci  for  reception  of  tibia. 

Kirkaldyia  Montandon. 
EE.    Front  femur  with  two  equal  sulci  for  reception  of  tibia. 

Lethocerus  Mayr. 
(See  p.  198) 

Key  to  Species  of  Lethocerus  Mayr  (Western  Hemisphere) 

(Note. — In  this  key  the  interocular  space  is  measured  at  its  narrowest  place,  and  measure- 
ments on  tibia  and  tarsus  do  not  include  fringe  of  hair.) 

A.    Under  side  of  posterior  tibia  with  inner  apical  angle  produced  into  a  sharp  point.f 
B.    Posterior  tibia  half  as  wide  as  front  femur. 

C    Upper  margin  of  eye  straight  as  viewed  from  the  front .  .  .  .L.  collosicus  Stal. 

(See  p.  205) 
CC.    Upper  margin  of  eye  rounded  as  viewed  from  the  front. 

D.    Lateral  margins  of  pronotum  broadly  foliareous  (thin). 

L.  camposi  Montandon. 
(See  p.   204) 


t  Species  from  Africa,  Madagascar  and  the  Orient  belong  in  this  first  section.  The  outer 
margin  of  hind  tibia  is  nearly  straight  in  Lethocerus  niloticum  Stal  from  Africa  and  in 
Lethocerus  insulanum  Mont,  from  New  Guinea  and  New  Caledonia,  while  in  Lethocerus 
indicus  L.  and  a  species  from  Madagascar,  L.  oculatus  (Mont.),  the  hmd  tibia  is  broader  and 
the  outer  margin  is  curved.  Montandon  has  designated  a  species  from  Queensland,  Australia, 
under  the  name  B.  edcntulum.  that  is  like  his  B.  insulanum  except  it  does  not  have  the  inner 
apical  angle  of  the  posterior  tibia  produced  into  a  sharp  point. 


CuMMiNGs:  The  Giant  Water  Bugs  199 

DD.    Lateral  margins  of  pronotuin  not  broadly  foliaceous.  .L.  angnstipes  Mayr. 

(See  p.  202) 
BB.    Posterior  tibia  at  least  two-thirds  as  wide  as  front  femur. 

C.    Anterior  and  posterior  femora  equal  in  length,  median  longitudinal  fuscous 

stripe  on  front  of  head  narrow L.   grandis  Linnaeus. 

(p.  208) 
CC.    Anterior  femur  longer  than   posterior   femur,   median   longitudinal   fuscous 

stripe  on  front  of  head  wide L.   largus  Ciininiings 

(p.  210) 
AA.    Under  side  of  posterior  tibia  with  inner  apical  angle  not  produced  into  a  sharp  point. 

B.    Interocular  space  equal  to  width  of  eye L.  americanus  Leidy 

(p.   201) 
BB.    Interocular  space  not  over  three-fourths  width  of  eye. 

C.    Interocular  space  greater  than  width  of  hind  tarsi. 

D.    Metaxyphus  not  sharply  pointed  posteriorly L.  truncatus  sp.  nov. 

(p.  212) 
DD.    Metaxyphus  sharply  pointed  posteriorly. 

E.    Two  longitudinal  fuscous  stripes  on  aljdominal  venter. 

L.  annulipes  Herrich-Schaffer. 
(p.  203) 
EE.    No   fuscous  stripes  on  abdominal   venter. 

F.    Interocular  space   nearly   equal   to   width    of   hind   tarsus, 
abdomen  tesselate  with  fuscous  spots. 

L.  mello-leitaoi  De  Carlo. 
(p.  211) 
FF.    Interocular    space    one-fifth     greater    than    hind    tarsus. 

Abdomen  reddish,  not  tesselate L.  uhleri  Montandon. 

(p.  213) 
CC.    Interocular  space  less  than  width  of  hind  tarsus. 

D.    Abdomen  with  fuscous  margin,  anterior  tarsi  white.  .L.  dilatus  sp.  nov. 

(p.   208) 
DD.     Abdomen  without  fuscous  margin,  anterior  tarsi  not  white. 

E.    Outer  margin   of  hind   tibia   more   curved    than   inner   margin, 

under  surface  of  tibia  mostly  light  brow-n.  .  .L,  del  pontei  De  Carlo. 

(p.  206) 
EE.    Outer  margin  of  hind  tibia  no  more  curved  than  inner  margin, 

under  surface  of  tibia  mostly  fuscous.  .  .  .L.  mello-leitaoi  De  Carlo. 

(p.   211) 

The  Old  World  species  of  Lethocerus  known  to  me  may  be  sep- 
arated by  the  following  key: 

A.    Under  side  of  posterior  tibia  with  inner  apical  angle  produced  into  a  sharp  point. 
B.    Posterior  lobe  of  pronotum  with  distinct  continuous  median  longitudinal  carina. 
C.    Width    of    interocular    space    in    front    a    little    more    than    Vs    transver.se 
diameter  of  an  eye. 
D.    Divergent  yellow  lines  on  anterior  lobe  of   pronotum  broad.      Eyes 

elongate,  nearly  twice  as  long  as  wide L.  indicus  (L.). 

DD.    Divergent  yellow  lines  on  anterior  lobe  of  pronotum  slender.     Eyes 
less  elongate,  only  IVz  times  as  long  as  wide. 

L.  niloticus  (Stal.)  and  var.  persicus  (Mont.). 
CC.    Width  of  interocular  space  in  front  less  than  one-half  transverse  diameter 

of  an  eye.     Eyes  large   /.-.  octdatus  (Mont.). 

BB.    Posterior  lobe  of  pronotum  without   a  continuous   distinct  median   longitudinal 

carina    I^-   insulanum   (Mont.). 

AA.    Under   side   of   posterior   tibia   with    inner   apical    angle   not   produced    into   a   sharp 

spine ^-  edentulum  (Mont.). 


200  The  University  Science  Bulletin 

Benacus  griseus  (Say),  1832 

1832.  Betostonia  griseus  Say.  Heter.  New  Harmony,  p.  37,  (Fitch  reprint,  p.  809). 
Compl.  Writ.,  I,  p.  365. 

1863.  Belostoma  griseus  (Say)  Dufour.     Ann.  Soc.  Ento.  Fr.,  ser.  4,  III,  p.  400. 

1876.  Benacus  griseus  (Say)  Uhler.     Bui.  U.  S.  Geol.  Geog.  Surv.,  I,  p.  337. 

1878.  Benacus  griseus  (Say)  Uhler.     Proc.  Bost.  Soc.  Nat.  Hist.,  XIX,  p.  441. 

1884.  Benacus  griseus  (Say)  Uhler.     Stand.  Nat.  Hist.,  II,  p.  256. 

1894.  Benacus  griseus  (Say)  Van  Duzee.     Bui.  Buf.  Soc.  Nat.  Sci.,  V,  p.   185. 

1905.  Benacus  griseus  (Say)  Howard.     Insect  Book,  p.  248,  fig.  4. 

1905.  Benacus  griseus  (Say)  Bueno.     Jl.  N.  Y.  Ento.  Soc,  XIII,  p.   434. 

1907.  Benacus  griseus  (Say)  Bueno  and  Brimley.     Ento.  News,  XVIII,  p.   434. 

1908.  Benacus  griseus  (Say)  Bueno.     Jl.  N.  Y.  Ento.   Soc,  XVI,  p.  237. 

1909.  Benacus  griseus  (Say)  Van  Duzee.     Bui.  Buf.  Soc  Nat.  Sci.,  IX,  p.   184. 
1914.  Benacus  griseus  (Say)  Barber.     Bui.  Am.   Mus.   Nat.   Hist.,  XXXIII,  p.   498. 
1919.  Benacus  griseus   (Say)  Hungerford.     Sci.  Bui.  U.  of  Kans.,  vol.  XI,  col.  pi.  II,  fig.  5. 
1926.  Benacus  griseus  (Say)  Blatchley.     Het.  of  East.  N.  A.,  p.  1042. 

The  following  also  refer  to  Benacus  griseus  Say: 

1847.    Belostoma  haldemanus  Leidy.     Jl.  Acad.  Nat.  Sci.  Phil.,  ser.   2,  I,  pp.   59,  66. 

1853.  Belostoma  impressus  Haldermann.     Proc.  Acad.  Nat.  Sci.  Phila.,  VI,  p.  364. 

1854.  Belostoma  harpax  Stal.     Of.  Vet.  Akad.  Forh.,  XI,  p.  240. 

1856.  Belostoma  angustatus  Guerin.  Sagra,  Hist,  de  Cuba,  VII,  p.   176. 

1861.  Benacus  haldemanus  (Leidy).  Stal,  Of.  Vet.   Akad.  Forh.,  XVIII,  p.   205. 

1863.  Benacus  haldemanus  (Leidy).  Dufour,  Ann.  Soc.  Ento.  Fr.,  ser.  4,  III,  p.  363. 

1871.  Benacus  haldemanus  (Leidy).  Mayr,  Verh.  Zool.  Bot.  Ges.  Wien,  XXI,  p.  428. 

Size.  Medium-sized  species.  Length,  47  mm.  to  64  mm.;  width, 
18.5  mm.  to  25  mm. 

Shape.  Body  elongate  oval  with  lateral  margins  of  hemelytra 
slightly  dilated.    Pronotum  convex,  with  lateral  margins  straight. 

Color.  General  color  medimn  brown.  Anterior  lobe  of  pronotmn 
fuscous,  margined  on  front  and  sides  with  brown;  a  broad  median 
brown  stripe  arising  on  anterior  margin  and  continuing  to  posterior 
margin  of  lobe.  Scutellum  with  large  fuscous  rectangular  area. 
Hemelytra  brown  with  distal  third  lighter.  Abdomen  medium 
brown.    Legs  medium  brown  and  unmarked. 

Structural  Characteristics.  Front  of  head  without  a  longitudinal 
median  carina.  Pronotum  smooth.  Length  of  first  segment  of  front 
tarsus  equal  to  second  on  outer  margin.  Under  side  of  posterior 
tibia  with  inner  apical  angle  not  produced  into  a  sharp  point.  Female 
operculum  minutely  bidentate  at  apex,  male  operculum  not  biden- 
tate.  Metaxyphus  sharply  pointed  posteriorly.  Width  of  hind 
tarsus  greater  than  interocular  space. 

Comparative  Notes.  This  species  differs  from  those  of  the  genus 
Lethocerus  in  not  having  grooves  in  the.  front  femora  for  reception 
of  tibiae. 

Types.    Are  unknown. 

Data  on  Distribution.  Kansas:  Douglas  Co.,  William  Hoffman, 
July   8,   1920;    May   24,    1918;    Douglas   Co.,   H.    B.    Hungerford. 


CuMMiNGs:  The  Giant  Water  Bugs  201 

Texas:  Galveston,  May,  F.  H.  Snow;  Sutton  Co.,  7:16:28,  R.  H. 
Beamer.  Florida:  Palm  Beach;  Inverness,  8:1:30,  Paul  W.  Oman. 
Cuba:  Rio  Almendares,  Marinao  Pro  v.-  Havana,  May  24,  1922,  P. 
J.  Bermudez;  Nov.,  1932,  P.  J.  Bermudez. 

Lethocerus  americanus  (Leidy),  1847 

1847.  Belostoma  americunum  Leidy.  J.  Acad.  Nat.  Sci.  Phila.,  VI,  pp.  58,  Cfi. 

1876.  Belostoma  americanum  Leidy.  Uhler,  Bui.  U.  S.  Geol.  Geog.  Surv.,  I,  p.  337. 

1878.  Belostoma  airiericanum  Leidy.  Uliler,  Proc.  Bost.  Soc.  Nat.  Hist.,  XIX,  p.   441. 

1884.  Belostoma  americanum  Leidy.  Uhler,  Stand.   Nat.   Hist.,  II,  p.   256. 

1894.  Belostoma  americanum  Leidy.  Van  Duzee,  Bui.  Buf.   Soc.  Nat.  Sci.,  V,  p.   18.'). 

1895.  Belostoma  americanum  Leidy.  Gillette  and  Baker,  Hemip.  Colo.,  p.  63. 

1896.  Belostoma  americanum  Leidy.  Montandon,   Ann.   Soc.   Ent.    Belg.,   XL,    p.    512. 
1905.  Amorgius  americanum  (Leidy).  Bueno,  Jl.  N.  Y.  Ent.  Soc,  XIII,  p.   44. 

1908.  Lethocerus  americanus  (Leidy).     Bueno,  Jl.  N.  Y.  Ent.   Soc.,  XVI,  p.   237. 

1914.  Lethocerus  americanus  (Leidy).  Barber,  Bui.  Am.  Mus.  Nat.  Hist.,  XXXIII, 
p.  498. 

1914.  Lethocerus  am-ericanus  (Leidy).     Parshley,   Psyche,   XXI,   p.    140. 

1925.  Lethocerus  americanus  (Leidy).     Conistock,   Intr.   to  Ent.,   p.   366. 

1926.  Lethocerus  americanus  (Leidy).     Blatchley,  Het.   of  East.  N.   A.,  p.   1043. 

1928.    Lethocerus  americanus  (Leidy).     Metcalf  &  Flint,  Dest.  Use.  Insects,  p.  190,  fig.  114. 

The  following  also  refer  to  L.  americanus  Leidy: 

1861.  Belostoma  griseum  Stal.     Of.  Vet.  Akad.  Forh.,  XVIII,  p.   206. 

1863.  Belostoma  litigiosum  Dufour.     Ann.  Soc.  Ent.  Fr.,  ser.   4,  III,  p.   383. 

1863.  Belostoma  obscurus  Dufour.     Ann.  Soc.  Ent.  Fr.,  ser.  4,  III,  p.  383. 

1871.  Belostoma  griseum  Dufour.     Mayr,  Verh.  Zool.  Bot.  Ges.  Wien,  XXI,  p.   427. 

1886.  Belostoma  grande  Dinimock.     Mass.  Fish  and  Game  Comm.,  p.  69. 

1896.  Belostoma  obscurum  Dufour.     Montandon,  Ann.  Soc.  Ent.  Belg.,  XL,  p.   512. 

1905.  Amorgius  obscurus  (Dufour).     Bueno,  Jl.  N.  Y.  Ent.  Soc,  XIII,  p.   44. 

1905.  Benacus  griseus  Say.     Howard,  Insect  Book,  p.  266,  p.  xxix,  fig.   36. 

1915.  Lethocerus  obscurus  (Dufour).     Bueno,  Psyche,  XXII,  p.  17. 

1926.    Lethocerus  obscurus  (Dufour).     Blatchley,  Het.   of  East.   N.   A.,  p.    1044. 

Size.  Medium-sized  species.  Length,  40.5  mm.  to  61.5  mm.; 
width,  16.5  mm.  to  24  mm. 

Shape.  Body  elongate  oval,  with  lateral  margins  of  hemelytra 
slightly  dilated.    Thorax  convex  with  lateral  margins  almost  straight. 

Color.  General  color  medium  brown.  Pronotum  brown  with  a 
median  pair  of  light-brown  lines  arising  on  anterior  margin  and  ex- 
tending back  about  one-third  the  length  of  anterior  lobe.  Scutellum 
with  large  fuscous  rectangular  area.  Hemelytra  brown  with  distal 
third  lighter.  Abdomen  medium  brown.  Middle  and  hind  femora 
with  three  transverse  fuscous  stripes ;  front  legs  and  middle  and  hind 
tibiae  usually  indistinctly  marked. 

Structural  Characteristics.  Front  of  head  with  indistinct  longi- 
tudinal median  carina.  Interocular  space  at  narrowest  place  about 
equal  to  width  of  eye.  First  segment  of  front  tarsus  in  ratio  of 
10  to  13  to  second  segment  measured  on  outer  margin.  Under  side 
of  posterior  tibia  with  inner  apical  angle  not  produced  into  a  sharp 


202  The  University  Science  Bulletin 

point.  Female  operculum  with  two  sharp  points  at  apex;  male 
operculum  without  points.    Metaxyphus  sharply  pointed  posteriorly. 

Comparative  Notes.  This  species  differs  from  other  species  of 
Lethocerus  in  having  the  interocular  space  at  narrowest  place  equal 
to  width  of  eye.  L.  obscurus  (Dufour)  has  been  applied  to  small 
males  of  L.  americanus  (Leidy)  and  is  not  a  good  species. 

Types.    Location  not  known. 

Data  on  Distribution:  Michigan:  Mich.,  Chickering,  May  3, 
1930;  Douglas  Lake,  Bryant  Bog,  Aug.  3,  1923,  and  Aug.  17,  1923, 
H.  B.  Hungerford;  Douglas  Lake,  Sedge  Point  Pool,  1932,  also 
July  3,  1923,  and  July  6,  1923,  H.  B.  Hungerford;  Douglas  Lake, 
July  20,  1926,  H.  B.  Hungerford.  New  York:  White  Plains,  May 
14,  1907;  West  Point,  Sept.  11,  1926,  W\  Robinson;  Queens,  L.  I., 
F.  M.  Schott;  Ithaca,  H.  L.  Zabristic;  Long  Island.  New  Jersey: 
Brown's  Mills,  Patterson.  Washington,  D.  C:  Washington,  D.  C, 
D.  H.  demons.  Wisconsin:  Cranmoor,  C.  W.  Hooker.  Delaware: 
Del.  Water  Gap.  Ohio:  Geauga  Co.,  Oct.  Kansas:  Lawrence, 
F.  H.  Snow;  Harvey  Co.,  Aug.  27,  1917;  Lawrence,  May  9,  1920, 
H.  B.  Hungerford;  Lawrence,  W.  Hoffman.  Nevada:  Meapa,  G.  F. 
Knowlton,  April  24,  1923.  Utah:  Logan,  Aug.  15,  1910;  May  6. 
1910;  Nov.  1, 1922;  Oct.  11,  1909;  Lehi,  May  27,  1930;  G.  F.  Knowl- 
ton; Vineyard,  Tom  Spaulding.  Oregon:  Lost  River,  Alene.  Cali- 
fornia: Sacramento,  April  29,  1908,  E.  D.  Ball;  Eureka,  H.  S. 
Barber;  San  Jose. 

Lethocerus  angustipes  (Mayr),  1871 

1871.    Belostoma  angustipes   Mayr.  Verh.   Zool.-Bot.   G'es.    Wien,  XXI,  p.   427. 

1896.    Belostoma  angustipes  Mayr.  Montandon,  Ann.  Soc.  Ent.  Belg.,  XL,  p.   511. 

1901.  Belostoma  angustipes  Mayr.  Champion,  Biol.  Centr.  Am.,  Heter.,  II,  p.  368,  pL 
22,   fig.  3. 

1909.    Lethocerus  angustipes  Mayr.  Kirkaldy  &  Bueno,  Proc.  Ent.  Soc.  Wash.,  X,  p.  188. 

1917.    Lethocerus  angustipes  Mayr.  Van  Duzee,  Cat.   of  Hemiptera,  p.   466. 

Size.  Medium  sized  species.  Length,  47  mm.  to  68  mm.;  width, 
21.5  mm.  to  28  mm. 

Shape.  Body  elongate  oval  with  lateral  margins  of  hemelytra 
almost  parallel.  Thorax  convex  with  lateral  margins  slightly 
curved. 

Color.  General  color  medium  brown.  Anterior  lobe  of  pronotum 
fuscous,  margined  on  front  and  sides  with  light  brown;  a  median 
pair  of  light-brown  lines  arising  on  anterior  margin  and  extending 
back  about  one-third  the  length  of  lobe.  Scutellum  with  large  rec- 
tangular fuscous  area  which  is  divided  on  front  half  by  a  slender 


CuMMiNGs:  The  Giant  Water  Bugs  203 

median  longitudinal  line.  Hemelytra  brown  with  distal  third 
lighter.  Abdomen  mostly  fuscous  with  outer  margin  light  brown. 
Markings  on  legs  often  indistinct,  broad  fuscous  longitudinal  streak 
on  upper  side  of  front  femur  which  is  nearly  immaculate  beneath.  ^ 

Structural  Characteristics.  Eyes  broadly  rounded  on  upper  mar- 
gin. Front  of  head  not  medianly  carinate.  Surface  of  pronotum 
smooth,  with  lateral  margins  thin  at  extreme  edge,  but  not  broadly 
foliaceous.  Under  side  of  posterior  tibia  with  inner  apical  angle 
produced  into  a  sharp  point.  Width  of  posterior  tibia  equal  to  in- 
terocular  space  at  narrowest  place,  and  equal  to  one-half  the  width 
of  front  femur.  Lateral  margins  of  posterior  tibia  nearly  straight. 
Posterior  end  of  metaxyphus  sharply  pointed  posteriorly,  with  an- 
terior end  not  abruptly  elevated  into  a  point.  Apex  of  female 
operculum  feebly  bidentate,  male  operculum  not  bidentate. 

Co7nparative  Notes.  This  species  differs  from  L.  camposi  Mon- 
tandon  in  being  smaller  and  in  having  the  lateral  margins  of  the 
pronotum  not  foliaceous,  and  from  L.  niloticum  Stal  and  L.  in- 
sulanum  Montandon  in  having  the  hind  tibia  equal  to  the  interocular 
space. 

Types.    Doctor  Hungerford  says  there  are  specimens  in  the  Mu 
seum  at  Vienna,  evidently  studied  by  Mayr. 

Data  on  Distribution.  United  States:  Death  Valley,  Nev.,  T  S 
Galmer.  Mexico;  Durango,  Dr.  E.  Palmer;  Mex.  R.  Muller; 
Zitacuaro,  Michoacan,  8-5-1932;  Dist.  Fed.,  por  Ancona;  Colima. 

Lethocerus  annulipes  (Herrich-Schaffer) ,  1846 

1846.    Belostoma  annulipes  H.-S.  Wanz.  Ins.,  VIII,  p.   28,  figs.   803,   804. 

1865.    Belostoma  annulipes  H.-S.  Mayr,  Hemip.,  pp.   185,  186. 

1871.    Belostoma  annulipes  H.-S.  Mayr,  Verh.  Zool.  Bot.  Ges.  Wien,  XXI,  p.  427. 

1930.  Lethocerus  annulipes  H.-S.  De  Carlo,  Rev.  de  la  Soc.  Ent.  Arg.,  p.  107,  pi.  VI, 
figs.  21  and  23. 

The  following  also  refer  to  L.  annulipes  H.-S.: 

1863.  Belostoma  reficeps  Dufour.     Ann.  Soc.  Ent.  Fr.,  ser.  4,  III,  p.   382  (?). 

1863.  Belostoma  signoreti  Dufour.     Ann.  Soc.  Ent.  Fr.,  ser.  4,  III,  p.  382  (?). 

1863.  Belostoma  distinctum  Dufour.     Ann.   Soc.  Ent.  Fr.,  ser.   4,  III,  p.   382   (?). 

1896.  Belostoma  mayri  Montandon.     Ann.  Soc.  Ent.  Belg.,  XL,  p.   514. 

&ize.  Medium-sized  species.  Maximum  length,  75  mm.;  maxi- 
mum width,  261/2  mm. ;  minimum  length,  53  mm. ;  minimum  width, 
21  mm. 

Shape.  Body  elongate  oval  with  lateral  margins  of  hemelytra 
nearly  parallel.  Thorax  convex  with  lateral  margins  distinctly 
curved. 

Color.    General  color  medium  brown.     Pronotum  brown  with  a 


204  The  University  Science  Bulletin 

median  pair  of  light-brown  lines  arising  on  anterior  margin  and  ex- 
tending back  about  one-third  the  length  of  anterior  lobe.  Scutellum 
with  large  fuscous  rectangular  area.  Hemelytra  brown  with  distal 
third  lighter.  Abdomen  light  brown;  a  pair  of  fuscous  stripes  be- 
ginning on  the  anterior  margin  of  second  segment  and  ending  on 
sixth.  Middle  and  posterior  tibise  and  femora  each  with  three  trans- 
verse fuscous  stripes  on  ventral  side.    Front  legs  indistinctly  marked. 

Structural  Characteristics.  Front  of  head  without  a  longitudinal 
median  carina.  Pronotum  smooth.  Width  of  interocular  space  at 
narrowest  place  is  greater  than  width  of  hind  tarsus.  Length  of 
first  segment  of  front  tarsus  one-half  that  of  second  on  outer  mar- 
gin. Underside  of  posterior  tibia  with  inner  apical  angle  not  pro- 
duced into  a  sharp  point.  Female  operculum  minutely  bidentate  at 
apex,  male  operculum  not  bidentate.  Metaxyphus  sharply  pointed 
posteriorly. 

Comparative  Notes.  This  species  differs  from  other  species  of 
Lethocerus  in  having  two  longitudinal  fuscous  stripes  on  lower  side 
of  abdomen. 

Types.  Described  from  South  America,  location  of  type  unknown. 

Data  on  Distribution.  United  States:  Palm  Beach,  Florida  (W. 
Robinson  bequest).  Brazil:  Ypirango,  S.  Paulo,  S.  A..  Rbt.  Spitz, 
1-20-26,  1-27-24,  2-7-24;  St.  Catharina;  Buenos  Ar.  Rep.,  Arja, 
Brazil,  S.  A.,  Alfredo  Faz,  1921;  Brazil.  E.  C.  Green.  Argentina: 
Icano-Rio  Salado,  Santiago  del  Estero,  S.  A.,  Fives  Lille,  Santa  Fe, 
A.  Bruch,  1923;  Bs.  Aires,  C.  S.  Reed;  Buenos  Aires,  Argentine 
Republic,  S.  A.,  H.  T.  Martin.  British  Guiana:  Plantation  Ecceles, 
E.  Bank  Demerara  River,  S.  Harris,  4-4-32.  French  Guiana:  St. 
Laurent,  Guyana.  Paraguay:  Villarrica,  Paraguay,  S.  A.,  Fran. 
Schade,  10-29-29,  9-10-23.  Colombia:  Colombia,  S.  A.  Venezuela: 
Caracas,  H.  Pittier.    Porto  Rico:   Porto  Rico,  E.  A.  Wagerrin. 

Lethocerus  camposi  (Montandon),  1900 

1900.    Amorgius  camposi  Montandon.     Bui.   de  la  Soc.  des  Sci.  de  Buc,  p.   561. 

Size.  Large  species.  Length,  77  mm.  to  85  mm. ;  width,  30  mm.  to 
32  mm. 

Shape.  Body  elongate  oval  with  lateral  margins  of  hemelytra 
almost  parallel.    Pronotum  convex  with  lateral  margins  curved. 

Color.  General  color  medium  brown.  Anterior  lobe  of  pronotum 
fuscous,  margined  on  front  and  sides  with  brown;  a  median  pair  of 
brown  lines  arising  on  anterior  margin  and  extending  back  about 


Cum  MINGS :  The  Giant  Water  Bugs  205 

one-third  the  length  of  lobe;  a  pair  of  conspicuous  brown  oval  spots. 
Scutellum  with  large  rectangular  fuscous  area.  Hemelytra  brown 
with  distal  third  lighter.  Abdomen  brown  with  margin  lighter  and 
tesselate  with  fuscous  spots.  Middle  and  hind  femora  with  three 
transverse  fuscous  stripes  on  ventral  side;  legs  otherwise  not  dis- 
tinctly marked. 

Structural  Characteristics.  Eyes  broadly  rounded  on  upper  mar- 
gin. Front  of  head  with  longitudinal  median  carina.  Lateral  mar- 
gins of  pronotum  broadly  foliaceous;  posterior  lobe  rugose,  anterior 
lobe  rough,  but  not  rugose.  Underside  of  posterior  tibia  with  inner 
apical  angle  produced  into  a  sharp  point.  Width  of  posterior  tibia 
less  than  half  that  of  anterior  femur.  Lateral  margins  of  hind  tibia 
nearly  straight.  Metaxyphus  sharply  pointed  posteriorly  with 
anterior  end  not  abruptly  elevated  into  a  point.  Apex  of  female 
operculum  feebly  bidentate,  male  operculum  not  bidentate  at  apex. 

Comparative  Notes.  This  species  differs  from  L.  collosicus  Stal 
in  being  lighter  in  color  and  in  having  the  upper  margin  of  eye 
rounded;  L.  niloticum  Stal,  L.  insulanum  Montandon,  and  L.  angus- 
tipes  Mayr  in  having  the  lateral  margins  of  pronotum  broadly 
foliaceous. 

Types.  Described  from  Guayaquil,  Ecuador.  In  Montandon's 
collection,  Bucharest. 

Data  on  Distribution.  Ecuador:  Guayaquil,  Duran,  F.  Campos, 
(Specimen  compared  with  cotype)  ;  Guayaquil,  F.  W.  Coding; 
Guayaquil,  F.  Campos. 

Lethocerus  collosicus  (Stal) ,  1854 

1854.  Belostoma  collosicus  Stal.     Ofv.   Vet.-Ak.  Forh.,  XI,  p.   239. 

1861.  Belostoma  collossicum  Stal.     Ofv.  Vet.-Ak.  Forh.,  XVIII,  p.   205. 

1863.  Belostoma  colossicum  Stal.     Dufour,  Ann.   Soc.  Ent.  Fr.,  sen   4,  III,  381. 

1865.  Belostoma  colossicum  Stal.     Mayr,  Hemiptera,  p.   184. 

1871.  Belostoma  colossicum  Slal.     Mayr,  Verh.  Zo6!.-Bot.    Ges.  Wien,  XXI,  pp.  423,  425. 

1884.  Belostoma  coUosicum  Stal.     Uhler,  Stand.   Nat.   Hist.,  II,  p.   256. 

1895.  Belostoma  colossicum  Stal.     Mont.,  Ann.   Soc.   Ent.   Belg.,  pp.   472-477,  fig.   2. 

1901.  Belostoma  colossicum  Stal.     Champion,  Biol.    C.-Am.,  Hem.-Het.,  II,  pi.  22,  fig.  4. 

1906.  Amorgius  colossicum  (Stal).     Bueno,   Ent.   News,  Feb.,  p.   55. 

1909 — Lethocerus  colossicus  (Stal).     Kirkaldy  and  Bueno,  Proc.  Ent.  Soc.  Wash.,  X,  p.  188. 

Size.  Large  species.  Length,  72  mm.-85mm.;  width,  31  mm.-34 
mm. 

Shape.  Body  elongate  oval  with  lateral  margins  of  hemelytra 
somewhat  dilated.  Head,  short,  as  seen  from  above.  Thorax  con- 
vex with  lateral  margins  widely  rounded. 

Color.  General  color  nearly  black.  Lateral  margins  of  heme- 
lytra coarsely  tesselate  with  light  brown.     Median  and  posterior 


206  The  University  Science  Bulletin 

tibia?  and  femora  each  with  two  narrow  transverse  light-brown 
stripes. 

Structural  Characteristics.  Front  of  head  with  a  longitudinal 
median  carina.  Eyes  triangular  in  appearance  when  viewed  from 
the  front,  with  upper  margin  of  eye  straight.  Pronotum  rugose  with 
lateral  margins  broadly  foliaceous  (thin).  Underside  of  posterior 
tibia  with  inner  apical  angle  produced  into  a  sharp  point.  Front 
femur  more  than  twice  as  wide  as  hind  tibia,  which  has  its  lateral 
margins  almost  straight.  Metaxyphus  sharply  pointed  posteriorly, 
with  anterior  end  not  abruptly  elevated  into  a  point.  Posterior  end 
of  female  operculum  minutely  bidentate  at  apex,  male  operculum  not 
bidentate  at  apex. 

Comparative  Notes.  This  species  differs  from  L.  camposi  Mon- 
tandon  in  being  darker  in  color  and  in  having  the  upper  margin  of 
eye  straight. 

Types.    Types  in  collection  of  Signoret. 

Data  on  Distribution.  Cuba:  Santiago,  Cuba;  Cuba,  Nov.  1932, 
P.  J.  Bermudez;  Central  San  Antonio,  Madruga,  Prov.  Havana, 
Cuba,  P.  J.  Bermudez,  6-8-32.  Mexico:  Campeche,  E.  P.  Creaser; 
Mexico,  W.  T.  M'Donald.  Yucatan:  Yucatan,  E.  P.  Creaser. 
Jamaica:  Jamaica,  J.  H.  Andrews.  Panama:  Panama,  J.  W. 
Green. 

Lethocerus  del  pontei  De  Carlo,  1930 

1930.  Lethocerus  del  pontei  De  Carlo.  Rev.  de  la  Soc.  Ent.  Arg.,  No.  13,  p.  108,  pi. 
VII,  fig.  24. 

1932.    Lethocerus  del  pontei  De  Carlo.     Re\-.  de  la  Soc.  Ent.  Arg.,  No.  3,  p.  217. 

The  following  also  refer  to  Lethocerus  del  pontei  De  Carlo: 

1876.  Belostoma  annuUpes  H.-S.  Uhler,  Bui.  U.  S.  Geol.  Geog.  Surv.,  I,  p.  337. 

1894.  Belostoma  annulipes  H.-S.  Uhler,  Proc.  Cal.  Acad.  Sci.,  ser.  2,  IV,  p.  291. 

1895.  Belostoma  annulipes  H.-S.  Gillete  and  Baker,  Hemip.  Colo.,  p.  63. 

1896.  Belostoma  annulipes  H.-S.  Montandon,  Ann.  Soc.  Ent.  Belg.,  XL,  p.  516. 

1901.  Belostoma  annulipes  H.-S.  Champion,  Biol.  Centr.  Am.,  Heter.,  II,  p.  367,  pi.  22, 
fig.  3. 

1906.    Amorgius  anmdipes  H.-S.     Bueno,  Ent.     News.  Feb.,  p.  55. 

Size.  Medium-sized  species.  Maximum  length,  71  mm.,  maximum 
width,  27I/2  mm.;  minimum  length,  52  mm.,  minimum  width,  21  mm. 

Shape.  Body  elongate  oval  with  lateral  margins  of  hemelytra 
slightly  dilated.  Pronotum  convex  with  lateral  margins  nearly 
straight. 

Color.  General  color  medium  brown.  Pronotum  brown  with 
lateral  margins  lighter;  a  median  pair  of  light-brown  lines  arising 
on  anterior  margin  and  extending  back  about  one-third  the  length 
of  anterior  lobe.     Scutellum  with  large  fuscous  rectangular  area. 


CuMMiNGS:  The  Giant  Water  Blgs  207 

Hemelytra  brown  with  distal  tliird  lighter.  Abdomen  reddish- 
brown,  thinly  tesselate  with  fuscous  spots.  Mesosternum  with  large, 
irregular  fuscous  spot  on  midline,  fuscous  markings  not  prominent 
on  either  side  of  midline.  Anterior  tibia  with  three  transverse  fuscous 
stripes  on  both  sides;  middle  and  posterior  femora  and  tibise  with 
three  transverse  fuscous  stripes  on  lower  side;  markings  on  legs 
often  indistinct  or  absent. 

Structural  Characteristics.  Front  of  head  without  a  longitudinal 
median  carina.  Pronotum  smooth.  Width  of  eye  nearly  twice  that 
of  interocular  space  which  is  not  as  wide  as  hind  tarsus.  Length  of 
first  segment  of  front  tarsus  one-half  that  of  second  on  outer  mar- 
gin. Underside  of  posterior  tibia  with  inner  apical  angle  not  pro- 
duced into  a  sharp  point.  Female  operculum  with  two  sharp  points 
at  apex,  male  operculum  without  points.  Metaxyphus  sharply 
pointed  posteriorly.  Outer  margin  of  hind  tibia  more  curved  than 
inner  margin. 

Comparative  Notes.  This  species  differs  from  L.  dilatus  Cum- 
mings  in  having  the  outer  margin  of  the  abdomen  not  fuscous,  and 
from  L.  mello-leitaoi  De  Carlo  in  having  different  color  markings 
on  lower  side  and  in  having  outer  margin  of  hind  tibia  more  curved 
than  inner  margin. 

Types.  National  Museum  of  Natural  History,  Buenos  Aires. 
Holotype  from  Argentina. 

Data  on  Distribution.  Arizona:  Santa  Cruz  Co.,  8-4-1927,  L.  D. 
Anderson;  Patagonia,  8-4-27,  R.  H.  Beamer;  Chiricahua  Mts.,  7-8- 
32,  R.  H.  Beamer;  S.  W.  edge  Tuscon,  7-20-32,  R.  H.  Beamer; 
Nagales.  Texas:  Jim  Wells  Co.,  7-24-28,  R.  H.  Beamer;  Valentine, 
July  13,  1927,  R.  H.  Beamer;  Victoria,  J.  D.  Mitchell;  Brownsville. 
Cuba:  Rio  A'mendares,  Marinao  Prov.  Havana,  May  24,  1932,  P. 
J.  Bermudez.  Havana,  Cuba,  Palmer  and  Riley;  Guantanamo. 
Jamaica:  Alligator  Pond,  Jamaica;  Claremont,  E.  Perkins.  Porto 
Rico:  San  Juan.  San  Salvador:  San  Salvador,  C.  Caderon.  Costa 
Rica:  Costa  Rica,  Heinrich  Schmidt,  1931.  Honduras:  Honduras, 
F.  H.  Dyer.  Panama:  Panama.  Mexico:  Yucatan,  G.  F.  Gaumer; 
50  m.  S.  Victorias,  San  Luis  Potosi,  6-10-32,  Hobart  Smith;  Colima, 
Encero  Vera  Cruz,  S.  of  Jalapa,  7-17-32,  H.  Smith;  Mexico,  A. 
Duges;  Cordoba,  Fred  Knab.  Ecuador:  Guayaquil,  W.  F.  Coding; 
Guayaquil,  F.  Campos.  Paraguay:  Villarrica,  S.  A.,  1-6-24,  Fran. 
Schade;  Villarrica,  4-11-23,  Fran.  Schade.  Brazil:  Ypirango,  S. 
Paulo,  R.  Spitz,  1-20-26 ;  Blumenau,  Espir  Santo. 


208  The  University  Science  Bulletin 

Lethocerus  dilatus  sp.  nov. 

Size.  Large  species.  Length,  74  mm. -85  mm.,  width,  29.5  mm.- 
31.5  mm. 

Shape.  Body  elongate  oval  with  lateral  margins  of  hemelytra 
dilated.  Pronotum  comparatively  small,  convex,  and  with  lateral 
margins  straight. 

Color.  General  color  medium  brown.  Anterior  lobe  of  pronotum 
dark  without  a  pair  of  light-brown  lines  arising  on  anterior  margin. 
Scutellum  with  large  fuscous  rectangular  area.  Hemelytra  uniform 
medium  brown  throughout.  Abdomen  medium  brown  with  outer 
margin  fuscous.  Anterior  tibia  fuscous  with  tarsus  white,  middle 
and  posterior  femora  and  tibiae  with  three  transverse  fuscous  stripes 
on  lower  side. 

Structural  Characteristics.  Front  of  head  without  a  longitudinal 
medium  carina.  Anterior  lobe  of  pronotum  rough,  but  not  rugose. 
Width  of  eye  twice  that  of  interocular  space  which  is  not  as  wide  as 
hind  tarsus.  Length  of  first  segment  of  front  tarsus  one-half  that  of 
second  on  outer  margin.  Underside  of  posterior  tibia  with  inner 
apical  angle  not  produced  into  a  sharp  point.  Female  operculum 
minutely  bidentate  at  apex,  male  operculum  not  bidentate.  Meta- 
xyphus  sharply  pointed  posteriorly.  Outer  margin  of  hind  tibia 
more  curved  than  inner  margin. 

Comparative  Notes.  This  species  differs  from  L.  del  pontei  De 
Carlo  and  L.  mello-leitooi  De  Carlo  in  having  the  outer  margin  of 
the  abdomen  fuscous  on  lower  side. 

Types.  In  Francis  Huntington  Snow^  Entomological  Museum  of 
the  University  of  Kansas,  holotype    5    and  one  para  type    5  . 

Data  on  Distribution.  Bolivia:  Buena  Vista,  Dept.  Santa  Cruz 
S.  A.,  9-24-1924,  R.  T.  Steinbach. 

Lethocerus  grandis   (Linnaeus) ,  1758 

1758.  Nepa  grandis  Linn.     Syst.  Nat.,  Ed.  x,  p.  440. 

1794.  Nepa  grandis  Linn.     Fabricius,  Ent.  Syst.,  IV,  p.  61. 

1803.  Nepa  grandis  Linn.     Fabricius,  Syst.   Rhyn.,  p.   106. 

1835.  Belostomum  grande  (Linn.).     Burmei.ster,  Handb.   der  Ent.,  II,  p.   195. 

1847.  Belostomvm  grande  (Linn.).     Leidy,  J.  Acad.  Nat.  Sci.  Phil.,  VI,  pp.  58,  66. 

1856.  Belostoma  grande  (Linn.).     Guerin,  Sagra,  Hist,  de  Cuba,  VII,  p.  175. 

1863.  Nepa  grandis  Linn.     Dufour,  Ann.  Soc.  Eht.  Fr.,  ser.  4,  III,  p.  380. 

1871.  Belostoma  grande  (Linn.).     Mayr,  Verh.  Zool.-Bot.  Ges.  Wien.,  XXI,  pp.  423,  425. 

1865.  Belostoma  grande  (Linn.).     Mayr,  Hemiptera,  p.   184. 

1884.  Belostoma  grandis  (Linn.).     Uhler,  Stand.  Nat.   Hist.,  II,  p.   256. 

1930.  Lethocerus  grandis  (Linn.).  De  Carlo,  Rev.  de  la  Soc.  Ent.  Arg.,  No.  13,  p.  105, 
pi.  VI,  fig.  22. 

Size.  Largest  species  of  the  genus  Lethocerus  and  also  of  the 
hemiptera.    Length,  94  mm. -110  mm.;  width,  34  mm. -40  mm. 


CuMMiNGs:  The  Giant  Water  Bugs  209 

Shape.  Body  elongate  oval  with  lateral  margins  of  hemelytra 
almost  parallel.    Thorax  convex  with  lateral  margins  curved. 

Color.  General  color  medimn  brown.  Front  of  head  with  a 
narrow  fuscous  stripe  reaching  from  vertex  to  clypeus.  Anterior 
lobe  of  pronotum  fuscous  margined  on  front  and  sides  with  brown; 
a  medium  pair  of  brown  lines  arising  on  anterior  margin,  diverging 
on  their  posterior  third  and  ending  about  the  middle  of  the  lobe;  a 
pair  of  conspicuous  brown,  oval  spots  in  front  of  which  are  indefinite 
brown  markings.  Scutellum  with  large  rectangular  fuscous  area. 
Hemelytra  brown  with  distal  third  lighter,  and  with  light-brown 
maculations  throughout.  Abdominal  venter  fuscous,  fading  out  to 
light  brown  at  apex.  Underside  of  all  tibiae  and  femora  each  with 
three  broad,  transverse  fuscous  bands. 

Structural  Characteristics.  Eyes  large  and  broadly  rounded  on 
upper  margin.  Front  of  head  not  medianly  carinate.  Surface  of 
pronotum  smooth,  lateral  margins  moderately  explanate  and  edges 
thickened,  not  foliaceous.  Underside  of  posterior  tibia  with  inner 
apical  angle  produced  into  a  sharp  point.  Width  of  posterior  tibia 
at  least  two-thirds  that  of  anterior  femur.  Length  of  anterior  femur 
equal  to  hind  femur.  Outer  margin  of  hind  tibia  curved  and  inner 
margin  nearly  straight.  Metaxyphus  sharply  pointed  posteriorly, 
with  anterior  end  abruptly  elevated  into  a  point.  Apex  of  female 
operculum  with  two  sharp  points,  male  operculum  without  points. 

Comparative  Notes.  This  species  differs  from  L.  largus  Cummings 
in  having  the  front  and  hind  femora  equal,  and  in  having  different 
color  markings  on  face  and  abdomen. 

Type.  Doctor  Hungerford  says  there  are  some  twenty-six  trays 
of  insects  in  the  Museum  at  Upsala,  Sweden,  that  are  supposed  to 
have  been  studied  by  Linnaeus.  There  is  a  specimen  of  Lethocerus 
measuring  83  mm.  long  bearing  the  following  labels  "Mus.  Gust. 
Adolphi"  "typus"  "Nepa  grandis"  and  another  only  38  mm.  long 
labeled  "Nepa  grandis?"  "C.  U.  Mus.  Gust.  Adolphi"  "Grandis  /3." 
Linnaeus,  in  his  10th  ed.  of  Systema  Naturae,  list  his  specimen  from 
"M.  L.  U."  and  gives  three  references  to  literature.  Since  Gust. 
Adolphus  was  a  small  boy  in  1758  it  may  be  that  the  specimen 
labeled  "typus"  is  not  the  type,  and  therefore  it  seems  best  to  let 
the  species  described  above  and  also  recognized  by  Mr.  dc  Carlo 
stand  as  Lethocerus  grandis  (Linn.). 

Data  on  Distribution.  Brazil:  Igarape  Assu,  1912.  Paraguay: 
Villarica,  8,  31,  Fr.  Schade.  Colombia:  Villa vicencio,  Ost.  Colom- 
bia, 400  m,  coll.  Fassl. 


210  The  University  Science  Bulletin 

Lethocerus  largus  sip.  nov. 

Size.  Large  species,  second  in  size  only  to  L.  grandis  Linnaeus. 
Length,  84  mm.  to  94  mm.;  width,  30  mm.  to  33  mm. 

Shape.  Body  elongate  oval  with  lateral  margins  of  hemelytra 
almost  parallel.    Pronotum  convex  with  lateral  margins  curved. 

Color.  General  color  medium  brown.  Front  of  head  with  broad 
fuscous  stripe  reaching  from  vertex  to  clypeus.  Anterior  lobe  of 
pronotum  fuscous,  margined  on  front  and  sides  with  light  brown ;  a 
median  pair  of  brown  lines  arising  on  anterior  margin,  diverging  on 
their  posterior  third  and  ending  about  the  middle  of  lobe;  a  pair  of 
conspicuous  brown,  oval  spots,  in  front  of  which  are  indefinite  brown 
markings.  Scutellum  with  large  rectangular  fuscous  area.  Hemely- 
tra brown  with  distal  third  lighter,  and  without  light-brown  macu- 
lations.  Abdomen  fuscous  with  outer  margin  light  brown.  Front 
legs  not  transversely  marked,  middle  and  hind  tibiae  and  femora 
each  with  three  broad,  transverse,  fuscous  bands  on  lower  side. 

Structural  Characteristics.  Eyes  large  and  broadly  rounded  on 
upper  margin.  Front  of  head  not  medianly  carinate.  Surface  of 
pronotum  smooth,  lateral  margins  moderately  explanate  and  edges 
thickened,  not  foliaceous.  Underside  of  posterior  tibia  with  inner 
apical  angle  produced  into  a  sharp  point.  Width  of  posterior  tibia 
at  least  two-thirds  that  of  anterior  femur.  Length  of  anterior  femur 
greater  than  that  of  hind  femur.  Outer  margin  of  hind  tibia  curved 
and  inner  margin  nearly  straight.  Metaxyphus  sharply  pointed 
posteriorly,  with  anterior  end  abruptly  elevated  into  a  point.  Apex 
of  female  operculum  with  two  sharp  points,  male  operculum  with- 
out points. 

Comparative  Notes.  This  species  differs  from  L.  grandis  (Lin- 
naeus) in  having  the  front  femora  longer  than  hind  femora  and  in 
having  different  color  markings  on  face  and  abdomen. 

Types.  Francis  Huntington  Snow  Entomological  Collection  of 
University  of  Kansas,  ^  holotype,  and   $   allotype. 

Data  on  Distribution.  Brazil:  Ypirango,  S.  A.,  S.  Paulo,  2-7-24, 
R.  Spitz;  Sao  Paulo,  Brazil,  S.  A.,  1922,  Alfredo  Faz;  S.  Paulo, 
Brazil. 


CuMMiNGs:  The  Giant  Water  Bugs  211 

Lethocerus  mello-leitaoi  de  Carlo,  1933 

1933.  Lethocerus  mcllo-leitaoi  de  Carlo.  Boletim  do  Muscu  Nacional,  Vol.  IX,  N.  1,  pp. 
93-95. 

Original  description  has  not  been  printed  at  the  date  of  this 
writing. 

Size.  Medium-sized  species.  Length,  60  mm. -71  mm.;  width, 
22.5  mm.-27  mm. 

Shape.  Body  elongate  oval  with  lateral  margins  of  hemelytra 
nearly  straight.  Pronotum  convex  with  lateral  margins  slightly 
curved. 

Color.  General  color  medium  brown.  Pronotum  brown,  a  median 
pair  of  light-brown  lines  arising  on  anterior  margin  and  extending 
back  about  one-third  the  length  of  anterior  lobe.  Scutellum  with 
large  fuscous  rectangular  area.  Hemelytra  brown  with  distal  third 
lighter.  Abdomen  light  brown  and  thickly  tesselate  with  fuscous 
spots.  Mesosternum  with  large  irregular  fuscous  spot  on  midline, 
and  large  irregular  fuscous  markings  on  either  side  of  median  spot. 
Front  legs  not  marked,  middle  and  hind  femora  each  with  three 
broad  transverse  fuscous  stripes  on  ventral  side;  middle  and  hind 
tibiae  almost  entirely  fuscous  on  ventral  side. 

Structural  Characteristics.  Front  of  head  without  a  longitudinal 
median  carina.  Pronotum  slightly  rough,  but  not  rugose.  Width 
of  interocular  space  about  two-thirds  width  of  eye.  Interocular 
space  is  about  equal  to  width  of  hind  tarsus.  Length  of  first  seg- 
ment of  front  tarsus  one-half  that  of  second  on  outer  margin. 
Underside  of  posterior  tibia  with  inner  apical  angle  not  produced 
into  a  sharp  point.  Female  operculum  with  two  sharp  points  at 
apex,  male  operculum  without  points.  Metaxyphus  sharply  pointed 
posteriorly.  Outer  margin  of  hind  tibia  no  more  curved  than  inner 
margin. 

Comparative  Notes.  This  species  differs  from  L.  uhleri  Mon- 
tandon  in  having  the  interocular  space  about  equal  to  width  of  hind 
tibia,  and  from  L.  del  pontei  de  Carlo  in  having  the  outer  margin 
of  hind  tibia  no  more  curved  than  inner  margin,  and  in  having 
different  color  markings  on  lower  side  of  abdomen,  tibia,  and  meso- 
sternum. 

Types.  Holotype  male  and  allotype  female  in  Francis  Hunting- 
ton Snow  collection  of  the  LTniversity  of  Kansas.  One  paratype 
in  National  Museum  of  Natural  History,  Buenos  Aires. 

Data  on  Distnbution.  Brazil:  St.  Catharina;  Blumenau,  Brazil. 
Paraguay:    Villarica,  1931,  Fr.  Schade. 


212  The  University  Science  Bulletin 

Lethocerus  truncatus  sp.  nov.* 

Size.    Medium-sized  species.    Length,  74  mm. ;  width,  28  5  mm. 

Shape.  Body  elongate  oval  with  lateral  margins  of  hemelytra 
slightly  dilated.  Pronotum  convex  with  lateral  margins  slightly 
curved. 

Color.  General  color  medium  brown.  Anterior  lobe  of  pronotum 
slightly  darker  than  posterior  lobe,  with  no  distinct  markings 
present.  Scutellum  medium  brown.  Hemelytra  of  uniform  me- 
dium tan  throughout.  Under  surface  of  a  uniform  medium  tan 
throughout.  Middle  and  hind  femora  each  with  three  transverse 
fuscous  bands  on  lower  side. 

Structural  Characteristics.  Front  of  head  without  a  longitudinal 
median  carina.  Pronotum  smooth.  Width  of  interocular  space  is 
greater  than  that  of  hind  tarsus  and  is  equal  to  three-fourths  width 
of  eye.  Length  of  first  segment  of  front  tarsus  two-thirds  that  of 
second  on  outer  margin.  Underside  of  posterior  tibia  with  inner 
apical  angle  not  produced  into  a  sharp  point.  Female  operculum 
minutely  bidentate  at  apex.  Metaxyphus  with  posterior  margin 
straight  and  not  pointed.  Lateral  margins  of  posterior  tibia  nearly 
straight. 

Comparative  Notes.  This  species  differs  from  all  other  species 
of  Lethocerus  in  having  the  posterior  end  of  metaxyphus  not 
pointed. 

Types.  In  Francis  Huntington  Snow  Entomological  Collection  of 
the  University  of  Kansas  (single  specimen,  holotype  female). 

Data  on  Distribution.  Argentina:  Argentina,  S.  A.,  Fives  Lille, 
Santa  Fe,  1923,  A.  Bruch. 

*  Since  Mr.  Cummings  died  I  have  received  a  paper  by  Mr.  Jose  A.  de  Carlo  in  which  he 
describes  a  specimen  takin  by  Mr.  Carlos  Bruch  from  State  of  Parana,  Brazil,  under  the 
name  L.  bruchi.  His  single  specimen  is  a  male  67  mm.  long  and  26  mm.  wide.  The  descrip- 
tion is  published  in  "Revista  de  la  Sociedad  Entomologica  Argentina  (No.  4,  Julio  31  de 
1931)"  pp.  217-218  and  photograph.  The  description  fits  Mr.  Cummings'  species  very  well. 
Mr.  de  Carlo  does  not  mention  the  unusual  shape  of  the  metaxyphus  described  by  Cunmiings 
so  for  the  present  I  permit  the  species  to  stand. — H.   B.   Hungerford. 


CuMMiNGs:  The  Giant  Water  Bugs  213 

Lethocerus  uhleri  (Montandon),  1896 

1896.  Belostoma  uhleri  Mont.     Ann.  Soc.  Ent.  Belg.,  XL,  p.  513. 

1905.  Belostoma  uhleri  Mont.     Howard,  Insect  Book,  pi.   29,  fig.   25. 

1907.  Belostoma  uhleri  Mont.     Bueno  and  Brimley,  Ent.  News,  XVIII,  p.   434. 

1909.  Amorgius  uhleri  (Mont.).     Van  Duzee,  Bui.  Buf.  Soc.  Nat.  Sci.,  IX,  p.   184. 

1914.  Amorgius  uhleri  (Mont.).     Barber,  Bui.  Am.  Mus.  Nat.  Hist.,  XXXIII,  p.  498. 

1926.  Lethocerus  uhleri  (Mont.).     Blatchley,  Het.  of  East.  N.  A.,  p.  1043. 

The  following  also  refer  to  Lethocerus  uhleri  Montandon: 

187G.    Beloi-toma  americanum  Uhler.     Bui.  U.  S.  Geol.  Geog.  Sun-.,  I,  pi.  21,  fig.  38.    1876. 

Size.  Small  species.  Maximum  length,  53  mm.,  maximum  width, 
181/2  mm.;  minimum  length,  40  mm.,  minimum  width,  141/2  mm. 

Sha'pe.  Body  elongate  oval  with  lateral  margins  of  hemelytra  al- 
most parallel.    Thorax  convex  with  lateral  margins  slightly  curved. 

Color.  General  color  medium  brown.  Pronotum  brown  with  a 
median  pair  of  light-brown  lines  arising  on  anterior  margin  and 
extending  back  about  one-third  the  length  of  anterior  lobe.  Scutel- 
lum  with  large  fuscous  rectangular  area.  Hemelytra  brown  with 
distal  third  lighter.  Abdomen  light  tan.  Middle  and  posterior 
tibiae  and  femora  each  with  three  transverse  fuscous  stripes  on  ven- 
tral side.    Front  legs  indistinctly  marked. 

Structural  Characteristics.  Front  of  head  not  medianly  carinate. 
Interocular  space  equal  to  two-thirds  width  of  eye.  Length  of  first 
segment  of  front  tarsus  one-half  that  of  second  on  outer  margin. 
Interocular  space  one-fifth  greater  than  the  width  of  hind  tarsus. 
Underside  of  posterior  tibia  with  inner  apical  angle  not  produced 
into  a  sharp  point.  Female  operculum  with  two  sharp  points  at 
apex;  male  operculum  without  points.  Metaxyphus  sharply  pointed 
posteriorly. 

Comparative  Notes.  This  species  differs  from  L.  melloleitaoi  de 
Carlo  and  L.  del  pontei  de  Carlo  in  being  smaller  and  in  having  the 
interocular  space  one-fifth  greater  than  width  of  hind  tarsus. 

Data  on  Distribution.  Massachusetts:  Sanborn.  Georgia:  Way- 
cross,  11-9-09.  Kansas:  Douglas  Co.,  May,  H.  B.  Hungerford; 
Douglas  Co.,  July;  Douglas  Co.,  May  21,  1920,  H.  B.  Hungerford; 
Douglas  Co.,  W.  Hoffman,  May  21,  1919;  Riley  Co.  Florida: 
Archer,  R.  H.  Beamer,  7-31-30;  Vero  Beach,  E.  M.  Becton,  Sept.  26, 
1927.  Louisiana:  New  Orleans,  Chickering,  4-30-28.  Texas:  Colo- 
rado Co.,  Mrs.  Grace  Wiley,  5-6-1922,  5-10-22;  Galveston.  F.  H. 
Snpw,  May.    Mexico:   Metamoras.  J.  L.  Leconte. 


15—3482 


214  The  University  Science  Bulletin 

Key  to  Species  of  Diplonychus 

There  are  evidently  some  undescribed  species  of  this  genus  which 
need  further  study.  Mr.  Bueno  has  given  the  following  table  for 
separating  the  species : 

A.    Anterior  tarsus  furnished  with  two  claws  of  equal  length D.  rectus  Mayr. 

(W.  Africa.) 
AA.    Anterior  tarsus  furnished  with  two  claws  of  unequal  length. 

B.    Disk  of  prothorax  punctate,  with  two  pronounced  round  fovese,  hemelytra  more 

or  less  punctate D.   punctatus   Stal. 

(Madagascar.) 
BB.    Disk   of  prothorax  slightly  punctate,   with   two   shallow   fove»   and   two   sulci 
converging  posteriorly  toward  the  transverse  sulcus,  hemelytra  impunctate. 

D.  columbim  Spinola. 
(Africa.) 

Mr.  Bueno  gives  D.  colurnbioe  as  from  Africa  and  questions  west- 
ern records.  Doctor  Hungerford,  who  examined  the  specimens  in 
the  Museum  at  Vienna,  says:  "Under  Hydrocyrius  colnmbioe  Spin, 
there  are  19  specimens  from  Africa  and  two  others,  one  labeled  "C. 
Ainer.  Ca  884"  and  the  other  "Mex.  Col!.  Sign."  "herculeus  det. 
Dufour."  "ColumbisB  det.  Mayr."  These  two  are  the  same  as  those 
from  Dutch  East  Africa.  They  have  two  claws  on  front  tarsus  and 
are  75  mm.  long.  The  outer  claw  is  a  little  more  than  half  as  long 
as  the  other. 


216  The  University  Science  Bulletin 


PLATE  XVIII* 

Fig.  1.  Lethocerus  delpontei  de  Carlo.  Under  side  of  posterior  tibia  and 
tarsus.  It  illustrates  "the  posterior  tibia  with  the  inner  apical  angle  not  pro- 
duced into  a  sharp  point."    Compare  with  figure  6. 

Fig.  2.  Lethocerus  sp.  Drawn  to  show  "upper  margin  of  eye  rounded  as 
viewed  from  the  front." 

Fig.  3.  Lethocerus  sp.  Drawn  to  show  "upper  margin  of  eye  straight  as 
viewed  from  the  front." 

Fig.  4.  Lethocerus  truncatus.  The  metaxyphus  distinguishes  this  species 
from  the  others  which  are  like  figure  5. 

Fig.  5.   Lethocerus  angustipes.    The  metaxyphus  is  of  usual  form. 

Fig.  6.  Lethocerus  angustipes.  Under  side  of  posterior  tibia  and  tarsus.  It 
illustrates  "the  posterior  tibia  with  the  inner  apical  angle  produced  into  a 
sharp  point. 

*  The  drawings  of  this  plate  were  prepared  for  me  by  Dr.  Kathleen  Doering. — H.  B. 
Httngerpord. 


CuMMiNGs:  The  Giant  Water  Bugs 
PLATE  XVni 


217 


1.  L.  delpontei — under  side  of  posterior  leg. 


2.  Head — eyes  rounded. 


3.  Head — eyes  straight. 


4.  L.  truncatus— metaxyphus. 


5.  L.  angustipes— metaxyphus 


6.  L.  angustipes — under  side  of  posterior  leg. 


218  The  University  Science  Bulletin 


PLATE  XIX 

The  drawings  on  this  plate  illustrate  the  shape  of  the 
last  ventral  abdominal  segment  of  females  of  Lethocerus. 


CuMMiNGs:  The  Giant  Water  Bugs  21U 


PLATE  XIX 


L,  americanus. 


2.  L.  delpontei. 


3.  L.  annulipe? 


4.  L.  uhlen. 


6.  L.  laigus. 


7  L.  camposi 


10.  L.  graiidis. 


5.  L  angustipes. 


».  L.  truncatus. 


y.  L.  callosicus. 


U.  L.  dilatus. 


THE  UNIVERSITY  OF  KANSAS 

SCIENCE  BULLETIN 

Vol.  XXL]  March,  1933.  [No.  3. 


A  Study  of  the  Genus  Brachymetra 
(Hemiptera — Gerridae) 

J.  GILBERT  SHAW,  Lawrenre,  Kansas* 


Abstract:    This  paper  gives  a  key  for  separation  of  five  species  and  one 
subspecies.    Three  species  and  the  subspecies  are  described  as  new. 


THE  family  Gerridae  is  divided  into  two  subfamilies— the  Ger- 
rinai  and  the  Halobatina.     Among  the  numerous  genera  of  the 
latter  belongs  the  genus  Brachymetra,  a  new  world  genus. 

DESCRIPTION  OF  THE  GENUS 

All  members  of  the  genus  Brachymetra  possess  a  triangular  and 
convex  head.  The  almost  oval  eyes  extend  obliquely  laterad  of 
anterior  angles  of  the  pronotum.  The  rostrum  extends  well  beyond 
anterior  coxse.  The  antennsB  are  over  one-half  the  length  of  the 
body,  all  segments  round  and  straight  except  the  basal,  which  is 
definitely  arched,  basal  segment  longer  than  the  head,  second  seg- 
ment shorter  than  the  head,  third  segment  invariably  longer  than 
the  second  segment,  and  apical  segment  of  about  same  length  as 
second.  Pronotum  possesses  medially  near  anterior  margin  a  shal- 
low horseshoe-shaped  impression;  single  faint  carination  runs  ob- 
liquely and  posteriorly  of  each  humeral  angle.  Lateral  margins  of 
mesothorax  strongly  divergent  in  both  sexes.  The  prothoracic  legs 
are  shorter  than  the  body  and  have  rather  stout  femora,  which  bear 
a  few  stout  hairs  beneath,  while  the  meso-  and  metathoracic  legs  are 
very  long  and  slender  with  the  frail  femora  alone  about  as  long  and 
longer,  respectively,  than  the  body.    The  elytra  exceed  the  abdomen. 

*  Contribution  from  the  Department  of  Entomology,  University  of  Kansas. 

(221) 


222  The  University  Science  Bulletin 

Abdomen  moderately  short,  sixth  segment  unarmed,  apical  margin 
on  ventral  side  of  male  emarginate,  while  that  of  female  is  almost 
straight.  Claspers  inserted  on  each  side  of  genital  capsule  of  male. 
Body  covered  by  a  dull  pubescence.  All  the  species  possess  on  meso- 
and  metathoracic  acetabula  a  heavier  pubescence,  which  is  generally 
silvery. 

Key  to  the  Species  of  Brachymetra 

A.     Species  with  first  antennal  segment  as  long  or  longer  than  the  two  following;    pro- 
notum  striped  with   black. 

B.    Pronotum  with  median  lioe  black unca  sp.  n. 

BB.    Pronotum  median  line  not  black  but  bordered  by  a  pair  of  black  bands. 

vittata  sp.  n. 
AA.    Species    with    first    antennal    segment    shorter    than    the    two    following;    pronotum, 
except  margin,  more  or  less  unicolorous. 

B.    Apex  of  pronotum  acutely  angulate,  extending  well  beyond  tip  of  mesothoracic 

acetabula   kleopatra  Kirk. 

BB.    Apex    of    pronotum    rather    bluntly    angulate,    not    extending    beyond    tip    of 
mesothoracic  acetabula. 
C.    Anterior  femur  only  slightly  larger  tlian  intermediate  femur;    rear  margin 

of  eye  definitely  greater  than  interocular  space lata  sp.   n. 

CC.    Anterior  femur  at  least  one  and   one-half  times  larger  than  intermediate 
femur ;   rear  margin  of  eye  equal  to  or  less  than  interocular  space. 
D.    First    antennal    segment    plainly    surpassing    rear    lateral    margin    of 
eye    when    turned    back;    first    abdominal    tergite    shallowly    incised 

behind    albinerva   (A.    &   S.). 

DD.  First  antennal  segment  just  about  attaining  rear  lateral  margin  of 
eye  when  turned  back ;  first  abdominal  tergite  deeply  incised 
behind    albinerva    incisa   sub-sji.    n. 

Brachymetra  unca  sp.  n. 

(PI.  1,  figs.  2,  2a,  2b,  2c) 

Apterous  Form 

Size.  Length  of  males  6.5  mm.,  females  6.1mm.;  width  of  males 
2.5  mm.,  females  2.8  mm.  at  widest  place. 

Color.  Chocolate  brown  above,  venter  yellowish  brown,  dull 
pubescence  with  a  varnished  background.  A  median  dark  line  ex- 
tending throughout  the  length  of  the  pronotum,  metanotum  and  the 
first  abdominal  segment.  Margins  of  pronotum  and  connexiva 
dark.  Caudal  margin  of  last  abdominal  segment  of  male  dark,  with 
the  genital  segments  somewhat  dark;  caudal  margin  of  last  ab- 
dominal segment  and  genital  segments  of  female  dark.  Meso-  and 
metathoracic  acetabula  dark  above.  Tip  of  rostrum,  eyes,  antennae, 
and  legs  dark,  except  the  anterior  coxae,  trochanter  and  basal  half 
of  the  femora,  which  are  about  same  shade  as  venter. 

Structural  Characteristics.  Antennal  fonnula  of  male:  1st  :  2d  : 
3d  :  4th  : :  30  :  11  :  18*  :  15;  female  :  26  :  10  :  16.5  :  14.  Eyes  rather 

*  Includes  tiny  basal  ring  segment. 


Shaw:  The  Genus  Brachymetra  223 

small  in  comparison  to  size  of  body,  less  than  one-half  the  length 
extending  caudad  of  the  posterior  angle  of  the  head.  Interocular 
space  about  four-fifths  the  length  of  eye.  Sides  of  prothorax  almost 
parallel  with  the  sides  of  mesothorax  inflated.  Length  of  pronotum 
on  median  line  is  to  length  of  metanotum  exposed  as  6.8  :  1  (both 
sexes).  Anterior  femur  as  long  as  the  pronotum  on  its  median 
dorsal  line;  in  the  male  this  femur  is  fully  twice  as  large  as  inter- 
mediate femur  and  distinctly  arched,  while  in  the  female  it  is  about 
one  and  one-half  times  as  large  as  intermediate  femur  and  quite 
straight;  tibia  slightly  shorter  than  femm-,  somewhat  curved  on 
distal  third,  its  tip  slightly,  if  any,  produced  beyond  base  of  tarsus; 
first  tarsal  segment  one-half  as  long  as  the  second.  Intermediate 
femur  a  trifle  shorter  than  body;  tibia  about  one-seventh  longer 
than  the  femur  and  over  four  times  as  long  as  tarsus;  first  tarsal 
segment  three  times  as  long  as  the  second.  Posterior  femur  about 
one-fifth  longer  than  intermediate  femur  and  almost  twice  as  long 
as  tibia;  tibia  almost  four  times  as  long  as  its  tarsus;  first  tarsal 
segment  over  one  and  one-half  times  as  long  as  the  second.  In 
the  female  the  last  abdominal  segment  surpasses  the  apex  of  pos- 
terior trochanter.  In  the  male  the  genital  segments  are  one-half  to 
three-fifths  the  length  of  the  abdomen,  the  abdomen  surpassing  the 
posterior  trochanter  by  about  one-third  the  length  of  the  last  seg- 
ment; the  last  ventral  abdominal  segment  as  long  as  the  two  pre- 
ceding combined  in  the  male  and  as  long  or  longer  than  the  three 
preceding  combined  in  the  female.  In  the  male  the  claspers  are  so 
long  and  curved  that  they  overlap  slightly  on  the  dorsal  side  of  the 
second  genital. 

Described  from  a  series  of  twenty  specimens  labeled,  "Trinidad, 
B.  W.  I.,  Port  of  Spain,  Nov.  5,  1931,  W.  E.  Broadway,"  and  the 
same  data  "near  St.  Patrick  Oct.  25,  1931."  Holotype,  allotype 
and  paratypes  in  the  Francis  Huntington  Snow  Entomological 
Museum,  University  of  Kansas,  Lawrence,  Kansas. 

Macropterous  Form 
Size.    Length  to  tip  of  elytra,  7.3  mm.;  width,  2.5  mm. 
Color.    Same  color  as  in  apterous  form.    A  median  dark  line  on 

pronotum,  extending  from  anterior  margin  to  beyond  the  middle. 

Elytra,  including  veins,  black  with  apices  somewhat  fuscous. 
Structural  Characteristics.    Lateral  margins  of  pronotum  back  of 

humeral  angles  more  depressed  than  in  related  species,  depressed 

area  at  apex  about  equals  the  head  in  length.    Anterior  femur  one- 


224  The  University  Science  Bulletin 

fifth  shorter  than  pronotum.    Elytra  exceed  abdomen  by  over  half 
the  length  of  the  head. 

Described  from  a  single  male  specimen  labeled,  "Trinidad,  B.  W. 
I.,  Port-  of  Spain,  Nov.  5,  1931,  W.  E.  Broadway."  Holomorphotype 
deposited  in  the  Francis  Huntington  Snow  Entomological  Museum. 
University  of  Kansas,  Lawrence,  Kansas. 

Brachymetra  vittata  sp.  n. 

(PI.  1,  figs.  3,  3a) 

Maceopterous  Form 

Size.  Length  of  male,  9.3  mm.,  female,  9  mm.;  width  of  male, 
3.3  mm.,  female,  3.3  mm. 

Color.  A  striped  species;  clay-yellow  with  head  almost  reddish- 
brown,  venter  yellowish.  Heavy  black  band  bordering  lateral  mar- 
gins of  pronotum  and  continuing  slightly  inside  the  margin  from 
the  humeral  angles  to  the  apex,  a  pair  of  heavy  black  bands  ex- 
tending from  anterior  end  two-thirds  the  way  along  the  median  line. 
Margins  of  connexiva  dark.  Anterior  acetabulum  with  black  spot 
on  dorsal  side,  a  black  spot  on  lateral  side  of  prothorax  dorsad  and 
caudad  of  this.  An  oblique  black  band  running  slightly  ventrad 
along  lateral  side  of  mesothorax,  with  slight  break  which  is  followed 
by  a  broader  band  on  the  mesothoracic  acetabulum.  A  solid  band 
also  originates  posterior  to  the  mesothoracic  spiracle  and  continues 
unbroken  to  apex  of  metathoracic  acetabulum.  Two  black  spots  are 
just  below  the  spiracles  of  each  abdominal  segment  of  female.  A 
black  spot  on  vertex  next  to  each  eye  continuous  with  median  bands 
of  pronotum,  a  third  spot  on  median  line  about  half  way  between 
these  and  rostrum.  Eyes,  antennae  and  tip  of  rostrum,  dark.  An- 
terior femur  with  apex  and  ventral  side  dark,  tibia  and  tarsus  also 
dark.  Meso-  and  metathoracic  legs  somewhat  dark  with  apex  of 
mesothoracic  femur  black  preceded  preapically  by  light  brown. 
Elytra  including  veins  black;  caudal  half  of  costal  margin  light 
brown. 

Structural  Characteristics.  Antennal  formula  of  female:  1st  :  2d  : 
3d  :  4th  : :  44  :  18  :  23  :  20  (no  antennae  on  male  specimen).  Eyes 
quite  small  in  comparison  to  size  of  body,  less  than  one-half  the 
length  extending  caudad  of  the  posterior  angle  of  the  head.  Inter- 
ocular  space  three-fourths  the  length  of  the  eye.  Apex  of  pronotum 
blunter  than  in  other  species,  margin  narrowly  depressed.  Anterior 
femur  not  as  stout  as  in  B.  unca  but  arched  apically  (no  prothoracic 
or  right  meso-  and  metathoracic  legs  present  on  the  male),  equal 
in  length  to  the  pronotum  on  its  median  line;  tibia  almost  as  long 


Shaw:  The  Genus  Brachymetra  225 

as  femur  and  somewhat  curved  apically,  its  tip  produced  slightly 
beyond  base  of  tarsus;  first  tarsal  segment  one-half  to  three-fifths 
as  long  as  the  second.  Intermediate  femur  a  trifle  longer  than  body 
in  female,  and  about  one-fourth  longer  in  the  male;  tibia  slightly 
shorter  than  the  femur  and  almost  three  times  as  long  as  tarsus; 
first  tarsal  segment  about  three  times  as  long  as  second.  Posterior 
femur  barely  longer  than  intermediate  femur  and  two-thirds  longer 
than  the  tibia;  tibia  over  four  times  as  long  as  tarsus;  first  tarsal 
segment  about  one-fourth  longer  than  second.  In  the  female  the 
last  two  abdominal  segments  exceed  tip  of  posterior  trochanter. 
In  the  male  the  genital  segments  only  one-half  longer  than  the 
posterior  abdominal  segment,  the  abdomen  surpassing  the  posterior 
trochanter  by  about  the  length  of  the  last  segment.  The  last  ven- 
tral abdominal  segment  about  one  and  one- fourth  times  as  long  as 
preceding  one  in  the  male  and  over  one  and  one-half  times  as  long  as 
preceding  one  in  the  female.  In  the  male  the  claspers  do  not  exceed 
the  posterior  margin  of  the  first  genital  and  are  scarcely  visible  be- 
tween the  first  and  second  genital.  Elytra  exceed  abdomen  by  over 
one-half  the  length  of  head. 

Described  from  single  male  and  female  specimens  labeled,  "Mari- 
anie  Port  au  Prince,  Haiti,  July  10,  1930,  R.  M.  Bond."  Holotype 
and  allotype  in  the  Francis  Huntington  Snow  Entomological  Mu- 
seum, University  of  Kansas,  Lawrence,  Kansas. 

Apterous  Form 

Size.    Length  of  females,  8.4  mm. ;  width,  3.3  mm. 

Color.  Color  markings  mostly  as  in  the  macropterous  forms. 
Median  bands  on  pronotum  almost  continuous  to  base.  Wide,  black 
band  originating  laterad  of  median  bands  of  pronotum  and  extend- 
ing posteriorly,  bifurcating  preapically  at  the  base  of  the  pronotum, 
one  branch  continuous  with  the  black  margin  of  connexivum  and  the 
other  running  around  apex  of  pronotum  and  meeting  band  from 
other  side  at  the  median  line  of  the  tergum  of  the  metathorax. 
Terga  of  abdominal  segments  mostly  black. 

Structural  Characteristics.  Same  as  in  the  macropterous  forms 
with  the  exception  that  the  apex  of  the  pronotum  is  less  blunt,  ex- 
posing only  the  posterior  margin  of  the  mesothorax;  lateral  margins 
more  parallel  and  lacking  the  rather  acute  humeral  angles. 

Described  from  four  female  specimens  labeled,  "Marianie  Port  au 
Prince,  Haiti,  July  10,  1930,  R.  M.  Bond."  Holomorphotype  and 
paramorphotypes  in  the  Francis  Huntington  Snow  Entomological 
Museum,  University  of  Kansas,  Lawrence,  Kansas. 


226  The  University  Science  Bulletin 

Brachymetra  kleopatra  Kirk. 

(PI.   1,  fig.  5) 
Brachymetra  kleopatra  Kirkaldy.     Bolletino  dei  Musei  di   Zoologia  Anatomia  Comparata, 
Torino,  vol.  XIV,  No.  350,  1899,  p.  3. 

Kirkaldy's  original  description  of  this  species  fits  the  single  female 
I  have  on  hand  with  the  following  exceptions:  (1)  The  specimen 
which  I  have  before  me  measures  8.2  mm.  long  and  3  mm.  wide, 
while  he  gives  measurements  of  9%  mm.  long  and  2.5  mm.  wide. 
(2)  "Head  and  pronotum  very  smooth  and  polished."  I  find  both 
to  be  quite  pubescent  except  for  smooth  spot  on  vertex  between  the 
eyes  (apparently  having  been  rubbed  smooth). 

The  specimen  at  hand  is  lacking  antennal  segments  and  anterior 
legs.    I  will  quote  Mr.  Kirkaldy's  descriptions  of  these  parts. 

Color.  Reddish-brown  above,  venter  tan.  Tip  of  meso-  and 
metathoracic  acetabula  dark.  Two  darkened  spots  just  below  spir- 
acle of  each  abdominal  segment.  Eyes  dull  wine,  rostrum  tip  dark, 
and  the  legs  dark  brown.  Elytra  velvety  brown  with  veins  grayish 
brown. 

Structural  Characteristics.  "First  segment  of  the  antennae  shorter 
than  the  second  and  third  together,  first  segment  rather  more  than 
twice  as  long  as  the  second,  second  and  fourth  subequal,  third 
one-third  longer  than  the  second."  Eyes  rather  small  in  comparison 
to  size  of  body,  about  one-half  the  length  extending  caudad  of  the 
posterior  angle  of  the  head.  Interocular  space  about  four-fifths  the 
length  of  eye.  The  apex  of  pronotum  most  widely  depressed  and 
most  acutely  produced  of  all  the  species  of  the  genus,  being  pro- 
duced well  beyond  the  apices  of  metathoracic  acetabula.  "Anterior 
tibia  one-fourth  longer  than  the  femur,  three  and  a  quarter  times 
longer  than  tarsus,  second  segment  of  the  latter  two-thirds  longer 
than  the  first."  Intermediate  femur  about  one-tenth  shorter  than 
body;  tibia  slightly  longer  than  femur  and  three  times  as  long  as 
tarsus ;  first  tarsal  segment  about  five  times  as  long  as  second.  Pos- 
terior femur  equals  the  length  of  the  body  and  one-tenth  longer  than 
intermediate  femur  and  almost  twice  the  length  of  the  tibia;  tibia 
three  times  as  long  as  tarsus;  first  tarsal  segment  over  twice  the 
second.  The  last  two  abdominal  segments  exceed  the  posterior  tro- 
chanter. Last  ventral  abdominal  segment  almost  as  long  as  two 
preceding.  Elytra  exceed  abdomen  by  one-half  the  length  of  head. 
The  single  female  specimen  on  hand  is  labeled,  "British  Guiana, 
S.  A.,  near  New  Amsterdam,  July  30,  1923,  F.  X.  Williams." 


Shaw:  The  Genus  Brachymetra  227 

Brachymetra  lata  sp.  n. 

(PI.  XX,  figs.  4,  4a,  4b) 

Apterous  Form 

Size.  Length  of  males,  7  mm.,  females,  7  mm.;  width  of  males, 
2.5  mm.,  and  females,  2.7  mm. 

Color.  Reddish-brown  above,  venter  a  lighter  shade  of  brown. 
Margins  of  pronotum  dark,  median  line  light.  Margins  of  con- 
nexiva  dark.  Genital  segments  mostly  dark,  caudal  half  of  the 
last  abdominal  tergum  of  male  dark.  Tip  of  metathoracic  acetabula 
dark  while  that  of  the  mesothorax  is  light  with  a  brown  spot  borne 
preapically.  Eyes  brown  mottled  with  dark  spots,  tip  of  rostrum, 
antennae  and  legs  dark,  except  anterior  coxse,  trochanters  and  basal 
half  of  anterior  side  of  femora. 

Structw'al  Characteristics.  Antennal  formula  of  male :  1st  :  2d  : 
3d  :  4th  : :  29  :  15  :  22.5  :  16;  female:  27  :  14  :  20  :  15.  Eye  about 
one  and  one-half  times  longer  than  wide,  larger  in  comparison  to  size 
of  body,  over  one-half  the  length,  with  its  rear  margin  slightly 
greater  than  interocular  space,  extending  caudad  of  the  posterior 
angle  of  the  head.  Interocular  space  about  three-fifths  the  length 
of  eye.  Head,  including  eyes,  considerably  wider  than  prothorax. 
Pronotum  with  apex  bluntly  produced,  length  on  median  line  is  to 
length  of  metanotum  exposed  as  5.6  :  1  (both  sexes).  Anterior 
femur  not  much  stouter  than  intermediate  femur,  but  somewhat 
arched  apically,  slightly  longer,  if  any,  than  pronotum  on  its  median 
line.  Tibia  shorter  than  femur  and  barely  curved  on  its  distal  end, 
its  tip  produced  slightly  beyond  base  of  tarsus;  first  tarsal  segment 
about  two-thirds  as  long  as  second.  Intermediate  femur  about  equal 
in  length  to  body  of  male,  but  almost  one-tenth  shorter  in  the  fe- 
male; tibia  equal  to  length  of  femur  in  male,  but  slightly  longer  in 
female,  and  five  and  one-half  to  six  times  as  long  as  tarsus;  first 
tarsal  segment  three  times  as  long  as  second  in  male  and  three  and 
one-half  times  as  long  in  female.  Posterior  femur  one-seventh 
longer  than  intermediate  femur  and  over  twice  the  length  of  its 
tibia;  tibia  four  and  one-half  times  longer  than  its  tarsus;  first 
tarsal  segment  about  one-third  longer  than  the  second.  In  the  fe- 
male the  last  two  abdominal  segments  exceed  tip  of  posterior  tro- 
chanter. In  the  male  the  genital  segments  are  about  one-fourth  the 
length  of  the  abdomen,  the  abdomen  surpassing  the  posterior  tro- 
chanter by  the  last  segment.  Last  ventral  abdominal  segment  about 
four-fifths  as  long  as  the  two  preceding  combined  in  the  male  and  as 


228  The  University  Science  Bulletin 

long  in  the  female.    A  pair  of  rather  straight  claspers  barely  exceed, 
if  any,  posterior  margin  of  first  genital. 

Described  from  a  series  of  160  labeled,  "Manacapimi,  S.  A., 
Amazonas,  Brazil,  Solimoes  river,  6.  26,  S.  M.  Klages."  Holotype, 
allotype  and  paratypes  in  the  Francis  Huntington  Snow  Entomo- 
logical Museum,  University  of  Kansas,  Lawrence,  Kansas. 

Macropterous  Form 

Size.  Length  of  males  to  tip  of  elytra,  7.4  mm.,  females,  7.4  mm. ; 
width  of  males,  2.5  mm.  and  females,  2.7  mm. 

Color.  Same  color  as  in  apterous  forms.  Margin  of  pronotum 
indistinctly  darkened.  Elytra  a  rich  velvety  brown,  paler  at  apices, 
veins  a  rich  yellow. 

Structural  Characteristics.  Pronotum  with  apex  about  as  blunt  as 
in  apterous  forms.  Anterior  femur  slightly  shorter  than  pronotum 
on  its  median  line.  Elytra  exceed  abdomen  by  about  half  the 
length  of  head  or  less. 

Described  from  a  series  of  114  with  same  data  as  for  apterous 
forms.  Holomorphotype,  allomorphotype  and  paramorphotypes  in 
the  Francis  Huntington  Snow  Entomological  Museum,  University  of 
Kansas,  Lawrence,  Kansas. 

Brachymetra  albinerva  (A.  &  S.) 

(PI.  XX,  figs.   1,  la,  lb,  Ic.) 
Brachymetra  albinerva  (Amyot  et  Serville).     Hist.   Nat.  Ins.  Hem.,   1843,  p.   412. 

Apterous  Form 

Size.  Length  of  males,  5.4-6.8  mm.,  females,  5.5-6.9  mm.;  width 
of  males,  2.3-2.7  mm.,  and  females,  2.5-3.1  mm. 

Color.  Dull  chocolate  brown  above,  venter  lighter.  Margins  of 
pronotum  and  connexiva  dark.  Genital  segments  dark.  Meso- 
thoracic  and  metathoracic  acetabula  darkened  above.  Eyes  and 
tip  of  rostrum  dark.  Antennae  and  legs  same  shade  as  the  body  or 
a  little  darker  except  the  anterior  femora,  which  are  lighter  on  the 
dorsal  basal  two-thirds. 

Structural  Characteristics.  Antennal  formula  of  male:  1st  :  2d  : 
3d  :  4th  : :  21  :  11  :  15  :  12;  female:  20  :  11  :  15  :  11.5;  in  the  larger 
specimens  it  measures  as  follows,  male:  25  :  14  :  18  :  14;  female: 
23  :  14  :  16  :  15.  Eye  with  its  rear  margin  about  equal  to  interocular 
space,  one-half  the  length  of  eye  extending  caudad  of  the  posterior 
angle  of  the  head.  Interocular  space  two-thirds  the  length  of  eye. 
Head,  including  eyes,  only  slightly  wider  than  prothorax.     Length 


Shaw:  The  Genus  Brachymetra  229 

of  pronotum  on  median  line  is  to  length  of  metanotum  exposed  as 
6.5  :  1  (both  sexes).  Anterior  femur  arched  and  at  least  one  and 
one-half  times  larger  than  intermediate  femur,  about  one-fifth 
shorter  than  pronotum  on  its  median  line;  tibia  slightly  shorter 
than  femur,  curved  on  distal  third,  its  tip  barely  produced  beyond 
base  of  tarsus;  first  tarsal  joint  three-fourths  as  long  as  second. 
Intermediate  femm'  one-ninth  shorter  than  body;  tibia  about  one- 
tenth  longer  than  femur  and  over  three  and  one-half  times  as  long 
as  tarsus;  first  tarsal  segment  slightly  more  than  three  times  as 
long  as  the  second.  Posterior  femur  about  one-tenth  longer  than 
intermediate  femur  and  about  twice  as  long  as  its  tibia.  Tibia 
over  three  times  as  long  as  its  tarsus.  First  tarsal  segment  about 
one  and  one-half  times  as  long  as  the  second.  In  the  female  the 
last  two  abdominal  segments  exceed  the  posterior  trochanter.  In 
the  male  the  genital  capsule  usually  only  slightly  longer  than  the 
posterior  abdominal  segment,  while  in  specimens  with  it  greatly  pro- 
truding it  may  be  one-third  the  length  of  abdomen,  the  abdomen 
surpassing  the  posterior  trochanter  by  about  the  length  of  the  last 
segment.  The  last  ventral  abdominal  segment  one  and  one-half 
times  as  long  as  preceding  in  the  male  and  twice  as  long  as  preceding 
in  the  female.  In  the  male  the  claspers  extend  beyond  first  genital 
and  are  strongly  curved  upward  and  laterad  of  second  genital. 

Specimens  are  at  hand  from  the  following  localities:  Banos,  Mera, 
Tena,  and  Huigra,  Ecuador,  and  Sao  Paulo,  Brazil,  South  America; 
Trinidad,  B.  W.  I.;  and  Fort  Clayton  and  Barro  Colo  Island,  Canal 
Zone. 

Macroptbeous  Form 

Size.  Length  of  males,  6  mm.,  females,  6.1  mm,;  width  of  males, 
2.3  mm.,  and  females,  2.4  mm. 

Color.  Same  color  as  in  apterous  foims.  Elytra  of  the  same 
shade  of  brown  as  the  body  or  darker,  veins  yellowish. 

Structural  Characteristics.  Anter'ior  femur  definitely  shorter 
than  pronotum  on  its  median  line.  Elytra  exceed  abdomen  by  at 
least  one-half  the  length  of  head. 

Specimens  on  hand  from  the  following  localities:  Mera  and 
Huigra,  Ecuador,  South  America:  and  Fort  Clayton  and  Barro 
Colo  Island,  Canal  Zone. 


16—3482 


230  The  University  Science  Bulletin 

Brachymetra  albincrva  incisa  subsp.  n. 
Apterous  Form 

Size.  Length  of  males,  5.4  mm.,  females,  5.6  mm. ;  width  of  males, 
2.4  mm.,  and  females,  2.5  mm. 

Color.  A  subspecies  which  is  yellowish  brown  above,  venter  paler, 
almost  yellowish.  Lateral  margins  of  pronotum  and  connexiva  dis- 
tinctly darkened  by  a  narrow  band,  as  well  as  the  posterior  margins 
of  abdominal  terga.  Meso-  and  metathoracic  acetabula  darkened 
on  apical  margins.  Eyes  and  tip  of  rostrum  dark.  Antennse  and 
legs  about  same  shade  as  the  body  or  a  little  darker,  except  the 
anterior  femora,  which  are  lighter  on  the  dorsal  basal  two-thirds, 
apices  of  femora  and  tibia  darkened. 

Structural  Characteristics.  Structurally  the  same  as  B.  albincrva, 
but  the  following  morphological  characteristics,  along  with  the  color, 
have  made  me  feel  justified  in  calling  this  series  a  subspecies.  An- 
tennal  formula  of  male:  1st  :  2d  :  3d  :  4th  : :  18.5  :  10.5  :  14.5  :  11.5; 
female:  18  :  19  :  14  :  11.  When  the  first  antennal  segment  of  B. 
albincrva  incisa  is  turned  back  it  about  attains  rear  lateral  margin 
of  eye,  while  in  B.  albincrva  it  plainly  surpasses  rear  lateral  margin 
of  eye.  The  first  abdominal  tergite  of  B.  albincrva  incisa  is  deeply 
incised  behind,  while  in  B.  albincrva  it  is  shallowly  incised. 

Macrofterous  Form 

Size.  Length  of  males,  6  mm.,  females,  6.4  mm.;  width  of  males, 
2.3  mm.,  and  females,  2.5  mm. 

Color.  Same  color  as  in  apterous  forms.  Elytra  of  the  same 
shade  of  brown  as  the  body  or  darker,  veins  yellowish. 

Structural  Characteristics.  Anterior  femur  about  one-fourth 
shorter  than  pronotum  on  its  median  line.  Elytra  exceed  abdomen 
by  at  least  one-half  the  length  of  head. 

Both  forms  described  from  a  series  of  37  specimens  labeled,  "Santa 
Cruz,  Bolivia,  S.  A.,  J.  Steinbach."  Holotype,  holomorphotype 
allotype,  allomorphotype,  paratypes,  and  paramorphotypes  in  the 
Francis  Huntington  Snow  Entomological  Museum,  University  of 
Kansas,  Lawrence,  Kansas. 


232  The  University  Science  Bulletin 


PLATE  XX 

Figs.  1,  la,  lb,  Ic.    Brachymetra  albinerva  (A.  &  S.). 

Fig.  1.    Head  and  pronotum  of  macropterous  form. 

Fig.  la.    Apical  end  of  abdomen  showing  genital  segments  and  claspers, 

ventral  view. 
Fig.  lb.   Lateral  view  of  fig.  la. 
Fig.  Ic.    Clasper. 

Figs.  2,  2a,  2b,  2c.   Brachymetra  unca  sp.  n. 

Fig.  2.    Head  and  pronotum  of  macropterous  form. 

Fig.  2a.    Apical  end  of  abdomen  showing  genital  segments  and  claspers, 

ventral  view. 
Fig.  2b.   Lateral  view  of  fig.  2a. 
Fig.  2c.   Clasper. 

Figs.  3,  3a.   Brachytnelra  vittata  sp.  n. 

Fig.  3.    Head  and  pronotum  of  macropterous  form. 

Fig.  3a.   Apical  end  of  abdomen  showing  genital  segments  and  claspers, 
ventral  view. 

P^GS.  4,  4a,  4b.    Brachymetra  lata  sp.  n. 

Fig.  4.    Head  and  pronotum  of  macropterous  form. 

Fig.  4a.    Apical  end  of  abdomen  showing  genital  segments  and  claspers, 

ventral  view. 
Fig.  4b.   Clasper. 

Fig.  5.   Brachymetra  kleopatra  Kirk.     Head  and  pronotum  of  macropterous 
form. 


Shaw:  The  Genus  Brachymetra 
PLATE  XX 


233 


4    lata 


3    vitlata 


5    kleopatra 


THE  UNIYERSITY  OF  KANSAS 

SCIENCE  BULLETIN 


Vol.  XXL] 


March,  1933. 


[No.  4. 


The  Genus  Chlorotettix  in  America  North  of  Mexico 
(Homoptera — Cicadellidse) 

WILLIAM  F.  BROWN,  Lawrence,  Kansas.* 

TABLE  OF  CONTENTS 


PAGE 

The  Genus  Chlorotettix    235 

Key  to  species    238 

C.  angustus    241 

C.    attenuatus    241 

C.  convexus    242 

C.  durus   242 

C.   fuscus    243 

C.   latifrons    243 


PAliK 

latus    244 

macvlosus    244 

pallidus    245 

rotundus    246 

rubidus 246 

sinuostts 247 

sordidus    248 


Abstract:  New  distributional  records  are  given  for  twenty-one  of  the  pre- 
viously know  species  of  the  genus,  in  a  number  of  cases  greatly  extending  their 
known  range.  Twelve  species  are  described  as  new  to  science  and  a  key  to  all 
the  known  species  is  provided. 


MOST  species  of  the  genus  Chlorotettix  are  found  in  North 
America,  north  of  Mexico.  The  genus  was  founded  in  1892 
by  Van  Duzee.  New  species  have  been  added  by  Van  Duzee, 
Osborn,  Ball,  Baker,  Crumb,  Sanders,  De  Long  and  the  author  until 
there  are  at  present  forty-six  species  and  one  variety  north  of 
Mexico.  This  paper  describes  twelve  new  species  and  the  male  of 
C.  latifrons. 

Most  species  of  the  genus  have  a  uniform  green  or  yellow  color; 
a  few  are  tinged  with  fuscous;  a  few  have  dark  spots  or  bands  on 
the  head,  pronotum  or  scutellum. 

The  vertex  is  usually  broad,  either  rounded  or  distinctly  angled 
and  rounding  with  the  front.    The  ocelli  are  on  the  margin  of  the 


•  Submitted  to  the  Department  of  Entomology  and  the  Faculty  of  the  Graduate  School 
of  the  University  of  Kansas  in  partial  fulfillment  of  the  requirements  for  the  degree  of  Master 
of  Arts,  September,  1932. 

(235) 


236  The  University  Science  Bulletin 

vertex  and  near  the  eyes.  The  elytra  have  three  antiapical  cells, 
are  longer  than  the  abdomen,  subhyaline,  with  a  distinct  appendix, 
the  nervures  usually  indistinct.  The  species  vary  in  length  from 
3.5  to  8  mm. 

The  members  of  this  group  usually  feed  on  grasses;  thus  they 
are  plentiful  as  well  as  of  some  economic  importance. 

Some  species  of  the  genus  have  a  wide  range,  while  others  are 
found  within  narrow  limits.  In  the  following  notes  are  additional 
records  of  the  distribution  of  a  number  of  species. 

C.  viridins  has  previously  been  reported  from  the  following  states: 
Connecticut,  New  York,  New  Jersey,  Maryland,  Tennessee,  Virginia, 
North  Carolina,  South  Carolina,  Florida,  Alabama,  Mississippi, 
Louisiana,  Texas,  Arkansas,  Kansas,  Ohio  and  Michigan.  In  the 
Snow  collection  are  also  specimens  from  Georgia.  This  species 
thus  seems  to  be  distributed  well  over  the  eastern  half  of  the  United 
States. 

C.  vividus  has  been  reported  for  Tennessee,  South  Carolina,  Ohio 
and  Kansas.  Specimens  are  at  hand  from  these  additional  states: 
Arizona,  Missouri,  Arkansas,  Mississippi,  Alabama,  Georgia,  Florida, 
and  North  Carolina. 

C.  tergatus  has  formerly  been  reported  for  New  York,  Maine, 
Connecticut,  Michigan,  Wisconsin,  Washington,  Iowa,  Illinois,  Ohio, 
New  Jersey,  Pennsylvania,  Virginia,  Tennessee,  South  Carolina  and 
Florida.  Specimens  are  at  hand  from  Minnesota.  Specimens  for- 
merly called  C.  tergatus  examined  from  the  southern  states  proved 
to  be  either  C.  fumidus  or  C.  divergens.  It  is  doubtful,  therefore, 
whether  the  records  of  C.  tergatus  from  such  states  as  Virginia, 
Tennessee,  South  Carolina  and  Florida  are  correct. 

C.  dentatus  was  described  by  Sanders  and  De  Long  from  speci- 
mens taken  near  Washington,  D.  C.  In  the  Snow  Entomological 
Collection  is  a  specimen  collected  at  Atherton,  Missouri. 

C.  lusorius  has  previously  been  reported  from  Iowa,  Ohio,  New 
York,  New  Jersey,  Pennsylvania,  New  Hampshire,  Connecticut, 
Virginia,  Wisconsin,  Colorado  and  Utah.  To  these  may  be  added 
Minnesota,  Illinois  and  Arizona. 

C.  limosus  was  described  from  specimens  collected  in  Pennsyl- 
vania. Before  us  are  also  specimens  from  Kansas,  Missouri,  Mis- 
sissippi, Alabama  and  New  York. 

C.  balli  has  been  reported  from  Iowa,  Ohio,  Tennessee,  South 
Carolina,  Virginia,  New  York  and  Connecticut.  Our  specimens 
have  been  taken  in  Kansas  and  Vineland,  Ontario,  also. 


Brown:  The  Genus  Chlorotettix  237 

C.  galhanatus  has  previously  been  taken  in  Iowa,  Ohio,  Kansas, 
Montana,  Wisconsin,  North  Dakota,  Connecticut,  Maine,  New  York, 
Pennsylvania,  Maryland,  Virginia,  North  Carolina,  South  Carolina, 
Tennessee,  Florida  and  Louisiana.  The  following  states  are  new: 
Missouri,  Arkansas,  Texas,  Mississippi,  Alabama,  Georgia,  and  New 
Jersey. 

C.  necopinus  is  a  southern  species  described  from  specimens  col- 
lected in  Mississippi.  It  has  been  collected  previously  from  Kansas, 
Tennessee,  North  Carolina,  South  Carolina  and  Florida.  It  has 
also  been  taken  in  Arkansas,  Texas,  Louisiana  and  Alabama. 

C.  spatulatus  has  a  wide  distribution,  having  been  reported  from 
Iowa,  Kansas,  Nebraska,  Colorado,  Ohio,  Wisconsin,  Connecticut, 
Virginia,  South  Carolina,  Tennessee  and  Florida.  New  localities 
are  Missouri,  New  Mexico,  Minnesota,  Texas,  Mississippi,  Alabama 
and  Georgia. 

C.  rugicollis  has  been  collected  in  Florida,  Texas,  Tennessee,  South 
Carolina  and  New  Jersey.  New  states  to  add  to  this  list  are:  Kan- 
sas, Arkansas  and  Alabama. 

C.  suturalis  type  specimens  are  from  Tennessee.  Our  collection 
shows  specimens  from  Mississippi,  Alabama,  Georgia  and  South 
Carolina. 

C.  minor  was  taken  in  Texas.  A  specimen  is  at  hand  from  Kan- 
sas.   This  species  is  evidently  very  uncommon. 

C.  nudatus  has  formerly  been  reported  from  Iowa,  Tennessee  and 
Connecticut.    Our  specimens  are  all  from  Douglas  county,  Kansas. 

C.  latijrons  was  described  from  specimens  taken  in  Virginia,  later 
found  in  Mississippi.    We  have  specimens  also  from  Alabama. 

C.  similis  was  originally  described  from  specimens  collected  in 
Oregon.    We  also  have  specimens  from  Washington  and  California. 

C.  fumidus  has  previously  been  reported  as  taken  from  Pennsyl- 
vania and  Tennessee.  The  writer  adds  to  the  list:  Missouri,  Mis- 
sissippi, Alabama  and  Georgia. 

C.  divergens  has  been  reported  for  Virginia  only,  but  we  have 
specimens  collected  in  Alabama,  Mississippi,  and  Arkansas. 

C.  productus  was  described  from  Virginia,  later  taken  in  Florida. 
The  following  states  are  added  to  the  list:  Louisiana,  Mississippi, 
Georgia  and  Texas. 

C.  unicolor  is  common  in  many  localities  and  thus  has  been  re- 
ported from  many  of  the  northern  states  and  Canada.  Specimens 
had  formerly  been  reported  for  Maine,  Connecticut,  New  Hamp- 
shire, New  York,  Wisconsin,  Ohio,  Iowa,  Nebraska,  Colorado,  Call- 


238  The  University  Science  Bulletin 

fornia,  Oregon,  Washington,  British  Columbia,  and  Ontario,  Canada. 
Specimens  are  at  hand  from  the  following  new  localities:  Michigan, 
Minnesota,  Montana,  Utah  and  Arizona. 

C.  tunicatus  is  widespread  over  the  southeastern  quarter  of  the 
United  States.  Specimens  have  been  reported  for  Kansas,  Missouri, 
Arkansas,  Texas,  Louisiana,  Tennessee,  Florida,  South  Carolina, 
North  Carolina  and  Virginia.  Alabama  and  Mississippi  are  here 
added  to  the  list. 

In  making  an  artificial  key  to  the  species,  the  most  outstanding 
characteristics  used  are:  shape  of  the  vertex,  male  valve  and  plates, 
and  the  last  ventral  segment  of  the  female.  Size  and  color  have 
been  used  in  a  more  general  way. 

Due  to  the  fact  that  many  species  are  very  similar  in  appearance, 
it  is  sometimes  difficult  to  distinguish  between  specimens  by  using 
only  the  above  characteristics.  Drawings  of  the  male  internal 
genitalia  of  the  new  species  are  included,  in  order  to  assist  in  de- 
termining specimens.  These  structures  furnish  definite  and  clear- 
cut  characters  for  the  determination  of  the  species. 

In  the  preparation  of  this  paper  the  writer  has  used,  for  examina- 
tion and  dissection,  specimens  from  the  Snow  Entomological  Col- 
lection. 

This  paper  has  been  worked  out  and  written  under  the  super- 
vision of  Dr.  Paul  B.  Lawson,  of  the  Department  of  Entomology  of 
the  University  of  Kansas.  Dr.  Dwight  M.  De  Long  and  Mr.  S.  E. 
Crumb  kindly  helped  with  the  comparison  of  some  specimens  and 
the  loan  of  material. 

Key  to  Species  of  Chlorotettix* 

1.  Vertex  with  anterior  margin  rounded,  length  at  middle  equaling  or  slightly  exceeding 

next  the  eye 2 

Vertex    with    anterior    margin    distinctly    but    bluntly    angulate,    distinctly    longer    at 

middle  than  next  the  eye 24 

2.  Size  small,  not  larger  than  5  mm 3 

Size  larger,  length  more  than  5.5  mm 5 

3.  Posterior   margin   of   female   last   ventral   segment   produced    at    middle,    but   without 

median    notch ;     male    valve    very    short,    plates    gradually    narrowed    to    produce 

fingerlike    apices    minor. 

Posterior   margin    of    female    last    ventral    segment    produced    at    middle,    but   with    a 
median  notch  ;   male  vah  e  longer,  plates  without  fingerlike  apices 4 

4.  Notch  in  female  last  ventral  segment  very  short ;    plates  with  lateral  margins  almost 

straight,  gradually  narrowed  to  rather  broad,  blunt  apices productus. 

Notch  in  female  last  ventral  segment  reaching  nearly  to  base ;    plates  gradually  and 
evenly  narrowed  to  sharp-pointed  tips minimus. 

5.  Vertex  not  longer  at  middle  than  next  the  eye 6 

Vertex  at  least  slightly  longer  at  middle  than  next  the  eye 7 

*  This  key  has  been  made  by  using  D.  M.  De  Long's  key  from  his  "A  synopsis  of  the 
Genus  Chlorotettix"  as  a  pattern. 


Brown:  The  Genus  Chlorotettix  239 

6.  Lateral   lobes   of    female   last   ventral    segment    distinctly    angulate;    and   male    plates 

narrow  and  at  most  but  slightly  upturned  apically latus. 

Lateral  lobes  of  female  last  ventral  segment  broadly  rounded,  and  male  plates  broad 
with  strongly  upturned  apices dozieri. 

7.  General  color  dark,  brownish   or  sordid   green 8 

General  color  light,  greenish  or  yellow 12 

8.  Size  large,  8  mm.;   pronotum,  scutellum  and  dorsal  portion  of  elytra  fuscus.  .  .fuscus. 
Size  smaller,  near  7  mm 9 

9.  Sordid  green  without  bands  or  stripes 10 

Vertex   pale  with   broad   brown  band   between   eyes,   elytra    dark   with    pale   nervuros, 

giving  it  a  striped  appearance necopimis. 

10.  Size  large,   7.5  mm.   or  more,  male  plates  short fallax. 

Size  small,   less  than   7.5   mm.,   often   appearing  almost   black    in  color;    male   plates 

of  medium  length H 

11.  Vertex  more  subangulate,  lighter  color tergatus. 

Vertex  more  rounded,  a  distinctly  blacker  color tergatus  var.   melanotus. 

12.  Vertex  or  elytra  with  red  or  dark  markings 13 

Vertex  and  elytra  greenish  or  yellowish 1-t 

13.  Margin  of  vertex  with  more  or  less  distinct  broad  red  or  orange  band  between  eyes, 

elytra   unmarked    rugicoUis. 

Margin  of  vertex  without  red  band  but  with  two  dark  spots,  elytra  yellowish   green 
with  a  more  or  less  distinct  dull,  three-lobed  fuscus  stripe  along  the  suture. 

surtwalis. 

14.  Head  subangulate,  body  quite  narrow,  a  cross  nervure  between  the  two   inner  veins 

of  clavus  occidentalis. 

Head  rounded,  body  usually  broad,  inner  veins  of  clavus  without  a  cross  nervure. ...      15 

15.  Female  last  ventral  segment  notched,  bearing  spatulate  process  from  its  apex;    male 

valve  broad  and  short,  plates  long,  gradually  tapering 16 

Female  last  ventral  segment  notched,  but  without  spatulate  process;  male  valve  and 
plates  variable   

16.  Spatulate  process  not  cleft  at  apex;  styles  not  extending  past  plates spatulatus. 

Spatulate    process    cleft    at    apex;    styles    extending    past    plates    and    visible    from 

below    limosus. 

17.  Female  segment  with  sides  of  notch  each  bearing  a  lateral  median  acute  tooth;  male 

plates  short   viridius. 

Female  segment  with  sides  of  notch  without  acute  tooth;   male  plates  longer 18 

18.  Size  large,  6.5  mm.  or  more 

Size  small,  6.5  mm.  or  less • 

19.  Color  greenish   yellow,   female  last  ventral  segment   with   median  notch   reaching   to 

base;    notch   and   posterior   margin  bordered   with   dark   brown;    male   valve   very 

short,  plates  broad  and  short,  about  three  times  length  of  valve latifrons. 

Color  uniform  pale  green,  female  last  ventral  segment  with  median  notch  reaching  not 

20 
more  than  one-third  distance  to  base 

20.  Female   last   ventral   segment  long,   with   a   broad,   shallow   notch,   narrowed   toward 

apex,  posterior  margin  of  lobes  sinuate;   male  plates  narrowed  at  half  their  length, 

then  produced    unicolor. 

Female  last  ventral  segment  longer  and  narrower,  narrow  median  notch  reaches  one- 
third  distance  to  base,  sides  convexly  rounded  and  often  overlap  along  median 
line,  posterior  margin  of  lobes  rounded;  male  plates  broader  and  longer  than  m 
unicolor ^'"'*''- 

21.  Head  broader,  1.75  mm.,  more  rounded,  body  more  robust 22 

Head  narrower,  1.5  mm.,  more  angulate,  body  narrower 23 

22.  Color   pale  yellow,   some   almost   colorless,    female   last    ventral    segment   with    lateral 

angles  rounded;    male  valve  with  apex  a  right  angle pallidus. 

Color  greenish  yellow,  female  last  ventral  segment  with  lateral  angles  broadly  obtuse; 
male  valve  with  apex  broadly  obtuse.  . rubidus. 

23.  Eyes  dark ;   color  dark  yellow ;   posterior  margin  of  female  last  ventral  segment  with 

deep  notch  with  sides  strongly  convex convexus. 

Eyes  red;  color  pale  yellow;  male  valve  broad  at  base,  rounded  at  apex;  plates 
narrower  than  valve,  about  three  times  the  length;  pygofer  one-half  longer  than 
plates   '^"™»- 


240  The  University  Science  Bulletin 

24.  Vertex  usually  with  distinct  transverse  band  across  middle  between  eyes .  .  .      25 

Vertex  without  definite  markings  in  the  form  of  either  spots  or  bands 26 

25.  Size  large,  7.5  mm.,  distinctly  reddish  brown  in  color hisorius. 

Size  smaller,  less  than  7  mm.,  color  yellowish  green scutellatus. 

26.  General  color  brownish  or  sordid  green 27 

General  color  light,  pale  green  or  yellowish 3^ 

27.  Vertex   subangulate  to   rounded rotundus. 

Vertex  angulate    28 

28.  Median  notch  in  female  last  ventral  segment  U-shaped 29 

Median  notch  in  female  last  ventral  segment  V-shaped 30 

29.  Sides  of  median  notch  in  female  last  ventral  segment  toothed. sordidus. 

Sides  of  median  notch  in  female  last  ventral  segment  not  toothed divergens. 

30.  Lateral  angles  of  last  ventral  segment  of  female  acute  and  prominent;   sides  of  notch 

not  tinged  with  brown iridescens. 

Lateral  angles  of  last  ventral  segment  of  female  obtuse;    sides  of  notch   tinged  with 
brown    jumidus. 

31.  Small,  not  more  than  6.5  mm 32 

Larger,  6.5  mm.  or  more 41 

32.  Vertex  distinctly  angled,  almost  twice  as  long  at  middle  as  next  the  eye,  sometimes 

conical    33 

Vertex  more  bluntly  angled,  not  more  than  one-half  longer  at  middle  than  next  the 
eye    34 

33.  At  least  5.5  mm.  in  length,  male  valve  sinuate,  apex  notched,  apices  of  plates  not 

produced  in  long  fingerlike  processes vividus. 

Length   4.5   mm.,   male  valve  obtusely   rounded,   apices   of   plates   produced   in   long 
fingerlike  processes  delta. 

34.  Small,  not  exceeding  5  mm.   in  length 35 

Large,  6  to  6.5  mm.  in  length 36 

35.  Female   last   ventral    segment   with    a    rather    broad    V-shaped    notch,    lateral    angles 

slightly  produced,   male  plates  broad borealis. 

Female  last  ventral  segment  with   a   very  narrow   notch,   lateral  angles  strongly   pro- 
duced;  male  plates  narrow  and  parallel  on  apical  half excultus. 

36.  Lateral  lobes  of  female  last  ventral  segment  broad  and  more  or  less  rounded 37 

Lateral  lobes  of  female  last  ventral  segment  angled  or  angularly  rounded 40 

37.  Notch  in  female  last  ventral  segment  extending  more  than  half  distance  to  base 38 

Notch  in  female  last  ventral  segment  extending  half  or  less  than  half  distance  to  base,     39 

38.  Eyes  red;    posterior  margin   of  last  ventral  segment  concavely  rounded capensis. 

Eyes  dark;    posterior  margin  of  last  ventral  segment  convexly  roundeA.  .  .galhanatus. 

39.  Dark  spot  on  median  line  of  base  of  last  ventral  segment maculosus. 

Without  dark  spot  on  last  ventral  segment angustus. 

40.  Vertex  about  one-third   longer  at  middle  than  next   the   eyes;    female   segment   with 

the  sides  of  notch  convexly  rounded vacunus. 

Vertex  longer,  one-half  longer  at  middle  than  next  the  eyes;    sides  of  female  notch 
concavely   rounded    distinctus. 

41.  Female  segment  notched,  bearing  a  spatulate  process  at  its  apex 42 

Female  segment  usually  notched  but  without  spatulate  process 44 

42.  Last  ventral  segment  of  female  with  spatulate  process  twice  as  long  as  wide.  .  .  .balli. 
Last   ventral   segment   of    female   with    spatulate   process   about    equal    in   length    and 

width    43 

43.  Lateral  angles  of  last  ventral  segment  acute  and  prominent,  sides  of  notch  concave; 

male  plates  long  and  slender  apically attennatus. 

Lateral   angles  of  last   ventral   segment   obtuse   and   not   prominent,   sides   of   notch 
sinuate sinuosus. 

44.  Last  ventral  segment  of  female  with  side  margins  short,  rounding  to  posterior  margin 

which  is  notched  giving  the  appearance  from  the  ventral  side  of  four  teeth. 

dentatus. 
Last  ventral  segment   of   female  with   side  margins   longer   than  at   center  of   median 
notch    45 

45.  Length,   7  mm Iftunicatus. 

t  C.  tunicatus  has  variation  in  shape  of  vertex  from  subangulate  to  rounded. 


Brown:  The  Genus  Chlorotettix  241 

Length,  more  than  7  mm 4g 

46.    Female  segment  slightly  emarginate  with  a  brown  spot  at  center;   male  valve  notch 

at  middle    stolatus. 

Female  segment  black  margined,  with  notch  at  center,  broadly,  shallowly  emarginate 

either  side;  male  valve  rounded,  without  notch nudatus. 

Chlorotettix  angustus  sp.  n. 

(PI.  XXI,  fig.  7;  PI.  XXII,  figs.  5,  5a;  PI.  XXIII,  figs.  8,  a-c) 

Resembling  galbanatus,  but  with  notch  in  last  ventral  segment  of 
female  excavated  only  one-half  distance  to  base  instead  of  nearly  to 
base;  male  genitalia  distinct.  Length,  6  to  6.5  mm.  Vertex  ob- 
tusely angled,  one-half  longer  at  middle  than  next  the  eye. 

Color.  Greenish-yellow.  Eyes  dark.  Elytra  subhyaline.  Ventral 
side  greenish-yellow.    Tarsal  claws  black. 

Genitalia.  Last  ventral  segment  of  female  twice  as  long  as  pre- 
ceding segment,  with  lateral  angles  obtusely  angled  and  prominent; 
posterior  margin  with  a  median,  narrow,  V-shaped  notch  reaching 
one-half  distance  to  base;  ovipositor  slightly  exceeding  pygofer. 
Male  valve  broad  at  base  and  obtusely  angled  on  posterior  margin, 
about  one  and  one-half  times  as  long  as  preceding  segment;  plates 
long,  slightly  less  than  three  times  length  of  valve,  broad  at  base, 
gradually  becoming  broader  for  one-third  distance  from  base  then 
gradually  narrowing  to  obtuse  apex ;  about  same  length  as  pygofer 
with  inner  margins  meeting  in  a  furrow. 

Holotype.  Female,  Batesburg,  S.  C.,  August  24,  1930,  L.  D.  Tut- 
hill. 

Allotype.    Male,  same  data. 

Types  deposited  in  Snow  Entomological  Collection. 

Chlorotettix  attenuatus  sp.  n. 

(PI.  XXI,  fig.  12;   PI.  XXII,  figs.  6,  6a;   PI.  XXIII,  figs.  5,  a-c) 

Resembling  balli,  but  with  shorter  spatulate  process  in  female  and 
with  male  plates  more  narrowed  in  apical  half.  Length,  6.75  to  7  mm. 

Vertex  one-half  longer  at  middle  than  against  the  eye,  subangulate. 

Color.  Yellowish-green.  Eyes  dark.  Elytra  subhyaline,  some- 
what smoky  apically.  Underside  uniformly  yellowish-green  except 
for  brownish  tarsal  claws. 

Genitalia.  Last  ventral  segment  of  female  with  lateral  angles 
prominent  and  distinctly  angulate;  posterior  margin  smoothly  ex- 
cavated to  about  one-half  the  length  of  the  segment  and  bearing  a 
short  spatulate  process  which  is  about  as  wide  as  long;  ovipositor 
slightly  exceeding  pygofer.    Male  valve  large,  subangulate;  plates 


242  The  University  Science  Bulletin 

broad  basally,  then  slightly  narrowed  to  long  and  acute  spine  which 
about  equals  the  pygofer;  pygofer,  viewed  laterally,  excavated  on 
posterior  margin  between  an  upper  and  a  larger  lower  lobe. 

Holotype.    Female,  Atherton,  Mo.,  July  2,  1922,  C.  F.  Adams. 

Allotype.    Male,  same  data. 

Paratype.    Male,  Atherton,  Mo.,  June  18, 1922,  C.  F.  Adams. 

Types  deposited  in  Snow  Entomological  Collection. 

Chlorotettix  convexus  sp.  n. 

(PI.  XXI,  fig.   4;    PI.   XXII,  fig.   11) 

Resembling  vacunus,  but  with  vertex  more  rounded  and  notched 
in  last  ventral  segment,  broader  at  apex.  Length,  6  to  6.25  mm. 
Vertex  rounded  and  one-third  longer  at  middle  than  next-  the  eye. 

Color.  Greenish-yellow.  Eyes  dark.  Elytra  subhyaline  tinged 
with  yellow.    Ventral  side  yellow.    Tarsal  claws  brown. 

Genitalia.  Last  ventral  segment  of  female  with  lateral  angles 
large,  distinctly  but  obtusely  angled;  posterior  margin  broadly  ex- 
cavated three-fourths  distance  to  base,  sides  of  notch  strongly  con- 
vex and  apex  tinged  with  brown;  ovipositor  slightly  shorter  than 
pygofer. 

Holotype.    Female,  Gulfport,  Miss.,  August  1,  1921,  C.  J.  Drake. 

Paratype.    Female,  Woodville,  Miss.,  July  25,  1921,  C.  J.  Drake. 

Holotype  deposited  in  Snow  Entomological  Collection;  paratype 
in  Doctor  Drake's  Collection. 

Chlorotettix  durus  sp.  n. 

(PI.   XXI,   5;    PI.   XXII,   fig.    12;    PI.   XXIII,   figs.    3,   a-e) 

Resembling  rugicollis,  but  with  distinct  genitalia  and  slightly 
smaller.  Length,  6  mm.  Vertex  rounded,  one-third  longer  at  middle 
than  next  the  eye. 

Color.  Greenish-yellow.  Eyes  red.  Elytra  subhyaline.  Ventral 
side  yellow.    Tarsal  claws  brown. 

Genitalia.  Male  valve  one-third  longer  than  preceding  segment, 
broad  at  base,  rounded  at  apex;  plates  narrower  at  base  than  valve, 
gradually  becoming  wider  to  middle  then  slowly  narrowing  to  a  very 
obtuse  or  nearly  truncate  apex,  nearly  three  times  as  long  as  valve ; 
pygofer  exceeding  plates  by  one-half  length  of  latter. 

Holotype.    Male,  Ft.  Myers,  Fla.,  August  14,  1930,  J.  Nottingham. 

Type  deposited  in  Snow  Entomological  Collection. 


Brown:  The  Genus  Chlorotettix  243 

Chlorotettix  fiiscus  sp.  n. 

(PI.  XXI,  fig.   6;   PI.  XXII,  fig.   13) 

Resembling  necopinus,  but  without  dark  transverse  band  between 
eyes.  Length,  8  mm.  Vertex  one-third  longer  at  the  middle  than 
next  the  eye,  obtusely  rounded. 

Color.  Pronotum,  scutellum  and  dorsal  portion  of  elytra  fuscous; 
vertex,  lateral  margins  of  pronotum  and  of  elytra  lighter.  Eyes 
dark.  Dorsal  and  lateral  sclerites  of  abdomen  mostly  black.  Ven- 
tral side  yellow.    Tarsal  claws  brown. 

Genitalia.  Last  ventral  segment  of  female  with  lateral  angles 
obtuse;  posterior  margin  slightly  concave  to  a  median,  V-shaped 
notch  reaching  two-thirds  distance  to  base;  ovipositor  slightly  ex- 
ceeding pygofer. 

Holotype.    Female,  Prattsville,  Ala.,  July  21, 1930,  R.  H.  Beamer. 

Paratypes.  One  female,  Prattsville,  Ala.,  July  21,  1930,  R.  H. 
Beamer;  2  females,  Dennis,  Miss.,  July  6,  1921,  C.  J.  Drake. 

Two  paratypes  deposited  in  Doctor  Drake's  Collection;  other 
types  deposited  in  Snow  Entomological  Collection. 

Chlorotettix  latifrons  (Sanders  and  De  Long) 

(PI.  XXI,  fig.   14;    PI.  XXII,  fig.   2;    PI.   XXIII,  figs.   7,  a-c) 

The  male  is  here  described  for  the  first  time.  Resembling  unicolor 
in  size  and  form,  but  greenish-yellow  in  color.  Length  to  tip  of 
pygofer  6  mm.*  Vertex  broadly  rounded,  about  the  same  length  at 
middle  as  next  the  eye. 

Color.  Greenish-yellow.  Eyes  dark.  Elytra  subhyaline.  Ven- 
tral side  yellow.    Tarsal  claws  brown. 

Genitalia.  Male  valve  slightly  more  than  one-half  the  length  of 
preceding  segment,  broad  at  base,  posterior  margin  very  broadly 
angled;  plates  broad  at  base  and  short,  broadening  slightly  at  first, 
then  narrowing  slowly  for  first  half,  then  rapidly  to  an  obtuse  or 
nearly  truncate  apex  which  does  not  quite  reach  end  of  pygofer; 
margin  fringed  with  spines;  end  of  pygofer  incurved. 

Allotype.    Male,  Okolona,  Miss.,  June  29,  1921,  C.  J.  Drake. 

Allotype  deposited  in  Snow  Entomological  Collection. 

*  Apex  of  wings  broken  off. 


244  The  University  Science  Bulletin 

Chlorotettix  latus  sp.  n. 

(PI.  XXI,  fig.  9;  PI.  XXII,  figs.  9,  9a;  PI.  XXIII,  figs.  2,  a-c) 

With  a  very  broad  and  parallel-margined  head,  resembling  dozieri, 
but  with  lateral  lobes  of  female  last  ventral  segment  more  pointed, 
and  the  male  plates  narrower  and  not  upturned  at  apex.  Length, 
5.75  to  6  mm. 

Head  distinctly  wider  than  pronotum.  Vertex  very  broadly 
rounded  and  about  the  same  length  at  middle  as  next  the  eye.  Front 
broad  and  short. 

Color.  Greenish-yellow.  Eyes  reddish-brown.  Elytra,  subhya- 
line,  tinged  with  green.  Ventral  side  greenish-yellow.  Tarsal  claws 
dark  brown. 

Genitalia.  Last  ventral  segment  of  female  with  lateral  angles 
strongly  produced  and  distinctly  angulate;  posterior  margin  with  a 
deep,  V-shaped  notch  two-thirds  distance  to  base ;  ovipositor  slightly 
longer  than  pygofer.  Male  valve  short,  obtusely  angled;  plates 
broad  at  base,  narrowing  to  blunt,  slightly  diverging  apices  which 
are  a  little  longer  than  pygofer. 

Holotype.    Female,  Hilliard,  Fla.,  August  19,  1930,  L.  D.  Tuthill. 

Allotype.    Male,  HilHard,  Fla.,  August  31,  1930,  Paul  W.  Oman. 

Paratypes.  Forty-two  females  and  42  males,  all  taken  at  Hilliard, 
Fla.,  August  19  and  31,  1930,  by  L.  D.  Tuthill,  Paul  W.  Oman,  R.  H. 
Beamer,  J.  Nottingham. 

Types  deposited  in  Snow  Entomological  Collection. 

Chlorotettix  maculosus  sp.  n. 

(PI.  XXI,  fig.  8;  PI.  XXII,  fig.  4) 

Resembling  balli,  but  smaller  and  without  spatulate  process. 
Length,  6  mm.  Vertex  subangulate,  one-third  longer  at  middle  than 
next  the  eye. 

Color.  Greenish-yellow.  Eyes  dark,  tinged  with  red.  Elytra 
subhyaline.    Ventral  side  yellow.    Tarsal  claws  light  brown. 

Genitalia.  Last  ventral  segment  nearly  three  times  as  long  as 
preceding  segment  with  lateral  lobes  broad  and  with  lateral  angles 
widely  obtuse;  posterior  margin  with  a  rather  narrow  median  V- 
shaped  notch  reaching  one-half  the  distance  to  base  which  has  a 
faint  brownish  spot  on  median  line;  ovipositor  slightly  exceeding 
pygofer. 

Holotype.  Female,  Polk  county.  Ark.,  August  21,  1928,  R.  H. 
Beamer. 


Brown:  The  Genus  Chlorotettix  245 

Pnratype.  Female,  Natchitoches  County,  La.,  August  16,  1928, 
A.  M.  James. 

Types  deposited  in  Snow  Entomological  Collection. 

Chlorotettix  pallidus  sp.  n. 

(PI.  XXI,  fig.   13;   PI.  XXII,  figs.   1,  la;    PI.  XXIII,  figs.   4,  a-c) 

Resembling  spatulatus  in  form  and  size  but  paler,  without  the 
spatulate  process.  Length,  5.75  to  6.5  mm.  Vertex  rounded  and 
slightly  longer  at  the  middle  than  next  the  eye. 

Color.  Pale  yellow.  Eyes  light  to  dark,  some  tinged  with  red. 
Elytra  subhyaline.    Ventral  side  light  yellow.    Tarsal  claws  brown. 

Genitalia.  Last  ventral  female  segment  with  lateral  angles  promi- 
nent and  bluntly  angled;  posterior  margin  with  a  broad,  deep,  V- 
shaped  excavation  reaching  almost  three-fourths  the  distance  to 
base,  excavation  rounding  at  the  apex;  ovipositor  about  the  same 
length  as  narrow  pygofer.  Male  valve  longer  than  preceding  seg- 
ment, apex  nearly  a  right  angle;  plates  broad  at  base,  roundingly 
narrowing  to  a  very  obtuse,  nearly  truncate  apex  which  exceeds 
pygofer,  margin  fringed  with  spines.  Pygofer  with  apex  blunt,  with 
a  pair  of  large,  straight  spines  directed  downward  and  backward 
from  the  ventral  posterior  margin. 

Holotype.  Female,  Cameron  Co.,  Texas,  August  3,  1928,  L.  D. 
Beamer. 

Allotype.  Male,  Cameron  Co.,  Texas,  August  3,  1928,  R.  H. 
Beamer. 

Paratypes.  Fifty  females  and  17  males,  Cameron  Co.,  Tex., 
August  3,  1928,  by  R.  H.  Beamer,  L.  D.  Beamer,  Jack  Beamer, 
A.  M.  James,  J.  G.  Shaw;  9  females  and  7  males,  Hidalgo  Co.,  Tex., 
July  28,  1928,  by  R.  H.  Beamer,  A.  M.  James;  1  female,  Hidalgo 
Co.,  Tex.,  July  31,  1928,  by  R.  H.  Beamer;  1  female,  Hidalgo  Co., 
Tex.,  August  2,  1928,  by  R.  H.  Beamer;  2  females  and  1  male. 
Brooks  Co.,  Tex.,  July  25,  1928,  by  A.  M.  James;  1  female,  Brazoria 
Co.,  Tex.,  August  10,  1928,  by  R.  H.  Beamer;  2  females  and  2  males, 
Brownsville,  Texas,  June,  F.  H.  Snow. 

Types  deposited  in  Snow  Entomological  Collection. 


17— 31S2 


246  The  University  Science  Bulletin 

Chlorotettix  rotundus  sp.  n. 

(PI.  XXI,  fig.  10;   PI.  XXII,  fig.  3;   PI.  XXIII,  figs.   1,  a-c) 

Resembling  tunicatus,  but  with  vertex  narrower,  slightly  browner 
in  color,  and  with  distinct  internal  male  genitalia.  Length,  6.5  mm. 
Vertex  one-half  longer  at  middle  than  nexi,  the  eye,  subangulate. 

Color.  Brownish-yellow.  Eyes  dark.  Elytra  siibhyaline.  Ven- 
tral side  yellow.    Tarsal  claws  brown. 

Genitalia.  Male  valve  slightly  longer  than  preceding  segment, 
broad  at  base,  posterior  margin  rounded  at  apex;  plates  broad  at 
base,  gradually  narrowing  to  a  very  obtuse  or  truncate  apex,  margin 
fringed  with  spines  below  and  fine  hairs  above,  slightly  exceeding 
pygofer.  Plates  of  rotundus  similar  to  those  of  tunicatus,  except 
that  in  tunicatus  plates  are  convex  below,  while  in  rotundus  plates 
are  slightly  concave  along  line  where  two  plates  meet;  apices  more 
acute  in  rotundv.s.  Processes  on  end  of  cedagus  in  rotundus  project 
straight  from  cedagus,  while  in  tunicatus  they  project  diagonally. 
Styles  vary  widely  in  shape. 

Holotijpe.    Male,  Polk  county,  Ark.,  July  21,  1928,  L.  D.  Beamer. 

Paratypes.  One  male,  Polk  county,  Ark.,  August  21,  1928,  R.  H. 
Beamer;  3  males,  Tuskegee,  Ala.,  July  22,  1930,  L.  D.  Tuthill;  2 
males,  Shuqualak,  Miss.,  July  16,  1930,  R.  H.  Beamer;  1  male,  Wal- 
nut, N.  C,  August  20,  1930,  Paul  W.  Oman;  1  male,  Scott  county, 
Ark.,  August  23,  1928,  R.  H.  Beamer;  1  male,  Prattsburg,  Ga.,  July 
25,  1930,  R.  H.  Beamer. 

Types  deposited  in  Snow  Entomological  Collection. 

Chlorotettix  ruhidus  sp.  n. 

(PI.  XXI,  fig.  1;   PI.  XXII,  figs.   8,  8a;   PI.  XXIII,  figs.   6,  a-c) 

Resembling  capensis,  but  with  a  less  pointed  vertex  and  lobes  of 
ventral  segment  more  rounded.  Length,  6  to  6.5  mm.  Vertex 
rounded,  slightly  longer  at  the  middle  than  next  the  eye. 

Color.  Greenish-yellow.  Eyes  dark  red.  Elytra  subhyaline. 
Ventral  side  greenish-yellow.    Tarsal  claws  reddish-brown. 

Genitalia.  Last  ventral  segment  of  female  with  lateral  angles 
broadly  obtuse;  posterior  margin  with  broad  median  V-shaped 
notch,  which  is  tinged  reddish  brown  at  apex  and  reaches  a  little 
more  than  two-thirds  the  distance  to  base;  ovipositor  slightly  ex- 
ceeding pygofer.  Male  valve  nearly  twice  as  long  as  preceding  seg- 
ment, very  broad  at  base  and  obtusely  angled;  plates,  three  times 
as  long  as  valve,  broad  at  base,  gradually  becoming  broader  for 


Brown:  The  Genus  Chlorotettix  247 

one-fifth  distance  from  base,  then  gradually  narrowing  to  veiy  ob- 
tuse or  truncate  apex,  margin  fringed  with  hairs;  pygofer  sHghtly 
exceeding  plates. 

Holohjpe.  Female,  Plant  City,  Fla.,  August  15,  1930,  J.  O.  Not- 
tingham. 

Allotype.    Male,  Hilliard,  Fla.,  August  19,  1930,  R.  H.  Beamer. 

Paratypes.  Two  females,  Sanford,  Fla.,  Sept.  19,  1929,  C.  0. 
Bare;  1  female,  Yankeetown,  Fla.,  July  31,  1930,  Paul  W.  Oman; 
1  female,  Natchitoches  county.  La.,  August  16,  1930,  A.  M.  James; 
1  female,  Orange  county,  Tex.,  August  14,  1928,  R.  H.  Beamer;  1 
male.  Plant  City,  Fla.,  June  5, 1926,  C.  0.  Bare;  1  female,  Cocoanut 
Grove,  Fla.,  August  9,  1930,  Paul  W.  Oman;  1  male,  Woodville, 
Miss.,  July  25,  1921,  C.  J.  Drake;  1  male,  Homestead,  Fla.,  August 
9,  1930,  J.  Nottingham. 

Types  deposited  in  Snow  Entomological  Collection. 

Chlorotettix  sinuosus  sp.  n. 

(PI.  XXI,  fig.  2;  PI.  XXII,  fig.  10) 

Resembling  balli,  but  with  a  shorter  spatulate  process  in  female 
and  last  ventral  segment  not  so  deeply  notched.  Length,  6.5  to  7 
mm.  Vertex  one-third  longer  at  the  middle  than  next  the  eye,  sub- 
angulate. 

Color.  Greenish-yellow.  Eyes  dark.  Elytra  subhyaline.  Ven- 
tral side  yellow.    Tarsal  claws  brown. 

Genitalia.  Last  ventral  segment  with  lateral  angles  roundingly 
angled;  posterior  margin  sinuated,  with  broad,  shallow,  median 
notch  reaching  one-third  distance  to  base  of  segment,  at  apex  of 
which  is  a  short,  emarginate,  spatulate  process  which  is  about  as 
broad  as  long;  ovipositor  slightly  exceeds  pygofer. 

Holotype.    Female,  Columbus,  Miss.,  July  24,  1921,  C.  J.  Drake. 

Paratype.    Female,  Columbus,  Miss.,  July  24,  1921,  C.  J.  Drake. 

Holotype  deposited  in  Snow  Entomological  Collection;  paratype 
in  Doctor  Drake's  Collection. 


248  The  University  Science  Bulletin 

Chlorotettix  sordidus  sp.  n. 

(PI.  XXI,  fig.  3;   PI.  XXII,  fig.   7) 

Resembling  tergatus,  but  smaller  and  with  vertex  more  angulate. 
Length,  7  mm.  Vertex  distinctly  angulate,  one-half  longer  at  middle 
than  next  the  eye. 

Color.  Sordid  greenish-yellow.  Eyes  dark.  Elytra  subhy aline 
with  a  smoky,  bro^\^lish  tinge.  Ventral  side  yellowish.  Tai-^al 
claws  brownish. 

Genitalia.  Last  ventral  female  segment  very  long  with  lateral 
angles  much  produced  and  distinctly  angulate;  posterior  margin 
with  a  broad,  V-shaped  excavation  reaching  seven-eighths  the  dis- 
tance to  base,  its  edges  tinged  with  brown;  ovipositor  longer  than 
pygofer. 

Holotype.  Female,  Caddo  county,  La.,  August  19,  1928,  L.  D. 
Beamer. 

Paratypes.  Two  females.  Port  Gibson,  July  20,  1921;  2  females, 
Port  Gibson,  July  21;  5  females.  Meridian,  August  14,  1921;  1 
female,  Yazoo  City,  July  6,  1921;  1  female,  Dennis,  July  6,  1921; 
1  female,  Tishomingo,  July  7,  1921;  all  paratypes  from  Mississippi 
taken  by  C.  J.  Drake. 

Types  deposited  in  Snow  Entomological  Collection. 


250 


The  University  Science  Bulletin 


1.  rubidus. 

2.  sinuosiis. 

3.  sordidus. 

4.  convexus. 

5.  durus. 


PLATE  XXI 
HEADS 

6.  fuscus. 

7.  angustus. 

8.  maculosus. 

9.  latus. 

10.  rotundus. 


11.  similis. 

12.  attenuatus. 

13.  pallidus. 

14.  latifrons. 


Brown:  The  Genus  Chlorotettix 


251 


PLATE  XXI 


1  rubidu<; 


2  sinuosus 


3  sordidus 


9  latus 


7  angustui 


1  durus 


8  maculosus 


6  fuscus 


10  rotundus 


12  attenuatus 


13  palhdus 


14  latifrons 


252  The  University  Science  Bulletin 


PLATE  XXII 
EXTERNAL  MALE  AND  FEMALE  GENITALIA 

L  pallidus — la.  6.  attenuatus — 6a  IL  convexus. 

2.  latifrons.  7.  sordidus.  12.  durus. 

3.  rotundus.  8.  rubidus — 8a.  13.  fuscus. 

4.  maculosus.  9.  latus — 9a.  14.  .similis — 14a. 

5.  angustus — 5a.  10.  sinuosus. 

1 — female.         la — male. 


Brown:  The  Genus  Chlorotettix  253 


PLATE  XXII 


254  The  University  Science  Bulletin 


PLATE  XXIII 
INTERNAL  MALE  GENITALIA 


1. 

rotundiis. 

la. 

lb. 

Ic 

2. 

latus. 

2a. 

2b. 

2c. 

3. 

durus. 

3a. 

3b. 

3c 

4. 

pallidus. 

4a. 

4b. 

4c 

5. 

attenuatus. 

5a. 

5b. 

5c 

6. 

rubidus. 

6a. 

6b. 

6c 

7. 

latifrons. 

7a. 

7b. 

7c 

8. 

angustus. 

8a. 

8b. 

8c 

9. 

similis. 

9a. 

9b. 

9c 

1 — Side  view  of  pygofer.  b — Dorsal  view  of  oedagus. 

a — Dorsal  view  of  style.  c — Lateral  view  of  oedagus. 


Brown:  The  Genus  Chlorotettix 


255 


PLATE  XXIII 


THE  UNIVERSITY  OF  KANSAS 

SCIENCE  BULLETIN 

Vol.  XXI.]  March,  1933.  [No.  5. 


A  New  Species  of  Lizard  from  Mexico 

EDWARD  H.  TAYLOR, 

Depart  mtrit  of  Zoology,   University  of  Kansas 


Abstract:  A  species  of  skink,  Eumeces  indubitus,  is  described,  related  to 
E.  dugesi,  but  differing  in  the  character  of  the  supraoculars  and  in  color.  The 
habitat  of  the  species  is  the  Mexican  states  of  Michoacan,  Mexico,  and 
Morel  OS. 


AMONG  the  lizards  obtained  in  southern  Mexico  in  the  summer 
of  1932,  by  Hobart  Smith  and  myself,  is  a  medium-sized  skink 
belonging  to  the  genus  Eumeces,  which  appears  to  be  new  to  science. 
The  form  is  represented  in  the  collection  by  a  series  of  thirty-three 
specimens,  all  from  the  southern  part  of  the  Mexican  highland. 

This  southern  Mexican  region  is  of  especial  interest  to  the  herpe- 
tologist,  since  the  fauna,  especially  the  lacertilian  part  of  it,  is  rich 
and  varied.  Many  of  the  genera  are  of  the  plastic  type,  and  these 
register,  as  external  scale  and  color  characters,  the  effects  of  the 
environment. 

Many  species,  too,  seem  to  be  of  limited  distribution.  The  fauna 
is  such  as  to  suggest  a  condition  as  might  obtain  if  the  species  from 
a  wide  territory  were  herded  together  and  driven  into  a  much 
smaller  teritory  by  some  impelling  environmental  factor.  In  Mexico 
in  the  region  south  of  the  Tropic  of  Cancer  are  to  be  found  not  less 
than  a  dozen  species  of  the  genus  Eumeces,  representing  most  of  the 
principal  groups  of  the  genus. 

(257) 


258  The  University  Science  Bulletin 

Eumeces  indubitus  sp.  nov. 

Type.  No.  1731,  Taylor-Smith  collection:  Collected  July  9, 
1932,  on  the  Mexico-Cuernavaca  highway,  about  forty  miles  south- 
east of  Mexico  City  (kilometer  63),  by  Edward  H.  Taylor  and 
Hobart  Smith. 

Diagnosis.  A  medium-sized,  robust  species;  four  supraoculars, 
the  three  anterior  in  contact  with  the  frontal;  the  parietals  inclos- 
ing small  interparietal;  one  postmental;  no  postnasal ;  the  subcaudals 
distinctly  widened;  seven  upper  labials,  seven  superciliaries ;  the 
seventh  upper  labial  broadly  in  contact  with  the  upper  secondary 
temporal;  primary  temporal  small,  widely  separated  from  lower 
secondary  or  tertiary;  24  scale  rows  about  middle  of  body;  57  to 
61  scales  from  occiput  to  above  anus.  Limbs  moderately  large,  but 
failing  to  touch,  even  in  young,  when  adpressed.  Color  above, 
olive  to  olive-brown  with  a  short  dorsolateral  light  line  from  rostral, 
the  line  disappearing  on  the  shoulder;  a  narrow  labial  light  line 
terminating  at  ear;  no  median  light  line  or  forking  lines  on  the  head. 

Description  of  the  Type.  Adult  male.  Portion  of  rostral  visible 
above,  equal  to  about  half  the  size  of  the  frontonasal;  internasals 
large,  broadly  in  contact;  frontonasal  hexagonal,  forming  sutures 
with  the  loreals,  and  narrowly  in  contact  with  the  frontal,  forming 
its  longest  sutures  with  the  prefrontals;  later  narrowly  separated, 
forming  sutures  with  first  superciliary,  both  loreals  and  the  anterior 
supraocular;  frontal  longer  than  its  distance  from  the  end  of  the 
snout,  obtusely  angular  anteriorly,  somewhat  rounded  posteriorly, 
somewhat  narrowed  in  the  middle,  and  only  a  little  wider  in  the 
anterior  part  than  in  the  posterior;  four  supraoculars,  the  anterior 
longer  than  wide  with  an  area  scarcely  less  than  the  fourth,  the 
three  anterior  bordering  the  frontal;  the  frontoparietals  larger 
than  prefrontals,  their  common  suture  less  than  half  their  length; 
interparietal  short  and  broad,  inclosed  behind  by  the  parietals  which 
are  more  than  twice  as  long  as  their  greatest  width;  two  pairs  of 
nuchals,  the  anterior  somewhat  the  larger;  nasal  of  moderate  size, 
divided,  the  anterior  part  not  as  large  as  the  posterior  part  with 
nostril;  anterior  loreal  distinctly  higher  than  long,  higher  than  the 
posterior,  which  is  considerably  longer  than  high ;  seven  superciliaries, 
the  anterior  less  than  one  and  one-half  the  size  of  the  second;  two 
subequal  presuboculars ;  four  posterior  suboculars;  primary  tem- 
poral less  than  one-fourth  the  size  of  the  upper  secondary  temporal; 
latter  very  broadly  in  contact  with  the  seventh  labial,  the  suture 


Taylor:  A  New  Species  of  Lizard  259 

more  than  half  its  length;  seven  upper  labials,  four  preceding  the 
subocular,  which  is  low  and  elongate;  seventh  nearly  double  the 
size  of  the  sixth  and  separated  from  the  ear  by  a  pair  of  small  post- 
labials;  tertiary  temporal  (the  lower  secondary  presumably  want- 
ing) small;  ear  surrounded  by  16  scales,  the  opening  no  larger  than 
the  first  upper  labial;  six  lower  labials,  the  last  elongate;  mental 
large,  deep,  with  a  distinctly  larger  labial  border  than  rostral;  one 
azygous  postmental;  three  pairs  of  chinshields,  only  one  in  contact; 
the  postgenial  large,  bordered  on  its  anterior  inner  edge  by  a  scale 
wider  than  long;  eye  small,  the  lower  eyelid  with  four  or  five  en- 
larged opaque  scales  separated  from  the  suboculars  by  at  least  three 
rows  of  granules;  two  or  three  median  palpebral  scales  directly  in 
contact  with  superciliaries,  others  separated  by  small  granular  scales. 
Scales  on  the  dorsal  surface  and  sides  about  equal  in  size,  24  rows 
about  the  middle  of  the  body;  29  rows  about  neck  behind  ear;  30 
about  the  constricted  portion  of  the  neck;  30  about  body  in  axillary 
region;  15  rows  about  the  base  of  the  tail;  57  scales  from  occiput 
to  above  the  anus;  scales  under  the  tail  two  and  one-half  to  three 
times  as  broad  as  long;  preanal  scales  large,  broad,  with  two  small, 
scarcely  differentiated,  scales  on  each  side,  the  outer  overlapping 
inner;  lateral  postanal  scale  not  or  scarcely  differentiated;  scales 
behind  ear,  about  insertion  of  arm  and  in  axillary  region,  on  posterior 
side  of  femur,  behind  insertion  of  hind  limbs,  and  along  side  of  anus, 
with  numerous  distinct  pits;  two  small  auricular  lobules. 

Legs  moderately  large,  separated  when  adpressed  by  a  length  of 
three  or  four  scales;  a  very  small  area  of  granular  axillary  scales; 
wrist  tubercle  fiat,  well  differentiated;  several  larger  rounded  tu- 
bercles on  palm  mixed  with  smaller  tubercles;  lamella  formula,  5  : 
8  :  11  :  10  :  7.  Heel  bounded  by  five  large,  flattened,  tubercular 
scales,  contiguous  or  overlapping  one  or  two  differentiated  tubercles 
on  sole;  lamella  formula,  5  :  9  :  11  :  13  :  9.  Teraiinal  lamella  on 
toes  not  tightly  bound  about  claws;  no  intercalated  series  of  scales 
along  the  side  of  the  fourth  toe. 

Color  in  Life.  Above  a  light  olive-brown,  the  head  somewhat 
browner;  darker  flecks  in  the  median  part  of  each  scale,  more  prom- 
inent posteriorly  and  tending  to  form  dotted  darker  lines;  a  dorso- 
lateral cream  line,  bordered  on  its  inner  edge  with  black,  begins  on 
rostral  and  continues  on  the  side  of  the  head  and  neck,  but  loses  its 
identity  on  the  shoulder;  the  two  median  scale  rows  are  a  shade 
darker  than  the  two  adjoining  rows  on  each  side;  beginning  on  the 


260  The  University  Science  Bulletin 

side  of  the  head  is  a  dark  bhickish  or  blackish-brown  stripe,  the 
color  not  uniform;  each  scale  with  light  bronze  areas,  the  black 
concentrated  on  the  anterior  part  of  the  scales  and  tending  to  form 
an  indistinct  line  on  each  scale  row ;  a  cream  line  beginning  on  the 
rostral  passes  along  the  lower  edge  of  the  first  four  labials  and 
through  the  middle  of  the  last  three,  the  edges  of  the  line  clearly 
demarcated,  terminates  in  the  lower  anterior  corner  of  the  ear;  be- 
low the  dark  lateral  stripe  the  ground  color  is  grayish,  and  the  scales 
have  darker  areas  forming  two  or  three  very  indistinct,  dotted  lines ; 
lower  labials  light,  bordered  with  darker;  chin,  lower  side  of  neck 
and  breast  light,  a  few  of  the  scales  with  darker  flecks;  hind  legs 
darker  than  forelegs,  each  scale  with  lighter  flecking;  tail  bluish 
gray,  lavender  blue  below ;  lamellae  under  toes  dark. 

Variation.  The  table,  giving  data  from  a  part  of  the  series  avail- 
able, shows  the  principal  variation  of  this  species  as  regards  meas- 
urements and  scale  variation. 

The  number  of  scale  rows  is  24  save  in  two  cases  where  there  are 
but  22  rows ;  the  number  of  upper  labials  is  constantly  7 ;  one  speci- 
men shows  the  third  and  fourth  partially  fused  on  one  side.  Only 
a  single  specimen  shows  the  parietals  separated,  and  this  only  very 
narrowly.  Scales  about  the  ear  vary  from  15  to  18,  the  numbers 
15  and  16  most  frequent;  the  higher  numbers  rarely  occur.  The 
scales  from  occiput  to  above  anus  vary  from  57  to  61,  59  being 
twice  as  frequent  as  the  other  numbers.  One  postmental  and  no 
postnasal  seem  to  be  invariable  characters.  The  seventh  labial  is 
invariably  the  largest,  frequently  double  the  size  of  the  sixth ;  sub- 
digital  lamellae  under  fourth  toe  11-14,  12  and  13  being  the  most 
usual  number.  The  number  of  the  supraoculars  is  invariable.  The 
temporals  are  surprisingly  stable  in  character. 

In  color,  the  ground  color  varies  in  shade  from  darker  to  lighter. 
In  younger  specimens  the  color  on  the  dark  lateral  stripe  may  be 
uniformly  black.  The  character  of  the  light  lines  does  not  vary, 
and  is  identical  in  the  very  young  (31  mm.)  and  in  adults.  In  the 
young  the  tail  is  a  bright  blue,  and  this  color  is  usually  retained  by 
the  adults,  but  with  blackish  or  gray  flecks  breaking  the  uniformity. 
There  is  no  trace  of  a  lateral  line  from  behind  the  ear  along  the 
side.  The  head  in  the  young  is  never  black.  The  dotted  dark  dorsal 
Tnes  are  more  distinct  in  some  specimens  than  in  others. 

Relationship.  The  relationship  of  this  species  is  with  Eumeces 
dugesi,  despite  the  very  striking  difference  in  the  character  of  the 
scales  of  the  top  of  the  head.     It  shows  a  parallel  development  with 


Taylor:  A  New  Species  of  Lizard 


261 


CO    K     >Tj     tr'    TJ     *x(    2; 


■3      £■     s- 


-     °      il 


5'    ;      "     3'    " 


a  *^   51   t-'   ^ 


ai    1-3    CO    CO 


^       O       >—       to       •t'       ^1 


to       to       I--       1—       !-■       I—       W       lo 
*.       O       ii       CO       to       1—  -  - 


^1      to     »     C-. 


M       O       I-- 


qj 


-1       2; 


C;<toi-'i—      i-'i-'i-'*-fOOC!:9j 
Oi—Wrfi^tD>(^CZ;lOMOOOOO>rf». 


02     ■'O  !    to 


a:       to      1—      !-•      t-- 


~)OH-10*>MO*>0 


q. 


CI        to        t-"        H-        "-■ 


O       -J      o 


)—      to     *•     ^J     oc      *. 


10     o     ^     oc     1-     o 


Or        to        I— 


^01-' 


tn  CO 


I  ^ 


K     CO      ^ 

»       (O      "       ^      c 


I  s 


t-"      to      it-      ^I      O       to      C      00      O:      ->      Ci      «      10      ~J      to 


18—3482 


262  The  University  Science  Bulletin 

that  which  has  taken  place  in  the  lynxe  group.  Furcirostris  with 
three  supraoculars  stands  to  lynxe  as  dugesi  does  to  indubitus.  How- 
ever, it  appears  that  in  the  two  latter  the  different  characters  have 
become  stabilized. 

This  new  form  may  readily  be  separated  from  Eumeces  dugesi  by 
the  character  of  four  supraoculars,  three  touching  the  frontal,  instead 
of  three  supraoculars,  with  only  two  touching  the  frontal.  The  con- 
trast of  color  between  the  dorsal  surface  and  the  sides  is  much  more 
pronounced  in  dugesi  than  in  indubitus;  the  former  often  becomes 
yellowish  bronze,  and  even  silvery  above.  In  all  the  specimens 
examined,  thirty -three  in  all,  there  is  no  evidence  that  the  characters 
separating  the  two  forms  overlap  or  intergrade. 

The  species  may  be  more  distantly  related  to  Eumeces  brevirostris 
Giinther  as  suggested  by  the  presence  of  a  large  seventh  labial  which 
makes  contact  with  the  upper  secondary  temporal. 

Remarks.  The  present  known  distribution  of  this  form  is  the 
Mexican  states  of  Morelos,  Mexico,  and  Eastern  Michoacan.  'So 
far  as  is  known,  it  is  a  high  mountain  form,  as  all  specimens  taken 
were  found  in  the  mountains  in  pine  forest,  under  rocks  or  logs. 

The  food  of  this  species,  judged  by  stomach  contents,  consists 
wholly  of  small  insects.  An  examination  of  the  reproductive  organs 
gave  no  clue  as  to  whether  the  form  is  oviparous  or  ovoviviparous, 
since  neither  the  ovaries  nor  uteri  contained  developing  eggs.  It  is 
highly  probable,  however,  that  this,  like  its  close  relatives,  dugesi,  is 
ovoviviparous. 

A  single  paratype  (No.  1672)  was  presented  to  the  Institute  de 
Biologia  in  Mexico  City. 


264  The  University  Science  Bulletin 


PLATE  XXIV 

Fig.  1.  Enlarged  drawing,  lateral  view  of  head  of  Eumecen  indulnius  sp.  nov. 
(No.  1727,  E.  H.  T.  and  H.  S.  Collection),  which  shows  slight  variation  in  scale 
characters  from  the  type.  Actual  length  of  head,  snout  to  end  of  parietals, 
10  mm. 

Fig.  2.   Same,  dorsal  view. 


Taylor:  A  New  Species  of  Lizard 


265 


PLATE  XXIV 


266  The  University  Science  Bulletin 


PLATE  XXV 

Fig.  a.   Photograph  of  Eumeces  indubitus  (No.  1674,  E.  H.  T.  and  H.  S. 
Collection)  from  type  locality.    About  actual  size. 

Fig.  B.   Photograph  of  the  type  specimen,  about  actual  size. 


Taylor:  A  New  Species  of  Lizard 
PLATE  XXV 


267 


Fig.  a. 


FiQ.  B. 


THE  UNIVERSITY  OF  KANSAS 

SCIENCE  BULLETIN 

Vol.  XXL]  March,  1933.  [No.  G. 


Observations  on  the  Courtship  of  Turtles 

EDWARD   H.   TAYLOR, 
Department   of  Zoology,   University   of  Kan.sas 

ON  October  2,  1932,  about  150  turtles  were  received  by  the  De- 
partment of  Zoology,  University  of  Kansas,  from  the  ponds  of 
the  State  Fish  Hatchery  at  Pratt,  Kansas.  On  receipt  they  were 
placed  in  a  water  tank,  4  feet  by  18  feet,  in  a  room  in  the  basement 
of  the  biology  building.  This  room  had  no  outside  source  of  light, 
and  was  kept  at  ordinary  room  temperature. 

A  few  days  after  they  arrived  they  were  given  a  large  feed  of 
cooked  table  refuse  consisting  of  fruits,  meats,  and  vegetables,  which 
they  ate  with  great  greediness.  A  census  was  taken  on  October  15, 
and  the  following  specimens  were  present:  3  Amy  da  mutica;  14 
Chelydra  serpentina;  12  Pseudemys  elegans;  35  Kinosternon  flavc- 
scens;  80  Chrysemys  marginata  belli.  Several  small  Chrysemys  had 
died,  or  had  been  killed;  and  several  had  been  partially  eaten  by 
the  larger  turtles.  At  this  time  a  second  large  feed  of  cooked  meat 
and  vegetables,  and  raw  lettuce  was  given  them.  All  seemed  to  eat 
to  the  point  of  satiation. 

On  October  16  I  observed  a  male  of  Chrysemys  behaving  in  an 
unusual  manner.  He  would  start  following  a  female  of  the  same 
species,  and  by  swimming  faster  would  quickly  overtake  her;  then, 
by  whirling  himself  about  in  the  water,  he  would  start  swimming 
backwards  just  in  front  of  the  female,  who  continued  swimming 
straight  ahead.  Then  the  male  would  slow  down,  and  as  the  female 
approached,  he  would  push  himself  forward  toward  her,  stretch 
out  his  arms  full  length  forward,  with  the  palms  turned  outward, 
the  claws  and  fingers  straightened,  and  vibrate  them  rapidly  against 
the  chin  and  lores  of  the  swimming  female.  The  fingers  would 
touch  the  female  from  five  to  seven  times  during  the  continued 
vibration,  which  lasted  perhaps  less  than  one  and  a  half  seconds. 

(269) 


270  The  University  Science  Bulletin 

He  would  then  withdraw  his  arms,  and  continue  swimming  back- 
ward ahead  of  the  female.  After  an  interim  of  from  four  to  five 
seconds,  this  same  action  would  be  repeated  in  practically  the  same 
way.  In  this  particular  manner  the  act  was  repeated  twelve  times 
before  they  were  interrupted  by  another  turtle  separating  them  by 
chance.  Then  the  male  turned,  sought  out  another  mate,  and  began 
the  same  type  of  courtship  with  her. 

The  turtles  were  watched  at  this  time  for  nearly  an  hour,  and 
eight  courtships  of  this  type  were  observed,  although  it  was  not  im- 
possible that  the  same  individuals  had  taken  part  in  more  than  one. 
The  activity  took  place  usually,  but  not  invariably,  at  the  surface 
of  the  water. 

On  October  17  the  tanks  were  drained  and  cleaned  and  filled  with 
fresh  tap  water,  after  which  courtship  began  again,  and  as  many  as 
eight  pairs  were  seen  performing  this  strange  series  of  maneuvers 
at  the  same  time.  It  is  probable  that  all  the  adult  specimens  were 
active  by  this  time.  The  females  seemed,  throughout  my  observa- 
tions, to  be  quite  indifferent  to  the  males,  paying  no  more  attention 
to  one  than  to  another  of  the  males  that  approached  them.  When 
the  lights  were  turned  out  the  turtles  all  seemed  to  become  quiescent; 
but  shortly  after  turning  on  the  lights,  the  whole  tank  would  begin 
movement,  and  the  courtships  would  be  continued  by  the  Chrysemys. 
The  other  species,  however,  remained  quiet  at  the  bottom  of  the 
tank.  On  the  succeeding  days  occasional  observations  showed  that 
the  activity  was  continuing  unabated. 

On  October  26  courtship  was  first  noticed  between  members  of 
the  Pseudemys  elegans  group ;  and  the  procedure  was  almost  identi- 
cal with  that  of  the  Chrysemys,  at  least  the  differences  are  not 
easily  described,  save  that  the  act  was  usually  not  repeated  more 
than  four  or  five  times  without  a  temporary  cessation  of  activity. 
Up  to  this  date  no  turtles  were  seen  in  copulation. 

On  October  27  a  group  of  Chrysemys  males  that  were  actively 
carrying  on  a  courtship  were  placed  in  a  separate  tank.  At  first 
they  kept  swimming  about  very  rapidly  as  if  seeking  females.  Their, 
mode  of  sex  recognition  could  not  be  'determined,  save  that  when  a 
male  was  approached,  and  the  arms  were  extended,  the  second  male 
would  thrust  his  arms  forward,  and  the  two  would  abandon  each 
other.  My  observations  were  interrupted,  and  I  left,  leaving  the 
room  lighted;  on  my  return  three  pairs  of  males  were  keeping  up 
the  play  with  each  other.  Each  kept  the  same  approximate  posi- 
tion, moving  back  a  few  inches;  then  each  advancing,  vibrated  the 


Taylor:  Courtship  of  Turtles  271 

fingers  against  the  fingers  of  the  other.  On  occasion  this  would  be 
kept  up  for  two  or  three  minutes. 

On  October  28,  shortly  after  the  room  was  lighted,  the  males  again 
became  active ;  and  after  a  considerable  amount  of  swimming  about, 
three  pairs  of  males  were  again  observed  continuing  the  courtship. 
When  three  females  were  placed  in  the  tank,  two  and  even  more 
males  might  attempt  to  engage  her  attention.  On  this  date  a  male 
Kinosternon  jiavescens  was  observed  attempting  copulation  with  a 
small  female  Chrysemys.  The  shell  of  the  female  was  held  by  all 
four  feet  of  the  Kinosternon  from  a  dorsal  position.  The  latter  would 
lift  his  body  and  strike  the  shells  together. 

On  October  29  a  pair  of  Kinosternon  were  observed  in  copulation, 
and  two  other  pairs  were  observed  clasping  and  apparently  tapping 
their  bodies  together.  Here  the  semiprehensile  tail  with  its  spine- 
like tip  serves  as  a  very  efficient  grasping  organ.  When  clasping,  the 
male  holds  the  female's  shell  by  all  four  of  his  feet  and  his  tail. 
In  copulation  the  hold  of  the  front  feet  is  loosened,  and  the  male 
stands  erect. 

Two  Chelydra  serpentina  were  seen  on  November  11,  their  heads 
close  together  in  shallow  water;  they  appeared  to  be  gulping  in 
water  and  then  forcing  it  through  their  nostrils,  causing  a  "boiling" 
of  the  water  in  two  areas  above  their  heads  at  the  surface.  This 
was  continued  for  about  ten  minutes,  their  heads  close  together, 
sometimes  touching.  After  cessation  of  this  behavior,  I  determined 
that  a  male  and  a  female  were  involved.  By  this  date  the  activity 
of  the  other  forms  had  practically  ceased,  and  no  others  were  ob- 
served in  copulation,  except  for  a  single  pair  of  Chelydra,  which 
were  found  in  copulation  November  4 ;  but  these  became  disengaged 
almost  as  soon  as  they  were  discovered,  and  before  accurate  ob- 
serv^ations  could  be  made.  A  week  later  all  the  specimens  were 
preserved.  The  Amy  da  mutica  were  all  females ;  no  sexual  activity 
was  observed  in  this  species. 

That  the  observed  behavior  of  these  forms  constitutes  the  typical 
courtship  behavior  can  scarcely  be  questioned,  even  though  it  may 
have  been  induced  by  high  temperature  and  heavy  feeding  at  a  sea- 
son of  the  year  when  it  does  not  normally  occur.  The  peculiar 
courtship  pattern  is  essentially  identical  in  Chrysemys  marginata 
belli  and  Pseudemys  elegans,  and  may  likewise  be  the  same  in  other 
aquatic  genera  of  the  family  Testudinidse. 

I  am  uncertain  whether  the  curious  behavior  of  Chelydra  ser- 
pentina, is  to  be  considered  as  a  courtship  activity  or  not.  It  was 
observed  only  on  a  single  occasion. 


THE  UNIVERSITY  OF  KANSAS 

SCIENCE  BULLETIN 

Vol.  XXL]  March,  1933.  [No.  7. 


A  New  Bassariscus  from  the  Lower  Pliocene  of  Nebraska 

CLAUDE  W.  HIBBARD, 

The  University  of  Kansas  Museum  of  Vertebratu  Paleontologj' 


Abstract:  A  new  carnivore,  Bassariscus  ogallalce,  is  described  from  a  speci- 
men in  the  University  of  Kansas  Museum  of  Vertebrate  Paleontology  from  the 
type  locality  of  the  Ogallala  formation,  Ogallala,  Nebraska.  Comparison  is 
made  with  other  known  fossil  species  of  the  genus. 


THROUGH  the  courtesy  of  Dr.  H.  H.  Lane,  acting  curator  in 
charge  of  the  Department  of  Vertebrate  Paleontology  of  the 
University  of  Kansas,  I  have  been  given  the  opportunity  of  studying 
and  describing  the  right  lower  jaw  of  a  new  species  of  Bassariscus. 

Fossil  remains  of  Bassariscus  are  rare,  since  there  have  been  re- 
ported heretofore  only  three  specimens  of  B.  antiquus  Matthew  and 
Cook;  one  from  the  Upper  Miocene  of  western  Nebraska,  one  from 
the  Upper  Miocene  of  western  Nevada,  and  another  from  the 
Pliocene  of  San  Joaquin  Valley  of  California.  There  is  but  a  single 
known  specimen  of  B.  parvus  Hall  from  the  Upper  Miocene  of  west 
central  Nevada. 

The  specimen  herein  reported  is  from  near  the  top  of  the  Lower 
Pliocene.  It  was  collected  on  the  Feldt  Ranch,  approximately  two 
miles  east  and  one-half  mile  north  of  the  town  of  Ogallala,  Nebraska. 

The  specimen  was  obtained  during  the  summer  of  1931  by  William 
K.  McNown  and  myself  while  making  a  survey  of  the  Lower 
Pliocene  through  southern  Nebraska,  western  Kansas  and  western 
Oklahoma  for  the  Department  of  Vertebrate  Paleontology  of  the 
Muesum  of  the  University  of  Kansas. 

(273) 


274  The  University  Science  Bulletin 

Bassariscus  ogallalce  sp.  nov. 

(Plate  XXVI;    figs.   1-3) 

Type.  No.  3749,  University  of  Kansas  Museum  of  Vertebrate 
Paleontology.  Right  lower  jaw  bearing  P3,  P4,  M^  and  Mg,  with 
alveoli  of  I3,  C,  Pi  and  P,,  lacking  angle,  condyle  and  coronoid 
process. 

Horizon  and  Type  Locality.  From  the  type  locality  of  the  Ogal- 
lala  formation  and  35  feet  below  the  top  of  the  local  section,  Lower 
Pliocene,  approximately  two  miles  east  and  one-half  mile  north  of 
the  town  of  Ogallala,  Keith  county,  Nebraska,  on  the  Feldt  ranch. 
(The  correlation  was  made  by  M.  K.  Elias,  member  of  the  Kansas 
Geological  Survey.) 

Diagnosis.  See  Table  2  of  Measurements  and  Ratios.  The  length 
of  Ml  and  Mo  are  the  same,  making  the  ratio,  length  of  Mi  to  the 
length  of  Mo,  100  or  1.  The  ratio,  length  of  the  talonid  of  Mi  to 
the  length  of  the  trigonid  of  Mi ;  and  the  ratio  length  of  Mi  to  the 
length  of  M2  are  greater  than  in  any  other  known  species  of  Bas- 
sariscus; the  length  of  the  trigonid  of  Mi,  and  the  ratio,  width  of 
M2  to  the  length  of  M2  are  less. 

Description  of  Type.  The  jaw  is  of  an  adult,  as  is  shown  by  the 
worn  cusps.  P^  has  a  well-developed  accessory  cusp  posterior  and 
lateral  to  the  protocone,  which  exceeds  in  size  any  examined  of  the 
living  species,  Basariscus  astutus  jiavus  Rhoads.  There  is  no  trace  of 
an  accessory  cusp  on  Py.  Mi  has  strong  and  well-developed  cusps. 
Mo  has  a  well-developed  paraconid,  greatly  exceeding  the  protoconid 
in  size.  The  entoconid,  hypoconulid,  and  hypoconid  are  distinct 
and  well  developed.  The  distance  across  M^  from  the  top  of  the 
entoconid  to  the  top  of  the  hypoconid  is  3.5  mm.  The  anterior 
mental  foramen  is  larger  and  is  located  more  posteriorly  than  that 
of  B.  astutus  flavus.  It  is  below  the  middle  of  P2,  while  in  B.  astutus 
flavus  it  is  below  the  anterior  root  of  P,.  The  posterior  mental 
foramen  is  smaller  than  that  of  B.  astutus  flavus  and  is  below  the 
posterior  root  of  P3  and  lies  slightly  dorsad  to  the  anterior  mental 
foramen,  a  condition  not  observed  in  B.  astutus  fiavu^.  The  inferior 
dental  foramen  is  larger,  and  is  situated  more  posterior  to  the  an- 
terior surface  of  the  ascending  ramus,  although  closer  to  M,  than  in 
B.  astutus  flavu£.  The  nearness  of  the  dental  foramen  to  Mo  is 
accounted  for  by  the  large  development  of  that  tooth.  The  jaw  is 
deeper,  thicker,  and  heavier,  though  shorter,  than  that  of  B.  astutus 
flavujs. 


Hibbard:  a  New  Bassariscus  275 

Remarks.  The  comparison  of  B.  ogallalce  with  that  of  other  fossil 
Bassariscus  shows  the  following  similarities  and  differences.  (See 
Table  1.)  Hall  ^  has  placed  Probassariscus  antiquus  matthewi 
Merriam-  in  synonomy  with  Bassariscus  antiquiLS  Matthew  and 
Cook.^  A  comparison  of  B.  ogallalce  with  B.  antiquus  shows  a 
slightly  smaller  M^  with  greatest  difference  in  the  width  of  the 
talonid,  which  is  an  average  of  0.6  mm.  smaller.  The  ratio,  width 
of  the  talonid  of  M^  to  the  length  of  M^  is  less  than  in  B.  ogallalce 
and  does  not  fall  within  the  limits  of  individual  variation  of  B. 
antiquum.  B.  ogallalce  may  be  distinguished  from  B.  antiquus  by 
the  size  of  Mo.  M^  and  M2  of  B.  ogallalce  are  the  same  length,  mak- 
ing the  ratio,  length  of  M^  to  the  length  of  M2,  100  or  1.  The  ratio, 
width  of  M2  to  the  length  M.  is  42.9,  the  smallest  known  ratio  of 
any  Bassariscus. 

Bassariscus  ogallalce  is  distinguished  from  Bassariscus  parvus 
Hall  ■*  by  the  ratio,  length  of  the  talonid  of  M^  to  the  length  of  the 
trigonid  of  M^,  which  is  10.7,  greater  in  B.  ogallalce.  Another  out- 
standing difference  between  B.  ogallalce  and  B.  parvus  is  the  crowded 
condition  of  the  premolars  of  the  latter,  which  does  not  occur  in 
B.  ogallalce. 

The  specimen  is  named  for  the  Ogallala  formation,  from  which  it 
was  collected. 


1.  Hall,  E.  R.,  Univ.  Calif.  Publ.  Bull.  Dtept.   Geol.,  Vol.   16,  No.  11,  p.   437,  March  17, 
1927. 

2.  Merriam,  J.  C,  Univ.  Calif.  Publ.  Bull.  Dept.  Geol.,  Vol.  6,  p.   246,  Part  2,  &ept.  16, 
1911. 

3.  Matthew  and  Cook,  Bull.  Am.  Mus.  Nat.  Hist.,  Vol.   27,  p.  337,  Sept.   3,  1909. 

4.  Hall,  E.  R.,  op.  cit.,  p.  435. 


276 


The  University  Science  Bulletin 


pq 


03 


Ratio,  width  of  M,  to 

C^  r-C 

0: 

length  of  Mj 

M 

■* 

(NOO 

M2  width 

rocq 

cc 

Ratio,  length  of  M,  to 

coo 

length  of  M2 

8 

00  t^ 

^lo  length 

*     " 

* 

"o 

"S 

o 
_2 

Ratio,  width  of  talonid  of 

roi^c 

a 

05 

M,,  to  length  of  M, 

0:  toot 

5?          d 

o         !:: 

l^iC  -X 

^ 

s     -s 

M,  width  of  talonid 

COMC 

■:' 

e<5 

d     .t; 

2    ^^ 

tS       o 

te  I 

onto 
a  of 

Ratio,  length  of  talonid 

■*T)<'* 

c 

of  Ml,  to  length  of 

'J'TfTr 

Tj 

tC 

2    -  ° 

trigonid  of  M^ 

t^t- 1^ 

c 

i^ 

V           ^ 
K       ®  — 

>    go 

Ml  length  of  trigonid 

■* 

n        «  >> 

on  1 
cry. 
Vert 
olog 

V3  -g      c 

c■^c•l^ 

t-                   1 

^k"^| 

M,  length  of  talonid 

wro:' 

0 

M 

,9-  SH 

O  g  3 

)rnia 

Natui 
Muse 
ite  of 

Ml  length 

l^^^  t~         to         t- 

.ill         '-' 

^'^  >>.ti 

Sgll 

5  ?"-' 

0." 

:  cj 

a           c 

o  a'g  ca 

a         0. 

V             0 

03  -■  li;  ^<-' 

t^  2 
0^ 

§        :2 

C.  V.  P.,  Uni 
.  N.  H.,  Ameri 
.  M.  V.  P.,  Ka 
T.  C.  V.  P.,  C 

13 
< 
0 

0 

-5 

at) 

^        . 

4    1 

)                ^ 

dSt>^ 

K^.    ;^> 

-.       ^'       i 

C    -4-^    -,- 

iD^Wd 

03  "  ; 

>     i 

>  0  c 

;    t    0 

§     :§ 

.t:  0  a 

c 

aj 

>Xti 

w     w 

■■f^  : 

■     ^   •  •  (li 

3 

5d^; 

«>  '^.• 

Mt! 

s 

^i> 

°B  c 

,     go    T?.  :     1 

rt  OS  0 

-  ■„-c 

)      a 

"Ji  fr-'c 

eo2. 

p-c 

g   OJ   0 

i^-^" 

P    8^    1 

ii° 

To  Or 

'Cot 

fw 

etc 

e 

^ 

CCICOO 

»r^ 

»T. 

2o)  race 

2c 

21' 

^rr? 

c  _ 
-     0?* 

£§!? 

S   3 


■5S 
O 

.  c 

IS  c 


"3  =3 


hh 


Hibbard:  a  New  Bassariscus  277 
Explanation  of  Measurements  (Hall)^ 

M],  length:    taken  at  cinguliini;   thus,  usually,  but  not  always,  the  greatest  length. 

Ml,  length  of  talonid :  taken  on  lateral  side  of  tooth  from  posterior  bas-e  of  protoconid  to  pos- 
terionnost  extension  of  talonid. 

Ml,  length  of  trigonid ;  taken  from  posterior  base  of  protoconid  to  anterior  end  of  tooth  at 
eingulum. 

Ml,  width  of  talonid;  taken  from  indentation  on  lateral  side  of  tooth  just  behind  the  pro- 
toconid to  opposite  side  of  tooth  and  perpendicular  to  lonsitudinal  axis  of  tooth. 

Ma,  length;  taken  from  eingulum  at  posterior  end  of  tooth  to  most  anterodorsal  point  of  tootli, 
usually  but  not  always,  greatest  length. 

Ma,  width;  taken  from  indentation  on  lateral  side  of  tooth  .iiist  behind  the  prntor(>ni<l  to 
opposite  .side  of  tooth  and  perpendicular  to  longitudinal  axis  of  tooth. 

TABLE  2 

Measurements  and  Ratios  of  Bassariscus  ogallalce 
The  ratios  given  in  the  table  are  the  actual  ratios  multiplied  by  100. 

Length  from  posterior  border  of  canine  aveoliis  to  posterior  mm. 

border  of  Mo 29 

Length  of  Pi  to  Mo,  inclusive 18.2 

Length  of  P3  to  M2,  inclusive 22 

Length  of  P.^  and  P4,  inclusive 8 

Length  of  P3 4 

Breadth  of  P3 2 

Breadth  of  P4 2.9 

Length  of  P4 4.5 

Length  of  Mi 7 

Breadth  of  Mi 3.5 

Length  of  M2 7 

Breadth  of  Mo 3 

Length  of  talonid  of  Mi 3 

Length  of  trigonid  of  Mj 4 

Ratio,  length  of  talonid  of  Mi  to  length  of  trigonid  of  Mi 75 

Width  of  talonid  of  Mi 3 

Ratio,  width  of  talonid  of  Mi  to  length  of  Mi 42.9 

Ratio,  length  of  Mi  to  length  of  Mo 100  or  ] 

Ratio,  width  of  Mo  to  length  of  M2 42.9 

Length  of  alveolus  of  canine 3 

Length  of  alveolus  of  P2 3.5 

Length  of  alveolus  of  Pi 2 

Breadth  of  alveolus  of  Pi 1.5 

Breadth  of  alveolus  of  Po 1.5 

Depth  of  jaw  beneath  Mi 8 

5.    Hall,   E.   R.,  Op.   Cit.   p.   445. 


19— a482 


278  The  University  Science  Bulletin 


BIBLIOGRAPHY 
Hall,  E.  Raymond 

1927.    Species    of    the    Mammalian    Subfamily    Bassari^cinae.      Univ.    Calif. 
Publ.  Bull.  Dept.  Geol..  \'ol.  16,  No.  11,  pp.  435-488;  March  17.  1927. 


1930.    A  Ba.ssarisk  and  a  New  Mustelid  from  the  Later  Tertiary  of  Cali- 
fornia.   Jour.  Mammalogy,  Vol.  11,  No.  1,  pp.  23,  24;  February,  1930. 

M.\TTHEW,  W.  D. 

1918.    Contributions  to   the   Snake   Creek  Fauna.     Bull.   Amer.   Mus.  Nat. 
Hist..  XXXVHI.  p.  185. 


1924.    Third  Contribution  to  the  Snake  Creek   Fauna.     Bull.   Amer.   Mus. 

Nat.  Hist.,  L,  p.  65. 
M.\TTHEW,  W.  D.,  and  Cook,  H.\rold 

1909.    A  Pliocene  Faima  From  Western  Nebraska.    Bull.  Amer.  Mus.  Nat. 

Hist.,  Vol.  27,  p.  377;  September  3,  1909. 

Merriam,  John  C. 

1911.  Tertiaiy  Mammal  Beds  of  Virgin  Valley  and  Thousand  Creek  in 
Northwest  Nevada.  Univ.  Calif.  Publ.  Bull.  Dept.  Geol.,  Vol.  6,  p. 
246,  pt.  2;  September  16,  1911. 


1916.  Tertiaiy  Vertebrate  Fauna  from  the  Cedar  Mountain  Region  of 
Western  Nevada.  Univ.  Calif.  Publ.  Bull.  Dept.  Geol..  Vol.  9,  pp. 
175,  176;  February  23,  1916. 

Rhoades,  S.  N. 

1894.  Geographic  Variation  in  Bassariscus  astutus,  with  Description  of  a 
New  Subspecies.    Proc.  Acad.  Nat.  Sci.  Phila.,  1893,  pp.  413-418. 


THE  UNIVERSITY  OF  KANSAS 

SCIENCE  BULLETIN 

Vol.  XXL]  March,  1933.  [No.  S. 


Two  New  Species  of  Coelacanthus  from  the  Middle 
Pennsylvanian  of  Anderson  County,  Kansas 

CLAUDE  W.  HIBBARD 

Kansas  University   Museum   of  Vertebrate   Paleontology 


Abstract:  Two  new  species  of  Ccelacanthid  fishes,  Cadacanthus  newelli  and 
CcBlacanthus  arcuatus,  are  described  from  specimens  in  the  University  of 
Kansas  Museum  of  Vertebrate  Paleontology.  Both  species  are  from  the 
Rock  Lake  shale,  Stanton  formation,  Missouri  series,  near  the  top  of  the 
Middle  Pennsylvanian  of  Mid-Continent.  Anderson  county,  Kansas.  A  com- 
parison is  made  with  other  known  species  of  the  genus  from  the  Pennsyl- 
vanian. 


THROUGH  the  courtesy  of  Dr.  H.  H.  Lane,  curator  in  charge  of 
the  Department  of  Vertebrate  Paleontology  of  the  Kansas  Uni- 
versity Museum,  I  have  been  given  the  opportunity  of  studying  and 
describing  the  remains  of  two  apparently  new  species  of  fish  of  the 
genus  Coelacanthus. 

The  remains  of  Coelacanthus  are  known  from  only  a  few  localities 
in  North  America.  C.  rohustus,  C.  ornatus,  and  C.  elegans  were 
described  by  Newberry^  from  the  Pennsylvanian  (Linton)  of  Ohio, 
U.  S.;  C.  exiguus,  by  Eastman^  from  the  Pennsylvanian  (Mazon 
creek)  of  Illinois,  U.  S.;  C.  welleri,  by  Eastman^  from  the  Missis- 
sippian  (Kinderhook)  of  Iowa,  U.  S.;  C.  banffensis,  by  Lambe* 
from  the  Triassic?  (Upper  Banff)  of  Alberta,  Canada. 

To  my  knowledge  the  above  records  include  all  localities  where 
Coelacanthus  has  been  found  to  occur  in  North  America,  except  the 

1.  Newberry.  Proctedings  of  the  Academy  of  Natural  Science  of  Philadelphia.  Vol. 
VIII,   1856,  p.   98. 

2.  Eastman,  C.  R.     Jour.   GeoL,  X,  p.   538. 

3.  Eastman.  C.   R.     .Jour.   Geol.,  XVI,  p.   358. 

4.  Laiiibe,  L.   M.      Trans.   Roy.   Soc.   Canada,   X,   sec.   IV,   p.   38. 

(279) 


280  The  University  Science  Bulletin 

specimen  reported  by  Eastman  in  the  Proceedings  of  the  U.  S.  Na- 
tional Museum,  vol.  52,  p.  271,  where  he  says,  "A  supposed  new 
form  has  recently  been  discovered  by  Prof.  E.  H.  Barbour  in  the 
Coal  Measures  of  Nebraska." 

The  specimens  herein  reported  are  from  near  the  top  of  the  Middle 
Pennsylvanian  of  Mid-Continent,  Stanton  formation,  Rock  Lake 
shale  member,  from  the  SW14  of  SEVt  of  sec.  32,  T.  19  S,  R.  19  E, 
on  the  Bradford  Chandler  farm,  Anderson  county,  Kansas.  (The 
Rock  Lake  shale  was  identified  by  the  Kansas  State  Geological 
Survey.) 

We  are  greatly  indebted  to  Mr.  Bradford  Chandler,  upon  whose 
farm  the  specimens  were  taken,  for  the  permission  to  collect,  and 
the  kindness  shown  us  while  working  at  the  quarry. 

Caelacanthus  newelli  sp.  nov. 

(Plate  XXA'II,  figs.   2-3) 

Type.  No.  786  F.  University  of  Kansas  Museum  of  Vertebrate 
Paleontology.  Nearly  complete  skeleton,  lacking  pectoral  fins,  with 
crushed  skull  and  only  a  few  complete  scales. 

Horizon  and  Type  Locality.  From  the  Rock  Lake  Shale,  Stanton 
formation,  Missouri  series,  near  the  top  of  the  middle  Pennsylvanian 
of  Mid-Continent.  Collected  on  the  SWi  of  SE^  of  sec.  32.  T.  19  S, 
R.  19  E,  of  the  Bradford  Chandler  farm,  Anderson  county,  Kansas. 

Diagnosis.  The  exposed  ridged  surface  of  the  scale  is  elongate, 
resembling  somewhat  a  truncated  cone.  It  is  1.74  mm.  long  and 
1.39  mm.  wide  at  the  base  of  the  exposed  surface.  The  scales  have 
an  average  of  9  ganoid  ridges  each.  The  ridges  do  not  meet  at  the 
posterior  end  in  an  acute  angle,  but  converge  slightly  toward  the 
median  ridge  near  the  posterior  end  and  then  straighten  out  again 
to  terminate  slightly  posterior  to  the  median  ridge. 

Description  of  Type.  The  specimen  is  nearly  complete.  The 
skull  is  crushed,  leaving  only  the  jugal  plate  in  place  and  perfect 
enough  for  detailed  study.  The  pectoral  fins  are  missing.  The  fish 
was  so  crushed  in  preservation  as  to  push  the  pelvic  fin  upward  onto 
the  side  of  the  body  and  to  bring  the  opposite  pelvic  fin  into  view 
and  nearly  into  the  original  position  of  the  first.  The  fish  was  de- 
posited upon  a  plant  (4)  four  branches  of  which  cross  the  skeleton 
at  right  angles.  Both  the  positive  and  negative  of  the  fish  were 
collected.  The  skull  is  missing  from  the  positive  of  the  shale. 
(PlateXXVII,  fig.  2.) 

The  length  of  the  fish  from  the  snout  to  the  tip  of  the  caudal  rays 
is  115  mm. 


HiBBARD :  New  Species  of  Coelacanthus  281 

Only  a  few  fragmentary  skull  bones  are  present.  Some  are  orna- 
mented with  short,  narrow,  ganoid  ridges;  others  with  closely  set 
tubercles.  There  is  no  trace  of  teeth,  probably  due  to  the  crushed 
condition  of  the  dentary  and  maxillary.  An  outline  of  the  opercular 
region  is  shown,  but  the  operculum  is  missing. 

The  jugal  plate  is  16  mm.  long  and  4.5  mm.  wide.  It  is  orna- 
mented with  fine,  unevenly  spaced,  ganoid  ridges  running  antcro- 
posteriorly,  but  having  a  tendency  to  meet  at  the  center  of  the  plate, 
as  though  radiating  from  this  central  point. 

The  posterior  portion  of  the  skeleton  is  not  distorted.  The  depth 
of  the  body  anterior  to  the  attachment  of  the  anal  fin  is  20  mm. 

The  anterior  dorsal  fin  is  badly  crushed,  bringing  the  rays  close 
together.  Only  6  rays  are  distinguishable.  These  are  twice  as  wide 
at  their  bases  as  the  rays  of  any  of  the  other  fins.  There  is  a  dis- 
tance of  26  mm.  from  the  anterior  base  of  the  anterior  dorsal  fin  to 
the  anterior  base  of  the  posterior  dorsal  fin.  The  posterior  dorsal 
fin  is  slightly  anterior  to  the  anal  fin.  Only  8  rays  can  be  counted. 
The  largest  is  11.5  mm.  long.  This  fin  furnishes  no  suggestion  of  a 
lobate  condition. 

The  caudal  fin  consists  of  24  principal  rays,  12  above  and  12  be- 
low. The  longest  rays  are  26.5  mm.  in  length.  The  supplementary 
caudal  fin  at  the  widest  is  4  mm.,  and  is  17  mm.  long.  There  are 
present  6  supplementary  caudal  rays,  3  above  and  3  below. 

The  anal  fin  is  nearly  perfect.  It  consists  of  8  rays,  the  longest 
being  7  mm.  The  anal  fin  is  the  smallest  of  the  fins  and  gives  no 
suggestion  of  being  lobate.  It  is  slightly  posterior  to  the  anterior 
attachment  of  the  posterior  dorsal  fin. 

The  pelvic  fin  is  composed  of  at  least  20  rays.  This  fin  shows  a 
strongly  lobate  condition.  The  length  of  the  lobe  of  the  pelvic  fin  is 
6  mm.  This  fin  is  located  posterior  to  the  anterior  attachment  of 
the  anterior  dorsal  fin.  The  distance  from  the  anterior  base  of  the 
anal  fin  is  25.5  mm.  Both  the  pectoral  and  pelvic  arches  appear  to 
be  missing. 

The  anterior  portion  of  all  the  fin  rays  present  is  jointed,  but 
shows  no  evidence  of  denticles. 

The  cycloid  scales  of  this  fish  are  covered  on  the  exposed  surface 
by  ganoid  ridges.  The  exposed  ridged  surface  of  the  scales  is  1.74 
mm.  long  and  1.39  mm.  wide  at  the  base  of  the  exposed  surface. 
(Plate  XXVII,  fig.  3.)  The  scales  seem  to  be  of  uniform  shape,  hav- 
ing the  same  pattern  of  ornamentation  where  present  through  the 
different   regions.     The   exposed   ridged   surface   of    the   scales    if" 


282  The  University  Science  Bulletin 

elongate,  resembling  somewhat  a  truncated  cone.  The  scales  have 
an  average  of  9  ganoid  ridges  each.  The  ridges  do  not  meet  at  the 
posterior  end  in  an  acute  angle,  but  converge  slightly  toward  the 
median  ridge  near  the  posterior  end  and  then  straighten  out  again 
to  terminate  slightly  posterior  to  the  median  ridge. 

Remarks.  The  comparison  of  Coelacanthus  newelli  with  the  other 
forms  of  Coelacanthus  from  the  Pennsylvanian  follows :  Coelacanthus 
newelli  differs  from  C.  ornatus  Newberry  in  that  the  raised  ganoid 
ridges  of  the  scales  are  not  parallel;  furthermore,  the  anal  fin  rays 
of  the  former  number  8  instead  of  6.  Newberry  ■"'  states,  "Coelacan- 
thus ornatus  may  be  readily  identified  by  its  small  size,  relatively 
large  cranial  tubercles,  and  very  thin,  delicate  scales  on  which  the 
raised  lines  are  parallel  and  do  not  converge  as  in  C.  elegans  and  C. 
robustu^."  Coelacanthus  newelli  differs  from  C.  elegans  Newberry 
and  C.  rohustus  Newberry  in  that  the  exposed  surface  of  the  scales 
is  not  circular  in  outline  and  is  not  ornamented  by  converging 
ganoid  ridges.  The  number  of  ridges  in  C.  newelli  averages  9,  while 
in  C.  elegans  the  average  is  at  least  20.  It  may  be  at  once  dis- 
tinguished from  C.  exigmts  Eastman  by  its  larger  size. 

The  Coelacanthus  here  called  newelli,  is  so  named  in  honor  of 
Noraian  D.  Newell,  m^io  was  a  graduate  student  in  the  Department 
of  Geology  at  the  University  of  Kansas  at  the  time  he  discovered 
the  fossil-bearing  pocket  in  the  Rock  Lake  Shale,  while  doing  field 
work  on  his  problem  in  the  summer  of  1931. 

Coelacanthus  arcuatus  sp.  nov. 

(Plate  XXVI,  fig.  8;    Plate  XXVII,  fig.   1) 

Type.  No.  787F.  University  of  Kansas  Museum  of  Vertebrate 
Paleontology.  Caudal  region  of  a  coelacanthid  fish  just  posterior 
to  anal  fin,  with  anterior  caudal  fin  rays  and  the  greater  part  of  the 
supplementary  caudal  fin  missing. 

Horizon  and  Type  Locality.  From  the  Rock  Lake  Shale,  Stanton 
formation,  Missouri  series,  near  the  top  of  the  middle  Pennsylvanian 
of  the  Mid-Continent.  Collected  on  the  SW14  of  SEI/4  of  sec.  32,  T. 
19  S.,  R.  19  E,  on  the  farm  of  Bradford  Chandler,  Anderson  county, 
Kansas. 

Diagnosis.  The  exposed  ridged  surface  of  the  scales  is  shaped 
like  a  Gothic  arch  in  outline.  It  is  ornamented  with  an  average  of 
12  to  14  ganoid  ridges  which  are  slightly  heavier  than  those  of  C. 

5.    Newberry,  Paleozoic  Fishes  of  North  America,   U.   S.   Geol.   Survey,  Vol.   XVI,  p.   227. 


Hibbard:  New  Species  of  Coelacanthus  283 

newelli.  The  ridges  converge  toward  the  median  ridge  at  the  pos- 
terior end  of  the  scale,  but  do  not  join  the  median  ridge.  The  ex- 
posed ridged  surface  of  the  scales  is  2.17  mm.  long  and  2.16  mm. 
wide.     (Plate  XXVII,  fig.  1.) 

This  Coelacanthus  may  at  once  be  distinguished  from  all  other 
forms  studied,  by  the  shape  of  the  scale  with  its  constant  pattern 
and  number  of  strong  ganoid  ridges. 

Description  of  Type.  The  specimen  herein  described  came  from 
the  Rock  Lake  Shale,  closely  associated  with  Coelacanthus  newelli, 
but  approximately  18  inches  above  the  former.  The  specimen  con- 
sists of  the  caudal  region  just  posterior  to  the  anal  fin,  with  the  tips 
of  the  caudal  rays  and  the  greater  part  of  the  supplementary  caudal 
fin  missing.  This  caudal  region  represents  a  fish  nearly  twice  as 
large  as  C.  newelli.    (Plate  XXVI,  fig.  8.) 

There  are  24  principal  caudal  fin  rays,  12  above  and  12  below. 
The  supplementary  caudal  fin  is  8  mm.  wide.  There  are  12  rows  of 
cycloid  scales  just  anterior  to  the  attachment  of  the  caudal  rays, 
running  parallel  to  the  median  line.  The  width  of  the  caudal  region 
just  anterior  to  the  attachment  of  the  caudal  rays  is  24  mm. 

Remarks.  The  significance  of  the  number  of  ganoid  ridges  and 
the  shape  of  the  scales  in  Coelacanthus  can  only  be  known  with 
certainty  when  sufficient  material  has  been  collected  from  one 
locality.  It  is  possible  that  age  and  relative  maturity  may  pro- 
foundly affect  the  number  and  arrangement  of  the  ridges,  as  well 
as  the  shape  of  the  scale.  A  large  series  of  specimens  will  be  neces- 
sary in  order  to  indicate  definitely  which  of  these  characters  are  due 
to  age,  and  which  to  speciation. 

The  figures  of  the  scales  of  C.  elegans  (Plate  XXVI,  fig.  7)  and  C. 
lepturus  (Plate  XXVI,  figs.  4,  5,  6  and  9)  have  been  given  for  com- 
parison. They  were  taken  from  the  Geological  Survey  of  the  United 
Kingdom,  Decade  XII,  1866,  after  Huxley. 

The  Coelacanthus  arcuatus  is  so  named  because  of  the  shape  of 
the  scale  which  resembles  so  closely  a  Gothic  arch  in  miniature. 

The  first  material  collected  from  the  Rock  Lake  Shale  by  the 
Department  of  Vertebrate  Paleontology  during  the  spring  and  sum- 
mer of  1932,  belongs  to  the  genus  Coelacanthus.  Besides  the  material 
described,  there  were  secured  many  unassociated  scales  and  fin  rays; 
also  one  pelvic  fin  and  many  skull  fragments. 


284  The  University  Science  Bulletin 


PLATE  XXVI 

Figs.  1-3.  Bassariscus  ogallalce.  Type :  No.  3749,  University  of  Kansas 
Museum  of  Vertebrate  Paleontology.  X  2.27  natural  size.  (1)  Lateral  view. 
(2)  Occlusal  view.    (3)  Lingual  view. 

Figs.  4,  5,  6  and  9.    Scales  of  Ccelacanthus  lepturus,  magnified.  After  Huxley. 

Fig.  7.   Scales  of  Ccelacanthus  elegans,  magnified.    After  Huxley. 

Fig.  8.  Ccelacanthus  arcuatus,  natural  size.  Type:  No.  786F,  University  of 
Kansas  Museum  of  Vertebrate  Paleontology.     Negative  by  Prof.  M.  K.  FJias. 


Hibbard:  New  Species  of  Coelacanthus 


285 


PLATE  XXVI 


.t%ir^#A 


286  The  University  Science  Bulletin 


PLATE  XXVII 

FiQ.  1.  Scales  from  the  caudal  region  of  the  type  No.  787F,  Ccelacanthus 
arcuatus.     X  H-5. 

Fig.  2.  Coelacanthus  newelli;  natural  size.  Type  No.  786F,  University  of 
Kansas  Museum  of  Vertebrate  Paleontology.    Negative  by  Prof.  M.  K.  Elias. 

Fig.  3.  Scales  from  region  near  midline  anterior  to  anal  fin,  of  the  type  No. 
786F,  Ccelacanthus  newelli.    X  11-5. 


Hibbard:  New  Species  of  Coelacanthus 


287 


PLATE  XXVII 


THE  UNIVERSITY  OP  KANSAS 

SCIENCE  BULLETIN 

Vol.  XXL]  March,  1933.  [No.  9. 


Cephalopods  of  the  Pierre  Formation  of  Wallace  County, 
Kansas,  and  Adjacent  Area 

MAXIM  K.  ELIAS 

TABLE  OF  CONTENTS 

Introduction 290 

Stratigraphic  range  of  the  described  species 290 

Methods  of  drawing  sutures  of  Baculites  and  other  Ammonoidea 290 

Description  of  the  species 294 

Baculites  297 

Heteroceras    313 

Scaphiles 314 

Acanthoscaphites    320 

Discoscaphites  323 

Placcnticeras    33 1 

Plates 334 


Abstract  :  The  paper  is  devoted  chiefly  to  the  description  and  to  the  discus- 
sion of  stratigraphic  vakie  of  some  of  the  Upper  Cretaceous  ammonoids  of  the 
Baculites  and  Scaphites  group  in  western  Kansas  and  adjacent  part  of  Colorado. 
The  following  new  names  are  introduced : 

Baculites  pseudovatus  Elias,  n.  sp. 

Baculites  pseudovatus  var.  A.  Elias,  n.  var. 

Bacidites  conipressus  var.  reesidei  Elias,  n.  var. 

Baculites  compressus  var.  corrugatus  Elias,  n.  var. 

Baculites  clinolohatus  Elias,  n.  sp. 

Baculites  meeki  Elias,  n.  sp. 

Discoscaphites  nicolleti  v^ar.  saltgrassensis  Elias  n.  var. 

The  specific  name  compressus  Say  is  very  narrowly  restricted  to  the  topotype 
of  the  species  Baculites  compressus  as  described  by  Meek  (1854,  1876).  Sca- 
phites nodosus  var.  plenus  Meek  is  removed  from  the  Acantoscaphites  nodosus 
group  and  is  elevated  to  the  rank  of  species  of  the  genus  Scaphites  sensu  stricto : 
Scaphites  plenus  (Meek). 

(289) 


290  The  University  Science  Bulletin 

INTRODUCTION 

AMMONITES  of  both  coiled  and  uncoiled  types  are  among  the 
most  abundant  fossils  in  the  Pierre  of  northwestern  Kansas. 
Some  genera  and  species  of  this  order  are  the  most  important  hori- 
zon-markers of  the  local  subdivisions  of  the  Pierre,  and  a  few  of 
them  are  of  great  importance  for  the  correlation  of  the  formation 
with  the  contemporaneous  beds  in  eastern  North  America  and 
Europe.  Owing  to  this  fact  the  writer  devoted  more  time  and  care 
to  the  study  of  these  invertebrates  than  of  the  other  marine  fossils 
of  the  formation.  It  is  intended  to  describe  the  latter  in  a  forth- 
coming separate  paper. 

The  writer  wishes  to  acknowledge  valuable  suggestions  and  criti- 
cism by  Dr.  R.  C.  Moore  and  Dr.  J.  B.  Reeside,  Jr.,  which  helped 
considerably  in  precise  identification  and  in  elaborating  the  descrip- 
tion of  the  species. 

STRATIGRAPHIC  RANGE  OF  THE  DESCRIBED  SPECIES 

In  the  following  tables  the  stratigraphic  distribution  of  the  am- 
monites herein  described  and  a  few  others  are  shown.  The  distri- 
bution in  space  and  time  of  the  most  important  of  these  fonns  is 
also  shown  on  the  stratigraphic  table  of  the  geological  report  on 
Wallace  county  ^( Bull.  18,  State  Geological  Survey  of  Kansas,  1931). 

METHOD  OF  DRAWING  SUTURES  OF  BACULITES  AND 
OTHER  AMMONOIDEA 

In  preparing  the  sketches  of  the  sutures  of  Baculites  and  other 
Aramonoidea  of  the  Pierre  two  methods  were  used.  The  larger 
sutures  were  traced  directly  on  cellophane,  which  was  wrapped 
around  a  portion  of  a  fossil.  In  order  to  add  visibility  and  avoid 
mistakes  in  tracing,  the  space  between  two  neighboring  sutures  was 
painted  with  black  India  ink.  A  fine  drafting  pen  was  used  for 
painting  the  minute  dentations  of  the  sutures.  Cellophane  is  pref- 
erable to  other  transparent  material  (celluloid  film,  wax  paper, 
tracing  cloth,  etc.)  because  it  is  perfectly  transparent,  and,  being 
soft  and  flexible,  can  be  pressed  more  tightly  against  the  uneven 
surface  of  the  variously  sculptured  molds  of  ammonoids.  The  India 
ink  sticks  to  cellophane  fairly  well,  and  fine  lines  can  be  drawn  on 
it.  However,  when  drawing  a  line  over  those  places  where  the 
cellophane  is  not  quite  firmly  pressed  against  the  surface  of  the 
fossil,  one  inevitably  scratches  the  soft  tissue  with  the  sharp  point  of 


Elias:   Cephalopods  of  Pierre  Formation 


291 


o  83  C3  C3 
»  "  m  2 
M"        6 


i-0 

a»  «  Q) 

iH'5'3 

C  (3  "> 


i  s 


££ 


03  ^ 

Is 


o  ® 

*  2 


a 

I 
o 


3 


oa  I  e; 


292 


The  University  Science  Bulletin 


2  ^ 

^  V 
02  2 


£6 


c5  g 


X      !«;      X 


'^ 


Elias:   Cephalopods  of  Pierre  Formation  293 

the  pen.  Owing  to  this  the  lines  soon  become  too  heavy  to  express 
the  finer  details  of  the  sutures.  The  writer  found  that  it  is  much 
safer  and  more  convenient  not  to  trace  on  cellophane  the  suture  lines 
themselves,  but  to  ink  in  the  space  on  one  side  of  a  suture. 

The  next  operation  is  the  final  tracing  from  the  cellophane  draw- 
ing. Due  to  slight  warping  of  the  cellophane  painted  with  India 
ink,  the  use  of  a  light-table  (table  with  glass  top  illuminated  from 
below)  is  recommended.  If  desirable  the  cellophane  copy  can  be 
enlarged  by  photography  and  the  final  tracing  of  the  suture  can  be 
made  from  the  photograph.  In  view  of  the  desirability  of  having 
the  drawings  two  times  larger  than  the  proposed  size  of  printed 
illustrations  the  enlargement  by  photography  is  ordinarily  preferable. 
For  the  tracing  of  very  small  and  complicated  sutures  the  method 
of  direct  tracing  on  cellophane  is  too  crude.  For  drafting  of  the 
sutures  of  juvenile  stages  of  Baculites  and,  also,  of  the  sutures  of 
smaller  coiled  scaphites,  the  writer  used  a  camera-lucida  attached 
to  a  microscope.  In  order  to  draw  an  undistorted  suture  on  a  plane, 
several  settings  of  the  specimen  must  be  made.  For  a  curved  surface 
of  about  180  degrees,  or  half  way  around  a  shell,  the  writer  made  six 
or  seven  settings.  After  a  part  of  a  suture  is  copied,  the  fossil  is 
turned  so  as  to  have  the  next  portion  of  the  suture  nearly  perpen- 
dicular to  the  axis  of  the  microscope.  In  order  not  to  depart  from  a 
constant  orientation  of  the  shell  one  must  revolve  it  around  the 
same  or  nearly  the  same  axis,  and  in  order  to  make  proper  con- 
nection of  two  neighboring  portions  of  a  suture  the  comparatively 
undistorted  edges  of  the  drawings  of  the  two  must  somewhat  overlap 
each  other.  The  smallest  magnification  of  ordinary  microscopes 
used  in  paleontological  laboratories  with  attached  camera-lucida  is 
about  10  times  and,  therefore,  the  drawn  suture  is  usually  too  large 
and  must  be  reduced.  This  can  be  done  by  photography  or  by  a 
pantograph. 

All  of  the  sutures  of  Baculites  shown  on  the  accompanying  plates 
were  drawn  by  the  direct  method  of  tracing  on  cellophane,  except 
Plate  XXXV,  figures  5d,  5e,  which  were  drawn  with  the  help  of 
camera-lucida  and  subsequently  reduced  by  pantograph.  All  the 
sutures  of  the  Scaphites  group  were  drawn  with  the  help  of  camera- 
lucida  and  reduced  by  photography. 


20—3482 


294  The  University  Science  Bulletin 

DESCRIPTION  OF  THE  SPECIES 
Class  CEPHALOPODA— Subclass  TETRABRANCHIATA 

Order  AMMONOIDEA— Suborder  EXTRASIPHONATA 

Family  LYTOCERATID^E 

Subfamily  MACROSCAPHITIN.E 

Genus  Baculites  Lamarck 

"Shell  with  minute,  closely  coiled  initial  stage  but  quickly  becoming  straight 
and  assuming  the  form  of  a  staff  which,  with  increasing  age,  increases  slowly 
in  diameter.  Cross  section  subtriangular,  oval,  or  subcircular,  though  usually 
more  or  less  compressed  laterally.  Living  chamber  large,  aperture  with  long, 
straight,  rounded  extension  on  the  siphonal  side  and  lateral  sinuses.  Surface 
smooth  or  with  low  rounded  ribs  that  are  parallel  to  the  aperture  and  as  a 
rule  are  distinct  only  on  the  flanks,  or  with  low  rounded  nodes  on  the  flanks. 
Suture  has  generally  six  saddles  and  six  lobes;  saddles  bifid;  lobes,  except  the 
antisiphonal  lobe,  bifid."  ^ 

Genotype.  Baculites  vertebralis  Lamarck  (Syst.  des  anim.,  p. 
103). 

The  remains  of  Baculites,  chiefly  in  the  form  of  internal  molds, 
are  perhaps  the  most  abundant  fossils  of  the  Pierre  shale  in  Wallace 
county  and  elsewhere.  The  common  smooth  specimens  from  this 
formation  are  ordinarily  referred  to  Baculites  ovatus  Say  and  B. 
compressus  Say,  which  were  instituted  on  material  from  the  Upper 
Cretaceous  of  New  Jersey  and  Upper  Missouri  river,  respectively. 
Unfortunately,  a  clear  conception  of  these  species  was  not  given  in 
the  original  descriptions,  and  considerable  confusion  as  to  which 
features  are  typical  for  each  species  was  thus  created.  In  addition, 
the  type  specimens  of  B.  compressus  are  lost.  For  the  clearing  up 
of  the  question  as  to  which  features  must  be  considered  typical  for 
each  of  the  two  species,  we  owe  much  to  F.  B.  Meek  and  J.  B. 
Reeside,  Jr.  The  latter  author  has  pointed  out  that  B.  compressus 
of  some  authors  (Morton,  Whitfield)  is  not  the  B.  compressus  of 
Meek,  but  is  a  slightly  compressed  variety  of  B.  ovatus.  He 
furthermore  has  suggested  acceptance  of  Meek's  clear  conception  of 
B.  compressus,  based  on  topotypes  of  the  species.  Reeside  proposes 
to  separate  the  forms  which  have  a  suture  closely  resembling  that  of 
B.  ovatus,  but  which  have  a  somewhat  compressed  cross  section  re- 
sembling that  of  B.  compressus,  as  Baculites  ovatus  var.  haresi 
Reeside. 

The  writer  collected  chiefly  from  the  middle  and  lower  part  of 

1.    After  Reeside,  1927,  p.  9. 


Elias:   Cephalopods  of  Pierre  Formation  295 

the  Pierre  and  from  the  topmost  beds  of  this  formation  and  found 
a  considerable  variety  of  forms  closely  related  to  B.  compressus  and 
B.  ovatus.    Among  these  are  several  specimens  with  the  ovate  cross 
section  of  the  B.  ovatus  type,  but  with  the  suture,  though  not  quite 
identical,  nevertheless  decidedly  of  the  B.  compressus  type.     He 
considers  it  preferable  to  give  this  form  the  new  specific  name  B. 
pseudovatus,  a  new  variety  of  which  {B.  pseudovatus  var.  A)  is  also 
provisionally  recognized  by  him.    The  writer  found  numerous  speci- 
mens which   are  identical  with   B.   compressus,   as   described   by 
Reeside,^  in  every  respect,  including  the  pronounced  and  regular 
corrugation  of  the  siphonal  edge.     At  the  same  time  the  writer 
collected  from  somewhat  different  horizons  of  the  Pierre  a  few  speci- 
mens indistinguishable  from  B.  compressus  as  described  by  Meek  and 
having  only  a  slightly  corrugated  siphonal  edge.  He  noticed,  further- 
more, that  the  sutures  of  the  B.  compressus  Meek  are  more  digitate 
than  those  of  B.  compressus  Reeside.     He  distinguishes  the  form 
described  by  Reeside  as  B.  compressus  var.  reesidei  and  recognizes 
in  his  material  from  Wallace  county  another  variety  with  still  more 
prominent  corrugation  of  the  siphonal  side,  which  he  designates  as 
B.  compressus  var.  corrugatus.    This  form  has  also  some  definite 
pecularities  of  the  suture  and  its  vertical  range  is  much  restricted. 
The  writer  is  aware  that  one  may  question  the  differentiation  of 
so  many  species  and  varieties  among  forms  which  until  recently 
were  referred  to  only  two  species:  B.  compressus  and  B.  ovatus. 
To  this  the  writer  answers  that,  besides  the  observed  differences  in 
form  and  suture,  the  definite  stratigraphic  position  of  these  forms 
in  the  local  Pierre  section  (as  shown  on  the  table,  p.  297)  justifies 
their  record  under  separate  names.    It  is  possible  that  some  of  these 
varieties  are  mere  responses  of  the  species  to  local  changes  in  the 
environments  of  the  Pierre  sea ;  but,  at  any  rate,  these  forms  appear 
to  be  of  local  stratigraphic  value  as  horizon  markers.    On  the  other 
hand,  some  of  the  forms  may  prove  to  have  wide  stratigraphic  im- 
portance and  may  help  considerably  in  the  distant  correlation  of 
subdivisions  of  the  Pierre   and   contemporaneous   formations,  not 
only  in  North  America  but  in  the  Upper  Cretaceous  formations  else- 
where.    It  is  significant  that  one  species  of  Baculites  from  the 
Beecher  Island  shale,  comparable  in  size,  cross  section,  smoothness 
of  the  shell  and  characters  of  the  suture  to  the  B.  ovatus  ^  and  B. 
compressus  in  the  broadest  sense,  was  found  to  be  still  more  similar 


2.  Reeside,  1927,  pp.  10-12. 

3.  In  fact  a  specimen  of  this   form  in  the  old   collection  in   the  Kansas  University   was 
identified  as  B.  ovatus  Say,  and  another  as  B.  anceps  Lamarck. 


296  The  University  Science  Bulletin 

and  possibly  identical  with  a  specimen  of  B.  anceps  var.  Leopoliensis 
described  by  Novak  from  the  Upper  Cretaceous  of  Poland.  The 
writer  feels  that  unless  the  forms  commonly  referred  to  B.  ovatus 
and  B.  compressus  are  studied  in  more  detail  and  are  separated  into 
species,  subspecies  and  varieties,  in  somewhat  the  same  way  as  has 
been  done  by  Reeside  for  the  Baculites  of  the  Eagle  Sandstone  and 
related  formations,^  these  widely  distributed  and  important  fossils 
will  be  of  very  little  help  as  horizon  markers,  but  will  be  merely 
characteristic  of  the  whole  Pierre.  Furthermore,  unless  the  various 
intermediate  forms  are  recognized,  the  types  themselves  can  be  hardly 
treated  as  distinct  species  on  account  of  the  proven  existence  of  a 
number  of  intermediate  forms  of  all  kinds ;  for  instance,  forms  with 
compressed  cross  section  and  moderately  digitate  suture  of  the  B. 
ovatus  type  and  forms  with  ovate  cross  section  and  intensely  digitate 
suture  of  the  B.  compressus  type.  The  following  is  a  key  for  identi- 
fication of  the  Baculites  of  the  Pierre  and  related  formations. 

For  convenience  of  reference  in  the  description  of  the  species  of 
Baculites  a  diagrammatic  sketch  of  a  typical  suture  of  the  genus  is 
appended  (PL  XL,  fig.  7).  The  suture  consists  of  siphonal  or 
ventral  lobe  (SL),  first  (lateral)  saddle  (IS),  first  lobe  (IL),  second 
saddle  (2S),  second  lobe  (2L),  third  (always  small)  saddle  (3S), 
and  antisiphonal  or  dorsal  lobe  (AL).  The  ventral  lobe  is  very 
complex  and  consists  of  two  lateral  tripartite  branches  which  are 
widely  separated  by  a  central  broad  saddle.  As  a  result  of  his  study 
of  material  from  Wallace  county,  with  the  addition  of  the  material 
illustrated  in  the  literature,  the  writer  concludes  that  the  form  and 
character  of  incision  of  this  central  part  of  the  ventral  lobe  is  of 
diagnostic  value  for  the  species  and  varieties  of  the  B.  ovatus  and 
B.  compressus  group.  Inasmuch  as  the  terminology  for  the  details 
of  this  part  of  the  suture  of  Baculites  is  not  worked  out,  he  proposes 
the  following  terms,  to  which  he  attaches  no  biological  or  ortho- 
genetic  significance,  but  which  are  introduced  for  descriptive  pur- 
poses alone.  The  whole  broad,  central  portion  of  the  ventral  lobe 
between  the  two  lateral  branches  is  designated  as  the  ventral  or 
siphonal  saddle  (a  term  used  already  by  some  authors).  This  saddle 
is  always  distinctly  subdivided  into  two  smaller  lateral  saddles  and 
a  median  saddle  (6)  between.  The  lateral  saddles  and  median 
saddle  are  separated  by  what  Meek  described  as  "smaller  digitate 
lateral  branches"  of  the  siphonal  lobe.  The  comparative  width  of 
the  siphonal  saddle,  the  relative  width  and  height  of  its  central 
branch  (b)  and  the  character  of  incision  of  the  latter  (uni,  bi-,  tri- 

i.    Reeside,  1927. 


KEY  FOR  THE  IDENTIFICATION  OF  THE  BACULITES  OF  THE  PIERRE  AND  RELATED  FORMATIONS 


Size,  (orm  and  sculpture. 

Character  of  Sutures. 

Sue  (Ions  dia, 
in  millimeters). 

Cross  section. 

Tapering 

of  lone 

length  oTshell 
in  millimeters). 

Sculpture. 

Degree  of 
dissection 

of  lobea. 

Stemlike  body 

branches 

of  first 

lateral  lobe. 

(See  pl.xl,  fig,  ?). 

Symmetrj;  and 

''rt'^sVom" 
lateral  lobe. 

Middle  branch  6  of  siphonal 
Binus  (see  pi.  XL,  Fig.  7). 

Of  flanks. 

Of  siphonal 
side. 

Height  in 

to  lateral 
branches. 

Dissection. 

B  ovalu»  Say 

Moderate,  to 
50  mm.  (pos- 
sibly larger). 

Elliptical  to 

Gradual.  1  to  2 

obscurely 
undulated. 

Smooth  to 
gently 
wrinkled. 

^•S,e. 

None 

Synunotrical, 

Even 

bipartite. 

Small, 
to  30  mm. 

Oblong  to 

Gradual, 

1  to  3  per  50. 

ob-curely 
undulated. 

Smooth  or 
gently 
wrinkled. 

Somewhat  more 
dissected  than 

None 

Symmetrical, 
slightly 

Tripartite 

B.  pneudotalus  EliaP.  n,  sp 

Moderate  to 

Elliptical     . 

Gradual, 

1  to  2  per  50 

Smooth.  , 

gently 
wrinkled. 

Moderately 
disaected. 

Aniiaiphonal 
side  reduced. 

Much  lower 

Bipartite. 

B.  ptnidovatut  var.  A.  EUrb,  n.  var 

Moderate, 
to  40  mm. 

Elliptical 

Gradual,  1  to 

Smooth 

Smooth 

Moderately 
dissected. 

Short 

Nearly 
symmetrical. 

Much  lower 

Unipartile  with 

tendency  to 
tripartite. 

Motlerate  to 
large,  to  50 
mm,  (poaeibly 

Oblong-ovate, 
subtrigonal 
in  Inter 
stages  (Meek). 

Moderate  to 
rapid  (par- 

the  young — 
Meek)  2  to  5 
per  50. 

large  shells 
broadly  un- 
dulated 
(Meek). 

Smooth  or 
gently 
wrinkled  (3 
to  4  wrinkles 
per  ID  mm. 

Much 
dissected. 

Long 

Symmetrical, 

Lower 

Bipartite 

B.  compre^niis  var.  reeaidei  Elias,  n.  var 

Moderate  to 

Elongately 

Moderate  to 

^"'"'''^ 

Wrinkled. 
2  wrinkles 

Moderately,..    , 
dissected. 

Long 

Antisiphonal 
side  reduced. 

Lower 

Tripartite. 

li.  comprcssus  var.  cormuatm  Eliaa,  n.  var.  . . 

Moderate  to 
lame,  to 

OblonE  to 
sublinear 

Moderate, 

^"'°°*'^ 

Prominently 
corrugated, 
1  wrinkle 

Moderately  to 
much  dis- 
sected. 

Long 

Symmetrical, 
inclined. 

Slightly 

Tri-tofive- 

B.  dinWot.a(.«  Eliaa.  n.  sp 

Moderate, 

Oblong  to 
elliptical. 

Moderate  to 
rapid,  3  to 

Smooth 

Smooth 

Moderately 
dissected. 

None 

Symmetrical, 
inclined. 

Lower 

Unipnrtite. 

B.  erandis  Hull  and  Meek 

Liirgc  to  very 
Inrae,  to 
100  mm. 

Ovate  tosub- 
cordiform. 

Gradual. 
1  per  50. 

Broadly  undu- 
lated, emoo'h 
in  early  stages 

Smooth 

Slightly 

dissected. 

None 

symmetrical. 

Lower 

Unipartitc  to 
irregularly 

B.  asp^r  Morion 

Small,  to 
30  mm. 

Elliptical 

Gradual, 
1  per  50. 

Round  nodes 
aiphonat  side. 

Numerous 
wrinkles, 

Moderately 
dissected. 

^'"'' 

Nearly 
symmetrical. 

Even  or 
slightly 

Unipartile. 

B.mrrA.-EUns.n.sp 

Small,  to 

Elliptical 

Moderate 

Oblique  ridges  ., 

Smooth,.      , 

Septit  unknown. 

Elias:   Cephalopods  of  Pierre  Formation  297 

or  quinquepartite)  seem  to  be  of  diagnostic  importance,  while  the 
shape  and  digitation  of  the  lateral  tripartite  branches  of  the  siphonal 
lobe,  between  which  the  siphonal  saddle  is  developed,  have  been 
found  fairly  constant  in  the  species  of  Baculites  here  described  and 
were  not  found  of  use  for  specific  distinction  of  the  species  and 
varieties  studied  by  the  writer. 

Among  the  other  features  of  the  suture  the  presence  or  absence  of 
a  stem-like  body  (a)  supporting  the  two  parallel,  slender  and  digi- 
tate terminal  branches  of  the  first  lateral  lobe  has  been  considered 
a  very  important  distinguishing  character  between  B.  compressus 
and  B.  ovatits  by  Meek  and  Reeside,  which  is  wholly  shared  by  the 
writer. 

Another  feature  of  the  suture,  which  is  used  for  specific  distinc- 
tion by  the  writer,  is  the  shape  of  the  second  lateral  lobe  and  its 
orientation  in  relation  to  the  rest  of  the  lobes  and  saddles  which  are 
always  fairly  parallel  to  each  other.  The  second  lobe  in  some  species 
or  varieties  is  distinctly  inclined,  with  its  apex  toward  the  siphonal 
side.  When  not  inclined  the  second  lobe  is  symmetrical  in  some 
species  while  in  others  it  is  much  reduced  on  the  antisiphonal  side, 
as  if  crowded  by  the  development  of  the  antisiphonal  saddle. 

Baculites  ovatus  Say 

(Plate  XXXIII,  figs.  3a,  b,  c) 

1820.    Baculites  ovata  Say,  Am.  Jour.  Sci.,  1  ser.,  vol.  2,  p.  41. 

1828.    B.  ovata  Morton,  Jour.  Acad.  Nat.  Sci.  Phila.,  1  ser.,  vol.  6,  p.  89,  pi.  5,  figs.  5,  6. 

1830.  B.  ovatus  Morton,  Am.  Jour.  Sci.,  1  sen,  vol.  17,  p.  280;  vol.  18,  p.  249,  pi.  1, 
figs.  6-8. 

1830.  B.  ovatxis  Morton,  Acad.  Nat.  Sci.  Phila.  Jour.,  1  ser.,  vol.  6,  p.  196,  pi.  8, 
figs.  6-8. 

1834.  B.  ovatus  Morton,  Synopsis  of  organic  remains  of  the  Cretaceous  group  in  U.  S. 
A.,  p.  42,  pi.  1,  figs.  6-8. 

1853.  B.  ovatus  Marcou,  Explanatory  te.xt  to  geologic  map  of  United  States  and  British 
provinces  of  North  America,  p.   46,  pi.  7,  fig.   5. 

1856.  B.  ovatus  Hall  and  Meek,  Am.  Acad.  Arts  and  Sci.,  Mem.,  new  ser.,  vol.  5,  p. 
399,  pi.  5,  figs,  la-c;    pi.   6,  figs.   1-7. 

1875.  B.  ovatus  White,  U.  S.  Geol.  Survey,  W.  100th  Mer.  Kept.,  vol.  4,  p.  199,  pi.  19, 
figs.  4a-5a-c  (not  figs.   4-b-c). 

1876.  B.  ovatus  Meek,  U.  S.  Geol.  Survey,  Terr.  Rept.,  vol.  9,  p.  394,  pi.  20,  figs,  la-b, 
2a-d. 

1889.  B.  ovatus  Whiteaves,  Contr.  Can.  Pal.,  vol.  1,  p.  181. 

1892.  B.  ovatus  Wliitfield,  U.  S.  Geol.  Survey,  Mon.  18,  p.  275,  pi.  46,  figs.  3-9. 

1907.  B.  ovatus  Weller,  New  Jersey  Geol.  Survey,  Pal.,  vol.  4,  p.  821,  pi.  109,  fig.  5. 

1910.  B.  ovatus  Grabau  and  Sihimer,  North  American  Index  Fossils,  p.  181,  figs.  1437, 
1438. 

1916.  B.  ovatus  Gardner,  Maryland  Geol.  Sun'ey,  Upper  Cretaceous,  pp.  375-377,  pi.  12, 
figs.  2,  3. 

1917.  B.  ovatus  Dowling,  Can.  Geol.  Surv.,  Mem.  95,  p.  31,  figs.  2,  2a. 

1926.  B.  ovatus  Wade,  U.  S.  Geol.  Survey,  Prof.  Paper  137,  p.  181,  pi.  60,  fig.  9. 

1927.  B.  ovatus  Reeside,  U.  S.  Geol.  Survey,  Prof.  Paper  151,  p.  9,  pi.  5,  figs.  12,  13; 
pi.  6,  figs.  1-4;   pi.  7,  figs.  1-8. 


208  The  University  Science  Bulletin 

The  species  may  be  recognized  by  its  ovate  to  elliptical  cross 
section;  smooth  or,  in  the  latest  stages,  obscurely  undulated  flanks; 
moderately  large  size;  relatively  simple,  little  incised  suture.  It 
differs  from  B.  compressus  Say  in  its  ovate  cross  section  and  much 
less  digitate  suture. 

This  species,  though  widespread  elsewhere,  has  not  been  found  as 
yet  in  Kansas,^  and  only  a  few  specimens  of  B.  ovatus  var.  haresi 
Reeside  were  collected  by  the  writer  in  Wallace  county.  According 
to  Reeside  B.  ovatus  is  found  in  the  Eagle  and  Telegraph  Creek 
formations  of  Montana  and  Wyoming,  in  the  lower  half  of  the 
Pierre  shale  of  the  western  Black  Hills  region  and  in  equivalent 
formations  of  the  midwestern  states ;  also  "in  the  later  parts  of  the 
Montana  group  up  to  the  base  of  the  Fox  Hills  sandstone,  and  in 
the  later  Cretaceous  formations  of  the  Atlantic  and  Gulf  Coastal 
Plain."  6  Meek  believed  that  the  species  "ranges  all  through  the 
Fort  Pierre  group,  and  up  into  the  Fox  Hills  beds  of  the  Upper 
Missouri  Cretaceous  series."  '^ 

The  writer  considers  it  quite  possible  that  with  separation  of  B. 
ovatus  in  the  restricted  sense  from  the  somewhat  similar  B.  pseudo- 
vatus  and  B.  clinolobatus,  the  stratigraphic  distribution  of  B.  ovatus 
will  be  reduced  and  will  possibly  be  limited  to  the  lowermost  portion 
of  the  Pierre,  to  the  Telegraph  Creek  and  Eagle  formations  and  to 
their  equivalents.  The  specimens  of  undoubted  B.  ovatus,  illustrated 
by  Reeside,  came  from  1,400  feet  below  the  top  of  the  Steele  shale 
of  Wyoming  (pi.  V,  figs.  12,  13)  and  from  the  Eagle  sandstone 
of  Montana  (pi.  VII,  figs.  3-5).^ 

Baculites  ovatus  var.  haresi  Reeside 

(Plate  XXXV,  figs.  3a,  b,  4a,  b) 

1927.  Baculites  ovatus  Say  var.  haresi  Reeside.  U.  8.  Geol.  Survey,  Prof.  Paper  151, 
p.  10,  pi.  6,  figs.  5-10;   pi.  7,  figs.  9,  10. 

According  to  Reeside  the  shell  is  smooth  with  dorsum  and  venter 
rounded,  and  it  has  a  relatively  simple  suture  of  the  B.  ovatus  type. 
It  differs  from  B.  ovatus  in  possessing  a  compressed  shell.  In  cross 
section  it  differs  from  B.  compressus  "in  lacking  the  tendency  to 
taper  toward  the  siphonal  side  and  in  the  details  of  the  suture" 


5.  B.  ovatus  was  recorded  in  the  Pierre  of  Kansas  by  Logan  (1897,  pp.  222,  230-231,  and 
1898,  p.  509),  but  inasmuch  as  this  record  was  made  at  a  time  when  the  clear  conception  of 
the  characteristics  of  the  species  was  not  established,  and  since  no  description  or  figures  of 
the  Kansas  form  is  given,  the  validity  of  the  record  may  be  questioned.  There  are  no  speci- 
mens of  B.  ovatus  in  the  old  collections  from  western  Kansas  at  the  University. 

6.  Reeside,  1927,  p.   10. 

7.  Meek,  1876,  p.  397. 

8.  Both  illustrated  by  Reeside,  1927. 


Elias:   Cephalopods  of  Pierre  Formation  299 

(Reeside,  1927,  p.  10).  The  specimens  from  Wallace  county,  which 
are  here  referred  to  the  variety  haresi,  have  the  shell  smooth  on  all 
sides  and  with  a  cross  section  of  the  haresi  type  (PL  XXXV,  fig.  4b). 
The  suture  of  Kansas  specimens  has  slightly  more  dissected  lateral 
sinuses,  but  the  lobes  are  designed  in  much  the  same  fashion  as  in 
haresi  from  Wyoming  (PI.  XXXV,  fig.  3a).  Though  the  dissected 
sinuses  of  Kansas  specimens  somewhat  resemble  the  corresponding 
parts  of  the  comjiressus  type  of  suture,  the  terminal  of  the  first 
lateral  lobe  is  decidedly  unlike  that  in  B.  compressus,  but  like  that 
in  B.  ovatus  and  its  variety  haresi.  The  second  lateral  lobe  is  some- 
what inclined  in  both  the  Wyoming  and  Kansas  specimens. 

Occurrence.  Upper  part  of  the  Upper  Weskan  shale  member  and 
base  of  the  Lake  Creek  shale  member  in  sec.  15,  T.  12  S.,  R.  38  W., 
Wallace  county,  Kansas. 

According  to  Reeside,  in  the  Telegraph  Creek  and  Eagle  sandstone 
formations  of  Montana  and  Wyoming;  in  the  lower  part  of  the 
Pierre  shale  on  the  western  and  northern  rim  of  the  Black  Hills;  in 
the  Steele  shale  of  Wyoming  and  the  upper  part  of  the  Mancos 
shale  in  Utah  and  New  Mexico.^ 

Baculites  compressus  Say 

1820.    Baculites  compressa  Say,  Am.  Jour.   Sci.,   1  ser.,  vol.   2,  pp.   41,   42. 

Not  1833.  Baculites  compressa  Morton,  Am.  Jour.  Sci.,  1  ser.,  vol.  23,  p.  291;  vol.  24, 
pi.  8,  fig.  1. 

Not  1834.  Baculites  compressus  Morton,  Synopsis  of  the  organic  remains  of  the  Cre- 
taceous group  in  the  United  States,  p.  43,  pi.  9,  fig.  1. 

Not  1892.  Baculites  compressus  Whitfield,  U.  S.  Geol.  Survey,  Men.  18,  p.  277,  pi.  46, 
figs.  1,  2. 

For  revision  of  the  conception  of  the  compressus  species  we  are 
indebted  to  J.  B.  Reeside,  Jr.,  whose  opinion  that  Morton  and  Whit- 
field described  under  the  name  compressus  a  variety  of  B.  ovatus  is 
wholly  shared  by  the  writer.  Since  the  type  specimen,  which  was 
inadequately  described  and  never  figured  by  Say,  is  apparently  lost, 
it  is  logical  to  accept  Meek's  clear  characterization  of  the  species, 
as  suggested  by  Reeside,  because  the  material  which  Meek  collected 
and  described  came  in  part  "from  the  region  where  Say's  specimen 
was  obtained."  Morton's  specimen,  which  was  erroneously  sup- 
posed to  be  the  type  of  Say's  species,  and  which  was  redescribed  and 
refigured  by  Whitfield,  has  septa  much  like  those  of  B.  ovatus  and 
can  be  justly  classified  with  B.  ovatus  var.  haresi  Reeside,  as  pro- 
posed by  Reeside.  For  a  complete  discussion  of  B.  compressus  of 
earlier  authors  see  Reeside,  1927  (pp.  10,  11). 


9.    Reeside.  1927,  p.  10. 


300  The  University  Science  Bulletin 

Though  the  writer  agrees  that  the  specimen  described  and  figured 
by  Reeside  and  referred  to  B.  compressus  is  similar  in  cross  section 
and  characters  of  the  sutures  with  B.  compressus  as  illustrated  and 
described  by  Meek,  there  are  some  differences  between  Meek's  and 
Reeside's  specimens  which  may  justify  the  separation  of  the  latter 
as  a  variety  of  B.  compressus  Meek.  This  conclusion  was  reached 
by  the  writer  after  comparative  study  of  numerous  specimens  of 
B.  compressus  from  Wallace  county,  among  wliich  both  the  speci- 
mens comparable  to  Meek's  and  to  Reeside's  forms  were  recognized. 
The  two  forms  were  found  in  somewhat  different  horizons  of  the 
Pierre  and  thus  proved  to  be  of  at  least  local  stratigraphic  value. 
The  writer  proposes  to  consider  the  form  described  by  Meek  as 
Baculites  compressus  s.  s.,  and  the  form  described  by  Reeside  as  the 
type  of  B.  compressus  var.  reesidei. 

Baculites  compressus  Say,  Meek,  s.  s. 

(Plate  XXVIII,  fig.  4  ;  Plate  XXXII,  figs.  3a,  b,  4a,  b,  c,  5a,  b) 

1854.  Baculites  compressus  Hall  and  Meek,  Am.  Acad.  Arts  and  Sci.,  Mem.,  new  ser., 
vol.  5,  pp.  400-402,  pi.  5,  figs.  2a,  b;   pi.  6,  figs.  8,  9. 

1876.  Baculites  compressus  Meek,  U.  S.  Geol.  Survey,  Terr.  Rept.,  vol.  9,  pp.  400-404, 
pi.  20,  figs.  30a-c;   text  figs.  55,  56. 

"Shell  attaining  a  large  size,  rather  rapidly  tapering,  particularly  in  the 
young,  or  near  the  smaller  extremity  of  adult  specimens,  strongly  compressed 
laterally  in  medium-sized  examples,  but  more  convex  in  the  young  and  toward 
the  larger  extremity  of  large  adults;  nonseptate  portion  of  fully  developed 
specimens  provided  with  large  broad  lateral  undulations;  lines  of  growth  gen- 
erally obscure;  siphonal  margin  sometimes  crossed  by  small  undefined  wrinkles; 
transverse  section,  like  the  outline  of  the  aperture,  varying  with  size  and  age, 
being  ovate  in  very  small  specimens,  strongly  compressed  in  medium  sized 
examples  and  proportionately  more  broadly  ovate  in  the  large  adult. 

"Septa  usually  crowded,  and  with  lobes  and  sinuses  deeply  divided  into 
slender  branches;  siphonal  lobe  nearly  twice  as  wide  as  long  and  provided 
with  two  widely  separated  tripartite  and  digitate  terminal  branches  and  one 
smaller  digitate  lateral  branch  on  each  side;  first  lateral  sinus  as  long  as  the 
siphonal  lobe,  but  not  much  more  than  half  as  wide  and  very  deeply  divided 
at  the  end  into  two  equal  tripartite  and  digitate  branches,  with  spreading 
subdivisions;  first  lateral  lobe  longer  than  the  siphonal  lobe,  and  about  half 
as  wide,  with  two  small,  parallel,  sharply  digitate  terminal  branches,  standing, 
as  it  were,  on  a  stem  formed  by  the  very  narrow  body  which  also  supports 
on  each  side  two  opposite,  sharply  digitate  lateral  branches,  one  pair  of  which 
shows  more  or  less  tendency  to  tripart  division;  second  lateral  sinus  usually 
a  little  longer,  but  otherwise  very  similar  to  the  first;  second  lateral  lobe 
shorter  and  broader  than  the  first,  and  provided  with  two  equal  tripartite  and 
digitate,  spreading  terminal  branches,  with  much  smaller,  irregular,  lateral 
branchlets;  third  lateral  sinus  usually  not  larger  than  one  of  the  main  ter- 
minal branches  of  the  others,  and  deeply  bifurcated  at  the  end,  the  divisions 


Elias:   Cephalopods  of  Pierre  Formation  301 

being  more  or  less  subdivided,  or  merely  digitate;  antisiphonal  lobe  generally 
only  about  half  as  long  as  the  second  lateral,  and  much  narrower,  with  two 
to  four  very  small  lateral  branches,  or  mere  digitations  on  each  side,  and  one 
small,  tridentate,  terminal  division."io 

In  his  material  from  AVallace  county  the  writer  has  observed  some 
specimens  which  are  identical  in  every  respect  with  the  specimens 
of  B.  compressus  as  illustrated  and  described  by  Meek,  and  which, 
in  some  minor  respects,  differ  from  the  more  numerous  specimens 
which  he  proposes  to  separate  under  the  name  B.  compressus  var. 
reesidei.  The  suture  of  B.  compressus  in  a  strict  sense  differs  from 
the  suture  of  reesidei  in  the  following  way : 

1.  The  median  saddle  of  the  siphonal  saddle  is  divided  into  two 
slightly  dentate  saddles  b-b  by  a  very  short  but  sharp  wedge-like 
siphonal  lobe,  while  the  corresponding  small  median  saddle  of  B. 
compressus  var.  reesidei  is  tripartite. 

2.  The  second  lobe  of  B.  compressus  s.  s.  is  wider  than  the  second 
saddle  and  about  as  symmetrical  as  the  latter,  while  the  second  lobe 
of  var.  reesidei  is  always  asymmetrical  with  reduced  branches  on  the 
antisiphonal  side. 

The  specimens  (PI.  XXVIII,  fig.  4,  and  PL  XXXII,  figs.  4a,  4b,  4c, 
5a,  5b)  of  the  collection  from  Wallace  county  can  be  referred  to  B. 
compressus  without  much  doubt.  Their  sutures  are  decidedly  of  the 
B.  compressus  s.  s.  type,  being  deeply  digitate  and  having  broad  and 
symmetrical  second  lobes.  The  small  median  saddle  {b)  of  the 
siphonal  lobe  is  bipartite.  The  cross  section  of  the  younger  of  the 
Wallace  county  specimens  (PI.  XXXII,  fig.  4c)  is  about  the  same 
size  and  is  as  much  compressed  as  Meek's  type  (PI.  XXXII,  fig.  3b) , 
though  there  is  a  slight,  but  hardly  important,  difference  in  the 
shape  of  the  cross  sections,  the  widest  part  of  the  Wallace  county 
specimen  being  about  in  the  middle  of  the  cross  section.  The  speci- 
men has  a  very  gentle  but  distinct  and  regular  plication  of  the 
siphonal  edge,  a  feature  in  agreement  with  Meek's  note  that  the 
siphonal  margin  is  "sometimes  crossed  by  small  undefined  wrinkles." 
The  regular  plication  of  the  siphonal  edge  of  our  specimens  is  cer- 
tainly very  gentle  and  wrinkles  are  quite  small,  the  distance  from 
one  depression  to  another  being  2  to  3  mm. 

The  largest  Wallace  county  specimen  is  a  fragment  incomplete  on 
the  siphonal  side  (PI.  XXXII,  fig.  5b).  The  antisiphonal  side  of 
this  specimen  is  abruptly  rounded,  much  like  the  cross  section  of 
the  medium-sized  B.  compressus,  as  shown  in  text  figure  55  on  page 
403  of  Meek's  monograph   (Meek,  1876).     Meek  observed  strong 

10.     Meek,   1876,  pp.   400,  401. 


302  The  University  Science  Bulletin 

lateral  undulations  on  the  nonseptate  part  of  a  large  specimen  of 
B.  compressus  (Meek,  1876,  text  figure  56,  p.  403),  but  the  non- 
septate  portion  of  large  B.  compressus  s.  s.  was  not  collected  in  the 
Pierre  of  Wallace  county. 

Occurrence.  Upper  part  of  the  Upper  Weskan  shale,  and  through 
the  Lake  Creek  shale  member  except  the  top  of  it  where  the  form 
becomes  scarce  or  is,  possibly,  absent.  According  to  Meek,  in  the 
PieiTe  exposed  at  the  Great  Bend  of  Missouri  river  below  Fort 
Pierre  and  from  "higher  positions  in  the  same  formation"  on  Sage 
creek,  at  the  Bad  Lands  and  on  Cheyenne  river  of  South  Dakota 
(Meek,  1876,  p.  403). 

The  specimens  described  by  Meek  were  collected  in  the  Upper 
Missouri  exposures  of  the  Pierre  formation,  where  also  the  type  of 
Say  was  found. 

Baculites  compressus  var.  reesidei  Elias,  n.  var. 

(Plate  XX^'III,  fig.  1;   Plate  XXXI,  fig.  3;   Plate  XXXII,  figs.  2a,  b,  c; 
Plate  XXXIII,  figs,  la,  b.  c;   2a,  b,  c) 

1927.  Baculites  compressus  Reeside,  U.  S.  Geol.  Survey,  Prof.  Paper  151,  pp.  10-12, 
pi.   9,  figs.  1-5. 

The  specimen  described  and  figured  by  Reeside  has  the  typical 
compressed  form  of  B.  compressus  and  a  much  digitate  suture  with 
the  characteristic  detail  of  the  compressus  type:  "Two  small, 
parallel,  sharply  digitate,  terminal  branches"  of  the  first  lateral  lobe 
"standing,  as  it  were,  on  a  stem  formed  by  the  very  narrow  body." 

The  differences  between  the  suture  of  var.  reesidei  and  that  of 
B.  compressus  s.  s.  are  as  follows: 

1.  The  suture  of  reesidei  is  all  in  all  comparatively  less 
digitate. 

2.  The  second  lateral  lobe  of  reesidei  is  comparatively  nar- 
rower than  the  corresponding  lobe  of  compressus  s.  s.,  and  besides 
is  distinctly  asymmetrical,  being  crowded  by  the  third  lateral 
saddle,  which  is  less  digitate  on  the  side  facing  the  second  lobe 
than  in  compressus  s.  s. 

3.  The  two  large  tripartite  terminal  branches  of  the  second 
lateral  saddle  are  standing  on  a  narrow  base  in  typical  B. 
compressus,  while  in  var.  reesidei  the  supporting  base  is  several 
times  wider. 

4.  The  small  median  saddle  (6)  of  reesidei  is  subdivided  into 
three  rounded  branches,  while  the  corresponding  portion  of  the 
siphonal  saddle  of  compressus  s.  s.  is  bipartite. 


Elias:   Cephalopods  of  Pierre  Formation  303 

Besides  the  difference  in  suture  the  var.  reesidei  is  differently 
ornamented  on  the  siphonal  edge  than  B.  compressus  s.  s.  The  edge 
of  the  type,  as  figured  by  Reeside,  is  pronouncedly  and  regularly 
plicated,  the  distance  between  plications  being  4  to  5  mm.,  or  about 
2  times  larger  than  between  the  far  less  pronounced  and  smaller 
wrinkles  of  B.  compressus  s.  s.  of  Wallace  county.  The  plications 
of  the  siphonal  edge  of  specimens  of  var.  reesidei  collected  by  the 
writer  have  a  distance  between  plications  of  4  to  5  mm.  on  the 
moderate-sized  types. 

Occurrence.  Lake  Creek  shale  member  and  lower  part  of  Salt 
Grass  shale  member  of  the  Pierre  shale,  Wallace  county,  Kansas. 
The  specimen  illustrated  by  Reeside  (1927,  PL  9,  figs.  1-5)  and 
reproduced  here  (PI.  XXXII,  fig.  2)  came  from  the  Eagle  sandstone 
of  Montana,  where  the  form  is,  however,  so  rare  that  Reeside  raises 
a  question  "whether  these  specimens  really  came  from  the  zone  con- 
taining the  Eagle  fauna,  and  it  must  be  left  for  future  experience  to 
determine  whether  B.  compressus  actually  does  belong  to  the  Eagle 
fauna  or  is  restricted  to  later  faunas."  ^^ 

Baculites  compressus  var.  corrugatu^  Elias,  n.  var. 

(Plate  XXVIII,  fig.  3;  Plate  XXX,  fig.  3;  Plate  XXXII,  figs,  la,  b,  c) 

Many  compressed  specimens  of  B.  compressus  with  a  still  more 
pronouncedly  corrugated  siphonal  side  than  in  var.  reesidei  have 
been  collected  in  the  upper  portion  of  Lake  Creek  shale  member. 
The  pronounced  wrinkles  of  the  siphonal  side  are  about  10  mm. 
apart.  The  typical  cross  section  of  this  variety  is  shown  on  Plate 
XXXII,  figure  Ic.  The  suture  is  decidedly  of  the  B.  compressus  type, 
being  deeply  dissected  and  having  terminal  branches  of  the  first 
lateral  lobe  standing  on  the  slender  stem-like  body  (a).  However, 
the  suture  of  this  variety  differs  from  that  of  B.  compressus  s.  s.  and 
of  var.  reesidei  in  having  an  inclined  second  lateral  lobe.  Further- 
more the  small  median  saddle  (6)  is  tripartite  with  a  further  splitting 
of  its  lateral  branchlets  into  two.  The  variety  corrugatus  differs 
from  both  typical  B.  compressus  and  var.  reesidei  in  having  a  very 
gradually  tapering  shell.  The  type  of  the  var.  corrugatus  (PL 
XXVIII,  fig.  3)  was  collected  and  presented  to  the  writer  by  Mr.  Joe 
De  Tilla,  of  Wallace. 

A  long  and  slender  specimen  of  Baculites  with  cross  section  and 
suture  of  the  compressus  (in  broad  sense)  type  was  collected  in  an 

H.  Reeside,  1927,  p.  12.  Reeside  is  now  convinced  that  the  species  "does  not  range 
so  low"  and  that  B.  compressus  is  wholly  upper  Pierre  form  (personal  notes  to  the  writer, 
April,  1931). 


304  The  University  Science  Bulletin 

irregular  body  of  ''Liicina  limestone"  at  the  base  of  the  Salt  Grass 
shale  member.  This  specimen  allows  study  of  the  change  of  suture 
with  growth  of  the  shell.  Though  the  initial  coiled  portion  of  the 
shell  is  not  preserved,  a  straight  portion  about  48  cm.  long  and  5 
to  31  mm.  wide  (in  the  long  diameter)  was  secured.  Several  stages 
of  the  suture  of  this  specimen  are  shown  on  Plate  XXXV,  figures 
5b,  5c,  5d,  5e.  The  comparison  of  these  sutures  with  those  of  B. 
conipressus  s.  s.  and  B.  comipressus  var.  reesidei  and  corrugatus  shows 
that  in  this  case  the  former  or  more  incised  {cojnpressus  s.  s.)  type  of 
suture  is  ancestral  to  the  less  incised  reesidei  and  corrugatus  types. 

It  is  interesting  to  recall  that  in  the  earliest  stages  of  the  straight- 
ened shell  of  B.  conipressus,  as  demonstrated  by  Brown  (1891,  p. 
159),  the  suture  develops  gradually  from  a  slightly  wavy  line  to  the 
complicatedly  curved  and  dissected  sutures  of  the  later  stages.  This 
is  also  a  generally  observed  fact  for  all  Ammonoidea,  which  ordinarily 
show  a  gradual  complication  and  increased  dissection  of  the  suture 
with  the  growth  of  the  shell.  Therefore,  the  subsequent  change  from 
a  more  complicated  suture  to  a  less  digitate  type,  as  now  demon- 
strated for  this  variety  of  B.  compressu^,  is  an  interesting  case  of 
simplification  of  suture  within  the  last  survivals  of  the  genus  Bacu- 
lites  of  the  North  American  Cretaceous  sea.  Compare,  also,  the 
return  to  a  less  dissected  type  of  suture  with  the  growth  of  the  shell 
in  B.  grandis  (p.  347  of  this  paper,  PL  XXXIV,  figs.  4  and  5a,  b) . 

The  evidence  of  a  later  appearance  of  the  reesidei  type  of  suture 
as  compared  with  the  typical  B.  compressv^s  suture,  agrees  with  field 
evidence  gathered  in  Wallace  county  that  reesidei  and  corrugatus 
appear  in  somewhat  higher  horizons  of  the  Pierre  formation  than 
typical  B.  compressus  s.  s. 

Occurrence.  Var.  corrugatus  is  perhaps  the  most  common  Bacu- 
lites  in  the  upper  part  of  the  Lake  Creek  shale  member  of  the  Pierre 
shale,  while  it  was  not  found  in  the  lower  half  of  the  member  or 
higher  than  at  the  base  of  the  Salt  Grass  shale  member  in  Wallace 
county. 

Bacidites  pseudovatus  Elias,  n.  sp. 

(Plate  XXIX,  figs,  la,  lb,  2 ;  Plate  XXXIII,  figs.  4a,  b) 

Among  the  specimens  of  Baculites  collected  from  the  Pierre  of 
Wallace  county  there  are  some  which  have  a  smooth  ventral  side 
and  a  cross  section  of  the  B.  ovatus  type,  but  the  suture  is  decidedly 
not  that  of  ovatus,  being  about  as  much  dissected  as  the  suture  of 
compressus.  The  writer  proposes  to  separate  these  forms  under  a 
new  specific  name,  pseudovatus.    The  forms  of  Baculites  which  have 


Elias:   Cephalopods  of  Pierre  Formation  305 

sutures  of  the  ovatiis  type  but  a  cross  section  approaching  that  of 
compressus  have  been  separated  already  by  Reeside  and  named  B. 
ovatus  var.  haresiP  This  author  states  that:  "An  examination  of 
a  large  number  of  specimens  from  various  horizons,  preserved  in  the 
collections  in  the  United  States  National  Museum,  showed  but  one 
specimen  combining  the  evenly  rounded  fonn  of  B.  ovatus  and  the 
suture  of  B.  compressus."  ^^  It  would  be  interesting  to  compare  the 
suture  line  of  this  specimen  with  that  of  B.  pseudovatus  from  Wal- 
lace county,  where  this  species  and  its  varieties  are  fairly  common. 

Description  of  the  pseudovatus  suture  line  is  as  follows:  Sutures 
moderately  closely  spaced;  lobes  and  saddles  not  as  deeply  sub- 
divided as  in  B.  compressus  s.  s.,  but  deeper  than  in  B.  compressus 
var.  reesidei;  siphonal  saddle  very  narrow,  being  only  one  and  a  half 
times  as  wide  as  long,  while  in  the  latter  two  forms  it  is  nearly  two 
times  as  wide  as  long.  The  tripartite  lateral  branches  of  the  siphonal 
lobe  are  cut  in  nearly  the  same  way  as  in  reesidei,  while  the  siphonal 
saddle  is  different  from  both  compressus  s.  s.  and  reesidei,  being  as 
wide  as  long,  or  two  times  narrower  than  in  the  latter  two;  the 
middle  incisions  of  the  first  and  second  saddles  are  sharply  pointed 
down  at  the  terminal,  while  in  the  latter  two  the  corresponding 
incisions  finger  out  at  the  terminals;  the  first  lateral  lobe  is  almost 
perfectly  round  in  the  outline,  while  the  corresponding  lobe  of  B. 
compressus  s.  s.  and  var.  reesidei  is  ovate  or  egg-shaped  with  the 
sharp  end  pointing  distad;  the  two  terminal  branches  of  this  lobe, 
which  in  B.  compressus  are  "standing,  as  it  were,  on  a  stem  formed 
by  the  very  narrow  body"  (a),  are  not  "parallel  and  sharply  digi- 
tate" as  in  B.  compressus  s.  s.  and  var.  reesidei,  but  bifurcate  each. 
The  third  saddle  and  the  antisiphonal  lobe  are  the  same  as  in  B. 
compressus  in  a  broad  sense. 

The  type  specimen  of  this  form,  showing  distinct  sutures,  was 
found  in  the  limestone  concretions  of  the  middle  zone  of  the  Upper 
Weskan  shale  member.  It  is  of  moderately  large  size,  having  a  long 
diameter  of  66  mm.  and  a  short  diameter  44  mm.  at  the  end  of  the 
septate  part  of  the  internal  cast. 

Numerous  individuals  of  smaller  size,  not  exceeding  40  mm.  in 
the  long  diameter  of  the  oval,  or  only  slightly  compressed  cross 
section  and  quite  smooth  on  the  outside,  are  very  common  in  the 
middle  part  of  the  Salt  Grass  shale  member.  Unfortunately  the 
sutures  of  these  numerous  molds,  when  preserved  at  all,  are  so  much 

12.  Reeside,  1927,  p.  10. 

13.  Ibid.,  p.  11. 


306  The  University  Science  Bulletin 

weathered  as  to  preclude  any  reliable  comparison.  However,  in  a 
very  few  cases  the  suture  was  sufficiently  preserved  to  allow  the 
writer  to  see  that  it  is  decidedly  not  of  the  ovatus  type,  but  is  much 
closer  to  that  of  B.  pseudovatus.  The  nearly  complete  suture  of 
the  best  specimen  is  illustrated  (PI.  XXXIII,  figs.  5a,  b).  Some 
differences  between  this  suture  and  that  of  the  typical  B.  pseudovatus 
(PL  XXXIII,  fig.  4a)  can  be  noticed  which  may  justify  the  separa- 
tion of  the  smaller  shell  as  a  distinct  variety  of  B.  pseudovatus.  This 
separation  has  support  in  the  fact  that  the  smaller  shell  was  found 
stratigraphically  higher  than  the  typical  B.  pseudovatus.  The 
smaller  form  is  provisionally  designated  here  as  B.  pseudovatus 
var.  A  (PL  XXVIII,  fig.  2,  and  PL  XXXIII,  figs.  5a,  5b). 

Occurrence.  The  type  specimen  of  B.  pseudovatus  was  found  in  the 
limestone  concretion  bed  (No.  8)  of  the  Upper  Weskan  shale  mem- 
ber. The  few  specimens  with  ovate  cross  section  but  without  any 
suture  line  which  were  collected  near  the  lower  part  of  Lake  Creek 
shale  member  most  probably  belong  to  this  species.  The  form  desig- 
nated here  as  B.  pseudovatus  var.  A  is  very  common  in  the  middle 
zone  of  the  Salt  Grass  shale  member,  where  other  fossils  are  rare. 

Intermediate  forms.  There  are  a  few  specimens  from  the  Pierre 
of  Wallace  county  which  may  be  considered  intermediate  forms  be- 
tween B.  pseudovatus  and  B.  compressus  var.  reesidei.  The  speci- 
men of  which  the  suture  and  cross  section  are  shown  on  Plate  XXXI, 
fig.  3,  and  Plate  XXXIII,  figs,  la,  lb,  is  close  to  B.  compressus  var. 
reesidei,  but  is  not  as  compressed  as  the  latter  and  has  only  very 
faint  wrinkles  on  the  siphonal  side.  This  specimen  came  from  the 
lower  half  of  Lake  Creek  shale  member. 

The  specimens  of  Plate  XXXV,  figs,  la,  lb  and  2a,  2b,  resemble 
B.  pseudovatus  var.  A,  but  are  comparatively  more  compressed.  The 
shell  of  both  is  perfectly  smooth  with  only  very  faint  wrinkles  on 
the  siphonal  side.  The  suture  is  of  intermediate  type  between  var. 
A  and  B.  compressus  var.  reesidei.  The  specimen  of  Plate  XXXV, 
figs,  la,  lb,  came  from  the  middle  zone  of  the  Salt  Grass  shale  mem- 
ber, and  the  specimen  of  Plate  XXXI,  figs.  2a,  2b,  came  from  the 
basal  zone  of  the  same  member. 


Elias:   Cephalopods  of  Pierre  Formation  307 

Baculites  grandis  Hall  and  Meek 

(Plate  XXXI,  figs,  la,  lb,  2a,  2b;   Plate  XXXIV,  figs,  i,  oa,  5b,  5c) 

1854.  Baculites  grandis  Hall  and  Meek,  Am.  Acad.  Arts  &  Sci.,  Mem.,  new  ser.,  vol.  5, 
p.  402,  pi.  7,  figs.  1,  2 ;   pi.  8,  figs.  1,  2 ;   pi.  6,  fig.  10. 

1876.    B.  grandis  Meek,  U.  S.  Geol.  Surv.  Terr.,  vol.  9,  p.   398,  pi.   33,  figs,  la-b-c. 
1926.    B.  grandis  Wade,  U.  S.  Geol.  Survey,  Prof.  Paper  137,  p.  182,  pi.  60,  figs.  8  and  12. 

This  species  is  recognized  chiefly  by  the  outstanding  large  diameter 
of  the  shell  (long  diameter  as  much  as  4  inches)  and  very  broad 
and  strongly  elevated  undulations  (Hall  and  Meek,  1854).  The 
suture  of  this  species  closely  resembles  that  of  B.  ovatus.  Hall  and 
Meek  describe  the  type  specimen  of  B.  grandis  as  follows  :^^ 

"Shell  elongate ;  section  varying  from  ovate  to  subcordiform ;  surface  of  cast 
marked  by  ^ery  broad  and  strongly  elevated  undulations,  which  commence  at 
the  dorsum  and  pass  obliquely  downward,  increasing  rapidly  in  size  and,  cross- 
ing the  sides  of  the  shell  in  a  broad  curve,  terminate  abruptly  on  the  ventro- 
lateral region.  Undulations  less  distinct  toward  the  smaller  extremity  and 
finally  become  obsolete.  Septa  very  deeply  lobed,  principal  divisions  scarcely 
divergent.  Dorsal  lobe  three-fourths  as  long  and  twice  as  wide  as  the  superior 
lateral  lobe;  terminated  on  each  side  by  a  narrow  elongated  branch,  which  is 
irregularl}^  sinuate  and  digitate  at  the  extremity.  Dorsal  saddle  shorter  and 
wider  than  the  superior  lateral  lobe,  formed  by  four  branches,  the  two  terminal 
ones  much  the  larger  and  each  of  them  bifid  at  the  extremity  by  a  small  sinus; 
the  whole  outline  more  or  less  sinuous  and  the  extremities  digitate.  Superior 
lateral  lobe  longer  by  one-fifth  than  the  inferior  lateral  lobe,  narrower  than 
the  ventral  saddle,  divided  at  its  extremity  bj^  a  deep  sinus  into  two  equal 
parts,  which  are  simply  digitate ;  above  these  are  two  unequal  branches  on  each 
side ;  terminal  sinus  much  deeper  than  the  lateral  ones.  Ventral  saddle  longer 
and  about  as  wide  as  the  dorsal  saddle,  more  deeply  divided  at  its  extremity 
by  the  auxiliary  lobe  into  two,  nearly  equal,  branches,  each  of  which  is  bifid 
and  the  extremities  digitate,  ventral  side  with  three  and  dorsal  side  with  two 
auxiliary  branches.  Inferior  lateral  lobe  shorter  and  broader  than  the  superior 
lateral  lobe,  divided  at  its  extremity  into  two,  nearly  equal,  branches,  the  one 
on  the  dorsal  side  bifid  at  the  tip  and  the  other  digitate,  with  an  auxiliary 
branch  on  the  ventral  side.  Ventral  lobe  as  long  as  the  auxiliary  lobe  of  the 
ventral  saddle,  but  wider  at  the  base,  digitate  at  its  extremity. 

"Angle  of  the  apex  about  5°.  Length,  as  deduced  from  the  measurement  of 
fragments,  by  the  convergence  of  the  dorsal  and  ventral  sides,  5^  feet  or  more. 
Longest  diameter  of  a  fragment  not  distorted  by  pressure,  3.7  inches;  shorter 
diameter  from  the  surface  of  undulations,  3.3  inches;  in  the  depressions  be- 
tween the  undulations,  2.95  inches. 

"The  species  is  nearly  related  to  B.  ovatus  of  Say,  from  which  it  differs  in  its 
much  greater  size,  larger  apical  angle,  much  stronger  and  more  extended  un- 
dulations, which  cross  the  entire  lateral  surface  of  the  shell.  The  section  is 
more  obtusely  ovate;  the  lobes  of  the  septa  are  much  deeper,  narrower,  and 
less  divergent  in  their  branches;  the  digitations  are  sharper  and  more  directly 
pointed  in  the  longitudinal  direction  of  the  shell.    The  auxiliary  lobe  of  the 

14.    Hall  and  Meek,  1854,  p.  402. 


308  The  University  Science  Bulletin 

ventral  saddle  is  longer  in  this  species,  while  the  extremities  of  the  terminal 
branches  are  less  deeply  bifid  than  in  B.  ovatus.  In  this  species  the  two 
terminal  branches  of  the  superior  lateral  lobe  are  simply  digitate,  while  in 
B.  ovatus  they  are  deeply  bifid,  with  obtuse  sinuosities.  Externally  in  its  un- 
dulations on  the  sides,  this  species  resembles  B.  anceps  of  Lamarck,  but  will  be 
readily  distinguished  by  the  absence  of  a  dorsal  carina  and  by  its  much  deeper 
lobes  with  less  divergent  divisions.  A  comparison  of  the  details  of  the  divisions 
of  the  lobes  and  saddles  shows  a  constant  difference  in  the  two  species." 

No  specimens  of  this  species  were  collected  in  Wallace  county, 
the  Pierre  shale  of  which  represents  horizons  of  the  formation  below 
those  in  which  this  species  has  been  found  elsewhere.  However,  the 
writer  collected  from  the  Pierre  shale  of  Beecher  Island,  Colorado, 
some  perfectly  preserved  internal  molds  of  B.  grandis,  the  longer 
diameter  of  which  varied  from  2  to  3%  inches.  The  molds  show 
strong  lateral  undulations  of  the  B.  grandis  type,  and  they  have  the 
sutures  figured  on  Plate  XXXIV,  fig.  4,  of  this  report. 

A  long  specimen  of  B.  grandis  showing  the  younger  stages  of  the 
shell  (but  not  the  very  earliest  stages  and  the  initial  coil)  was  col- 
lected in  the  Pierre  at  the  NWi/4,  sec.  2,  T.  2  S.,  R.  40  W.,  Cheyenne 
county,  Kansas.  The  changes  in  the  cross  section  of  the  shell  and 
in  the  suture  line  of  this  specimen  are  illustrated  on  Plate  XXXIV, 
figs.  5a,  5b,  5c.  The  most  evident  and  important  changes  are  the 
following: 

(1)  The  pronounced  lateral  undulations,  which  are  so  typical 
for  the  adult  shell,  do  not  exist  at  all  in  the  younger  stages, 
where  the  sides  of  the  shell  are  perfectly  smooth. 

(2)  Contrary  to  the  above  mentioned  change,  the  siphonal 
side  of  the  shell  is  smooth  in  the  adults,  but  in  the  young  stages 
there  appear  occasional  faint  to  moderately  strong  wrinkles 
spaced  about  4  mm.  apart. 

(8)  The  first  and  second  saddles  are  more  incised  in  younger 
stages  of  the  shell.  While  the  degree  of  dissection  of  these 
saddles  in  the  adult  sutures  of  the  shell  can  be  (and  always 
have  been)  compared  with  that  of  B.  ovatus,  the  degree  of  dis- 
section in  younger  stages  is  comparable  to  that  in  the  adult 
shells  of  B.  com'pressus  var.  reesidei  and  var.  corrugatus. 

(4)  The  central  incision  between  the  two  terminal  branches 
of  the  first  lateral  lobe  is  shorter  in  comparison  with  these 
branches  in  earlier  stages,  while  the  next  two  lateral  incisions 
of  the  terminal  have  a  tendency  to  come  closer  together  and 
build  the  slender  stem  (a)  supporting  the  terminal  branches, 
which  is  a  very  characteristic  feature  of  the  B.  compressus  type 
of  suture. 


Elias:   Cephalopods  of  Pierre  Formation  309 

(5)  In  the  younger  stages  the  second  lateral  lobe  is  inclined 
as  in  B.  clmolobatus  and  B.  compressus  var.  corrugatus. 

(6)  In  younger  stages  the  cross  section  is  elliptical  and  much 
compressed,  while  in  the  adult  shell  it  is  subtrigonal  to  oval. 

All  in  all  the  earlier  stages  of  the  shell  of  B.  grandis  resemble  in 
many  respects  the  shell  of  B.  compressus,  and  especially  B.  com- 
pressus var.  corrugatus.  B.  grandis  has  been  often  compared  with 
B.  ovatus  and  inasmuch  as  it  appears  later  in  the  Upper  Cretaceous 
than  the  latter  form,  one  is  tempted  to  surmise  that  ovatus  is  an 
ancestral  form  to  grandis.  However,  if  the  form  and  suture  of  the 
earlier  stages  of  B.  grandis  are  regarded  as  suggestive  of  the  general 
appearance  of  the  ancestral  form,  it  must  be  a  shell  of  the  B.  com- 
pressus type  from  which  B.  grandis  descended. 

Occurrence.  Several  localities  at  Black  Wolf  Creek,  Beecher 
Island,  Yuma  county,  Colorado,  chiefly  in  and  about  large  limestone 
concretions  at  the  base  of  the  Beecher  Island  shale.  From  the  same 
zone  in  Hackberry  creek  and  tributaries  to  Arikaree  river  in  north- 
western part  of  Cheyenne  county,  Kansas. 

B.  grandis  is  the  most  typical  baculite  of  the  topmost  beds  of  the 
Pierre,  but  originally  it  was  arbitrarily  referred  to  Fox  Hills  forma- 
tion by  Meek.^^  Later,  however,  a  belief  was  expressed  that  "neither 
Baculites  nor  Inoceramus  is  known  to  occur  in  the  Fox  Hills  sand- 
stone,"^^ and  this  is  now  generally  accepted.  Species  of  Baculites 
were  recorded  from  the  "Fox  Hills"  formation  at  Cooper  Creek  and 
at  Point  of  Rocks  of  Laramie  Plains,  Wyoming,^^  at  Bighorn  Moun- 
tains^^ and  in  eastern  Colorado. ^^  According  to  Reeside  none  of 
these  are  in  the  Fox  Hills,  but  rather  in  the  Upper  Pierre.  In  the 
Colorado  foothills  Fox  Hills  of  the  older  literature  is  mostly  Pierre.^" 
Recently  "some  of  the  concretions"  in  the  zone  about  1,120  feet 
above  the  base  of  the  "Fox  Hills  sandstone"  of  Fossil  Creek  were 
noticed  to  be  "crowded  with  Baculite s."'^'^  If  the  latter  identification 
is  correct,  the  survival  of  the  genus  Baculites  high  into  what  is 
usually  considered  by  field  geologists  as  Fox  Hills  formation  must 
be  considered  proved  unless  the  lithological  criterion  which  is  usually 
employed  to  distinguish  Fox  Hills  sandstone  from  the  Pierre  shale 

15.  Meek,  1876,  pp.  338-340. 

16.  Stanton,  1925,  p.  8. 

17.  Stanton.  1897,  p.   141  and  p.   147. 

18.  Stanton  in  Darton,   190.5,  p.   52. 

19.  Darton,  1905,  p.  109. 

20.  Personal  note  by  Reeside,  April,  1931. 

21.  Mather,  Gilluly  and  Lusk,  1928,  p.  96. 

21— 3IS2 


310  The  University  Science  Bulletin 

is  not  reliable  at  all  and  some  other  means  of  recognition  of  the  Fox 
Hills  sediments  must  be  adopted. 

The  simultaneous  disappearance  in  the  Upper  Cretaceous  seas  of 
North  America  of  Baculites  and  Inoceramus  can  be  arbitrarily 
chosen  as  the  end  of  the  Pierre  time,  but  this  feature  alone  is  only 
a  negative  characteristic  for  Fox  Hills  time.  According  to  Reeside,^^ 
the  genus  Sphenodiscus  is  restricted  to  the  Fox  Hills  and  is  our  best 
marker  for  the  formation. 

Baculites  dinolohatus  Elias,  n.  sp. 

(Plate  XXX,   figs.   1,   2;    Plate  XXXIV,   figs.   1,   2a,  2b,   3) 

1908.  Baculites  anr.eps  var.  leopoliensis  Novak,  Bull.  Acad.  Sci.,  Cracovie,  1908,  p.  326, 
text  fig.,  p.  331,  pi.  14,  fig.  10. 

Among  the  material  collected  from  the  Pierre  shale  at  Beecher 
Island,  Colorado,  there  are  several  perfectly  preserved  specimens 
with  smooth  surface,  moderately  compressed  oval  cross  section,  and 
a  very  interesting  suture  which  is  different  from  the  rest  of  the 
American  baculites  known  to  the  writer.  The  suture  line  is  cer- 
tainly more  dissected  than  those  of  B.  ovatus  and  B.  grandis  and  is 
about  as  digitate  as  that  of  B.  compressus  var.  reesidei.  There  are, 
however,  some  important  differences  which  do  not  permit  the  classi- 
fication of  the  Beecher  Island  species  with  the  latter  variety  of  B. 
compressiis  (or  with  the  typical  B.  compressus) .  Besides  the  smooth- 
ness of  the  siphonal  side  of  the  shell  and  the  only  slightly  com- 
pressed oval  cross  section,  the  design  of  the  suture  is  considerably 
different.  The  siphonal  lobe  is  as  wide  as  long.  The  first  lateral 
lobe  is  as  oval  in  outline  as  that  of  B.  pseudovatus  var.  A,  but  differs 
considerably  from  both  B.  pseudovatus  and  B.  compressus  by  having 
the  terminal  branches  not  standing  "on  a  stem  formed  by  the  very 
narrow  body,"  as  in  all  these  species.  They  are  here  much  separated 
by  a  prominent  and  long  incision,  which  cuts  the  lobe  deeply  into  two 
large  bipartite  terminal  branchlets.  The  second  lobe  is  not  parallel  to 
the  rest  of  the  lobes  and  saddles,  but  is  considerably  inclined  to 
the  axis  of  the  shell.  At  the  same  time  the  inclined  lobe  is  sym- 
metrical in  itself,  having  the  branches  of  each  side  of  equal  size  and 
sequence. 

It  is  interesting  to  note  that  the  suture  of  B.  dinolohatus  is  more 
similar  to  one  of  the  European  types  than  to  any  one  of  the  American 
species  so  far  described.  On  Plate  XXXIV,  fig.  3,  the  suture  line 
of  Baculites  anceps  var.  leopoliensis  Novak  is  shown  for  comparison 

22.    Unpublished   stratigraphic  chart  of   Upper  Cretaceous,   1930. 


Elias:   Cephalopods  of  Pierre  Formation  311 

with  the  Beecher  Island  species.  The  striking  similarity  of  the 
following  features  in  the  suture  line  of  the  American  and  European 
species  may  be  noted:  (1)  General  character  and  degree  of  dis- 
section; (2)  vertically  elongated  terminal  branches  of  both  the 
first  and  second  lateral  saddle;  (3)  perfectly  oval  outline  of  the 
first  lateral  lobes,  which  besides  are  dissected  in  the  very  same 
fashion;  (4)  inclination  of  the  second  lobes,  the  dissection  of  which 
is  again  very  similar;  (5)  small,  bipartite  and  not  digitate  median 
saddle  (6)  of  the  siphonal  saddle. 

Taking  into  account  the  character  and  the  degree  of  variability 
of  the  sutures  within  the  same  individual,  as  exhibited  in  our  speci- 
men from  Beecher  Island  (compare  the  two  neighboring  sutures  of 
Plate  XXXIV,  figs.  1  and  2a),  one  comes  to  the  conclusion  that 
there  is  not  the  slightest  difference  between  the  sutures  of  the  Ameri- 
can and  the  European  forms.  If  we  add  to  this  the  similarities  of 
the  size,  outline  and  smoothness  of  the  septate  portions  of  the  shells, 
we  have  to  confess  that  the  two  are  as  identical  as  two  individuals 
of  the  same  species  can  ever  be  expected  to  be,  and  therefore  they 
should  be  classified  under  the  same  specific  name.  The  writer,  how- 
ever, hesitates  to  apply  to  the  American  specimen  the  name  Baculites 
anceps  var.  leopoliensis,  because  the  other  sutures  of  the  same  va- 
rieties, as  shown  on  figures  5  to  9,  page  331,  of  Novak's  paper ,^3  are 
quite  different  from  that  of  figure  10,  page  331,  of  the  same  paper, 
which  is  compared  here  with  the  suture  of  the  Beecher  Island  form. 
These  other  sutures  of  the  var.  leopoliensis  Novak  are  much  like 
the  sutures  of  Baculites  grandis  or  B.  ovatus,  and  one  may  question 
the  identity  of  these  with  the  specimen  the  suture  of  which  is  shown 
on  figure  10,  page  331,  of  Novak's  paper,  unless  extreme  variability 
of  the  septa  of  var.  leopoliensis  is  admitted.  The  much  confused 
situation  in  regard  to  the  conception  of  the  European  species  anceps 
has  been  pointed  out  by  Meek,  who  judged  "from  published  figures 
and  descriptions  .  .  .  that  shells  belonging  to  more  than  one 
species  have  been  included  under  Lamarck's  name."^*  The  situation 
concerning  the  limits  of  anceps  has  not  improved  appreciably  since 
Meek's  time,  and  recently  again  it  has  been  pointed  out  that  there  is 
"a  considerable  variation  in  the  concept  of  the  species  held  by  Euro- 
pean students. "2^ 

It  is  advisable,  therefore,  to  give  the  American  forms,  which  are 


23.  Novak,   1908. 

24.  Meek,   1876,  p.   407. 

25.  Reeside,   1927,  p.   12. 


312  The  University  Science  Bulletin 

related  and  in  part  apparently  identical  with  some  specimens  referred 
to  B.  anceps  by  European  authors,  separate  specific  names,  as  was 
done  in  a  similar  case  by  Reeside.^*^  At  the  same  time  the  relation 
and  even  identity  of  the  American  foniis  with  this  or  that  particular 
European  form  described  under  the  name  B.  anceps  must  be  pointed 
out. 

Distribution.  Topmost  Pierre  or  transitional  to  Fox  Hills  beds  of 
shale  exposed  at  Black  Wolf  Creek,  near  Beecher  Island,  Yuma 
county,  Colorado.  Also  probably  from  the  same  zone  of  the  Pierre 
in  Cheyenne  county,  Kansas.  The  perfect  molds  of  Baculites  from 
this  county  in  the  old  collection  of  the  Geology  Department  of  the 
University  of  Kansas  are  labeled  B.  anceps.  They  are  indistinguish- 
able from  B.  clinolobatits  of  Beecher  Island. 

Baculites  meeki  Elias,  n.  sp. 

(Plate  XXX,  figs.  4a,  4b) 

1876.  Baculites  asperl  Meek,  Rept.  Invert.  Cret.  and  Tert.  Fossils,  U.  S.  Geol.  Survey, 
Terr.,   vol.   9,  p.   404,  pi.  39,  figs.   10b,   10c  (not  figs.   10a,   lOd). 

Meek  described  two  small  fragments  of  internal  molds  which  he 
doubtfully  referred  to  B.  asper  and  thought  to  have  the  same  strati- 
graphic  position  "near  the  top  of  the  Fox  Hills  group  of  the  Upper 
Missouri  Cretaceous  series. "^^  Recently  Reeside,  who  described  a 
much  better  B.  asper  from  Montana  and  other  mid  western  states, 
correctly  admitted  to  this  species  only  that  specimen  of  Meek  which 
was  illustrated  on  Plate  39,  figures  10a,  d.-^  That  the  two  frag- 
ments belong  probably  to  two  different  species  was  realized  by  Meek 
himself.  Speaking  of  the  fragment  shown  in  figures  10b  and  c,  he 
says,  "I  can  scarcely  doubt  now  that  it  represents  a  distinct  species, 
its  strong,  oblique,  nearly  straight  ridges  or  undulations  extending 
entirely  across  the  sides  being  a  very  marked  feature,  contrasting 
strongly  with  the  nodelike  prominence  on  the  other  specimen  rep- 
resented by  figures  10a,  d."  ^^  A  fragment  of  the  same  size,  cross 
section  and  sculpture  (PI.  XXX,  figs.  4a,  4b)  was  collected  by  the 
writer  in  the  Pierre  shale  at  Beecher  Island.  This  checks  to  some 
extent  the  stratigraphic  position  of  this  rare  and  interesting  form 
high  in  the  section  of  the  Upper  Cretaceous  and  makes  it  worthy  of 
designation  by  a  separate  name. 

26.  Ibid.,   p.    12,  Baculites  aquilcensis  and   its   varieties. 

27.  Meek,  1876,  p.  405. 

28.  Reeside,  1927,  p.   13. 

29.  Meek,  1876,  p.  405. 


Elias:   Cephalopods  of  Pierre  Formation  313 

The  names  asperoides  or  aspeiiformis,  proposed  by  Meek  (1876, 
p.  405),  were  intended  for  the  specimen  10a,  d  in  case  it  was  proved 
not  to  belong  to  B.  asper,  and,  therefore,  it  is  preferable  not  to  use 
these  for  Meek's  specimen,  figures  10b,  c.  Therefore,  the  new  name 
meeki  is  proposed  for  the  latter  and  for  the  specimen  from  Beecher 
Island  with  which  it  is  identical.  The  suture  of  B.  meeki  remains 
unknown. 

Meek's  specimen  10a,  d,  which  is  a  true  B.  asper,  possibly  did  not 
come  from  the  Fox  Hills  formation  at  all,  as,  according  to  the  new 
data,  the  species  appears  to  be  restricted  to  the  lower  part  of  the 
Pierre  and  to  related  formations. 

Occurrence.  In  Beecher  Island  shale  member,  Yuma  county,  Colo- 
rado. According  to  Meek,  near  the  top  of  the  Fox  Hills  in  South 
Dakota. 

Subfamily  Turrilitin^ 

Genus  Heteroceras  d'Orbigny 

Heteroceras  tortum  Meek  and  Hayden 

(Plate  XLII,  fig.   2) 

1858.  Helicoperas  tortum.  Meek  and  Hayden,  Proc.  Acad.  Nat.  Sci.  Phila.,  vol.  10,  p.  54. 

1864.  Heteroceras  tortum.  Meek,  Smithsonian  Check-List  N.  Am.  Cret.  Fossils,  p.  2.5. 

1876.  Heteroceras  tortum.  Meek,  U.  S.  Geol.  Survey  Terr.  Kept.,  vol.  9,  p.  481,  pi.  22,  figs. 
4a,  b,  c. 

The  writer  has  little  doubt  that  a  fragmentary  specimen  collected 
from  Wallace  county  belongs  to  this  species  of  Meek  and  Hayden. 
The  size  of  the  shell,  the  number  and  prominence  of  transverse  ribs, 
the  presence  of  one  or  possibly  of  two  rows  of  not  prominent  nodes 
and  the  wide  umbilicus  are  much  like  the  features  of  the  type 
described  and  figured  by  Meek.^°  The  nodes  are  spaced  as  widely  as 
in  Meek's  type,  every  second  or  every  third  rib  provided  with  a 
node.  Only  very  incomplete  sutures  can  be  observed  in  some  por- 
tions of  the  Wallace  county  specimen. 

Distribution.  Meek  refers  this  type  to  the  "lower  part  of  the 
Fort  Pierre  group."  ^^  The  Wallace  county  specimen  came  also  from 
the  basal  portion  of  the  Pierre  as  there  represented ;  from  the  top  of 
the  Sharon  Springs  shale  member,  where  it  was  found  in  a  limestone 
lens.  According  to  Recside  "this  type  of  shell  does  not  occur  in 
lower  Pierre."  ^^ 

30.  Meek,  1876,  p.  481,  pi.  22,  figs.  4a,  b,  c. 

31.  Meek,  1876,  p.   482. 

32.  Personal  note  to  the  writer  of  April,  1931. 


314  The  University  Science  Bulletin 

Family  COSMOCERATID.E 

Subfamily  Scaphitin^ 

Genus  Scaphites  Parkinson 

Scaphites  plenus  Meek 

(Plate  XXXVI,  figs,  la,  lb,  Ic,  2a,  2b,  2c;   Plate  XXXVII,  figs,  la,  lb,  Ic ;   Plate  XXXIX, 
figs,  la,  lb,  Ic;  Plate  XL,  figs.  3,  4,  5,  6) 

1860.  Scaphites  nodosus  var.  plenus.  Meek  and  Haydcn,  Proc.  Acad.  Nat.  Sci.  Phila.,  p. 
420. 

1876.  Scaphites  nodosus  var.  plenus  Meek,  U.  S.  Geol.  Survey  Terr.  Rept.,  vol.  9,  p.  429, 
pi.  26,  figs,  la,  b,  c. 

1885.  Scaphite.i  fsuhglohosus  Whiteaves,  pars.,  Geol.  Siin-ey  Canada,  Contr.  Can.  Paleon- 
tology, vol.   1,  p.   52,  pi.   8,  figs.  2,  2a.      (Not  pi.   7,  fig.   3;    pi.  8,  figs.   1,   la.) 

1905.  Scaphites  nodosus  var.  plenus.  Smith  (W.  D.),  Jour.  Geol.,  vol.  13,  pp.  638,  644, 
647-648;  fig.  1  (p.  641)  and  figs  3-7,  8,  10  (p.  645). 

1917.  Scaphites  subglobosus  Dowling,  pars.,  Geol.  Survey  Canada,  Mem.  93,  p.  32,  pi.  31, 
figs.  2,  2a.     (Not  pi.  31,  figs.  1,  la.     Reprints  of  Whiteaves'  drawings.) 

1927.  Acanthoscaphites  nodosus  var.  plenus,  Reeside,  U.  S.  Geol.  Survey  Prof.  Paper  150- 
B,  p.  32. 

The  following  is  the  complete  quotation  of  Meek's  description  of 
this  interesting  form :  ^^ 

"This  form  differs  from  the  typical  Scaphites  nodosus,  not  only  in  its 
proportionally  shorter  deflected  body-part,  but  also  in  its  much  more  gibbous 
form,  and  in  the  smaller  size  of  its  inner  rows  of  lateral  nodes,  which  are  placed 
nearer  the  umbilical  side.  It  also  sometimes  shows  a  slight  tendency  to  de- 
velop a  third  intermediate  series  of  very  small  lateral  nodes  about  midway 
between  the  other  rows,  such  as  I  have  not  seen  in  any  of  the  other  varieties. 

"This  tendency,  however,  is  only  manifested  by  a  scarcely  perceptible  swell- 
ing of  the  costae  at  this  point,  and  consequently  escaped  the  attention  of  the 
artist  in  drawing  the  figures;  while  even  this  faint  tendency  is  not  constant. 

"Compared  with  the  foregoing  forms  regarded  as  varieties  of  S.  nodosus, 
this  shell  also  presents  a  remarkable  contrast,  both  in  form  and- size.  So  great, 
indeed,  is  the  difference  that  it  seems  difficult  to  believe  that  it  can  be  properly 
regarded  as  even  a  marked  variety  of  the  same  species  as  that  represented  by 
our  figures  2,  a,  b,  of  plate  25;  and,  although  I  continue  here  to  range  it  pro- 
visionally as  a  variety  of  s.  nodosiis,  I  am  not  altogether  free  from  doubts  on 
this  point.  Still,  when  we  come  to  compare  it  with  Owen's  figure  of  the  typical 
s.  nodosus,  and  with  the  intermediate  forms  represented  on  our  plate  25,  little 
is  found  to  distinguish  it  from  the  same,  excepting  its  very  ventricose  form, 
a  character  not  alone  generally  reliable  for  distinguishing  species  in  this  and 
allied  groups  of  shells.  Its  septa  will  be  observed  to  agree  almost  exactly, 
excepting  in  mere  minute  individual  details,  with  those  of  the  variety  brevis, 
figured  on  plate  25,  and  to  present  the  same  differences  in  the  smaller  lobes 
from  the  variety  quadrangularis,  that  the  variety  brevis  does.  At  one  time  I 
was  rather  inclined  to  think  that  this  great  difference  in  the  convexity  of  these 
shells  might  be  merely  sexual;  but  the  fact  that  the  variety  or  species  plenus 
was  only  found  at  a  locality  on  Yellowstone  river  (where  it  occurs  associated 

33.    Meek,  1876,  pp.  429,  430. 


Elias:   Cephalopods  of  Pierre  Formation  315 

with  some  of  the  more  compressed  forms),  while  no  specimens  of  it  were 
found  at  numerous  other  localities  where  the  latter  forms  are  common,  does 
not  seem  to  sustain  this  view." 

The  specimen  from  Wallace  county  illustrated  here  (PI.  XXXVI, 
figs.  2a,  2b,  2c)  has  been  (and  still  is)  compared  by  the  writer  with 
the  smaller  specimens  of  s.  subglobosus  Whiteaves.  At  the  same 
time  he  came  to  a  conclusion  that  the  larger  specimen  of  Whiteaves' 
should  not  be  classified  with  the  smaller  ones  and  that  Whiteaves 
was  in  error  in  doing  so.  Inasmuch  as  the  species  established  by 
Whiteaves  was  compared  by  Stanton  =^^  with  S.  mullananus  Meek 
and  Hayden,  the  writer  felt  justified  in  restricting  the  name  sub- 
globosus to  the  smaller  specimens  (PI.  8,  figs.  2,  2a)  of  Whiteaves, 
which  certainly  do  not  resemble  mullananus.  Consequently,  he 
felt  justified  to  identify  the  specimen  from  Wallace  county  with  the 
smaller  specimens  of  S.  subglobosus.  However,  in  discussing  the 
matter  with  J.  B.  Reeside,  the  latter  has  advised  the  writer  that  he 
"examined  the  younger  stages  of  nodosus  var.  plenus  and  found 
them  to  duplicate  Whiteaves'  figures  of  the  supposed  young  of  sub- 
globosus." To  this  he  added  his  opinion,  which  confirms  the  writer's 
view,  that  "Whiteaves  was  in  error  in  supposing  his  small  specimens 
to  be  the  young  of  his  larger  specimens."^^  Subsequently  Reeside 
has  very  obligingly  supplied  the  writer  with  a  squeeze  of  the  inner 
whorl  of  a  large  specimen  that  agrees  exactly  with  subglobosus,  a 
cast  of  the  holotype  of  S.  nodosus  var.  plenus  M.  and  H.  and  two 
younger  individuals  of  the  latter  form  from  South  Dakota.  The 
examination  of  this  material  convinces  the  writer  of  the  correctness 
of  Reeside's  view  that  both  the  smaller  specimens  of  Whiteaves  and 
the  specimen  from  Wallace  county  represent  younger  individuals  of 
S.  plenus.  Consequently  the  name  subglobosus  must  be  restricted 
to  the  large  individual  of  Whiteaves'  species  only  (Whiteaves'  PI.  7, 
fig.  3,  and  PI.  8,  figs.  1,  la),  and  not  to  one  small  individual  de- 
scribed by  W-hiteaves  under  one  name,  subglobosus. 

Inasmuch  as  Meek  illustrated  only  one  large  adult  specimen  of 
plenus,  in  which  the  inner  volutions  are  hidden  inside  of  the  outer 
whorl,  it  is  appropriate  to  illustrate  and  describe  here  the  younger 
stages  of  the  shell,  which  seem  to  be  found  more  often  in  the  Pierre 
shale  than  the  complete  large  specimens  with  their  living  chamber 
preserved. 

One  of  the  most  important  results  of  the  study  of  these  earlier 
stages  of  plenus  is  the  disclosed  fact  that  the  shell  is  wider  than 

34.  Stanton,  1893,  p.  189. 

35.  Letter  to  the  writer,  Novomber  13,  1930. 


316  The  University  Science  Bulletin 

high  in  all  stages  of  growth.  In  this  respect  plenus  differs  con- 
siderably from  all  varieties  of  Acmithoscaphites  nodosus  (including 
the  typical  form),  the  shell  of  which  is  always  higher  than  wide. 
This  difference  alone  justifies  the  removal  of  plenus  from  Acan- 
thoscaphites  nodosus,  a  variety  of  which,  though  not  without  hesita- 
tion, it  was  considered  by  Meek. 

The  Wallace  county  specimen  (PL  XXXVI,  figs.  2a,  2b,  2c)   is 
round  and  much  inflated;  the  maximum  width  is  20  mm.;  the  larger 
diameter  is  35  mm.    Volutions  are  broadly  rounded  on  the  periphery 
and  on  the  lateral  side,  but  are  much  more  narrowly  convex  on  the 
inner  or  umbilical  side.     The  preserved  nonseptate  portion  of  the 
shell  equals  about  one-third  of  a  volution.     Ribs  are  prominent, 
slightly  curved  toward  aperture  on  the  sides  of  shell  and  nearly  or 
quite  straight  on  the  periphery.    The  ribs  increase  in  number  from 
the  umbilicus  to  the  periphery  by  intercalation  of  additional  ones, 
two  extra  ribs  being  usually  well  developed  at  the  row  of  peripheral 
nodes.    On  the  rounded  ventral  portion  of  the  shell,  between  the  two 
rows  of  the  peripheral  nodes,  an  extra  rib  is  added  at  each  node, 
which  brings  the  total  number  of  ribs  on  the  venter  to  about  34  on 
the  half  of  the  outer  volution,  as  against  9  at  the  corresponding 
part  of  the  shell  of  the  umbilicus.    The  nodes  occur  in  one  row  only 
on  each  side  and  are  moderately  prominent,  every  third,  or  rarely 
every  second,  of  the  lateral  ribs  being  provided  with  a  node.    Inner 
volutions  are  also  provided  with  the  peripheral  rows  of  nodes  which 
begin  to  appear  when  the  radius  of  volutions  reaches  about  5  mm. 
The  smallest  specimens  described  as  *S.  subglobosus  by  Whiteaves^^ 
resemble  in  every  respect  the  above-described  specimen  from  Wal- 
lace county.    The  ribs  of  Whiteaves'  specimen  are  as  much  curved 
and  nearly  as  densely  spaced  as  on  the  Kansas  specimen.    However, 
in  the  latter  shell  there  are  three  to  four  times  as  many  ribs  at  the 
periphery  as  at  the  umbilicus,  whereas  Whiteaves  observed  ''twice 
or  perhaps  three  times  as  many  on  the  center  of  the  periphery  as 
on  the  umbilical  margin     ...     in  half-grown  and  very  young 
shells"  of  the  Canadian  material.    However,  the  direct  comparison 
of  our  shell  with  the  sketch  by  Whiteaves  (PI.  8,  fig.  2)  shows  not 
much  difference,  if  any,  in  the  number  of  the  lateral  ribs.    The  num- 
ber of  ribs  counted  on  the  periphery  of  the  same  diameter  for  a  volu- 
tion is  about  34  on  the  half- volution  of  this  Kansas  form  and  about 
27  on  the  half-volution  of  the  Canadian  specimen  (counted  on  the 
sketch  on  the  left  half  of  the  outer  volution,  PI.  8,  fig.  2,  of  Whit- 

36.    Whiteaves,  1885,  pi.  8,  figs.  2,  2a. 


Elias:  Cephalopods  of  Pierre  Formatiox  317 

eaves'  paper) .  The  nodes  on  our  specimen  are  as  prominent  and  as 
distantly  spaced  as  in  the  Canadian  type,  in  average  about  every 
fourth  rib  being  provided  with  a  node.  The  peripheral  nodes  are 
also  noticed  on  the  inner  volutions  of  the  Kansas  specimen,  which 
agrees  with  the  characteristic  given  by  Whiteaves,  who  noticed  the 
nodes  in  the  specimens  "about  two  inches  in  their  greatest  diameter 
and  in  still  smaller  ones."  All  these  smaller  specimens  of  S.  sub- 
glohosiis  Whiteaves  must  be  now  considered  young  stages  of  S. 
plenus. 

The  specimen  (18907)  from  Pierre  of  the  Black  Hills  (PL  XXXVI, 
figs,  la,  lb,  Ic)  is  identical  in  almost  every  respect  with  S.  plenus 
from  Wallace  county,  but  it  has  a  somewhat  smaller  umbilicus  and 
same  minor  differences  in  sutures.  The  larger  specimen  from  west- 
ern South  Dakota  (PI.  XXXVII,  figs,  la,  lb,  Ic;  PI.  XXXIX,  figs, 
la,  lb,  Ic)  is  somewhat  more  gibbous  and  the  ribs  of  this  shell  are 
slightly  coarser  and  more  distantly  spaced  than  in  the  other  two 
shells  above  described.  For  instance,  there  are  only  26  peripheral 
ribs  to  the  half  volution  with  diameter  about  30  to  35  mm.,  while  in 
the  shell  from  Wallace  county,  which  has  the  same  diameter,  there 
are  34  ribs.  However,  this  difference  is  not  very  important  and, 
when  considered  alone,  hardly  justifies  separation  of  the  western 
South  Dakota  shell  as  a  variety  of  S.  plenus.  The  size  of  the  um- 
bilicus of  the  form  from  South  Dakota  is  comparable  to  that  of  the 
specimen  from  the  Black  Hills  (18907). 

Meek  illustrated  and  described  only  the  outer  part  of  the  suture 
of  plenus  (see  PL  XL,  fig.  6).  The  material  at  hand  permits  study 
of  the  complete  suture  of  this  species  on  all  three  individuals  (PL 
XL,  figs.  3,  4  and  5)  here  described.  The  sutures  of  these  individuals 
are  not  exactly  alike,  but  their  differences  do  not  seem  to  be  so  im- 
portant as  to  prevent  consideration  of  the  three  as  specifically 
identical.  Possibly  they  may  be  regarded  as  varieties  of  plenus, 
but  one  must  verify  the  constancy  of  those  differences  which  seem 
sufficient  for  a  separation  of  a  variety  on  more  material  than  that 
at  the  writer's  disposal.  The  general  design,  the  number  of  larger 
lobes  and  saddles  and  the  character  of  dissection  of  the  suture  of 
the  described  specimens  are  fairly  identical. 

Meek  pointed  out  the  similarity  of  the  sutures  of  plenus  with  those 
of  S.  nodosus  var.  brevis.  Though  admitting  the  similarities  as 
pointed  out  by  Meek,  the  writer  sees  the  following  common  features 
in  the  sutures  of  Meek's  type  of  plenus  and  of  the  shells  here  de- 
scribed which  may  serve  for  their  distinction  from  the  nodosus  groii]): 


318  The  University  Science  Bulletin 

(Ij  The  general  outline  of  the  first  lateral  saddle  is  nearly  square 
in  both  var.  brevis  and  var.  quadrangularis  of  nodosus  group  (see 
PI.  25,  figs.  Ic,  2c,  3c,  Meek,  1876).  A  square  outline  of  this  saddle 
is  shown  also  by  the  type  specimen  of  S.  nodosusP  Contrary  to  this 
the  first  lateral  saddle  of  S.  plenus  has  an  oval  outline  with  outer 
branch  much  shorter  than  the  larger  inner  branch ;  see  the  suture  of 
the  type  of  S.  plenus  (PI.  XL,  fig.  6)  and  the  sutures  of  all  speci- 
mens here  described  (PI.  XL,  figs.  3,  4  and  5). 

(2)  The  second  lateral  lobe  of  brevis  and  quadrangularis  is  fairly 
symmetrical  and  is  divided  into  two  equal  or  nearly  equal  branches, 
each  of  which  is  in  turn  distinctly  bipartite.  The  corresponding 
second  lateral  lobe  of  S.  plenus  has  its  outer  branch  always  more 
digitate  than  the  inner  branch  (see  all  sutures  of  the  species  here 
illustrated).  Owing  to  this,  the  second  lobe  has  an  appearance  of 
being  subdivided  into  three  branches  (see  the  sutures  of  the  speci- 
mens here  described,  PI.  XL,  figs.  3,  4  and  5). 

There  seem  to  be  no  particular  points  of  distinction  between  the 
inner  part  of  the  suture  of  S.  plenum  and  the  inner  part  of  the  suture 
of  the  specimens  of  A.  nodosus  of  the  writer's  collection.^^ 

Novak  placed  nodosus  Owen  and  its  varieties  as  established  by 
Meek  in  the  genus  Acanthoscaphites,  while  Reeside  expresses  his 
opinion  in  this  matter  in  the  following  way: 

"Whether  all  the  forms  customarily  placed  in  the  species  nodosus  Owen 
belong  to  Acanthoscaphites  may  be  doubted,  but  this  comprehensive  species 
needs  extensive  revision  before  much  can  be  said  of  it  with  confidence,  and 
in  the  meanwhile  it  may  be  left  as  Novak  assigned  it."^^ 

The  writer  did  not  make  any  revision  of  the  whole  nodosus  group, 
but  his  material  on  plenus,  supplemented  by  the  specimens  sent  to 
him  by  Reeside,  permits  him  to  conclude  that  plenus  does  not  belong 
to  Acanthoscaphites,  but  must  be  placed  in  H olcoscaphites  Novak, 
which  is  a  synonym  of  Scaphites  in  the  proper  sense.  The  following 
are  the  features  of  plenus  which  are  characteristic  of  the  latter 
genus : 

(1)  The  shell  is  wider  than  high,  which  holds  true  in  all  stages 
of  growth. 

(2)  In  the  young  stages  of  the  shell  the  outer  sculpture  is  pre- 

37.  Owen,  1852,  pi.  8,  fig.  4a.  Meek  .justly  remarks  that  Owen's  "figure  of  a  septum" 
of  nodosus  "is  evidently  not  drawn  with  sufficient  completeness  and  accuracy  of  detail,"  but 
the  first  lateral  saddle  of  Owen's  drawing,  which  shows  a  square  outline  comparable  to  that 
of  brevis  and  quadrangularis,  seems  to  be  more  accurately  drawn  than  the  rest  of  the  suture. 

38.  The  writer  does  not  know  of  any  published  illustrations  of  the  inner  part  of  the 
sutures  of  A.  nodosus  or  its  varieties. 

39.  Reeside,  1927,  p.  27. 


Eltas:  Cephalopods  of  Pierre -Formation  319 

cisely  of  the  type  which  is  considered  typical  for  Scaphites:  "the 
ribs  pass  without  nodes  from  the  umbilicus,  end  at  the  edges  of  the 
venter  with  a  nodose  thickening  and  then  split  into  weaker,  sec- 
ondary ribs."*" 

The  suture  of  plenus  is  of  the  type  of  S.  aequalis  Sowerby  except 
that  the  second  lateral  lobe  is  much  shorter  than  the  first  lateral 
lobe,  while  according  to  Novak  the  lobes  of  Holcoscaphites 
{=  Scaphites)  decrease  in  size  gradually.  However,  it  must  be 
added  that  in  the  suture  which  is  next  to  the  living  chamber  in  the 
specimen  of  S.  plenus  from  Wallace  county  (PI.  XL,  fig.  5),  the  first 
lateral  lobe  is  only  slightly  higher  than  the  second  lobe,  while  in 
the  next  suture  (not  illustrated  here)  the  first  lobe  is  much  higher 
than  the  second.  It  seems  as  if  the  difference  in  the  sutures  of  this 
sort  may  be  considered  of  lesser  importance  than  the  differences  in 
the  shape  and  sculpture  of  the  shells  of  Scaphites  group.  The  genus 
Yezoites,  based  by  Yabe  on  some  peculiar  characters  of  the  internal 
(dorsal)  part  of  the  suture  in  some  Japanese  species,  is  not  regarded 
valid  either  by  Novak  or  by  Reeside. 

In  conclusion  the  writer  would  say  that  inasmuch  as  he  did  not 
have  an  opportunity  to  revise  the  whole  group  nodosus  Owen  he 
leaves  open  to  question  the  generic  relation  between  plenus  and  the 
rest  of  the  shells  of  the  nodosus  group;  he  thinks,  however,  that 
there  is  enough  evidence  at  hand  to  make  plenus  a  distinct  species. 
The  possibility  of  such  separation  of  S.  plenum  Meek  from  the  rest 
of  the  nodosus  group  was  foreseen  by  Meek  himself,'*^  while  Reeside 
also  holds  the  view  that  "there  is  warrant"  to  do  so.*^ 

Occurrence.  The  specimen  of  S.  plenus  was  found  in  the  con- 
cretionary limestone  zone  at  the  base  of  Salt  Grass  shale  member 
in  Wallace  county.  The  type  specimen  of  S.  plenum  is  referred  by 
Meek  to  the  "upper  part  of  the  Fort  Pierre  group  of  the  Upper  Mis- 
souri Cretaceous  series."  *^  Canadian  specimens  are  referred  to  the 
Fox  Hills  sandstone  north  of  Wood  Mountain,  Alberta.'^*  It  appears 
as  if  S.  plenum  is  characteristic  of  somewhat  higher  horizons  of  the 
Pierre  than  A.  nodosus  and  its  varieties  brevis  and  quadrangularis. 

40.  Reeside,  1927,  p.  24,  definition  of  genus  Holcoscaphites  (=  Scaphites)  translated 
after  Novak. 

41.  Meek,   1876,  pp.   429,   430. 

42.  Letter  to  the  writer  of  December  9,  1930. 

43.  Meek,   1876,  p.   430. 

44.  Ann.  Rept.  Geol.  Survey  Canada,  vol.  1,  p.  46  C,  also  Dowling,  1917,  p.  82. 
According  to  Reeside  this  "Fox  Hills"  is  not  true  Fox  Hills,  but  a  sandy  Pierre  horizon 
(personal  note  of  April,  1931). 


320  The  University  Science  Bulletin 

Genus  Acanthoscaphites  Novak 

Acanthoscaphites  nodosus   (Owen)    (1852) 

This  species  of  Owen  and  its  varieties  hrevis  and  qiiadrangularis 
established  by  Meek  are  the  most  common  scaphites  of  the  Pierre 
in  Kansas.  However,  complete  and  undistorted  specimens  of  this 
form  are  very  rare  and,  in  nearly  every  case,  the  species  or  its 
varieties  were  identified  on  fragments  of  various  size.  In  spite  of 
the  fragmentary  occurrence,  this  important  index  fossil  of  the  Pierre 
can  be  nearly  always  identified  with  a  fair  degree  of  accuracy,  its 
comparatively  large  size,  coarseness  of  costse  and  prominence  of 
nodes  being  very  characteristic. 

Acanthoscaphites  nodosus  (Owen)  s.  s. 

(Plate  XXXVIII,  figs,    la,   lb,   2,   3) 

1852.  Scaphites  (Ammonites)  nodosus,  Owen,  Report  Geol.  Survey  Iowa,  Wis.  and  Minn., 
p.   581,  pi.   8,   fig.   4. 

1892.  Scaphites  nodosus,  Whitfield,  U.  S.  Geol.  Survey,  Mem.  18,  p.  261,  pi.  44, 
figs.   13,   14. 

1896.  Scaphites  7wdosus.  Gilbert,  U.  S.  Geol.  Survey,  17th  Ann.  Rept.,  pt.  2,  pi.  65, 
fig.   2. 

1905.    Scaphites   nodosus  Owen,   Smith   (W.   D.),  Jour.   Geol.,  vol.   13,  pp.   638,   648. 

1907.  Scaphites  nodosus  Owen,  Weller,  Rept.  Cret.  Pal.  New  Jersey,  Geol.  Surv.  New 
Jersey,  Pal.,  vol.  4,  p.  824,  pi.  107,  figs.  1,  2. 

1916.  Acanthoscaphites  nodosus  (Owen),  Novak,  Die  Bedeutung  von  Scaphites  fiir  die 
Gliederung  der  ober  Kreide :     K.    -k.   geol.   Reichsanstalt  Verh.,   Jahrg.   1916,  w.   3,  p.   63. 

1927.    Acanthoscaphites  nodosus,  Reeside,  U.   S.   Geol.   Survey,  Prof.  Paper,  150-B,  p.   32. 

Owen  describes  the  type  of  the  species  as  follows:  ^^ 

"Shell  large  and  ponderous.  Volutions  subcylindrical,  enlarging  gradually 
towards  the  terminal  chamber.  Surface  ornamented  with  sinous  costse,  most  of 
which  bifurcate  at  different  distances  from  the  umbilicus  and,  thus  multiplied, 
proceed  across  the  dorsum.  Two  rows  of  very  prominent  tubercles.  The  row 
near  the  periphery  especially  large  and  prominent,  and  from  one-half  to  three- 
quarters  of  an  inch  apart.  Aperture  subovate.  Greatest  diameter,  four  inches; 
greatest  thickness,  two  and  a  half  inches." 

The  following  characteristic  features  of  the  typical  form  as  com- 
pared with  its  variety  brevis  and  quadrangularis  were  used  by  the 
writer  in  identification  of  his  material  from  the  Pierre  of  Wallace 
county:  (1)  Larger  size;  (2)  costse  of  both  periphery  and  sides  are 
more  distant  than  in  brevis  and  quadrangularis ;  (3)  both  lateral 
nodes  and  peripheral  nodes  are  equally  prominent  and  nearly  equi- 
distantly  spaced;  (4)  the  row  of  nodes  corresponding  to  the  umbilical 
nodes  of  var.  brevis  and  var.  quadrangularis  in  the  typical  nodosus 
is  nearly  in  the  middle  of  the  flank  of  the  shell. 

45.    Owen,  1852,  p.  581. 


Elias:  Cephalopods  of  Pierre  Formation  321 

Occurrence.  In  the  upper  half  of  the  Upper  Weskan  member  and 
in  the  Lake  Creek  member  of  the  Pierre  shale  of  Wallace  county. 
Owen's  type  specimen  came  from  the  upper  part  of  the  Pierre  from 
near  Cheyenne  river,  South  Dakota.'"^  The  specimen  figured  by 
Gilbert  came  from  "Tepee  butte"-zone  of  the  Pierre  of  Arkansas 
river,  Colorado.  According  to  Reeside  ^^  the  species  belongs  to  the 
upper,  as  compared  to  the  lower,  part  of  the  Pierre. 

Aoanthoscaphites  nodosus  var.  brevis  Meek 

CPlate  XXXVI,  fig.  2  ;   Plate  XLI,  fig.  3) 

1876.  Scaphites  nodosus  var.  brevis.  Meek,  U.  S.  Geol.  Survey  Terr.  Kept.,  vol.  9,  p.  426, 
pi.  25,  figs.  ]a,  b,  c. 

Not  1905.  Scaphites  nodosus  var.  brevis  Smith  (W.  D.),  Jour.  Geo!.,  vol.  13,  pp.  638, 
640-649,  fig.  1-2  (p.  641). 

1927.  Aoanthoscaphites  nodosus  var.  brevis,  Reeside,  U.  S.  Geol.  Survey,  Prof.  Paper 
150-B,  p.  32. 

Meek  states  that:  ^^  "This  shell  differs  from  Doctor  Owen's  type 
of  his  S.  nodosus  in  having  its  nonseptate  part  deflected  portion  of 
the  last  volution  much  shorter,  and  its  inner  volutions  more  com- 
pressed, while  the  nodes  near  its  umbilicus  are  decidely  smaller." 

The  writer  w'ould  add  to  this  that  the  costse  of  var.  brevis  are  more 
closely  spaced  than  in  the  typical  A.  nodosus  and  that  the  lateral 
row  of  nodes  in  the  former  is  closer  to  the  umbilicus  than  in  the 
latter  form:  "along  about  one-third  the  height  from  the  umbilicus" 
in  var.  brevis.^^  Furthermore,  there  are  only  a  few  (three  on  fig. 
lb  of  Meek's  monograph)  and  smaller  nodes  at  the  umbilicus  of  var. 
brevis,  while  the  lateral  nodes  are  numerous  (8  or  more)  in  the 
typical  A.  nodosus  and  at  least  the  middle  nodes  of  the  row  are  not 
less  prominent  than  the  peripheral  nodes  of  the  species.^" 

Occurrence.  In  the  Lake  Creek  member  and  at  the  base  of  the 
Salt  Grass  member  of  the  Pierre  shale  in  Wallace  county;  possibly 
also  in  the  Upper  Weskan  shale  member.  The  type  of  the  variety 
came  from  the  upper  part  of  the  Pierre  on  Yellow^stone  river, 
Montana."'^     In  the  Upper  Pierre  according  to  Reeside. ^''^ 

46.  Meek,  1876,  p.   428. 

47.  Reeside,  1927,  p.  32. 

48.  Meek,  1876,  p.  427. 

49.  Meek,  1876,  p.  427. 

50.  Compare  Owen,  1852,  pi.  8,  fig.   4. 

51.  Meek,  1876,  p.  428. 

52.  Reeside,  1927,  p.  32. 


322  The  University  Science  Bulletin 

Acanthoscaphites  nodosus  var.  quadrangularis  Meek  and  Hayden 

(Plate  XXXVII,  fig.  3) 

1860.  Scnphites  nodos'.is  var.  quadrangularis  Meek  and  Hayden,  Proc.  Acad.  Nat.  Sci. 
Phila.,  vol.  12,  p.  420. 

1860.  Scaphites  nodosus  var.  exilis  Meek  and  Hayden,  Proc.  Acad.  Nat.  Sci.  Phila.,  vol. 
12,  p.  420. 

1876.  Scaphites  nodosus  var.  quadrangularis^  Meek,  U.  S.  Geol.  Survey  Terr.,  Rept.,  vol. 
9,  p.  428,  pi.  25,  figs.  2a,  b,  c ;  3a,  b,  c  and  4. 

Not  1905.  Scaphites  nodosus  var.  quadrangularis.  Smith  (W.  D.),  Jour.  Geol.,  vol.  13, 
pp.  638,  640-649,  fig.  1-3  (p.  641). 

1927.  Acanthoscaphites  nodosus  var.  quadrangularis  Reeside,  U.  S.  Geol.  Survey,  Prof. 
Paper  150-B,  p.  32. 

According  to  Meek  this  variety  differs  from  var.  brevis  "not  only 
in  its  usually  smaller  size,  but  also  in  having  its  periphery  flattened, 
its  umbilicus  rather  large,  and  its  body  part  narrower  in  its  vertical 
diameter,  as  well  as  less  straightened  along  its  upper  margin.  Its 
inner  row  of  nodes  will  also  be  seen  to  be  nearer  the  umbilical 
margin,  and  the  outer  rows  near  the  periphery;  the  latter  character 
being,  of  course,  due  to  the  flattening  of  the  periphery.  There  will 
also  be  seen  to  be  some  slight  differences  in  the  details  of  its  septa, 
particularly  in  the  form  of  its  third  lateral  lobe,  and  the  presence  of 
a  small  fourth  lateral. 

"Compared  with  Doctor  Owen's  typical  form  of  S.  nodosus  it  will 
be  seen  to  differ  even  more  strongly  than  the  last,  in  form  and  several 
other  respects." 

The  smaller  size,  the  presence  of  a  number  of  rather  prominent 
nodes  at  the  umbilicus  and  subquadrate  cross  section  are  the  chief 
characters  used  for  identification  of  this  variety  of  A.  nodosus  in  the 
Pierre  of  Wallace  county. 

Occurrence.  In  the  Upper  Weskan  and  in  the  Lake  Creek  shale 
members  of  Wallace  county  Pierre;  also  at  the  base  of  the  Salt  Grass 
shale  member.  In  the  upper  part  of  the  Pierre  on  Yellowstone  river, 
Montana,  according  to  Meek.^^  In  the  upper  Pierre  according  to 
Reeside.^'* 

Acanthoscaphites  (?)  reesidei  Wade 

1926.  Scaphites  reesidei  Wade,  U.  S.  Geol.  Survey,  Prof.  Paper  137,  p.  183,  pi.  61, 
figs.  3-7. 

1927.  Acanthoscaphites  (?)  reesidei,  Reeside,  U.  S.  Geol.  Survey,  Prof.  Paper  150-B, 
p.  33. 

A  good  and  nearly  complete  (except  living  chamber)  specimen 
of  this  rare  species  was  found  by  the  writer  in  Wallace  county  in 
1927.  Unfortunately,  the  specimen  is  not  now  at  his  disposal  and 
therefore  it  can  be  neither  illustrated  nor  described.    The  specimen 

53.  Meek,  1876,  p.   429. 

54.  Reeside,  1927,  p.  32. 


Elias:   Cephalopods  of  Pierre  Formatiox  323 

was  compared  by  the  writer  in  1928  with  the  figures  and  descriptions 
by  Wade  and  was  found  to  agree  with  the  figure  of  the  type  in 
every  respect,  including  the  character  of  the  suture. 

Occurrence.  In  Hmestone  concretionary  zone  at  the  base  of  the 
Salt  Grass  shale  member  in  AVallace  county.  The  type  specimen  of 
A.  reesidei  came  from  the  Coon  creek  locality  of  the  Ripley  formation 
in  Tennessee. 

Genus  Discoscaphites  Meek 

cL  Discoscaphites  (?)  comtrictus  Sov/erhy  var.  tenuistriatus  (Kner) 

(Plate  XXIX,  fig.  3  ;  Plate  XXXIX,  fig.  10) 

Compare  1908.  Scaphites  constrictv^  Sowerby,  Grossouvre,  Mem.  Mus.  Royale  D'  Hist. 
Nat.  Belgique,  p.  36,  pi.  11,  fig.  6. 

Compare  1911.  Hoploscaphites  coTistrictus -tenuistriatus  Novak,  Acad.  Sci.  Cracovie  Bull, 
intemat.,  ser.  B,  p.  585,  pi.  33,  figs.  13,  21  and  22. 

Though  the  writer  realizes  that  an  identification  of  European 
species  among  the  American  Upper  Cretaceous  forms  must  be  made 
with  caution,  he  nevertheless  compares  provisionally  fragmentary 
remains  from  Wallace  county  with  some  illustrated  shells  of  the 
constrictiis  type  from  Europe,  because  the  Wallace  county  remains 
differ  from  all  American  Upper  Cretaceous  amonities  so  far  de- 
scribed. Their  size  is  small,  they  are  considerably  compressed,  their 
surface  is  marked  by  fine  ribs  and  they  have  a  small  umbilicus. 

In  all  these  respects  the  remains  are  comparable  to  Discoscaphites 
nicolleti  Morton,  but  not  the  faintest  trace  of  tubercles  of  any  kind 
can  be  observed  on  the  specimens  from  Wallace  county.  It  is  in- 
teresting to  note  that  Meek  ^^  compared  D.  nicolleti  with  some 
tuberculate  varieties  of  Scaphites  constrictus  Sowerby  from  Europe, 
as  illustrated  by  d'Orbigny,^*^  and  according  to  Novak  both  tuber- 
culate and  nontuberculate  forms  exist  among  the  shells  which  he 
refers  to  S.  constrictus  var.  tenuistriatus,  with  which  the  specimens 
from  Wallace  county  have  so  much  in  common.  Thus  it  appears  as 
if  we  might  extend  the  conception  of  D.  nicolleti  so  as  to  include 
nontuberculate  forms,  which  otherwise  differ  little  or  not  at  all  from 
the  typical  nicolleti.  This,  however,  is  not  advisable  for  the  follow- 
ing reasons: 

(1)  The  presence  of  peripheral  tubercles  is  mentioned  in  the  origi- 
nal characterization  of  the  type  specimen  of  nicolleti  Morton.^^ 

(2)  The  stratigraphic  position  of  true  nicolleti  is  Fox  Hills  and  the 
upper  part  of  the  Pierre.     Though  the  writer  collected  this  species 

55.  Meek,  1876,  p.   436. 

56.  Paleont.  Fr.,  Terr.  Cret.,  I,  pi.   129,  fig.  8. 

57.  1842,  p.  209,  though  tubercles  are  not  shown  on  the  much  generalized  sketch,  pi  10 
fig.   3. 


324  The  University  Science  Bulletin 

in  Wallace  county  as  low  as  at  the  top  of  the  Salt  Grass  shale  mem- 
ber, which  is  about  in  the  middle  of  the  Pierre,  he  considers  the 
Salt  Grass  specimens  a  new  variety  of  nicolleti,  which  is  described 
below  (this  variety  is  tuberculate).  The  remains  here  compared 
with  the  nontuberculate  varieties  of  European  constrictus  have  been 
collected  as  low  in  the  Pierre  as  in  the  Lower  Weskan  shale  member, 
which  is  only  about  200  feet  above  the  base  of  the  formation. 

If,  in  the  course  of  time,  the  identity  of  nicolleti  with  European 
constrictus  is  proved,  there  would  be  more  reason  to  separate  the 
nontuberculate  European  variety  of  constrictus  as  a  distinct  form, 
and  not,  contrary  to  this,  mix  up  the  American  tuberculate  and  non- 
tuberculate forms  of  the  nicolleti  type,  which,  as  far  as  is  established 
at  present,  do  not  occur  together  in  the  stratigraphic  column  of  the 
North  American  Upper  Cretaceous. 

In  view  of  all  these  considerations  the  nontuberculate  remains 
here  described  should  be  given  a  new  name.  However,  due  to  the 
imperfection  of  the  remains  from  Wallace  county  and  the  badly 
preserved  suture,  the  writer  prefers  to  designate  them  temporarily 
as  cf.  D.  constrictus  var.  tenuistriatus. 

Occurrence.  Two  small  individuals,  the  better  of  which  is  shown 
on  Plate  XXXIX,  figure  10,  came  from  limestone  concretions  of  the 
Weskan  shale,  and  probably  from  the  Lower  Weskan  shale  member. 
The  larger  individual  of  Plate  XXIX,  figure  3,  was  collected  in 
the  Upper  Lake  Creek  shale  member.  All  specimens  were  collected 
in  Wallace  county. 

Discoscaphites  nicolleti  (Morton)  Meek 

1842.  Ammonites  nicolleti  Morton,  Acad.  Nat.  Sci.  Phila,  Jour.,  1  ser.,  vol.  8,  pt.  2,  p. 
209,  pi.  10,  fig.  3. 

1852.  Ammonites  nicolleti  Owen,  Rept.  U.  S.  Geol.  Surv.  Wisconsin,  Iowa  and  Minnesota, 
pi.  8,  fig.   1. 

1852.    Scaphites   (^ATnm^mites)  comprimis  Owen,   ibid.,   p.    580,   pi.   7,   fig.    4. 

1857.    Scaphites  nicolleti  Meek  and  Hayden,  Acad.  Nat.  Sci.  Phila.  Jour.,  vol.  8,  p.  281. 

1876.  Scaphites  (Discoscaphites)  nicolleti  Meek,  U.  S.  Geol.  Survey  Terr.,  Rept.,  vol.  9, 
p.   435,  pi.   34,  figs.   2  and  4. 

1927.  Discoscaphites  nicolleti  Reeside,  U.  S.  Geol.  Survey,  Prof.  Paper  150-B,  p.  31-32, 
pi.  9,  figs.  5-7. 

Meek  gives  the  following  definition  of  the  species: 

"Shell  oval-subcircular,  much  compressed;  volutions  so  deeply  embracing 
as  to  leave  only  a  small  umbilicus,  all  strongly  compressed  laterally,  inner  ones 
narrowly  rounded  on  the  periphery;  deflected  part  of  last  turn  so  very  short 
as  not  to  become  free  at  the  aperture,  narrowly  flattened  on  the  periphery 
below,  and  somewhat  widened  and  straightened  along  the  upper  margin  near 
the  umbilicus;  aperture  narrow-oval;  surface  ornamented  by  numerous  small, 
somewhat  flexuous  costse,  which  increase  by  division  and  intercalation  so  as 


Elias:   Cephalopods  of  Pierre  Formation  325 

to  number  about  five  time  as  many  around  the  periphery  as  at  the  inner  side; 
eostae  everywhere  without  tubercles  or  nodes,  excepting  a  single  row  along 
each  side  of  the  flattened  periphery  of  the  outer  volution,  all  crossing  the 
periphery  with  a  moderate  forward  curve. 

"Length,  2.31  inches;  height,  1.92  inches;  convexity,  about  0.62  inch."  =58 
The  suture  of  the  species  was  unknown  until  the  description  by 
Meek.  According  to  this  author  the  suture  of  the  species  is  much 
like  that  of  Scaphitcs  conradi  var.  intermedins,  which  "holds  almost 
an  exactly  intermediate  position,  as  it  were,  between  the  typical  .S. 
conradi  and  *S.  nicolleti  Morton  (=5.  comprimis  Owen),  both  in 
external  and  internal  characters. "'•'' 

Though  Meek  points  out  some  minor  differences  in  the  sutures  of 
S.  nicolleti  and  S.  conradi  var.  intermedins,  he  says  that  "these  dif- 
ferences in  the  details  of  the  lobes  and  sinuses  of  the  septa  are  not, 
it  must  be  confessed,  very  important,  or  even  not  greater  than  we 
may  frequently  see  between  those  of  different  individuals  of  the 
same  species."  *^°  Meek  furthermore  stresses  the  differences  in  the 
sculpture  between  the  conradi  and  nicolleti  groups  and  points  out 
the  "entire  absence  of  tubercles  on  the  sides  of  all  the  volutions, 
both  inner  and  outer"  ^^  of  the  nicolleti  form.  In  this  respect  the 
"shell  departs  from  the  Discoscaphites  group,"  but  "from  its  close 
general  relations  to  the  last,  however,  in  which  that  character  does 
occur,  both  in  form  and  in  the  details  of  the  septa,  it  seems  improper 
to  place  it  in  any  other  section."  ^- 

It  is  interesting  to  point  out  that  in  Owen's  Scaphites  comprimis, 
which  Meek  classifies  with  D.  nicolleti,  there  is  a  row  of  "more 
obscure  tubercles,  one-fourth  of  the  distance  from  the  inner  margin 
of  the  convolutions,"  ^^  which  are  also  shown  on  Owen's  sketch. 
Since  the  outer  sculpture  is  an  important  feature  in  classification 
of  the  species  of  the  Discoscaphites  group,  it  appears  that  Owen's 
type  may  possibly  be  a  variety  of  D.  nicolleti,  having  not  only  an 
additional  row  of  tubercles  but  also  a  somewhat  different  arrange- 
ment of  costse.  As  will  be  shown  below,  the  specimens  of  D.  nicolleti 
from  Wallace  county  have  more  distantly  spaced  marginal  costse 
than  in  the  specimens  described  by  Meek.  There  are  again  some 
differences  in  the  outer  sculpture  as  well  as  in  the  suture  line,  which 
permit  the  separation  of  this  as  a  new  variety  of  the  nicolleti  type. 

58.  Metk,   1876,  p.   435. 

59.  Meek,   1870,   p.    434. 

60.  Ibid.,   p.   434. 

61.  Ibid.,  p.   436. 

62.  Ibid.,   p.   436. 

63.  Owen,   18.52,  p.   580. 

22—3482 


326  The  University  Science  Bulletin 

Occurrence.  According  to  Meek  the  species  was  found  in  the  Fox 
Hills  group  in  South  Dakota  and  Montana.  The  specimen  illustrated 
by  Reeside  came  from  the  top  of  the  Pierre  shale  at  Linton,  North 
Dakota.  ^ 

Discoscaphites  nicoUeti  var.  saltgrassensis  Elias,  n.  var. 

(Plate  XXXVI,  figs.  3a,  3b,  3c;   Plate  XL,  fig.  1) 

The  variety  differs  from  D.  nicolleti  as  described  by  Meek  in 
having  the  marginal  tubercles  not  on  every  costse  as  shown  in  Meek's 
monograph  (PI.  34,  fig.  26).^'*  On  the  shell  of  var.  saltgrassensis 
the  tubercles  begin  to  appear  when  the  radius  of  the  volutions  ap- 
proaches 15  mm.  At  this  radius  each  fourth  of  the  peripheral  costae 
has  a  tubercle,  but  soon,  at  about  20  mm.  radius  of  the  volutions, 
each  third  costse  has  a  tubercle,  while  Meek  shows  a  tubercle  on 
every  peripheral  costa  at  23  to  25  mm.  radius.^^  The  number  of  the 
costae  of  the  specimens  as  figured  by  Meek  appears  to  be  about  the 
same  as  on  the  specimens  from  Wallace  county.  Meek  does  not 
give  any  figures  for  the  number  of  the  Costse  on  the  periphery,  which 
for  our  variety  is  10  to  11  per  1  cm.  at  the  radius  of  15  mm.  and  is 
about  7  per  1  cm.  at  the  end  of  the  next  half  of  a  complete  volution 
or  at  about  25  mm.  radius.  The  shape  of  the  variety  is  the  same  as 
in  the  specimens  figured  by  Meek,  the  periphery  of  the  volutions 
being  broadly  rounded  (as  on  Meek's  PI.  34,  fig.  46,  and  not  as  on 
PL  34,  fig.  26,  where  the  periphery  is  shown  to  be  nearly  acute  at 
the  end  of  the  preserved  volutions) . 

The  suture  of  the  variety  differs  in  one  respect  more  from  that  of 
D.  nicolleti  Meek  than  the  latter  differs  from  the  suture  of  D.  con- 
radd  var.  intermedius.  The  first  lateral  saddle  of  var.  saltgrassensis 
is  oval  in  outline  while  this  saddle  has  a  nearly  square  outline  in 
both  D.  nicolleti  Meek  and  D.  conradi  var.  intermedius  Meek.  In 
other  words,  while  the  three  branches  of  the  first  saddles  in  the  two 
latter  types  are  nearly  equal  in  height,  the  central  branch  of  the 
first  saddle  of  var.  saltgrassensis  is  the  highest,  while  the  lateral 
branches  and  branchlets  slope  down,  gradually  filling  a  perfectly 
oval  outline  of  the  saddle  (see  PI.  XL,  fig.  1,  of  this  report).  How- 
ever, this  and  some  other  minor  differences  in  the  suture  of  var. 
saltgrassensis  are  not  very  important  and,  if  not  supported  by  dif- 
ferences in  the  sculpture,  would  not  justify  the  separation  of  the 
variety  saltgrassensis  from  typical  D.  nicolleti. 

64.     Meek,  1876. 

6.5.     Ibid.,  pi.  34,  fig.   26. 


Elias:   Cephalopods  of  Pierre  Formation  327 

The  lower  stratigraphic  position  of  the  var.  saltgrassensis  at  the 
middle  of  the  Pierre  formation  is  another  point  in  favor  of  its  separa- 
tion as  a  distinct  variety  of  D.  nicolleti,  which  belongs  to  the  Fox 
Hills  beds  and  to  the  top  of  the  Pierre  shale. 

It  is  interesting  to  add  that  the  suture  of  D.  nicolleti  from  the  top 
of  the  Pierre  at  Linton,  South  Dakota,  as  figured  by  Reeside  (1927, 
PL  9,  fig.  7)  is  closer  to  that  of  var.  saltgrassensis  than  to  the  typical 
form  as  shown  by  Meek. 

Occurrence.  The  variety  was  found  in  a  lens  of  typical  "Lucina 
limestone"  at  the  top  of  the  Salt  Grass  member  of  the  Pierre  in 
Wallace  county  in  the  center  of  sec.  2,  T.  12  S.,  R.  42  W.;  thus 
stratigraphically  the  variety  belongs  to  the  middle  of  the  Pierre  as 
represented  in  northwestern  Kansas.  It  appears  that  the  first  ap- 
pearance of  the  Discoscaphites  nicolleti  in  the  Pierre  of  the  Central 
High  Plains  belongs  to  this  horizon.  At  the  base  of  the  Salt  Grass 
member,  or  only  about  50  to  60  feet  stratigraphically  below  the 
beds  with  D.  nicolleti  var.  saltgrassensis,  the  typical  representatives 
of  the  Acanthoscaphitcs  group,  such  as  A.  nodosus  var.  brevis  and 
var.  quadrangularis  and  also  Acanthoscaphites  (?)  reesidei  and 
Scaphites  plenus,  were  collected.  A  small  scaphite,  very  similar  to 
var.  saltgrassensis,  was  also  collected  in  the  "Lucina  limestone" 
cores  at  the  base  of  the  Salt  Grass  member,  but  these  specimens  were 
not  preserved  well  enough  for  precise  identification. 

Discoscaphites  conradi  (Morton) 

(Plate  XXXIX,  figs.   9a,   9b) 

1S34.  Ammo-nites  conradi,  Morton,  Synopsis  of  organic  remains  of  Cretaceous  group  of 
the  U.  S.  A.,  p.  39,  pi.  Iti,  figs.   1-3. 

1850.    Scaphites  conradi,  d'Orbigny,  Prodrome  de  Paleon.,  p.   214. 

1S71.    Scaphites   (.Discoscaphites)   conradi.  Meek,  Am.  Phil.  So.,  Proc,  vol.  11,  p.   429. 

1876.  Scaphites  (Discoscaphites)  conradi.  Meek,  U.  S.  Geol.  Survey  Terr.,  Rept.,  vol.  9, 
p.  430,  pi.  36,  fig.  2. 

1916.  Scaphites  conradi,  Gardner,  Marjland  Geol.  Survey,  Upper  Cretaceous,  p.  383,  pi. 
12,  fig.  1. 

1927.    Discoscaphiies  conradi,  Reeside,  U.  S.  Geol.  Survey,  Prof.  Paper  150-B,  p.   28. 

A  fragment  which  undoubtedly  belongs  to  this  form  was  found  in 
the  Beecher  Island  shale  and  identified  by  the  writer  in  1928.  The 
species  was  recognized  by  the  discoidal  form  and  by  the  presence 
of  three  to  four  rows  of  nodes  on  the  outer  part  of  the  flanks  of  the 
shell.  These  nodes  were  located  on  the  costse  of  the  outer  sculpture. 
The  size  of  the  shell  as  judged  from  the  fragment  is  suggestive  of 
either  D.  conradi  or  D.  cheyennensis  of  the  North  American  Dis- 
coscaphites, while  the  comparative  evenness  of  the  venter  resembles 


328  The  University  Science  Bulletin 

that  of  former  species.  Among  the  so-far  published  illustrations  of 
American  Discoscaphites  the  shell  from  Beecher  Island  i;fsembles 
most  that  figured  by  Gardner.^^  Another  fragment  of  a  smaller  in- 
dividual of  D.  conradi  was  found  by  the  writer  in  1930  and  is  here 
illustrated  (PL  XXXIX,  figs.  9a,  9b). 

Occurrence.  In  the  Beecher  Island  member  of  the  Pierre  shale 
at  Beecher  Island,  Colorado.  The  type  species  of  D.  conradi  came 
from  the  Upper  Cretaceous  beds  of  Alabama.*^^  According  to  Meek: 
'Tt  also  occurs  at  the  same  horizon  in  New  Jersey,"  ^^  but  no 
identical  or  related  forms  have  been  described  in  comprehensive 
monographs  of  the  Upper  Cretaceous  fauna  of  New  Jersey  by  later 
authors.^^  Stephenson  records  D.  conradi  in  the  Exogyra  costata 
zone  of  the  Eastern  Gulf  region  and  in  the  Liopistha  protexta  sub- 
zone  at  the  top  of  Selma  chalk  in  western  Alabama  and  east-central 
Mississippi.'^^ 

Meek  has  "seen  specimens  of  it  from  Saskatchewan,  British 
America,  in  Professor  Hind's  collections."  The  specimens  described 
by  Meek  came  from  the  "Fox  Hills  group  at  Fox  Hills,  Long  Lake, 
Moreau  river  and  near  the  eastern  base  of  the  Black  Hills, 
Dakota."  '^ 

Gardner  described  the  species  from  the  Monmouth  formation  of 
Maryland.'^^  According  to  Reeside  the  species  occurs  in  the  Fox 
Hills  sandstone,  which  he  compares  with  the  Maestrichtian  of 
Europe. 

Discoscaphites  abyssinus  (Morton) 

(Plate  XXXIX,  figs.   2,   3,   4,   5,  6a,  6b,  7,   8;    Plate  XL,  fig.   2) 

1841.  Ammonites  abyssinus  Morton,  .Jour.  Acad.  Nat.  Sci.  Phila.,  vol.  8,  p.  209,  pi.  10, 
figure  4. 

1856.  Scaphites  mandanensis?  Meek  and  Hayden,  Proc.  Acad.  Nat.  Sci.  Phila.,  vol.  8, 
p.  281. 

1860.    Scaphites  abyssinus  Meek  and  Hayden,  Proc.  Acad.  Nat.  Sci.  Phila.,  vol.   12,  p.  420. 

1864.  Scaphites  abyssi7uis  Meik  and  Hayden,  Smithsonian  Check-List  of  North  American 
Cretaceous  Fossils,  p.   23. 

1876.  Scaphites  (Discoscaphites)  abyssinus  Me^k,  Invertebrate  Cretaceous  and  Tertiary 
fossils  of  the  Upper  Missouri  country,  U.  S.  Geol.  Survey  Terr.,  Kept.,  vol.  9,  p.  441,  pi. 
35,  figs.   2a,  b  and   4. 

1927.  Discoscaphites  abyssinus  Reeside,  Scaphites,  an  Upper  Cretaceous  Ammonite  group, 
U.  S.  Geol.  Survey,  Prof.  Paper  150-B,  pp.   27,  36. 


06.  Gardner,  1916,  pi.   12. 

67.  Morton,   1834,  p.  39. 

68.  Meek,  1876,  p.  432. 

69.  Whitfield,  1886  and  1892;   Welkr,   1907 

70.  Stephenson,  1914,  pp.   36-37. 

71.  Meek,  1876,  p.   432. 

72.  Gardner,   1910,  p.   383,  pi.    H. 


Elias:   Cephalopods  of  Pierre  Formation  329 

Meek  gives  the  following  definition  of  the  species:  ^^ 

"Shell  short-oval,  or  subcircular,  much  compressed;  inner  volutions  deeply 
embraced  within  the  dorsal  groove  of  each  succeeding  turn ;  last  whorl  flattened 
on  the  sides  and  periphery,  having  a  more  or  less  distinct  subnodose  angle 
around  the  umbilicus,  and  on  each  side  of  the  periphery;  nonseptate  portion 
short,  not  widened  or  straightened  along  its  upper  margin,  and  deviating  so 
little  from  the  regular  curve  of  the  inner  whorls  as  to  become  but  slightly 
disconnected  from  them  at  the  aperture;  umbilicus  rather  small;  aperture 
oval  or  subcordate;  surface  ornamented  by  rather  distinct,  straight,  or  slightly 
flexuous  costae,  which  increase  chiefly  by  the  intercalation  of  shorter  ones 
between  the  longer,  so  as  to  number  from  two  to  three  times  as  many  at  the 
periphery  as  near  the  umbilicus;  those  on  the  inner  whorls  often  supporting 
some  three  or  more  rows  of  very  small  nodes  on  each  side,  in  addition  to  the 
larger  series  on  each  side  of  the  narrow,  flattened  periphery. 

"Septa  divided  into  three  or  four  lobes  and  as  many  sinuses  on  each  side 
of  the  siphonal  lobe,  which  is  of  an  oblong  form,  being  a  little  longer  than 
wide,  with  two  principal  branches  on  each  side,  the  two  terminal  of  which  are 
slender,  larger  than  the  others,  nearly  parallel,  obscurely  bifid,  and  provided 
with  a  few  obtuse,  short  digitations  or  crenulations;  while  the  lateral  branches 
are  short  and  spreading,  the  larger  pair  being  merely  obtusely  tridentate, 
and  the  others  simple;  first  lateral  sinus  as  long  as  the  siphonal  lobe,  and  near 
one-fourth  wider,  with  a  nearly  quadrangular  form,  and  provided  with  two 
large  unequal,  irregular  branches,  with  short,  obtusely  crenate  subdivisions; 
first  lateral  lobe  a  little  shorter  than  the  siphonal,  and  of  nearly  the  same 
breadth,  with  two  branches  on  each  side,  the  two  terminal  of  which  are  of 
moderate  length,  spreading,  bifid,  and  provided  with  obtusely  crenate  or 
dentate  margins,  while  the  two  lateral  branches  are  much  smaller,  directed  out 
at  right  angl&s  from  the  margins,  and  each  obtusely  tridentate  at  the  end; 
second  lateral  sinus  scarcely  half  as  large  as  the  first,  and  bearing  two  unequal, 
shortly  bipartite  or  tripartite,  deeply  sinuous,  and  obtusely  dentate  terminal 
branches,  and  one  or  two  unequal,  short,  nearly  simple,  lateral  branchlets; 
second  lateral  lobe  much  like  the  first,  but  scarcely  more  than  half  as  long 
and  wide;  third  lateral  sinus  about  one-third  as  large  as  the  second,  with  two 
short,  spreading,  obtusely  dentate,  terminal  branches;  third  lateral  lobe  very 
small  and  merely  obscurely  bilobate  at  the  end;  fourth  lateral  sinus  as  long 
as  the  third  lateral  lobe,  but  wider  and  faintly  bilobate  at  the  end;  fourth 
lateral  lobe  slightly  wider  and  longer  than  the  third,  but  more  distinctly 
trilobate  at  the  end. 

"Length,  2.09  inches;  height,  1.65  inches;  convexity,  0.63  inch. 

"The  angle  around  the  inner  side  of  the  nonseptate  part  of  the  outer 
volution  in  this  species  is  not  always  well  marked,  and  varies  a  little  in  its 
distance  from  the  umbilical  margin.  It  is  apparently  always  provided  with  a 
row  of  low  prominences,  like  transversely-elongated  nodes,  that  never  exist  on 
the  inner  volutions.  The  angle  on  each  side  of  the  narrow,  flattened  periphery, 
with  its  row  of  nodes,  seems  to  be  always  well  defined  on  the  outer  volution, 
excepting  near  the  aperture,  where  both  angles  and  nodes  usually  fade  away 
and  the  costse  become  finer  and  crowded.     On  the  inner  volutions,  also,  the 

73.     Meek,    1876,   pp.    441,    442. 


330  The  University  Science  Bulletin 

peripheral  angles  become  obscure  or  obsolete,  though  their  place  is  occupied 
by  the  rows  of  nodes  reduced  in  size.  The  little  nodes  seen  on  the  sides  of 
the  inner  volutions,  and  sometimes  on  the  inner. half  of  the  last  turn,  are 
placed  on  the  costse  so  as  to  form  about  three,  nearlj'  equidistant,  revolving 
rows  on  each  side  of  the  shell." 

The  specimens  which  were  collected  in  the  Upper  Pierre  shale  near 
Beecher  Island  and  in  the  corresponding  beds  in  northern  part  of 
Cheyenne  county,  Kansas,  are  somewhat  smaller  than  the  types 
figured  by  Morton  and  Meek  and  are  slightly  different  in  some  other 
respects,  which,  however,  do  not  seem  to  be  so  important  as  to 
warrant  separation  of  the  form  collected  from  these  localities  from 
the  typical  species. 

The  fragment  of  Plate  XXXIX,  figure  8,  represents  the  largest 
individual  of  the  collection,  which  approaches  in  size  Meek's  types, 
while  the  nearly  complete  specimen  (PI.  XXXIX,  fig.  3)  is  only 
slightly  smaller.  It  is  interesting  to  note  that  in  both  these  indi- 
viduals the  "low  prominences  like  transversely  elongated  nodes" 
(Meek)  are  so  insignificant  as  to  be  nearly  absent.  However,  they 
are  quite  prominent  on  smaller  individuals  (PL  XXXIX,  figs.  2,  4, 
7a).  The  small  lateral  nodes  of  the  inner  volutions  seem  to  be 
absent  in  all  specimens  of  the  collection  with  the  exception  of  one, 
in  which  only  one  row  of  lateral  nodes  (instead  of  three  rows  as 
according  to  Meek)  is  developed  (PI.  XXXIX,  fig.  4).  However, 
the  presence  of  lateral  nodes  is  not  a  constant  feature  of  the  species. 
When  comparing  his  material  with  Morton's  type  Meek  remarked 
that  the  type  specimen  itself  "showed  none  of  the  little  nodes  on  the 
costsB  of  the  sides  of  the  shell"  and  that  "these  are  not  always 
present"  on  the  material  described  by  Meek.  The  suture  of  the 
form  here  described  does  not  differ  much  from  that  shown  by  Meek 
for  D.  abyssinus  (the  suture  of  Morton's  type  is  unknown).  The 
main  features  which  permit  classification  of  the  specimens  from 
Beecher  Island  shale  member  with  Morton's  species  are  the  general 
similarity  in  form  and,  to  a  certain  extent,  in  size  of  the  shells;  the 
presence  of  prominent  nodes  on  the  periphery  and  at  the  umbilical 
margin  of  the  outer  volution;  the  change  from  rather  prominent 
regular  costae  to  much  finer  and  crowded  costse  at  the  outer  part  of 
the  aperture;  the  very  narrow  umbilicus.  The  suture  is  quite  typical 
for  the  species  (see  PI.  XL,  fig.  2). 

To  the  characterization  of  the  species  given  by  Meek  may  be 
added  that  the  living  chamber,  as  noted  on  the  material  here  de- 
scribed, occupies  a  little  more  than  one-half  of  a  volution. 


Elias:   Cephalopods  of  Pierre  Formation  331 

Distribution.  In  the  Beecher  Island  shale  member  of  the  Pierre 
in  sec.  8,  T.  2  S.,  R.  43  W.,  Yuma  county,  Colorado,  and  in  the 
corresponding  beds  in  NEi^  sec.  9,  T.  2  S.,  R.  42  W.  and  in  NE^A 
sec.  5,  T.  IS.,  R.  39  W.  of  Cheyenne  county,  Kansas. 

According  to  Meek  the  specimens  from  Moreau  river,  South 
Dakota,  belong  to  the  "Fox  Hills  group." 

Subfamily  Placenticeratin^ 
Genus  Placenticeras  Hyatt 
Placenticeras  meeki  Boehm 

(Plate  XLI,   figs,   la,  lb,   2;    Plate  XLII,   figs,   la,   lb) 

1870.  Placenticeras  placenta.  Meek,  U.  S.  Geol.  Survey  Terr.,  vol.  9,  p.  465,  text  fig.  6.') ; 
pi.  24,  fig.  2. 

1898.  Placenticeras  meeki  Boehm,  Deutsche  Geol.  Gesell.  Zeitschr.,  vol.  50,  p.  200  (foot- 
note). 

1903.  Placenticeras  whitfieldi  Hyatt,  U.  S.  Geol.  Survey,  Mon.  44,  p.  221,  pi.  45,  figs. 
3-16;   pi.  46;   pi.  47,  figs.  1-4. 

1910.  Placenticeras  whitefieldi  Grabau  and  Shimer,  North  American  Index  Fos.sils,  p.  218, 
figs.  1493,  1494. 

1927.  Placenticeras  meeki,  Reeside,  U.  S.  Geol.  Survey,  Prof.  Paper  ].->!.  p.  20,  pi.  22, 
figs.  5-7;   pi.   23;   pi.  24;   pi.  25,  figs.  1-2. 

This  large  Placenticeras  is  common  in  the  Pierre  of  Wallace 
county,  where  no  other  species  of  the  genus  have  been  found  by  the 
writer.  The  species  was  recognized  by  its  large  size,  smooth  non- 
tuberculate  sides,  compressed  volutions,  venter  narrow  in  youth  and 
moderately  rounded  in  old  age,  narrow  umbilicus  and  a  very  typical 
suture.  For  the  detailed  characteristics  of  the  species  see  Hyatt  "^ 
and  Reeside.'^^ 

The  remains  collected  in  Wallace  county  represent  shells  of  large 
to  very  large  size.  A  nearly  complete  shell  (PL  XLI,  figs,  la,  lb,  2 
and  PI.  XLII,  figs,  la,  lb)  approaches  18  inches  in  diameter  with 
part  of  the  living  chamber  preserved.  The  living  chamber  shows 
broadly  undulated  venter  (PI.  XLII,  fig.  la) . 

Some  fragments  from  Wallace  county,  which  seem  to  belong  to  P. 
meeki,  indicate  that  the  species  attained  as  large  a  size  as  31/2  feet 
in  diameter. 

Occurrence.  In  the  lower  and  upper  Weskan  shale  member  and 
in  the  lower  part  of  Lake  Creek  shale  member  of  the  Pierre  in 
Wallace  county.  According  to  Reeside '«  this  species  ''has  been 
found  widely  distributed  in  the  Pierre  shale  and  equivalent  fonna- 
tions  of  the  Western  Interior  province  of  the  American  Cretaceous." 

74.  Hyatt,  1903,  p.  221. 

75.  Reeside,  1927,  p.  29. 

76.  Reeside,  1927,  p.  30. 


332  The  University  Science  Bulletin 


LTST  OF  WORKS  TO  WHICH  REFERENCE  IS  MADE 

Darton,  N.  H.     1905.    Preliminaiy  Report  on  the  Geology  and  Underground 

Water  Resources  of  the  Central  Great  Plains:    U.  S.  Geol.  Survey,  Prof. 

Paper  32. 

Gardner,  Julia  Anna.    1916.    Upper  Cretaceous.    Maryland  Geological  Survey. 

Rall,  James,  and  Meek,  F.  B.     1S56.    Descriptions  of  new  species  of  fossils 

from  the  Cretaceous  formations  of  Nebraska     .     .     .    Am.  Acad.  Arts  and 

Sci.,  Mem.  n.s.,  vol.  5,  p.  379. 

Hyatt,  Alpheus.    1903.    Pseudoceratites  of  the  Cretaceous :   U.  S.  Geol.  Survey 

Mon.  44. 
Logan,  W.  N.     1897.     Upper  Cretaceous  of  Kansas:    Univ.  Geol.  Survey  of 

Kansas,  vol.  II,  p.  195. 
Logan,  W.  N.     1898.    Invertebrates  of  the  Benton,  Niobrara  and  Fort  PieiTe 

Groups:   Univ.  Geol.  Survey  of  Kansas,  vol.  IV,  p.  431. 
Mather,  K.  F.,  Gilluly,  J.,  and  Lusk,  R.  G.    1928.    Geology  and  Oil  and  Gas 
Prospects  of  Northeastern  Colorado:    U.  S.  Geol.  Survey  Bull.  796-B,  p.  65. 
Meek,  F.  B.    1876.    Report  on  the  Invertebrate  Cretaceous  and  Tertiary  Fos- 
sils:  U.  S.  Geol.  Survey  Terr.,  vol.  9. 
Morton,  S.  G.     1834.    Synopsis  of  Organic  Remains  of  Cretaceous  Group  of 

the  U.  S. 
Novak,  J.    1908.   Untersuchungen  liber  die  Cephalopoden  der  Oberen  Kreide 

in  Polen.    I  Teil,  Bull.  Acad.  Sci.  Cracovie,  serie  B,  April,  1908,  p.  324. 
d'Orbigny.    1850.    Prodrome  de  Paleon. 

Owen,  D.  D.    1852.    Report  Geol.  Survey  Wisconsin,  Iowa  and  Minnesota. 
Reesidb,  J.  B.,  Jr.     1927.     Cephalopods  of  the  Eagle  Sandstone  and  Related 

Formations:     U.  S.  Geol.  Survey,  Prof.  Paper  151. 
Stanton,  T.  W.    1893.    Colorado  Formation  and  Its  Invertebrate  Fauna:    U. 

S.  Geol.  Survey,  Bull.  106. 
Stanton,  T.  W.     1897.     Stratigraphy  and  Paleontology  of  the  Laramie  and 
Related  Formations  in  Wyoming:    Bull.  Geol.  Soc.  America,  vol.  8,  p.  127. 
Stanton,  T.  W.    1925.   "Fossil  Content"  in  W.  L.  Russell's  Well  Log  in  North- 
ern Ziebach:    South  Dakota  Geological  and  Natural  History  Survey,  Cir- 
cular 18,  p.  8. 
Stephenson,  L.  W.    1914.    Cretaceous  deposits  of  the  eastern  Gulf  region  and 
species  of  Exogyra  from  the  eastern  Gulf  region  and  the  Carolinas :    U.  S. 
Geol.  Survey,  Prof.  Paper  81. 
Weller,  Stuart.    1907.    Report  on  the  Cretaceous  Paleontology  of  New  Jersey: 

New  Jersey  Geol.  Survey,  Pal.  Series  No.  4. 
Whiteaves,  J.  F.    1885.    Report  on  the  Invertebrates  of  the  Laramie  and  Cre- 
taceous Rocks  of  the  Vicinity  of  the  Bow  and  Belly  Rivers  and  adjacent 
Localities  in  the  Northwest  Territory:    Canadian  Geol.  Survey,  Contr.  to 
Paleontology,  No.  1. 


Elias:   Cephalopods  of  Pierre  Formation  333 

Whitfield,  R.  P.     1885.     Brachiopoda  and  Lamellibranchiata  of  the  Raritan 

Clays  and  Greensand  Marls  of  JS'ew  Jersey:   New  Jersej^  Geol.  Survey,  Pal. 

Series  No.  1.    Also,  1886,  U.  S.  Geol.  Survey  Mon.  9. 
Whitfield,  R.  P.     1892.     Gastropoda  and  Cephalopoda  of  the  Raritan  Clays 

and  Greensand  Marls  of  New  Jersey :   New  Jersey  Geol.  Survey,  Pal.  Series 

No.  2.    Also,  1892,  U.  S.  Geol.  Survey  Mon.  18. 


334  The  Uni^^rsity  Science  Bulletin 


EXPLANATION  OF  PLATES 


PLATE  XXVIII 

Fig.  1.  Baculitefi  cotnpressus  var.  reesidei  Elias,  n.  var.  Side  view.  Natural 
size.  For  cross  section  and  suture  of  this  specimen  see  PI.  XXXIII,  figs. 
2b,  2c.  From  Lake  Creek  shale  member,  Pierre  formation,  center  sec.  35, 
T.  lis.,  R.  39  W.,  Wallace  county,  Kansas. 

Fig.  2.  Baculites  pseudovatiis  var.  A  Elias,  n.  var.  Side  view  of  holotype. 
Natural  size.  For  cross  section  and  suture  of  this  specimen  see  PI.  XXXIII, 
figs.  5a,  5b.  From  Baculites  zone  of  Salt  Grass  shale  member,  Pierre  forma- 
tion, E.  sec.  2,  T.  12  S.,  R.  42  W.,  Wallace  county,  Kansas. 

Fig.  3.  Baculites  compressus  var.  corrugatus  Elias,  n.  var.  Side  view  of  tj'pe 
specimen.  Natural  size.  For  cross  section  and  suture  of  this  specimen  see 
PI.  XXXII,  figs,  la,  lb,  Ic.  From  Lake  Creek  shale  member,  Pierre  formation, 
SEi/4  sec.  29-,  T.  11  S.,  R.  39  W.,  Wallace  county,  Kansas. 

Fig.  4.  Baculites  compressus  Say  (in  restricted  sense).  Side  view.  Natural 
size.  For  cross  section  and  suture  of  this  specimen  see  PI.  XXXII,  figs.  4a, 
4b,  4c.  From  basal  part  of  Lake  Creek  shale  member,  Pierre  formation, 
W.  sec.  7,  T.  12  S.,  R.  38  W.,  Wallace  county,  Kansas. 


Elias:   Cephalopods  of  Pierre  Formation 


335 


PLATE  XXVIII 


386  The  University  Science  Bulletin 


PLATE  XXIX 

Fi(;s.  la,  lb.  Baculites  pf^eudovatuR  Elias,  n.  sp.  Cotype.  Reduced  to  %. 
From  upper  Weskan  shale  member,  Pierre  formation,  NW^/i  sec.  18,  T.  13  S., 
R.  41  W.,  Wallace  county,  Kansas,  la,  side  view;  lb,  siphonal  view.  For 
cross  section  of  this  specimen  see  PI.  XXXIII,  fig.  4b. 

Fig.  2.  Baculites  pseudovatm  Elias,  n.  ,sp.  Cotype.  Natural  size.  From 
same  locality.  For  suture  and  cross  section  of  this  specimen  see  PI.  XXXIII, 
figs.  4a,  4b. 

Fig.  3.  ci.  Discoscaphites  con  strict  us  var.  temiistriatus  (Kner).  Natural  size. 
From  upper  half  of  Lake  Creek  shale  member,  Pierre  formation,  NE^/i  NE^A 
sec.  8,  T.  11  S.,  R.  38  W.,  Wallace  county,  Kansas. 


l<]iJAs:   Cephalopods  of  Pierre  Formation 


337 


PLATE  XXIX 


338  The  University  Science  Bulletin 


PLATE  XXX 

Fig.  1.  Bacidites  clinolobalus  Elias,  n.  sp.  Cotype.  Side  view.  Reduced 
75.  For  suture  and  cross  section  of  this  specimen  see  PI.  XXXIV,  figs.  2a,  2b. 
From  Beecher  Island  shale  member,  Pierre  formation,  one-half  mile  northeast 
of  Beecher  Island,  Yuma  county,  Colorado. 

F^G.  2.  Baculites  clinolobatus  Elias,  n.  sp.  Cotype.  Side  view.  Reduced  %. 
The  fragment  represents  younger  stage  of  the  species.  Found  close  to  the 
specimen  illustrated  on  figure  1. 

Fig.  3.  Baculites  compressus  var.  corrugatus  Elias,  n.  var.  Side  of  non- 
septate  portion  of  a  large  individual.  Reduced  %.  From  upper  part  of  Lake 
Creek  shale  member,  Pierre  formation,  NW%  SW^^  sec.  5,  T.  13  S.,  R.  41  W., 
Wallace  county,  Kansas. 

Figs.  4a,  4b.  Baculites  meeki,  n.  sp.  Holotype.  Natural  size.  From 
Beecher  Island  shale  member,  Pierre  formation,  2  miles  northwest  of  Beecher 
Island,  Yuma  county,  Colorado.    4a,  side  view;  4b,  cross  section. 


Elias:  Cephalopods  of  Pierre  Formation 


339 


PLATE  XXX 


340  The  University  Science  Bulletin 


PLATE  XXXI 

Fi(is.  la,  lb.  Baculites  grandis  Hall  and  Meek.  Reduced  %.  From  Beecher 
Island  shale  member,  Pierre  formation,  1  mile  northwest  of  Beecher  Island, 
Yuma  county,  Colorado,    la,  side  view;  lb,  siphonal  view. 

Figs.  2a,  2b.  Baculites  grandis,  Hall  and  Meek.  Reduced  Va.  From  Beecher 
Island  shale  member,  Pierre  formation,  Hackberry  creek,  Cheyenne  county, 
Kansas.  2a,  side  view;  2b,  antisiphonal  view.  For  sutures  and  cross  section 
of  this  specimen  see  PI.  XXXIV,  figs.  5a,  5b,  5c. 

Fig.  3.  Bacidites  compressvs  cf.  var.  reesidei  Elias,  n.  var.  Reduced  %. 
Side  view.  From  Lake  Creek  shale  member.  Pierre  formation,  center  sec.  35, 
T.  lis.,  R.  39  W.,  Wallace  county,  Kansas. 


Elias:  Cephalopods  of  Pierre  Formation 


341 


PLATE  XXXI 


23-  3482 


342  The  University  Science  Bulletin 


PLATE  XXXII 

Figs,  la,  lb,  Ic.  Baculites  compressus  var.  corrugatus  Elias,  n.  var.  Holo- 
type.  Natural  size.  From  Lake  Creek  shale  member,  Pierre  formation,  SE% 
sec.  29,  T.  US.,  R.  39  W.,  Wallace  county,  Kansas,  la,  sutures;  lb,  siphonal 
margin;  Ic,  cross  section. 

Figs.  2a,  2b,  2c.  Baculites  compressus  var.  reesidei  Elias,  n.  var.  Type. 
Natural  size.  From  Eagle  sandstone  in  sec.  34,  T.  16  N.,  R.  28  E.,  Fergus 
county,  Montana.  After  J.  B.  Reeside,  1927,  PI.  9,  figs.  1.  5.  2a,  suture;  2b, 
siphonal  margin  (modified  after  Reeside's  photograph) ;  2c,  cross  sections. 

Figs.  3a,  3b.  Baculites  covipressus  Say  (in  restricted  sense).  From  Pierre 
formation  in  Upper  Missouri  exposures.  After  F.  B.  Meek,  1876,  PI.  20,  figs. 
30a,  30c.  Suture  reduced  by  the  writer  to  suit  the  cross  section,  which  is 
natural  size. 

Figs.  4a,  4b,  4c.  Baculites  compressus  Say  (in  restricted  sense).  Natural 
size.  From  basal  part  of  Lake  Creek  shale  member,  Pierre  formation,  W. 
sec.  7,  T.  12  S.,  R.  38  W.,  Wallace  county,  Kansas.  4a,  suture;  4b,  siphonal 
margin;  4c,  cross  section. 

Figs.  5a,  5b.  Baculites  com])rrssus  Say  (in  restricted  sense).  Natural  size. 
From  upper  part  of  Lake  Creek  shale  member,  Pierre  formation,  in  NW^/4 
SWi/4  sec.  5,  T.  13  S.,  R.  41  W.,  Wallace  county,  Kansas.  5a.  suture;  5b,  cross 
section. 


Elias:  Cephalopods  of  Pierre  Formation  343 


PLATE  XXXII 


Living  chamber 


40  mm. 


Living  chamber 


4a 


f^-r^ 


30  mm. 


4b 


344  The  University  Science  Bulletin 


PLATE  XXXIII 

Figs,  la,  lb.  Baculites  compressus  cf.  var.  reesidei  Elias,  n.  var.  Natural 
size.  From  Lake  Creek  shale  member,  Pierre  forniation,  center  sec.  35,  T.  11  S., 
R.  39  W.,  Wallace  county,  Kansas,    la,  suture;  lb,  cross  section. 

Figs.  2a,  2b,  2c.  Baculites  compressus  var.  reesidei  Elias,  n.  var.  Natural 
size.  From  Lake  Creek  shale  member,  Pierre  formation,  center  sec.  35,  T.  11  S., 
R.  39  W.,  Wallace  county,  Kansas.  2a,  sutures;  2b,  siphonal  margin;  2c,  cross 
section. 

Figs.  3a,  3b,  3c.  Baculites  ovatus  Say.  Natural  size.  3a,  3b  from  Eagle 
sandstone,  near  top,  in  sec.  34,  T.  16  N.,  R.  28  E.,  Fergus  county,  Montana. 
3c  from  Elk  Basin  sandstone  member  of  Telegraph  Creek  formation  in  T. 
57  N.,  R.  98  W.,  Park  county,  Wyoming.  After  J.  B.  Reeside,  1927,  PI.  7, 
figs.  3,  5,  and  PI.  6,  fig.  3. 

Figs.  4a,  4b.  Baculites  pseudovatm  Elias,  n.  sp.  Cotypes.  Natural  size. 
From  upper  Weskan  shale  member,  Pierre  formation,  NW^/4  sec.  18,  T.  13  S., 
R.  41  W.,  Wallace  county,  Kansas.  4a,  suture;  4b,  cross  sections  of  two 
specimens. 

Figs.  5a,  5b.  Baculites  pseudovatus  var.  A  Elias,  n.  var.  Holotype.  From 
Baculites  zone  of  Salt  Grass  shale  member,  Pierre  formation,  E.  sec.  2,  T.  12  S. 
R.  42  W.,  Wallace  county,  Kansas.    5a,  suture;  5b,  cross  section. 


Elias:  Cephalopods  of  Pierre  Formation 
PLATE  XXXIII 


345 


Living  chamber 


30  mm. 


lb 


chamber  ^ij 


i 


346  The  University  Science  Bulletin 


PLATE  XXXIV 

Figs.  1,  2a,  2b.  Baculites  clinolobatus  Elias,  n.  sp.  Cotypes.  Natural  size. 
From  Beecher  Island  shale  member,  Pieri'e  formation,  one-half  mile  northeast 
of  Beecher  Island,  Yuma  county,  Colorado.  1,  sutures;  2a,  sutures  of  an- 
other specimen;  2b,  cross  sections  of  the  latter. 

Fig.  3.  Baculites  anceps  var.  Icopoliensis  Novak.  Natural  size.  Suture 
after  Novak,  1908,  p.  331,  fig.  10. 

Fig.  4.  Baculites  grandis  Hall  and  Meek.  Suture  of  large  individual. 
Natural  size.  From  Beecher  Island  shale  member,  Pierre  formation,  one-half 
mile  northeast  of  Beecher  Island,  Yuma  county,  Colorado. 

Figs.  5a,  5b,  5c.  Baculites  grandis  Hall  and  Meek.  Natural  size.  From 
Beecher  Island  shale  member,  Pierre  formation,  Hackberry  creek,  Cheyenne 
county,  Kansas.  5a,  5b,  sutures  of  younger  stages  of  the  shell;  5c,  cross 
sections  of  various  stages  of  growth  of  the  shell.  Note:  The  larger  cross 
section  belongs  to  the  specimen  illustrated  on  PI.  XXXI,  figs,  la,  lb. 


Elias:  Cephalopods  of  Pierre  Formation 
PLATE  XXXIV 


347 


348  The  University  Science  Bulletin 


PLATE  XXXV 

Figs,  la,  lb.  Bacvlites  cf.  pseiidovatus  Elias,  n.  sp.  Natural  size.  From 
Baculites  zone,  Salt  Grass  shale  member,  Pierre  formation,  SW^/4  sec.  6,  T. 
13  S.,  R.  41  W.,  Wallace  county,  Kansas,    la,  sutures;  lb,  cross  section. 

Figs.  2a,  2b.  Baculites  compressus  cf.  var.  reesidei,  Elias,  n.  var.  Natural 
size.  From  basal  part  of  Salt  Grass  shale  member,  Pierre  formation,  NW^/4 
NEi^  sec.  3,  T.  12  S.,  R.  42  W.,  Wallace  county,  Kansas. 

Figs.  3a,  3b.  Baculites  ovalus  var.  haresi  Reeside.  Natural  size.  From 
Cody  shale,  250  feet  below  top,  in  NEVi  SW^  sec.  5,  T.  50  N.,  R.  92  W.,  Big 
Horn  county,  Wyoming.    After  J.  B.  Reeside,  1927,  PI.  6,  figs.  9,  10. 

Figs.  4a,  4b.  Baculites  ovatus  var.  haresi  Reeside.  Natural  size.  From 
basal  part  of  Lake  Creek  shale  member,  Pierre  formation,  S.  sec.  15,  T.  12  S, 
R.  38  W.,  Wallace  county,  Kansas.    4a,  suture;  4b,  cross  section. 

Figs.  5a,  5b,  5c,  5d,  5e.  Baculites  compressus  cf.  var.  corrugatus  Elias,  n.  var. 
Different  stages  of  growth  of  a  young  shell.  From  basal  part  of  Salt  Grass 
shale  member,  Pierre  formation,  NW^i  NEi/4  sec.  3,  T.  12  S.,  R.  42  W.,  Wal- 
lace county,  Kansas.  5a,  cross  sections,  natural  size.  5b,  5c,  sutures,  natural 
size;  5d,  5e,  sutures,  magnified  3  times.  Note:  The  four  sutures  correspond 
to  the  four  cross  sections  of  figure  5a. 


Elias:  Cephalopods  of  Pierre  Formation 
PLATE  XXXV 


349 


350  The  University  Science  Bulletin 


PLATE  XXXVI 

Figs,  la,  lb,  Ic.  Scaphites  plenus  Meek.  J>ont,  side  and  rear  views.  Nat- 
ural size.    From  Pierre  formation  of  Black  Hills. 

Figs.  2a,  2b,  2c.  Scaphites  pJenus  Meek.  Rear,  side  and  front  views.  Nat- 
ural size.  From  basal  part  of  Salt  Grass  shale  member,  SW^/4  SW%  sec.  16, 
T.  13  S.,  R.  41  W..  Wallace  county,  Kansas. 

Figs.  3a,  3b,  3c.  Discoscaphiles  nicolleti  var.  saltgrassensis  Elias,  n.  var. 
Front,  rear  and  side  views.  Very  slightly  enlarged.  From  basal  part  of  Salt 
Grass  shale  member,  SE^  sec.  2,  T.  12  S.,  R.  42  W.,  Wallace  county,  Kansas. 


Elias:  Cephalopods  of  Pierre  Formation 


351 


PLATE  XXXVI 


352  The  University  Science  Bulletin 


PLATE  XXXVII 

Figs,  la,  lb,  Ic.  Scaphites  plenus  Meek.  Back,  side  and  front  views. 
Natural  size.    From  Pierre  shale  of  western  South  Dakota. 

Fig.  2.  Acanthoscaphites  nodosus  var.  hrevis  Meek.  Imperfect  specimen  in 
rock.  From  basal  part  of  Lake  Creek  shale  member,  Pierre  formation,  NW^ 
NEi/4  sec.  8,  T.  13  S.,  R.  41  W.,  Wallace  county,  Kansas. 

Fig.  3.  Acanthoscaphites  nodosus  var.  quadrangidaris.  Fragment  of  an 
adult  shell.  Natural  size.  Upper  part  of  Lake  Creek  shale  member,  sec.  12, 
T.  12  S.,  R.  39  W.,  Wallace  county,  Kansas. 


Elias:  Cephalopods  of  Pierre  Formation  353 

PLATE  XXXVII 


354  The  University  Science  Bulletin 


PLATE  XXXVIII 

Figs,  la,  lb,  2.  Acanthoscaphites  nodosus  Say  (in  restricted  sense).  Slightly- 
reduced.  From  upper  part  of  Upper  Weskan  shale  member,  Pierre  formation, 
NEi/4  sec.  8,  T.  13 S.,  R.  41  W.,  Wallace  county,  Kansas,  la,  side  view;  lb, 
rear  view;  2,  side  of  internal  volution  of  the  same  specimen. 

Fig.  3.  Acanthoscophites  nodosus  Say  (in  restricted  sense).  Slightly  reduced. 
From  middle  part  of  Lake  Creek  .shale  member,  Pierre  formation,  N.  sec.  13, 
T.  13S.,  R.41  W. 


Elias:  Cephalopods  of  Pierre  Formation  355 

PLATE  XXXVIII 


356  The  University  Science  Bulletin 


PLATE  XXXIX 

Figs,  la,  lb,  Ic.  Scaphites  plenus  Meek.  Internal  volution  of  the  specimen 
illustrated  on  PI.  X,  figs,  la,  lb,  Ic.    Magnified  4/3. 

Figs.  2-5,  6a,  6b,  7,  8.  Discoscaphites  abyssinus  (Morton).  Natural  size, 
except  fig.  5,  which  is  magnified  2  times.  From  Beecher  Island  shale  member, 
Pierre  formation.  2,  3,  5,  7,  8,  from  2  miles  northwest  of  Beecher  Island,  Yuma 
county,  Colorado.  4,  from  NE^/4  sec.  9,  T.  2  S.,  R.  42  W.,  Cheyenne  county, 
Kansas.    6a,  6b,  from  NE%  sec.  5,  T.  1  S.,  R.  39  W.,  Cheyenne  county,  Kansas. 

Figs.  9a,  9b.  Discoscaphites  conradi  cf.  var.  gulosus  (Morton).  From 
Beecher  Island  shale  member,  Pierre  formation,  2  miles  northwest  of  Beecher 
Island,  Yuma  county,  Colorado. 

Fig.  10.  Discoscaphites  cf.  constrictus  ,var.  tenuistriatus  (Kner).  Natural 
size.  From  Upper  Weskan  shale  member,  Pierre  formation,  center  sec.  18,  T. 
13  S.,  R.  40  W.,  Wallace  county,  Kansas. 


i']i,iA.s:  Cephalopods  of  Pierre  Formation  357 

PLATE  XXXIX 


6b 


'^'^^Mi 


24— 31S2 


358  The  University  Science  Bulletin 


PLATE  XL 

Fig.  1.  Discoscaphites  nicolleti  var.  saltgrassensis  Elias,  n.  var.  Suture  of 
the  specimen  illustrated  on  PI.  XXXVI,  figs.  3a,  3b,  3c.    Magnified  3  times. 

Fig.  2.  Discoscaphites  abyssinus  Morton.  Suture  of  the  specimen  illustrated 
on  PI.  XXXIX,  fig.  5.    Magnified  2  times. 

Fig.  3.   Scaphites  plenus  Meek.     Suture  of  the  specimen  illustrated  on  PI. 

XXXVI,  figs,  la,  lb,  Ic.    Slightly  reduced. 

Fig.  4.   Scaphites  plenus  Meek.     Suture  of  the  specimen  illustrated  on  PI. 

XXXVII,  figs,  la,  lb,  Ic.    Slightly  reduced. 

Fig.  5.  Scaphites  plenus  Meek.  Suture  of  the  specimen  illustrated  on  PI. 
XXXVI,  figs.  2a,  2b,  2c.    Magnified  3  times. 

Fig.  6.  Scaphites  plenus  Meek.  Suture  of  the  holotype  after  Meek.  Prob- 
ably somewhat  magnified. 

FtG.  7.  Schematic  sketch  of  a  suture  of  Baculites  compressus  and  allied 
species.  SL,  Siphonal  lobe;  IL,  1st  lateral  lobe;  2L,  2d  lateral  lobe;  AL,  anti- 
siphonal  lobe;  SS,  siphonal  saddle;  IS,  1st  lateral  saddle;  2S,  2d  lateral  saddle; 
3S,  3d  lateral  saddle;  a,  stemlike  body  supporting  the  terminal  branches  of 
the  1st  lateral  lobe;  b,  median  saddle  of  the  siphonal  saddle. 


Elias:  Cephalopods  of  Pierre  Formation  359 


PLATE  Xr> 


360  The  University  Science  Bulletin 


PLATE  XLI 

Figs,  la,  lb,  2.  Placenticeras  meeki  Boehm.  Reduced  M2-  From  basal  part 
of  Upper  Weskan  shale  member,  Pierre  formation,  SW^/i  NW^/4  sec.  18,  T.  13 
S.,  R.  41  W.,  Wallace  county,  Kansas,  la,  lb,  side  and  dorsal  veiws  of  septate 
volution.    2,  side  view  of  inner  volution. 

Fm.  3.  Acanthoscaphites  nodosus  var.  brevis  Meek.  Reduced  %.  Side  view. 
From  lower  part  of  Lake  Creek  shale  member,  Pierre  formation,  SEi/4  sec.  33, 
r    11  S  ,  R.  39  W..  Wail;  re  cruntv,  Kan.^as. 


Elias:  Cephalopods  of  Pierre  Formation 
PLATE  XLI 


361 


•^r^v 


8G2  The  University  Science  Bulletin 


PLATE  XLII 

Figs,  la,  lb.  Placenticeras  meeki  Boehm.  Part  of  living  chamber  which 
overlaps  umbilicus.  Reduced  Vi.  From  basal  part  of  Upper  Weskan  shale 
member,  Pierre  formation,  SWV^  NW^/i  sec.  18,  T.  13  S.,  R.  41  W.,  Wallace 
county,  Kansas,  la,  side  view;  lb,  dorsal  view.  The  septate  volution  of  this 
specimen,  which  is  here  overlapped  by  living  chamber,  is  illustrated  on  PI. 
XLI,  figs,  la,  lb. 

Fig.  2.  Heteroceras  torlum  Meek  and  Hayden.  Natural  size.  From  top 
of  Sharon  Springs  shale  member,  Pierre  formation,  sec.  2,  T.  14  S.,  R.  40  W , 
Wallace  county,  Kansas. 


Elias:  Cephalopods  of  Pierre  Formation  363 

PLATE  XL  1 1 


THE  UNIVERSITY  OF  KANSAS 

SCIENCE  BULLETIN 

Vol.  XXL]  March,  1933.  |  No.  10. 


A  New  Megasecopteron  from  the  Carboniferous  of 

Kansas 


By  F.  M.  CARPENTER 


Abstract:  The  order  Megasecoptera  has  previously  been  represented  in  the 
American  Carboniferous  by  only  two  fragmentary  wings  of  doubtful  affinities. 
A  new  fossil,  Parabrodia  carboniara,  n.  sp.,  found  in  the  Upper  Carboniferous 
of  Garnett,  Kan.,  and  belonging  to  a  new  family,  ParabrodiidsB,  is  a  typical 
member  of  the  order,  closely  related  to  the  European  Brodia.  The  type  is  in 
the  Geological  Museum  of  the  University  of  Kansas. 


'T^HE  Carboniferous  members  of  the  extinct  order  Megasecoptera 
-*-  are  Icnown  almost  exclusively  from  European  deposits.  Only 
two  fossils  ^  from  the  North  American  Carboniferous  can  be  fitted 
into  the  order,  and  both  of  these  are  so  fragmentary  and  aberrant 
that  they  were  placed  there  only  tentatively  by  Handlirsch.  But 
a  new  fossil,  recently  found  by  M.  K.  Elias  in  the  Carboniferous 
of  Kansas,  consists  of  a  nearly  complete  wing  typical  of  the  Euro- 
pean Megasecoptera  of  this  horizon.  To  Mr.  Elias  and  the  author- 
ities of  the  Geological  Department  of  the  University  of  Kansas  I 
am  indebted  for  the  opportunity  of  studying  this  insect. 

As  we  might  expect  from  its  unique  geographical  position,  the 
fossil  belongs  to  a  new  family: 

PARABRODIID^,  New  Family 

Allied  to  the  family  Brodiidae,  from  the  English  Carboniferous 
(Westphalian) ,  but  differing  in  the  possession  of  a  definite  coales- 
cence of  MA  with  Rs. 

1.  Raphidlopsis  diversipenva  Scud.,  from  Rhode  Island,  and  ProchoTcyptera  calopteryx 
Ilandl.,  from  Mazon  creek,  Illinois. 

(365) 


366 


The  University  Science  Bulletin 


PARABRODIA,  New  Genus 

Medium-sized  insects;  wings  slender,  probably  subpetiolate ;  Sc 
extending  well  beyond  the  middle  of  the  wing;  Rs  arising  about  the 
middle  of  the  wing,  almost  immediately  coalescing  with  MA;  Rs 
2-branched;  M  free  from  R  at  the  base  of  the  wing;  MA  un- 
branched;  MP  forked;  Cul  and  Cu2  unbranched;  Cul  probably 
coalescing  with  M  at  the  base  of  the  wing. 

Genotype:    Parabrodia  carbonaria  n.  sp. 

Parabrodia  carbonaria  n.  sp. 

(Figure  ]) 

Length  of  wing,  24mm.;  width,  7  mm.;  apex  pointed;  Sc  straight; 
R  with  a  slight  bend  at  the  origin  of  Rs ;  Rl  remote  from  the  margin 
of  the  wing  distally ;  MA  fused  with  Rs  for  a  distance  about  as  long 
as  its  free  part  proximad  of  the  coalescence;  MP  diverging  apically 
of  the  origin  of  Rs  and  proximad  of  the  divergence  of  MA  from  Rs ; 
Cul,  Cu2,  and  lA  nearly  parallel.    No  cross  veins  are  preserved. 

Holotype.  In  Geological  Museum,  University  of  Kansas;  col- 
lected by  M.  K.  Elias,  six  miles  northwest  of  Gamett,  Kan.^  Horizon 
is  the  late  middle  Pennsylvanian. 


Text  Figure  1 

The  wing  is  preserved  on  a  large  slab  of  light-brown  shale,  con- 
taining remains  of  Walchia  and  other  plants,  one  of  which  overlies 
a  part  of  the  anterior  margin.  The  veins  are  only  faintly  preserved, 
and  in  making  the  accompanying  figure  I  have  used  to  advantage 
an  excellent  photograph  provided  by  Mr.  Elias. 

As  I  have  mentioned  above,  Parabrodia  is  especially  interesting 
because  it  is  the  first  typical  Mcgasecopteron  to  be  found  in  a  satis- 
factory state  of  preservation  in  the  Carboniferous  of  North  America. 

2.  The  locality  is  mentioned  in  the  abstract  "Conifer  Forest  of  Late  Middle  Pennsylvania 
Time,"  by  Mr.  Elias,  published  in  the  Proceedings  of  the  Paleontological  Society:  Bull.  Geol. 
Soc.   Amer.,   43:28,');    1932. 


Carpenter:  A  New  Megasecopteron  367 

The  shape  of  the  wing  is  closest  to  that  of  the  members  of  the  family 
Brodiidae,  but  in  the  latter  the  anterior  media  (MA)  does  not  quite 
coalesce  with  Rs,  and  the  radial  sector  (Rs)  has  five  branches.  If 
my  interpretation  of  the  formation  of  Rs,  i.  e.,  its  division  into 
R2  -f  3  and  R4  -f-  5,  is  correct,  then  the  structure  of  the  vein  ap- 
proaches that  of  Raphidiopsis  and  the  Permian  Protohymenoptera. 
But,  unfortunately,  the  convexity  and  concavity  of  the  veins  are  not 
preserved,  and  it  is  possible  that  the  vein  which  I  have  named  MA 
really  consists  of  a  compound  vein  formed  by  the  fusion  of  MA  and 
R4  -f  5,  and  that  the  independent  branches  of  Rs  are  R2  and  R3.  In 
either  case  the  venation  as  a  whole  could  easily  be  derived  from 
that  of  the  Brodiidae  or  some  closely  related  family. 


THE  UNIVERSITY  OP  KANSAS 

SCIENCE  BULLETIN 

Vol.  XXI. 1  March,  1933.  INo.  II 


Comparative  Anatomical  Research  Within  the  Genus 

Ribes  * 


JAMES  C.  BATES 


Abstract:  The  results  obtained  from  a  detailed  anatomical  and  histological 
study  of  fifteen  species  of  Ribes  are  summarized  and  some  conclusions  drawn 
as  to  their  probable  genealogical  relationships. 

The  species  showed  considerable  differences  in  the  following  features: 
Amount  and  distribution  of  trichomes  of  the  leaf  epidermis;  absence  or 
presence  of  glands  and  kinds  of  glands;  shape  of  leaf  epidennal  cells;  shape 
of  leaf  tip;  margin  and  midrib;  number  of  layers  of  palisade  cells;  number 
of  vascular  bundles  entering  the  petiole;  presence  or  absence  of  trichomes  of 
the  stem  epidermis;  presence  or  absence  of  bast  fibers  in  the  petiole;  stem 
armed  or  unarmed;  shape  and  size  of  stem  epidermal  cells;  cutinization  of 
radial  walls  of  stem  epidermal  cells;  kinds  of  collenchyma;  number  of  rings 
of  peridermal  tissue  in  the  two  years'  growth;  number  of  protoxylem  points; 
length  of  xylem  elements;  and  the  distribution  of  starch,  tannin  and  cluster 
crystals  of  calcium  oxalate  in  the  tissues  of  the  stem  and  leaf. 

Suggestions  as  to  the  genealogical  relationships  of  the  fifteen  species  are 
based  on  their  similarities  and  differences  in  anatomical  structure  and  cell 
contents.  Some  of  the  highest  degrees  of  anatomical  relationship  foimd  were 
between  American  and  European  species,  and  some  of  the  lowest  between  some; 
American  species  and  between  American  and  European  species. 

The  paper  includes  8  tables,  five  plates  and  1  figure. 


INTRODUCTION 

'T^HE  writer  has  attempted  to  summarize  the  results  obtained 
■*-  from  a  detailed  anatomical  study  of  the  following  fifteen 
species  of  Ribes,  and  to  offer  a  few  suggestions  as  to  their  probable 
genealogical  relationships:  R.  aureum  Pursh.,  R.  alpinum  L.,  R. 
americanum  Mill.,  R.  cynobasti  L.,  R.  diacantha  Pall,  R.  fascicii- 
latum  Sieb.  &  Zucc,  R.  hirtellum  Michx.,  R.  holosericeum  Otto  & 

*  This  work  was  carried  out  in  the  Botany  Department  of  the  University  of  Kansas  under 
the  direction  of  Prof.  W.  C.  Stevens.  The  writer  desires  to  make  his  appreciative  acknowl- 
edgment to  Professor  Stevens  for  his  help  and  encouragement  during  the  preparation  of  the 
work    here   presented. 

(369) 


370  The  University  Science  Bulletin 

Dietr.,  R.  luridum  Hook.  f.  &  Thorns.,  R.  missouriense  Nutt.,  R. 
Nigrum  L.,  R.  sativum  Syme.,  R.  stenocarpum  Maxim.,  R.  tenue 
Jancz.,  and  R.  vilmorinii  Jancz. 

These  species,  brought  together  from  their  native  places  in  differ- 
ent parts  of  the  earth,  were  growing  in  the  Arnold  Arboretum, 
Boston,  and  the  differences  which  they  show  are  therefore  inherent 
and  not  due  to  the  influence  of  different  environments.  This  is  con- 
sidered a  vital  fact  for  this  research. 

SUMMARY  OF  PUBLISHED  ANATOMICAL  RESEARCH 

A  comparatively  small  amount  of  anatomical  work  on  Rihes  has 
been  published  by  Petit,  Reinke,  and  Holle,  and  summarized  by 
Solereder  (1908)  in  his  "Systematic  Anatomy  of  the  Dicotyledons," 
Vol.  I. 

According  to  Holle  (1893)  the  cuticle  is  thin  in  all  species  of  Rihes 
examined.    The  hairs  are  long,  simple,  and  conically  pointed. 

Solereder  claims  the  unicellular  trichomes  are  not  calcified  and 
that  external  glands  have  been  observed  only  on  the  leaves  of  the 
genus.  The  glandular  trichomes  on  the  leaf  and  stipules  of  Rihes, 
according  to  Solereder,  are  shaggy  hairs  with  a  multiseriate  stalk  of 
variable  length  and  a  spherical  or  peltate  head,  the  center  of  which 
is  occupied  by  isodiametric  cells,  while  the  epidermal  cells  are  elon- 
gated radially  like  a  palisade. 

According  to  Holle  the  glandular  hairs  differ  in  various  species 
and  are  divided  by  him  into  two  classes:  (a)  Wedge-formed  glandu- 
lar hairs  with  a  rounded  multicellular  head  and  a  rather  long, 
several-celled  stalk.  In  this  group  he  includes  R.  alpinum,  R. 
aureum,  R.  cynobasti,  and  R.  fasciculatum.  (h)  Sessile  glands  with 
a  relatively  large  multicellular  head  on  a  short  multicellular  stalk 
submerged  in  the  epidermis.  In  this  group  he  includes  R.  ameri- 
canum  and  R.  nigrum. 

The  spines  of  the  species  of  Rihes  are,  according  to  Delbrouck 
(1875),  essentially  periblem  structures. 

Occasionally  the  leaf-teeth  in  species  of  Rihes  have  a  glandular 
function,  according  to  Reinke  (1876). 

Holle  (1893)  attributes  the  following  features  to  Rihes:  Epi- 
dermal cells  with  mostly  straight  side  walls,  as  seen  in  surface  view; 
the  possession  of  fairly  small  pairs  of  guard  cells  of  nearly  circular 
outline  and  always  surrounded  by  many  associate  cells;  from  one  to 
three  layers  of  palisade  cells;  the  larger  vascular  bundles  in  the 
leaf  with  mostly  thin-walled  strengthening  tissue  extending  to  the 


Bates:  Study  of  the  Genus  Ribes  371 

epidermis  of  both  surfaces ;  and  isolated  sclerenchyma  fibers  entirely 
lacking  in  both  leaf  and  stem. 

According  to  Petit  (1887),  all  investigated  species  of  Ribes  have 
three  vascular  bundles  pass  into  the  petiole. 

The  following  account  of  conditions  found  in  the  cortex,  phloem 
and  xylem  are  given  by  Holle. 

Cortex.  The  two  to  three  outer  layers  of  the  primary  cortex  are 
thickened  collenchymatously,  the  walls  of  the  outer  layer  becoming 
thickened  all  around.  Aside  from  collenchyma  this  genus  is  lack- 
ing in  all  sclerotic  elements  in  both  primary  and  secondary  cortex, 
unless  some  walls  of  the  cork  cells  become  sclerotic.  Very  charac- 
teristic also  are  the  tangential  rings  and  bands  of  cells,  radially  one 
cell  broad,  bearing  small  rosette  crystals.  Also  in  the  primary  cortex 
and  pith  of  this  genus  are  rosette  crystals  of  considerably  larger  size. 
The  contents  of  the  cortical  parenchyma  cells  are  often  brown  in 
color  and  possessed  of  tannin,  especially  in  the  outer  cortex  layers 
of  Ribes.  Cork  originates  in  the  innermost  part  of  the  cortex,  and 
the  cork  cells  are  somewhat  radially  elongated. 

Phloem.  In  the  phloem  are  one  to  four  regular  transverse  rows  of 
contiguous,  small  cluster  crystals,  the  number  of  rows  being  charac- 
teristic of  the  species. 

Xylem.  The  medullary  rays  reach  as  high  as  seven  cells  in 
breadth.  The  elements  of  the  tracheal  tubes  have  in  the  end  walls 
scalariform  perforations  with  few  thick  strips.  The  wall  of  a  vessel 
where  it  is  in  contact  with  the  parenchyma  of  the  medullary  rays 
bears  bordered  pits  only.  The  wood  prosenchyma,  which  mostly 
has  thick  walls,  bears  bordered  and  simple  pits.  The  lumen  of  the 
simple-pitted  wood  fibers  in  Ribes  is  septate. 

Holle  (1893)  divides  the  species  into  two  classes  and  gives  some 
of  their  distinguishing  characteristics  as  follows: 

(a)  With  Stalked  Glandular  Hairs.  R.  alpinum  L.  (Fursten- 
stein,  ex  Herb.  G.  A.  Pritzel.)  Epidermal  cells  with  undulating  side 
walls.  Palisade  tissue  one-rowed.  Cluster  crystals  in  the  palisade 
tissue.  Three  to  four  regular  transverse  rows  of  cluster  crystals  in 
the  phloem. 

R.  aureum  H.  (Hort.  Botan.  Monac.)  Upper  epidennal  cells  with 
straight  side  walls,  somewhat  bulging  outward.  Palisade  tissue  two- 
to  three-rowed.  Cluster  ciystals  few  in  the  leaf.  An  almost  con- 
tinuous row  of  cluster  crystals  in  the  phloem,  and  outside  this  are 
sometimes  groups  of  a  second  row. 

R.  cynobasti  L.  (Ohio;  Doctor  Frank.)    Epidermal  cell  with  un- 


372  The  University  Science  Bulletin 

dulating  side  walls.  Palisade  tissue  one  rowed.  Only  one,  often 
interrupted,  regular  transverse  row  of  cluster  crystals  in  the  phloi?ra. 

R.  fasciculatum  S.  et  Z.  Leg.  (Burger,  Japan.)  Four  to  five 
almost  continuous  rows  of  cluster  crystals  in  the  phloem. 

(6)  With  Sessile  Glandular  Hairs.  R.  americanum  Mill. 
(Americ.  Septentr,  Ehrhard,  Select.  14.)  Epidermal  cells  v^dth  un- 
dulated side  walls.  Stomata  oval.  Only  the  larger  vascular  bundles 
are  extended  from  surface  to  surface  by  weak  strengthening  tissues. 
Palisade  tissue  in  one  row.  Spongy  parenchyma  compact.  One  to 
two  almost  uninterrupted  rows  of  small  cluster  crystals  in  the 
phloem. 

R.  nigrum  L.  (In  Silva  Caspat.  Comm.,  Kitaibel.)  This  is  dis- 
tinguished from  R.  multijloram  in  that  only  isolated  groups  make 
up  the  two  rows  of  cluster  crystals  in  the  phloem. 

SUMMARIZED  DISCUSSION  OF  MICROCHEMICAL  TESTS 
IN  STEMS,  PETIOLES  AND  MIDRIBS 

The  distribution  of  starch,  tannin  and  cluster  crystals  of  calcium 
oxalate  in  the  tissues  of  the  stem,  petiole,  midrib  and  blade  are 
given  in  Tables  I,  II  and  III.  These  substances  are  represented 
by  the  symbols  s,  t  and  *,  respectively,  and  their  relative  abundance 
is  shown  by  the  number  of  symbols  occurring  together.  Thus:  "s" 
represents  a  small  amount  of  starch,  "ss"  a  moderate  amount,  and 
"sss"  an  abundance  of  starch.  The  figures  in  the  third  column  of 
Table  I  indicate  the  number  of  transverse  rows  of  cluster  crystals 
of  calcium  oxalate  in  the  phloem. 


Bates:  Study  of  the  Genus  Ribes 


373 


TABLE  I. — Showing  the  distribution  of  starch,  tannin  and  cluster  crystals  of  calcium  oxalate 

in  the  tissues  of  the  stem. 


Phello- 
derm. 

Inner 
Cortex. 

Phloem. 

Xylem 
rays. 

Xvlem 
Par. 

Pith. 

sss 

tt 

sss 

tt 

* 

sss 

tt 

*2-3 

ss 

ttt 

ss 

ttt 

*l-2 

ss 

ttt 

sss 

ttt 

*l-2 

sss 
ttt 
*l-3 

sss 

t 

*4-5 

sss 

ttt 

*3-4 

sss 

ttt 

*l-3 

s 

ttt 
*1 
tt 

*l-2 

s 
ttt 

* 

sss 

ttt 

*l-4 

s 
tt 

tt 

* 

ss 

ttt 

sss 

ttt 

ttt 

ss 

ttt 

* 

ss 

ttt 

sss 

ttt 
** 

ss 

ttt 

** 
sss 

t 

tt 

ss 
ttt 

ttt 

8 

ttt 

* 

sss 

ttt 

sss 

ttt 

ss 

ttt 

sss 

ttt 

sss 

ttt 

ttt 

«* 

R.  nigrum 

sss 

ttt 

sss 

ttt 

ttt 

** 

sss 
t 

sss 

t 

sss 

t 

sss 

tit 

sss 

ttt 

sss 

ttt 

s 

sss 

ttt 

* 

sss 

ttt 

sss 

ttt 

* 

s 

ttt 

ttt 

ttt 

* 

ttt 

sss 

ttt 

ttt 

* 

tt 

tt 

** 

s 

ttt 

t 

tt 

** 

s 

ttt 

s 

ttt 

s 

ttt 

* 

R.  missouriense 

sss 
t 

sss 

t 

** 

ss 
tt 

S 

sss 
tt 

** 

s 

t 

tt 

ttt 

* 

tt 

* 

ttt 
*l-2 

t 
*2-4 

ttt 

tt 

H.* 

t 

t 

*# 

25—3482 


374 


The  University  Science  Bulletin 


TABLE  U.— Showing  the  distribution  of  starch,  tannin  and  cluster  crystals  of  calcium  oxalate 
in  the  tissues  of  the  petiole. 


K.  holusericeum . 
R.  lenuf, 


Epidermis. 


R.  aureum . 


R.  diacanlha. . . . 
R.  americanum . 


R.  nigrum 

R.  fasciculatum . 
R.  alpinum .  .  .  . 


R.  sativum 

R.  luridum 

R.  stenocarpum. 
R.  mlmorinii .  .  . 
R.  missouriensp . 
R.  cynohasti . .  .  . 
R.  hirtellum. .  .  . 


Cortex. 


Phloem. 


Xylem. 


Bast. 


When  tannin  and  starch  are  present,  they  are  always  found  in  the 
phelloderm  and  pith  and  nearly  always  in  the  inner  cortex,  phloem, 
and  phloem  and  xylem  rays.  These  two  substances  are  quite  uni- 
formly distributed  throughout  the  pith  cylinder,  with  the  exception 
of  R.  missouriense,  which  has  a  larger  amount  of  starch  in  the 
peripheral  pith  cells.  These  two  substances,  when  both  present, 
are  usually  found  occurring  together  within  the  cell.  A  small 
amount  of  starch  is  found  in  the  xylem  parenchyma  cells  of  R. 
aurem,  R.  alpinum,  R.  missouriense,  and  a  relatively  large  amount 
in  R.  jasciculatum.  Cluster  crystals  of  calcium  oxalate  were  found 
scattered  throughout  the  cortex  and  pith,  and  in  regular  transverse 
rows  in  the  phloem  of  the  majority  of  species  studied.  Tannin  is 
always  present,  and  starch  is  found  in  all  species  except  R.  steno- 
carpum and  R.  hirtellum,  but  varies  in  amount  in  the  different 
species.  Table  III  shows  that  both  tannin  and  starch  may  be 
present  in  abundance;  either  may  be  in  excess  of  the  other,  or  tannin 
may  be  present  in  small  or  moderate  amount  with  starch  absent. 
Thus  the  different  species  may  be  grouped  according  to  their  tannin 
and  starch  content  as  follows:     (1)  Abundance  of  both  tannin  and 


Bates:  Study  of  the  Genvs  Ribes 


375 


TABLE  III. — Showing  the  distribution  of  starch,  tannin  and  cluster  crystals  of  calcium  oxalate 
in  the  tissues  of  the  midrib  and  blade. 


Midrib. 

Mesophyll. 

Epidermis. 

Paren- 
chyma. 

Phloem. 

Xylem. 

Palisade. 

Spongy. 

R.  holosericeum 

R.  tenue 

tt 
tt 

tt 

t 

8 
tt 

tt 

** 

R.  diacantha 

tt 
tt 
tt 
tt 

R.  americanum 

tt 
tt 

ss 

tt 

tt 

t 

t 

tt 

tt 

tt 

t 

t 

tt 
tt 

tt 

t 

tt 

tt 

tt 

t 

t 

R.  luridum . 

R.  stenocarpum .     . 

R.  vilmorinii 

R.  misfouriense 

R.  cynohasti 

R.  hirtellum 

starch,  R.  aureum,  R.  diacantha,  R.  americanum,  R.  nigrum,  R.  al- 
pinum,  and  R.  sativum;  (2)  abundance  of  tannin  and  a  moderate 
amount  of  starch,  R.  tenue;  (3)  abundance  of  tannin  and  a  small 
amount  of  starch,  R.  luridum,  R.  vilmorinii,  and  R.  cynohasti;  (4) 
moderate  amount  of  tannin  and  abundance  of  starch,  R.  holoseri- 
ceum and  R.  missouriense ;  (5)  small  amount  of  tannin  and  abun- 
dance of  starch,  R.  fasdculatum ;  (6)  a  small  to  moderate  amount  of 
tannin  with  starch  absent,  R.  stenocarpum  and  R.  hirtellum.  The 
distribution  of  starch  and  tannin  in  the  petiole  and  midrib  is  given 
in  Tables  II  and  III,  only  two  classes  of  frequency  being  indicated. 
Table  II  shows  for  the  petiole  that  starch  was  found  only  in  the 
cortex,  phloem,  and  xylem  of  four  species:  R.  aureum,  R.  ameri- 
canum, R.  nigrum,  and  R.  fasdculatum.  Tannin  is  always  present 
in  the  epidermis,  cortex,  phloem,  and  xylem  rays  and  usually  in 
the  bast  fibers,  the  amount  varying  somewhat  in  the  different  species. 
Relatively  small  amounts  of  tannin  were  found  in  R.  fasdculatum, 
R.  sativum,  R.  stenocarpum,  and  R.  vilmorinii.  The  remaining 
species  contained  relatively  large  amounts  of  tannin  in  the  phloem 
and  xylem;  however,  the  amount  was  small  in  the  epidermis,  cortex. 


376  The  University  Science  Bulletin 

and  bast  fibers  of  some  species.  Cluster  crystals  of  calcium  oxalate 
were  found  in  the  cortex  of  R.  tenue  and  R.  alpinum.  Starch  was 
found  in  the  midrib  and  mesophyll  of  R.  aureum  and  R.  fasdcula- 
tum.  All  of  the  tissues  of  the  leaf  contained  relatively  large  quanti- 
ties of  tannin.  Cluster  crystals  of  calcium  oxolate  were  found  in  the 
palisade  tissue  of  R.  holosericeum  and  R.  hirtellum. 

Tests  were  made,  also,  for  sugar,  glucosides,  protein,  volatile  oils, 
resins,  calcium  carbonate,  and  mucilage,  but  only  negative  results 

were  obtained. 

SUMMARY  OF  STEMS 

The  following  features  are  common  to  the  stems  of  all  of  the  fif- 
teen species  studied:  The  outer  wall  of  each  epidermal  cell  is  cuti- 
nized  and  projecting.  The  epidermal  cells  are  elongated  vertically, 
but  vary  considerably  in  the  amount  of  elongation  in  the  different 
species;  radial  walls  are  straight  and  pitted  (Plate  XLIII,  figs. 
14-28).  The  absence  of  both  endodermis  and  bast  fibers.  The 
formation  of  cork  within  the  innermost  part  of  the  cortex,  and  the 
sloughing  off  of  the  epidermis  and  outer  part  of  the  cortex  soon 
after  their  formation.  A  relatively  small  amount  of  phloem  con- 
sisting of  phloem  parenchyma,  sieve  tubes,  and  companion  cells. 
Protoxylem  points  project  into  the  pith  cylinder,  giving  it  a  stellate 
appearance  as  seen  in  cross  section  (Plate  XLIV).  The  protoxylem 
consists  of  tracheal  tubes  with  spirally  thickened  walls.  The 
metaxylem  is  composed  of  tracheal  vessels  with  bordered  pits  and  a 
small  amount  of  xylem  parenchyma.  The  secondary  xylem  has 
fiber  tracheids  and  wood  fibers,  in  addition  to  the  elements  found  in 
the  metaxylem,  and  its  elements  are  radially  arranged.  Xylem  ele- 
ments from  1-10  rows  wide  alternate  tangentially  with  xylem  rays 
from  1-7  cells  wide.  The  tracheal  vessels  have  oblique  cross  walls 
with  few  thick  strips,  reticulate  pitting,  and  scalariform  perfora- 
tions in  the  end  walls  of  their  elements.  The  xylem  rays  consist 
of  vertically  elongated  cells  with  the  vertically  longest  cells  occurring 
at  the  upper  and  lower  extremities  of  the  ray.  A  relatively  large 
cylinder  of  homogeneous  pith.  The  cells  at  the  center  of  the  cyl- 
inder have  larger  cross  diameters  and  relatively  thinner  walls  than 
those  bordering  the  xylem.  The  presence  of  tannin  in  the  phello- 
derm,  cortex,  phloem,  xylem  rays  and  pith. 

Trichomes  are  present  in  9  of  the  15  species  studied.  Those  with- 
out trichomes  are  R.  aureum,  R.  diacantha,  R.  alpinum,  R.  luridum, 
R.  missouriense,  and  R.  hirtellum.  There  is  considerable  variation 
in  amount  and  length  of  trichomes  in  the  various  species   (Plate 


Bates:  Study  of  the  Genus  Ribes  377 

XLIII,  figs.  1-6).  R.  holosericeum  and  R.  sativum  have  a  small 
amount  of  short  trichomes  with  an  average  length  of  .07  mm.  R. 
vilmorinii,  R.  americanum,  and  R.  tenue  have  a  moderate  amount 
with  an  average  length  of  .06,  .12,  and  .20  mm.,  respectively.  The 
remaining  four  species  have  a  dense  amount  with  an  average  length 
of  .04  mm.  in  R.  nigrum,  and  .16  mm.  in  R.  fasciculatum,  R.  luriduin, 
and  R.  cynobasti.  The  radial  walls  of  the  epidermal  cells  are  cutin- 
ized  in  all  except  the  following  six  species:  R.  tenue,  R.  sativum, 
R.  vilmorinii,  R.  missouriense,  R.  cynobasti,  and  R.  hirtellum.  Mul- 
ticellular sessile  glands  (Plate  XLIV,  fig.  3)  were  found  with  the 
epidermis  of  R.  americanum  and  R.  nigrum,  and  multicellular 
stalked  glands  with  the  epidermis  of  R.  alpinum  and  R.  cynobasti. 
According  to  Holle  (1893)  external  glands  have  been  observed  on 
the  leaves  only  of  this  genus. 

There  is  also  considerable  variation  in  the  vertical  elongation  of 
the  epidermal  cells,  which  ranges  from  very  slight  in  R.  americanum, 
R.  nigrum,  R.  cynobasti,  and  R.  hirtellum  to  considerable  elongation 
in  R.  fasciculatum,  R.  sativum,  and  R.  missouriense.  Typical  col- 
lenchyma  (Plate  XLIV,  Fig.  2)  was  found  lignified  in  R.  steno- 
carpum,  R.  missouriense,  and  R.  cynobasti.  The  cork  cells  are  tan- 
gentially  elongated  in  all  of  the  fifteen  species  except  R.  aureum, 
R.  americanum,  and  R.  luridum.  According  to  Solereder  (1908)  the 
cork  cells  are  somewhat  radially  elongated.  R.  missouriense  differs 
from  all  other  species  in  having  three  rings  of  peridermal  tissue  in 
the  two  years'  growth.  Table  IV  shows  the  number  of  protoxylem 
points,  diameter  of  pith  cylinder,  and  number  of  tracheal  tubes  per 
square  mm.  of  xylem  area  in  each  of  the  fifteen  species  studied.  The 
number  of  protoxylem  points  varies  from  6-16  in  the  different 
species.  There  is  also  considerable  variation  in  diameter  of  pith 
cylinders,  which  ranges  from  .43  mm.  to  1.92  mm.  There  is  little 
or  no  correlation  between  diameter  of  pith  cylinder  and  number  of 
protoxylem  points.  The  number  of  tracheal  tubes  per  square  mm. 
varies  from  690  in  R.  fasciculatum  and  R.  luridum  to  1725  in  R. 
americanum.  Table  V  shows  the  average  length  of  tracheal  ele- 
ments, tracheids,  and  wood  fibers  in  the  different  species.  The 
average  length  of  tracheal  elements  varies  from  .22  mm.  in  R.  sati- 
vum and  R.  hirtellum  to  .46  mm.  in  R.  holosericeum.  The  average 
length  of  tracheids  shows  a  similar  variation,  ranging  from  .21  mm. 
in  R.  diacantha,  R.  alpinum,  and  R.  cynobasti  to  .46  mm.  in  R.  hir- 
tellum. The  greatest  variation  occurs  in  the  length  of  wood  fibers, 
which  ranges  from  .22  mm.  in  R.  sativum  to  .55  mm.  in  R.  tenue. 


378 


The  University  Science  Bulletin 


TABLE  IV. — Showing    the    number    of    protoxylem    points,    diameter    of    pith    cylinder,    and 
number  of  tracheal  tubes  per  square  mm.  of  xylem  area. 


R.  Iiolosericeum . 

R.  tenue 

R.  aureum.  .  .  . 
R.  diacantha. . . 
R.  americanum 

R.  nigrum 

R.  fasciculaturn 
R.  alpinum. . . . 
R.  sativum .... 
R.  hiridum.  .  . 
R.  stenocarpum 
R.  vilmorinii .  . 
R.  missouriense 
R.  cynobasti .  .  . 
R.  hirtellum.  .  . 


Number 

protoxylem 

points. 


1.5 
11 
12 

8 
12 
12 

6 
16 
13 
13 
12 
13 
13 
16 
14 


Diameter 

pith 
cylinder. 


mm. 
1.20 

.52 

.43 

.90 

.50 

1.44 
.72 

1.20 

1.92 
.96 
.56 
.56 
.72 
OS 

1  44 


Number 

tracheal 

tubes. 


1,480 

1,380 

1,104 

1,025 

1,725 

725 

690 

828 

800 

690 

1,173 

828 

800 

1,035 

866 


TABLE  V. — Showing  the   average  length   in   millimeters   of   tracheal   elements,   tracheids,   and 
wood  fibers  in  the  different  species. 


R,  holosericeum . 

R.  tenue 

R.  aureum 

R.  dixcantha  .    . 
R.  americanum . 

R.  nigrum, 

R.  Jasciculatum . 

R.  alpinum 

R.  sativum  .... 
R.  luridum.  .  .  . 
R.  stenocarpum 
R.  vilmorinii.  .  . 
R.  missouriense . 
R.  cynobasti.  .  . 
R.  hirtellum.  .  .  . 


Average 
length  of 
tracheal 
element. 


mm 
0. 


4<) 
.35 
.26 
.37 
.31 
.36 
.28 
.25 
.22 
.41 
.32 
.25 
.30 
.37 
.22 


Average 

length 

tracheid. 


mm. 
0.43 

.32 

.20 

.21 

.22 

25 

.36 

.21 

.28 

.23 

.40 

.22 

.28 

.21 

.46 


Average 
length 
wood 
fiber. 


mm. 
0.41 

.55 

.27 

.47 

.27 

.40 

.37 

.30 

.22 

.28 

.37 

.46 

.41 

.50 

.32 


Bates:  Study  of  the  Genus  Ribes  379 

There  is  little  or  no  correlation  in  length  of  any  two  of  these  ele- 
ments in  all  of  the  different  species.  There  is  little  or  no  variation 
in  size  or  shape  of  the  xylem  parenchyma  cells  in  the  different 
species.  These  cells  measure  approximately  .025  mm.  in  cross 
diameter  and  .062  mm.  in  vertical  length.  There  is  considerable 
variation  in  the  amount  and  distribution  of  starch,  tannin  and 
cluster  crystals  of  calcium  oxalate  in  the  tissues  of  the  stem  as 
shown  in  Table  I. 

SUMMARY  OF  LEAVES 

The  leaves  of  all  of  the  fifteen  species  of  Ribes  studied  have  the 
following  features  in  common:  Three  to  five  lobed,  palmately  veined 
leaves  with  dentate  margins;  three  main  veins  extending  from  the 
base  of  the  blade  to  the  tips  of  the  lobes,  giving  off  numerous  side 
branches  (Plate  XLVII) ;  the  upper  epidermal  cells  (Plate  XLVI) 
the  larger  and  with  walls  thicker  than  those  of  the  lower  epidennis; 
the  outer  wall  of  the  epidermal  cells  of  the  blade  and  petiole  pro- 
jecting and  overlaid  by  a  thin  cuticle;  stomata  occurring  only  in 
the  lower  epidermis  in  all  species  except  R.  aureum;  no  bast  fibers 
in  the  leaf  blade ;  the  midrib  with  relatively  thin-walled  strengthen- 
ing tissue  extending  from  epidermis  to  epidermis;  from  1  to  3 
layers  of  palisade  cells  composing  from  %  to  I/2  of  the  mesophyll 
volume  (Plate  XLVI) ;  either  a  single  3-lobed  vascular  bundle  or 
3  separate  vascular  bundles  entering  the  petiole ;  the  lobes  or  bundles 
tending  to  fuse  on  passing  to  the  blade  (Plate  XLV,  figs.  16-30). 

Table  VI  shows  the  variation  in  average  length  of  leaves  and 
average  width  and  thickness  of  petioles  in  the  different  species.  The 
leaf  varies  in  width  from  less  than  14  its  length  in  R.  stenocarpum 
and  R.  diacantha  to  nearly  equal  its  length  in  R.  missouriense.  There 
is  a  fair  correlation  between  the  length  of  leaf,  width  of  leaf,  and 
length  of  petiole.  There  is  considerable  variation  in  the  thickness 
of  the  leaf  blade,  which  ranges  from  .12  mm.  in  R.  americanum  to 
.26  mm.  in  R.  diacantha.  The  variation  in  thickness  of  petioles 
ranges  from  .50  mm.  in  R.  vilmorinii  to  1.28  mm.  in  R.  sativum. 
(Plate  XLV,  figs.  16-30).  The  radial  walls  of  the  epidermal  cells 
(Plate  XLIII,  figs.  29-54)  are  slightly  undulated  in  all  except  the 
following  five  species:  R.  holosericeuni,  R.  aureum,  R.  diacantha, 
R.  stenocarpum,  and  R.  cynobasti.  All  have  trichomes  (Plate  XLIII, 
figs.  7-13)  except  R.  aureum  and  R.  diacantha;  however,  there  is 
considerable  variation  in  amount,  length,  and  distribution  of  tri- 
chomes in  the  different  species.  Trichomes  are  sparse  in  the  following 
species:    R.  holosericeum,  R.  tenue,  R.  nigrum,  R.  alpinum,  and  R. 


380 


The  University  Science  Bulletin 


TABLE  VL — .Sliowing  the  average  length  of  leaves,  width  and  thickness  of  blades,  and  length 

and  thickness  of  petioles. 


Leneth 
of 

leaf. 

Width 
of 

leaf. 

Thickness 

of 

blade. 

Lcntrth 

of 
petiole. 

Thickness 

of 

petiole. 

R.  holosrnreiim 

mm. 
53 

mm. 
40 

mm. 
0.24 

mm. 
17 

mm. 
0.92 

R.  tenue 

2.5.5 

17.5 

.2fi 

6 

.64 

R.  aureum 

40 

24.5 

.17 

IS 

.56 

R.  diacantha 

39.5 

14 

.26 

8 

.74 

R.  americnnum 

50 

37.5 

.12 

18 

.64 

67 . 5 

40 

.16 

23.5 

.90 

R.  fasciculatum 

42.5 

33 

.15 

12 

.68 

IS .  5 

12 

.24 

7 

.64 

R.  sativum    .  .  .  -  .     . 

5-i.5 

44   5 

.14 

16.5 

1.28 

R.  luridum 

55 

20 

.21 

21 

1.12 

R.  stenncarpum 

27  5 

10.5 

.20 

10.5 

.69 

R.  vilmorinii 

27,5 

19 

.16 

11.5 

.50 

R.  missouriense.  . 

31 

30 

.18 

10 

.96 

R.  cynobasti 

.37.5 

2f) .  5 

.17 

10 

.80 

32 

1.S.5 

.16 

16 

.64 

luridum;  moderate  in  number  in  R.  americanum  and  R.  hirtellum; 
and  abundant  in  the  remaining  species.  The  average  length  of 
trichomes  in  mm.  in  the  various  species  is  as  follows:  R.  americanum 
.05;  R.  alpinum,  R.  luridum,  and  R.  vilmorinii  .10;  R.  hirtellum 
and  R.  nigrum  .14  and  .16,  respectively;  R.  stenocarpum,  R.  tenue, 
R.  fasdcullatum,  R.  missouriense,  and  R.  cynobasti  .23,  .24,  .26,  .28 
and  .29,  respectively;  J^.  holosericeum  .35  and  R.  sativum  .36. 
Trichomes  are  found  on  both  the  upper  and  lower  epideiTais  of  the 
blade  and  on  the  petiole  of  the  following  species:  R.  holosericeum, 
R.  americanutn,  R.  nigrujn,  R.  fasciculatum,  R.  sativum,  R.  steno- 
carpum, and  R.  vilmorinii.  R.  alpinum  has  a  few  trichomes  mostly 
at  the  margin.  R.  luridum  has  a  small  amount  of  trichomes  with 
the  epidermis  of  the  petiole,  margin  and  upper  epidermis  of  the 
blade.  R.  missouriense  has  a  moderate  amount  with  the  epidermis 
of  the  petiole,  margin  and  upper  epidermis  of  the  blade.  R.  cyno- 
basti and  R.  hirtellum  have  trichomes  with  both  the  upper  and 
lower  epidermis  of  the  blade.  R.  tenue,  R.  americanum,  R.  mis- 
souriense, and  R.  cynobasti  have  multicellular  trichomes  and  R. 
sativum  multicellular,  branched  trichomes  along  tiie  borders  of  the 
groove  of  the  petiole.     There  are  multicellular  se.  sile  glands  with 


Bates:  Study  of  the  Genus  Ribes 


381 


the  epidermis  of  R.  americanum  and  R.  nigrum,  and  multicellular 
stalked  glands,  with  a  several-celled  stalk  with  the  epidermis  of 
R.  alpinum  and  R.  cynobasti.  Table  VII  shows  the  number  of 
stomata  and  palisade  cells  per  square  mm.  of  leaf  area  and  the 
average  length  of  palisade  cells  in  the  different  species.  There  is  a 
slight  correlation  between  the  number  of  stomata  and  palisade  cells 
per  square  mm.  of  leaf  area.  The  ratio  of  palisade  cells  to  stomata 
varies  from  14  to  1  in  R.  fasciculatum  to  30  to  1  in  R.  tenue.  The 
length  of  palisade  cells  varies  from  .03  mm.  in  R.  tenue  and  R. 
aureum  to  .06  mm.  in  R.  alpinum  (Plate  XLVI) .  It  is  interesting  to 
note  that  R.  tenue  and  R.  aureum,  with  the  highest  ratio  of  palisade 
cells  to  stomata,  have  the  shortest  palisade  cells.  The  number  of 
meshes  per  square  mm.  of  leaf  area  varies  from  12  in  R.  tenue  to 
35  in  R.  hirtellum.  The  number  of  veinlet  ends  terminating  within 
the  mesophyll  per  square  mm.  of  leaf  area  varies  from  5  in  72.  tenue 
and  R.  alpinum  to  26  in  R.  hirtellum  and  R.  vilmorinii.  There  is 
some  correlation  between  the  number  of  meshes  and  veinlet  ends 
in  the  different  species.  There  is  considerable  variation  in  size  and 
shape  of  the  midribs  and  vascular  bundles  as  seen  in  cross  sections 
(Plate  XLV,  figs.  1-15).     The  midribs  vary  in  thickness  from  .24 


table  Vll.-^Showing  the  number  of  stomata  and  palisade  cells  per  square  mm.  of  leaf  area 
and  the  average  length  of  palisade  cells. 


Number  of 
stomata. 

Number 

of 

palisade 

cells. 

Average 

lensrth  of 

palisade 

cells. 

342 

o .  635 

0,05 

207 

24  .T 

6,210 
6,831 

.03 

R.  aureum 

.03 

R.  diacanlhn 

201 

3,726 

.05 

432 
276 

6,240 
.5,. ^80 

.01 

R.  nigrum 

.01 

R.  jascicuhtum ... 

242 

3,405 

.04 

R.  alpinum ... 

207 

5,000 

.06 

208 
276 

3,033 

5 ,  589 

04 

R.  luridum 

.  055 

R.  stennrarpum ■  . 

2.50 

4,96/ 

.05 

R.  Tilmorinii . 

414 

6,240 

.04 

R.  mtssourievse .  .  .                                                               .    . 

167 

3,519 

.04 

R.  cynohasH 

208 

5,. 589 

.04 

R.  hirtellum 

310 

5,616 

.04 

382  The  University  Science  Bulletin 

mm.  in  R.  aureum,  R.  diacantha,  R.  americanum,  and  R.  steno- 
carpum  to  .54  mm.  in  R.  nigrum.  The  upper  surface  of  the  midrib 
varies  in  shape  from  nearly  flat  to  decidedly  V-shaped,  with  the 
lower  surface  rounded.  The  midrib  with  relatively  thin-walled 
strengthening  tissue  extends  from  epidermis  to  epidermis.  The  leaf 
margin  is  rounded  in  all  species  except  R.  missouriense,  R.  vilmorinii 
and  R.  cynobasti,  which  have  pointed  margins,  as  seen  in  cross 
section  (Plate  XLVI).  The  palisade  cells  extend  to  the  margin 
in  all  species  except  R.  holosericeum.  In  most  species,  cells  the 
size  and  shape  of  the  upper  epidermal  cells  extend  around  to  the 
lower  surface  where  there  is  an  abrupt  transition  in  size  and  shape 
to  that  of  the  typical  lower  epidennal  cells.  There  is  considerable 
variation  in  size  and  shape  of  petioles  in  the  different  species  (Plate 
XLV,  figs.  16-30).  The  upper  surface  of  the  petiole  is  concave  in 
most  species  and  ranges  from  nearly  flat  to  distinctly  V-shaped  with 
the  lower  surface  rounded.  The  upper  surface  is  slightly  convex 
in  R.  hirtellum,  R.  diacantha,  R.  missouriense,  and  R.  aureum. 
The  petiole  is  triangular  in  outline  in  R.  nigrum  and  inverted  dome- 
shaped  in  R.  sativum.  Bast  fibers  bordering  the  outer  edge  of  the 
phloem  were  found  in  all  species  except  R.  alpinum  and  R.  steno- 
carpum,  the  number  of  cell  layers  varying  from  1  to  6  among  the 
different  species  (Plate  XLV,  figs.  16-30).  There  are  3  separate 
vascular  bundles  entering  the  base  of  the  petiole  in  all  species 
except  R.  americanum,  R.  stenocarpum,  and  R.  missouriense,  which 
have  a  single  3-lobed  bundle.  The  lobes  of  the  single  bundle  tend 
to  disappear  on  passing  to  the  blade,  and  the  three  separate  bundles 
fuse  into  one  in  all  species  except  R.  diacantha  and  R.  sativum. 
There  is  some  variation  in  the  rapidity  and  amount  of  fusion  of 
vascular  bundles  in  the  different  species. 

KEY  TO  STEMS 

I.    CoUenchyma  typical. 

A.  More  than   one   line  of  peridernial   tissue  in   2   years'    growth,   st^'ni  unarmed. 

R.   missouriense. 

B.  One  rjne  of  peridernial   tissue  in  2   years'  growth,   stem  armed. 

1.  Radial  walls  of  the  epidermal  cells  cutinized R.   stenocar-pum. 

2.  Radial  walls  of  the  epidermal  cells  not  cutinized R.  cynobasti. 

II.    CoUenchyma  not  typical. 

A.    Trichomes  present. 

1.    Radial  walls  of  the  epidermal  cells  cutinized. 

a.  Multicellular,  sessile  glands  present. 

(1)  Average  cross  diameter  of  tracheal  tubes  less  than  .025  mm. .  .i?.   americanum. 

(2)  Average  cross  diameter  of  tracheal  tubes  more  than  .025  mm....R.  nigrum. 

b.  Glands  absent. 

(1)  Average  less  than  865  tracheal  tubes  per  square  mm.  of  xylem  area. 

R.  fasciculatum. 

(2)  Average  more  than  865  tracheal  tubes  per  square  mm.  of  xylem  area. 

R.  holosericeum. 


Bates:  Study  of  the  Genus  Ribes  383 

2.     Radial  walls  of  epidermal  cells  not   cutinizfd. 

a.  Average  more  than  1050  tracheal  tubes  per  square  nan.  of  xylem  area.  .Ji.  tenue. 

b.  Average  less  than  1050  tracheal  tubes  per  square  mm.  of  xylem  area. 

(1)  Peridermal  tissue  fonned  at  outer  edge  of  phloem R.  vilmorinii. 

(2)  Peridermal  tissue  formed  in  midst  of  cortex R.  sativum. 

B.    Trichomes  absent. 

1.  Radial  walls  of  epidermal  cells  cutinized. 

a.    Outer  wall  of  epidermal  cells  hardly  at  all   projecting R.   alpinum. 

h.    Outer  wall   of   epidermal   cells   decidedly   projecting. 

(1)  Largest  tracheal  tubes  less  than   .022  mm.   in  cross  diameter. .  .fl.  diacantha. 

(2)  Largest  tracheal  tubes  greater  than  .022  mm.  in  cross  diameter.  .R.  luridum. 

2.  Radial  walls  of  epidermal  cells  not  cutinized. 

a.  Less  than  13  protoxylem  points  in  stem  cross  section R.  aureum. 

b.  More  than  13  protoxylem  points  in  stem  cross  section R.  hirtellum. 

KEY  TO  PETIOLES 

Bast  fibers  absent. 

A.  Vascular  bundle  fan-shaped   as  seen  in  cross   section R.   alpinum. 

B.  Vascular  bundle  circular  as  seen  in  cross  section R.  stenocarpum. 

Bast  fibers   present. 

A.    One  vascular  bundle  at  base  of  petiole. 

1.  Sclerenchyma    thick   walled    with    small    intercellular   space R.    missouriense. 

2.  Sclerenchyma  relatively  thin-walled  with   relatively  large  intercellular  space. 

a.  Vascular  bundle  in  the  shape  of  a  circle,  3-lobed,  and  fan-shaped  as  seen 
in  a  cross  section  midway  between  leaf  base  and  leaf  blade.  .  .  .R.  americanum. 

h.    Vascular  bundle  bent  in  the  shape  of  a  semicircle  and  not  3-lobed  as  seen 

in  a  cross  section  midway  between  leaf  base  and  leaf  blade.  .  .  .R.  fasciculatum. 
h.    Three  vascular  bundles  at  base  of  petiole. 

1.  Two  or  more  distinct  bundles  at  apex  of  petiole. 

a.  A  bundle  system  of  three  equal -sized  triangularly  arranged  bundles,  with 
entire  outer  edge  of  phloem  bordered  by  sclerenchyma R.  sativum. 

b.  A  bundle  system  consisting  of  unequal -sized  bundles  not  triangularly  ar- 
ranged, with  only  part  of  outer  edge  of  phloem  Vjordered  by  scleren- 
chyma   R.  diacantha. 

2.  One  bundle  at  the  apex  of  petiole. 

a.    Bundle  cross  section  3-lobed  at  apex  of  the  petiole. 

(1)  Petiole  cross  section  is  triangular  in  outline  at  a  point  mid\\ay  between 
leaf  base  and  leaf  blade R.   nigrum. 

(2)  Petiole    cross    section    is    inverted    dome-shaped    at    a    point    midway 
between  leaf  base  and   leaf  blade R.    holosericeum. 

h.    Bundle  cross  section  not  3-lobed  at  apex  of  petiole. 

(1)  Bundle  cross  section  at  apex  of  petiole  is  only  slightly  bent. 

(a)  Three  separate  bundles  bordered  by  relatively  thick-walled  scleren- 
chyma midway  between  leaf  base  and   leaf  blade R.   luridum. 

(b)  One  bundle  bordered  by  relatively  thin-walled  sclerenchyma  mid- 
way between  leaf  base  and  leaf  blade. 

1.  Bundle  cross  section  midway  between  leaf  base  and  blade  is 
circular  in  outline R-  aureum. 

2.  Bundle  cross  section  midway  between  leaf  base  and  blade  is 
3-lobed    and    fan-shaped R.    vilmorinii. 

(2)  Bundle   cross    section   at    apex    of    petiole    is   bent    in    the    shape    of    a 
semicircle. 

(a)  One   fan-shaped   bundle   seen   in   a    cross   section   midway   between 
leaf  base   and   blade R-    cynobasti. 

(b)  Three  separate  bundles  midway  between  leaf  base  and  blade. 

1.  Sclerenchyma  relatively  thin-walled  with  relatively  large  inter- 
cellular  space    R.    hirtellum. 

2.  Sclerenchyma   relatively   thick-walled   with   relatively   small   in- 
tercellular space    R-   tenue. 


384  The  University  Science  Bulletin 

KEY  TO  MIDRIBS 

I.    Upper  surface  of  midrib  U-shaped R-  nigrum. 

II.  Upper  surface  of  midrib  narrow  to  broadly  V-shaped. 

A.  Palisade  cells  one  layered  throughout  the  leaf R.  missouriense. 

B.  Palisade  cells  ranging  from  one  to  two  layers  througliout  the  leaf. 

a.  Vascular  bundle  fan-shaped  as  seen  in  cross  section. 

1.  Lower  epidermis  beset  with  multicellular,  sessile  glands R.  americanum. 

2.  Glands  absent   R-  sativum. 

b.  Vascular  bundle  nearly  a  circle  in  outline  as  seen  in  cross  section. 

1.  Leaf  blade  sharply  decreasing  in  thickness  on  approaching  the  midrib. 

R.  holosericeum. 

2.  Leaf  blade  not  decreasing  in  thickness  on  approaching  the  midrib. 

(a)  Vertical  thickness  of  midrib  more  than  twice  that  of  the  blade. 

R.  vilmorinii. 

(b)  Vertical  thickness   of  midrib   less  than   one   and   a   half   that   of  the 
jjl^jjg    R.   stenocarpum. 

C.  Palisade  cells  two  layered  throughout  the  leaf. 

a.    Two-thirds  of  the  midrib  projecting  below   the  leaf  blade R.   hirtellum. 

h.    About  one-third  of  the  midrib  projecting  below  the  leaf  blade R.  tenue. 

III.  Upper  surface  of  midrib  nearly  flat. 

A.  Midrib  decidedly  projecting  below  the  leaf  blade R.   cynobasti. 

B.  Midrib  not  decidedly  projecting  below  the  leaf  blade. 

a.  Outer  walls  of  the  upper  epidermal  cells  decidedly  projecting R.  aureum. 

b.  Outer  walls  of  the  upper  epidermal  cells  only  slightly  projecting. 

1.  Tangential  diameter  of  vascular  bundles  greater  than  vertical  diameter. 

R.  luridum. 

2.  Vertical  diameter  of  vascular  bundle  greater  than  tangential  diameter. 

R.  diacantha. 

IV.  Upper  surface  of  midrib  slightly  concave. 

A.  Outer  walls  of  upper  epidermal  cells  decidedly  projecting R.  fasciculatum. 

B.  Outer  walls  of  upper  epidermal  cells  only  slightly  projecting R.  alpinum. 

KEY  TO  MARGINS 

I.    Blade  cross  section  rounded  at  the  margin. 

A.  Blade    cross    section    less   than    .11    mm.    thick    at    a    point    .125    mm.    from    the 
margin    R-   americanum. 

B.  Blade  cross  section  between  .11  and  16  mm.  thick  at  a  point  .125  mm.  from  the 
margin. 

a.    Margin  roughened R-   hirtellum. 

h.    Margin  not  roughened. 

1.  Outer  walls  of  epidermals  cells  relatively   thick R.   fasciculatum. 

2.  Outer  walls  of  epidermal  cells  relatively  thin. 

(a)  Palisade  cells  not  extending  to   the  margin R.   holosericeum. 

(b)  Palisade  cells  extending  to  the  margin. 

(1)  Lower  epidermis  beset  with  multicellular,  sessile  glands.  .  .  .R.  nigrum. 

(2)  Glands  absent. 

*Mostly  two  layers  of  palisade  cells R.  stenocarpum. 

**Mostly  one  layer  of  palisade  cells R.  sativum. 

C.  Blade  cross  section  greater  than  .16  mm.  at  a  point  .125  mm.   from  the  margin. 

a.  Average  length  of  cells  in  the  upper  palisade  layer  greater  than   .035  mm. 

1.  Blade  cross  section  less  than   .21   mm.   thick   at   a   point   .125   mm.    from 

the   margin    R-    alpinum. 

2.  Blade  cross  section  greater  than   .21   mm.   at  a   point   .125   mm.    from   the 
margin. 

(a)  Outer  walls  of  epidermal  cells  relatively  thick R.   diacantha. 

(b)  Outer  walls  of  epidermal   cells  relatively   thin R.   luridum. 

b.  Average  length  of  cells  in  the  upper  palisade  layer  less  than  .035  mm. 

1.  Stomata  present  in  the  upper  epidermis R-  aureum. 

2.  Stomata  absent  in  the  upper  epidermis R-  tenue. 


Bates:  Study  of  the  Genus  Ribes  385 

II.    Blade  cross  section  pointed  at  the  margin. 

A.  Upper  epidermal   cells   relatively   thick R.   cynobasti. 

B.  Upper  epidermal  cells  relatively  thin. 

a.  Upper  surface  of  margin  curved  and  lower  surface  straight R.   7nissouriense 

b.  Margin   wedge-shaped R.    vilmorivii 

SUMMARY  AND  CONCLUSION 

An  examination  of  the  data  given  in  the  summary  of  stem  and 
leaf  features  leads  one  to  the  conclusion  that  all  fifteen  species 
agree  in  the  general  plan  of  their  anatomical  structure.  It  is  also 
evident  that  species  which  show  a  close  agreement  in  the  details 
of  their  anatomical  structure  also  show  a  close  agreement  in  their 
larger  morphological  features. 

In  comparing  a  large  number  of  species  which  are  similar  in  some 
structural  features  and  dissimilar  in  others  we  may  divide  them 
into  two  or  more  groups  on  the  basis  of  this  situation.  By  using 
many  such  features  a  large  number  of  different  groupings  may  be 
obtained.  Assuming  the  inheritance  of  these  different  features  to 
be  of  equal  persistence,  and  knowing  the  plants  have  been  raised 
under  the  same  environmental  conditions,  then  the  frequency  with 
which  any  two  species  fall  within  the  same  group  may  be  used  as 
an  indication  of  the  closeness  of  their  genealogical  relationship. 
Table  VIII,  based  on  eighteen  anatomical  features  not  common  to 
all  species,  shows  at  a  glance  the  frequency  with  which  each  species 
falls  in  the  same  group  with  each  of  the  other  species.  The  eighteen 
following  features  were  used  in  making  the  table:  Amount  and  dis- 
tribution of  trichomes  of  the  leaf  epidermis;  absence  or  kinds  of 
glands  of  the  epidermis  present;  shape  of  leaf  epidermal  cells;  shape 
of  leaf  tip;  shape  of  leaf  margin;  shape  of  midrib;  number  of 
layers  of  palisade  cells;  number  of  vascular  bundles  entering  the 
petiole;  presence  or  absence  of  trichomes  of  the  stem  epidermis; 
presence  or  absence  of  bast  fibers  in  the  petiole;  stem  armed  or 
unarmed;  shape  and  size  of  stem  epidermal  cells;  cutinization  of 
radial  walls  of  stem  epidermal  cells;  kinds  of  collenchyma;  number 
of  rings  of  peridermal  tissue  in  the  two  years'  growth;  number  of 
protoxylem  points ;  length  of  xylem  elements ;  and  the  distribution  of 
starch,  tannin  and  cluster  crystals  of  calcium  oxalate  in  the  tissues 
of  the  stem  and  leaf.  There  are  many  features  that  are  common  to 
all  species  given  in  the  summary  of  stems  and  leaves.  The  table 
indicates  that  there  are  many  instances  of  similarities  of  anatomical 
features  between  American  and  European  species.  The  three  highest 
frequencies  obtained  are  between  some  American  and  European 
species,  and  the  lowest  between  some  American  species  and  between 
American  and  European  species. 


386 


The  University  Science  Bulletin 


TABLE  Vlll. — Showing  the  frequency  with  which   any  two  species  fall  into  the  same  group 
on  the  basis  of  their  similarity  of  features 


103  holosericeum .  .  1 

110     tenue 2 

127  *aureum 3 

123     diacantha 4 

103  *americanum. . .  5 

108     nigrum 6 

119  fagciculatum. .  7 

118     alpinum 8 

116  sativum 9 

114     luridum 10 

74  stenocarpum .  .  11 

106  vilmorinii  ....  12 

66  •missouriense  . .  13 

18  *cjTiobasti 14 

117  *hirteUum 15 


8 

8 

16 

8 

6 

9 

8 

8 

7 

11 

10 

14 

9 

8 

12 

10 

10 

10 

6 

10 

11 

10 

8 

8 

7 

10 

8 

]0 

9 

7 

9 

11 

7 

7 

11 

10 

10 

7 

8 

10 

10 

4 

7 

7 

6 

5 

4 

4 

6 

8 

9 

7 

7 

5 

5 

8 

6 

5 

4 

3 

3 

5 

3 

6 

3 

.5 

6 

5 

7 

3 

4 

5 

6 

9 

10 

10 

11 

8 

7 

9 

10 

9       9 


4 

3  9 

8  10 

6  10 


9     10     11     12     13     14     15 


*  Indicates  American  species. 

Nos.   1-15  outside  the  square  are  used  as  symbols  for  the  different  species. 

The  numbers  inside  the  square  represent  frequencies. 

The  numbers  preceding  the  names  are  total  frequencies. 

Figure  I  is  a  map  of  the  world  showing  the  relative  position  of  the 
various  continental  masses  as  they  appear  to-day.  The  name  of  each 
of  the  fifteen  species  appears  on  the  particular  continent  or  part  of 
a  continent  to  which  the  species  is  native.  Lines  are  drawn  con- 
necting species  between  which  a  relatively  high  degree  of  anatomical 
relationship  was  found  in  Table  VIII.  The  table  in  figure  I  shows 
the  degree  of  anatomical  relationship  of  each  of  the  fifteen  species 
with  all  other  species  and  is  obtained  from  Table  VIII  by  combin- 
ing the  frequencies  with  which  each  species  falls  within  the  same 
group  with  each  of  the  other  species. 

Figure  I  shows  at  a  glance  that  there  are  numerous  instances  of 
a  relatively  close  anatomical  relationship  between  species  isolated 
on  different  continents.  It  also  shows  that  there  are  several  species 
{R.  cynobasti,  R.  stenocarpum,  and  R.  missouriense)  which  show  but 
little  anatomical  relationship  to  other  species,  even  to  those  found 
on  the  same  continent.  For  example,  the  two  highest  degrees  of 
anatomical  relationship  found  were  between  R.  aureum  (West  N. 
A.)  and  R.  diacantha  (North  Asia)  and  between  R.  americanum 
(East  N.  A.)  and  R.  nigrum  (North  Eu).  The  question  arises,  how 
could  species  isolated  on  such  remote  parts  of  the  earth  come  to 


Bates:  Study  of  the  Genus  Ribes 


387 


.   < 


aureun 127  . 

dlacantha 123  ; 

fasciculatum  119; 

hlrteHum 117; 

satlTTOT 116  •• 

lurldum 114 

tenue 110 

nigrum 108 


alplnuD— 108 

Tilmorlnll —  108 

amerloanuiD--  103 

holoserlceum  103 

cynobastl---  78 

stenocarpuiD-  74 

mlssourlense  6a 


have  such  a  close  degree  of  anatomical  correspondence?  It  cannot 
be  explained  by  hybridization,  and  it  is  very  improbable  that  they 
may  have  mutated  in  the  same  direction.  A  more  suitable  explana- 
tion is  furnished  by  assuming  a  former  connection  of  these  con- 
tinents, either  by  land  bridges  or  by  their  being  formerly  collected 
together  into  a  single  continental  mass  which  later  separated  and 
its  fragments  drifted  apart.  Such  a  theory  is  furnished  by  Koppen 
and  Wegener  (1924),  and  is  favorably  regarded  by  Zimmermann 
(1930)  and  Seward  (1931).  According  to  this  theory  the  Eastern 
coast  of  North  America  was  in  former  contact  with  Europe  and  the 
Western  coast  of  North  America  was  in  former  contact  with  Asia, 
which  accounts  for  the  close  anatomical  correspondence  between  R. 
americanum  and  R.  nigrum  and  between  R.  aureum  and  R.  dia- 
cantha.  The  numerous  lines  on  the  map  connecting  R.  aureum  with 
many  other  species  on  both  continents  and  the  high  degree  (127)  of 
anatomical  relationship  to  all  other  species  shown  in  the  accompany- 
ing table  below  the  map,  indicates  that  R.  aureum  either  belongs  to 
or  is  closely  related  to  the  original  hypothetical  stock  of  Rihes  from 
which  all  other  species  descended.  It  is  also  observed  that  the 
fifteen  species  are  listed  in  the  table  in  a  descending  order  from 
the  original  stock  (i  e.,  the  smaller  the  number  the  less  anatomical 
correspondence  with  other  species  and  the  further  the  descent  from 
the  original  stock) .  The  small  degree  of  anatomical  correspondence 
of  R.  cynobastl,  R.  stenocarpum,  and  R.  missouriense  with  other 
species  would  seem  to  indicate  that  they  have  probably  originated 
by  mutation. 


iiHH 


The  University  Science  Bulletin 


PLATE  XLIII 

TRICHOMES     (X  135) 


l^G.  1. 
Fig.  2. 
Fig.  3. 
Fig.  4. 
Fig.  5. 
Fig.  6. 
Fig.  7. 
Fig.  8. 
Fig.  9. 
Fig.  10. 
Fig.  11. 
Fig.  12. 
Fig.  13. 


Fig.  14. 
Fig.  15. 
Fig.  16. 
Fig.  17. 
Fig.  18. 
Fig.  19. 
Fig.  20. 
FIG.2L 
Fig.  22. 
Fig.  23. 
Fig.  24. 
Fig.  25. 
Fig.  26. 
Fig.  27. 
Fig.  28. 
Fig.  29. 
Fig.  30. 
Fig.  31. 
Fig.  32. 
Fig.  33. 
Fig.  34. 
Fig.  35. 
Fig.  36. 
Fig.  37. 
Fig.  38. 
Fig.  39. 
Fig.  40. 
Fig. 41. 
Fig.  42. 
Fig.  43. 
Fig.  44. 
Fig.  45. 
Fig.  46. 
Fig.  47. 
Fig.  48. 
Fig.  49. 
Fig.  50. 
Fig.  51. 
Fig.  52. 
Fig.  53. 
Fig.  54. 


Stem  trichome  of  -R, 
Stem  trichome  of  R. 
Stem  trichome  of  R 
Stem  trichome  of  R 
Stem  trichome  oi  R. 
Stem  trichome  oi  R 
Leaf  trichome  of  R. 
Leaf  trichome  of  ii!. 
Leaf  trichome  oi  R. 
Leaf  trichome  oi  R. 
Leaf  trichome  oi  R. 
Leaf  trichome  oi  R. 
Leaf  trichome  of  R. 


.  amencanum. 

vilmorinii. 
.  stenocarpum. 
.  holosericeum. 

sativum, 
.jasciculatum. 
missouriense. 
nigrum, 
jasciculatum . 
cynobasti. 
stenocar'pum. 
vilmorinii. 
hirtellum,. 


EPIDERMISES     (X  135) 


Stem  epiderm 
Stem  epiderm 
Stem  epiderm 
Stem  epiderm 
Stem  epiderm 
Stem  epiderm 
Stem  epiderm 
Stem  epiderm 
Stem  epiderm 
Stem  epiderm 
Stem  epiderm 
Stem  epiderm 
Stem  epiderm 
Stem  epiderm 
Stem  epiderm 
fep 
fep 
f  ep 
fep 
fep 
fep 
fep 
fep 
fep 
fep 
fep 
fep 
fep 
fep 
fep 
fep 
fep 
fep 
fep 
fep 
fep 
fep 
fep 
fep 
fep 
fep 


Lower 

leaf 

Upper 
Lower 

eaf 
leaf 

Upper 
Lower 

leaf 
leaf 

Upper 

leaf 

Lower 

leaf 

Upper 
Lower 

leaf 
leaf 

Upper 

leaf 

Lower 

leaf 

Lower 

leaf 

Upper 
Lower 

leaf 
leaf 

Upper 
Upper 
Lower 

leaf 
leaf 
leaf 

Lower 

leaf 

Upper 
Lower 

leaf 
leaf 

Upper 
Upper 
Upper 
Upper 
Lower 

leaf 
leaf 
leaf 
leaf 
leaf 

Lower 

leaf 

s  oiR.  tenue. 
s  of  R.  jasciculatum. 
soi  R.aureum. 
soi  R.  hirtellum. 
soi  R.  sativum, 
soi R.  missouriense. 
soi R.  vilmorinii. 
s  oi  R .  alpinu7n . 
s  of  R.  stenocarpum. 
soi  R.  holosericeum. 
soi  R.  nigrum. 
soiR.  luridum. 
s  oi  R.  cynobasti. 
s  of  R.  diacantha  (female) . 
s  of  jB.  americanum. 
dermis  of  R.  luridum. 
dermis  oi  R.  hirtellum. 
dermis  oi  R .  stenocarpum. 
dermis  oi  R.  stenocarpum. 
dermis  of  R.  americanum. 
dermis  of  R.  americanum. 
dermis  of  R.  holosericeum. 
dermis  oi  R.  luridum. 
dermis  of  R.  hirtellum. 
dermis  of  jB.  cynobasti. 
dermis  of  R.  cynobasti. 
dermis  oi  R.alpinum. 
dermis  oi  R.alpinum. 
dermis  of  R.  jasciculatum. 
dermis  of  R.  holosericeum. 
dermis  of -R.  vilmorinii. 
dermis  oiR.  vilmorinii. 
dermis  oiR. sativum. 
dermis  of  R.  nigrum. 
dermis  of  R.  nigrum. 
dermis  of  R.  jasciculatum. 
dermis  of  R.  diacantha. 
dermis  of  R.  sativum. 
dermis  of  R.  missouriense. 
dermis  of  -R.  missouriense. 
dermis  of  R.  diacantha. 


Bates:  Study  of  the  Genus  Ribes 


389 


PLATE  XLIII 


v><->^-"-^      54 


26-3482 


390  The  University  Science  Bulletin 


PLATE  XLIV 

TYPICAL  STEM  CROSS  SECTIONS     (X  48) 

Fig.  1.   Stem  section  of  R.  vilmorinii  in  the  primary  condition. 
Fig.  2.   Stem  section  of  R.  missouriense  taken  from  current  year's  growth. 
Fig.  3.    Stem  section  of  R.  nigrum  taken  from  current  year's  growth. 
Fig. 4.   Stem  section  of  R.  hirtclhim  taken  from  two  years'  growth. 


Bates:  Study  of  the  Genus  Ribes 


:v.)] 


PLATE  XLIV 


'^A 


I 


'^^^ 


fe'v:.:^' 


yd  ■■y 


^£&?S 


392 


The  University  Science  Bulletin 


PLATE  XLV 
MIDRIB  CROSS  SECTIONS 


(X48) 

Fig.  1.   R.  missouriense.  Fig.   9.  R.  americanum. 

Fig.  2.   R.  tenue.  Fig.  10.  R.  stenocarpum. 

Fig.  3.   R.  fasciculatum.  Fig.  11.  R.  diacantha. 

Fig. 4.   R.  aureum.  Fig.  12.  R.  hiriellum. 

Fig.  5.   R.  vilmorinii.  Fig.  13.  R.  alpinum. 

Fig.  6.   R.  nigrum.  Fig.  14.  R.  sativum. 

Fig. 7.   R.luridum.  Fig.  15.  R.  holosericeum. 
Fig.  8.    fl.  cynobasli. 

PETIOLE  CROSS  SECTIONS 

cynobasti.  Fig.  24. 


Fig.  16. 
Fig.  17. 
Fig.  18. 
Fig.  19. 
Fig.  20. 
Fig.  21. 
Fig.  22. 
Fig.  23. 


R. 
R. 
R. 
R. 
R. 
R. 
R. 
R. 


vilmorinii, 

holosericeum. 

luridum. 

alpinum. 

nigrum. 

hiriellum. 

diacantha. 


Fig.  25. 
Fig.  26. 
Fig.  27. 
Fig.  28. 
Fig.  29. 
Fig.  30. 


(X48) 

R.  aureum. 
R.  tenue. 
R.  americanum, 
R.  stenocarpum. 
R.  jasciculatum. 
R.  sativum. 
R.  missouriense. 


Bates:  Study  of  the  Genus  Ribes 


393 


PLATE  XLV 


304  The  University  Science  Bulletin 


PLATE  XLVl 

EPIDERMISES     (X  135) 

Fig.  1.  Lower  leaf  epidermis  of  i?.  auretim. 

Fig.  2.  Upper  leaf  epidermis  of  72.  awrewm. 

Fig.  3.  Lower  leaf  epidermis  of  R.  tenuc. 

Fig.  4.  Upper  leaf  epidermis  of  i?.  fenwe. 

MARGINS  OF  THE  LEAVES     (X  165) 

Fig.    5.  Leaf  margin  of  i^.americarjum. 

Fig.   6.  Leaf  margin  of  i?.nigrrum. 

Fig.    7.  Leaf  margin  oi  R.  vilmorinii. 

Fig.   8.  Leaf  margin  oi  R.  hirtellum. 

Fig.   9.  Lea,i  margin  oi  R.stenocarpum. 

Fig.  10.  Leaf  margin  of  R.  fasciculatvm. 

Fig.  11.  Leaf  margin  of  R.  missouriense . 

Fig.  12.  Leaf  margin  oi  R.  alpinum. 

Fig.  13.  Leaf  margin  of  R.  holosericeum . 

Fig.  14.  Leaf  margin  of  R .  cynobasti. 

Fig.  15.  Leaf  margin  oi  R.  diacantha. 

Fig.  16.  Leaf  margin  of  R.  sativum. 

Fig.  17.  Leaf  margin  of  R.  aureum. 

Fig.  18.  Leaf  margin  of  jK.  luridum. 

Fig.  19.  Leaf  margin  of  .R.  ^erme. 


Bates:  Study  of  the  Genus  Ribes 


395 


PLATE  XLVI 


396  The  University  Science  Bulletin 


PLATE  XLVII 

TYPICAL  LEAF  VENATION     (X  28) 

Fig.  1.  Leaf  venation  of  R.  alpinum. 

Fig.  2.  Leaf  venation  oiR.stenocarpum. 

Fig.  3.  Leaf  venation  of  R.  americanum. 

Fig.  4.  Leaf  venation  of  R.  hirtellum. 

Fig.  5.  Leaf  venation  of  R.  sativum,. 


Bates:  Study  of  the  Genus  Ribes 


397 


PLATE  XLVII 


398  The  University  Science  Bulletin 


BIBLIOGRAPHY 

Bailey,  L.  H.    Hortus.    The  Macmillan  Company,  New  York.    1930. 

Bailey,  L.  D.    Manual  of  Cultivated  Plants.    The  Macmillan  Company,  New 

York,  1924. 
Brooks,  C.  E.  P.    Climate  Through  the  Ages.    R.  V.  Coleman,  New  York,  192C. 
Delbrouck.    Pflanzenstach.,  in  Hanstein,  Bot.  Abh.,  Bd.  ii,  1875. 
HoLLE.    Beitrage  zur  Anatomie  der  Saxifragaceen  und  deren  Systematischen 

verwertung.    Botanishes  Centralblatt,  Band  53,  No.  1,  1893. 
KorPEN,  W.,  und  Wegener,  A.    Die  Klimate  der  geologischen  Vorzeit.    Berlin, 

G.  Borntraeger,  1924. 
Petit.     Petiole.     Mem.  Soc.  Phys.  et  Nat.  de  Bordeaux,  Ser.  3,  t.  Ill,  1887, 

pp.  318-20. 
Reinke.    Sekretionsorg.,  in  Pringsheims  Jahrb.,  Bd.  x,  1876,  pp.  145-9. 
Rehdee,  Alfred.     Manual  of  Cultivated  Trees  and  Shrubs.     The  Macmillan 

Company,  New  York,  1927. 
Seward,  A.  C.    Plant  Life  Through  the  Ages.    Cambridge,  at  the  University 

Press,  1931. 
Solereder,  Hans.     1860.     Systematic  Anatomy  of  the  Dicotyledons.     Tr.  by 

L.  A.  Boodle  and  F.  E.  Fritsch;  rev.  by  D.  H.  Scott.    2  v.,  il,  0.    Oxford, 

1908. 
Stevens,  W.  C.    Plant  Anatomy.    P.  Blakiston's  Son  &  Co.,  Philadelphia,  1924 
W.  A.  J.  M.  van   Waterschoot  van    der   Gracht.     Theory   of   Continental 

Drift.    Tulsa,  Okla.,  The  American  Association  of  Petroleum  Geologists; 

London,  T.  Murby  &  Co.,  1928. 
Zimmermann,  W.     Die  Phylogenie  der  Pflanzen.     Jena,  Verlag  Von  Fisher, 

1930. 


THE  UNIVERSITY  OF  KANSAS 

SCIENCE  BULLETIN 

Vol.  XXT.l  March,  1933.  [No.  12 


Morphology  and  Anatomy  of  Mollugo  verticillata  L. 

SISTER   M.   ANTHONY  PAYNE,  O.   S.   B. 


Abstr.act:  The  probable  relation.=;hip  of  the  Aizoaceae  to  the  Phytolaccaceae 
is  analyzed,  as  well  as  its  relationship  to  the  Chenopodiacese,  Portulacacese 
and  Caryophyllacese.  Mollugo  verticillata,  the  "carpet  weed,"  belongs  to  the 
family  Aizoaceae,  which  is  principally  tropical,  containing  18  genera  and  600 
species. 

Mollugo,  found  in  temperate  climes,  has  twelve  to  eighteen  radiating  stems 
which  crown  a  tapering  root  and  lie  prostrate  on  the  ground.  The  leaves  are 
in  small  whorls,  and  the  inconspicuous  flowers  are  borne  on  slender  pedicels. 
They  are  perfect,  but  incomplete.  Three  stamens  alternate  with  the  locules 
of  the  three-chambered  ovary,  which  contains  about  thirty  kidney-shaped 
seeds. 

The  thickened  spatulate  leaf  has  stomata  on  both  surfaces  and  vestigial 
trichome  bases  on  the  upper  epidermis.  The  mesophyll  is  about  eight  layers 
in  thickness.  Numerous  translucent,  simple  crystals  abound  in  the  mesophyll 
of  the  leaf  and  in  the  cortical  tissue  of  the  stem.  The  crystals  are  probably 
calcium  oxalate,  for  they  refract  interference  colors  of  the  third  order.  The 
structure  of  older  portions  of  stems  varies  between  nodes  and  intenodes, 
while  pseudoannulation  occurs  in  cross  sections  of  both  stem  and  root.  The 
units  which  form  the  tracheal  tubes  are  heavily  pitted.  The  dense  cylindrical 
arrangement  of  tracheal  tubes  and  lignified  tracheids,  followed  by  a  cylinder 
of  less  compactly  disposed  tubes  and  fibrous  tracheids,  given  the  impression  of 
annual  rings. 

Drawings  and  photomicrographs  illustrate  various  points  in  the  paper. 


CLASSIFICATION  AND  DISTRIBUTION 

BEFORE  discussing  the  morphology  and  anatomy  of  an  unfamiliar 
plant  it  seems  well  to  classify  it,  for,  although  "carpet  weeds" 
may  cover  the  pathway,  it  is  probable  that  few  of  them  attract 
curious,  much  less  quizzical,  glances.  In  Rendle's  "Classification 
of  Flowering  Plants"  (1925)  Mollugo  verticillata  L.  is  pigeonholed 
as  follows:  Genus  and  species,  Mollugo  verticillata,  of  the  family 
Aizoaceae,  order  Centrospermae,  and  grade  Monochlamydese. 

(399) 


400  The  University  Science  Bulletin 

In  the  few  lines  devoted  to  it  by  various  authors  Mollugo  has  been 
described  by  comparison  with  related  forms  rather  than  by  specific 
description.  Well-known  related  families  are  the  Chenopodiacese, 
the  Phytolaccaceae,  the  Portulacaceae  and  the  Caryophyllaceae. 
These  families  possess  in  common  a  characteristically  curved  embryo 
surrounding  a  floury  or  mealy  endosperm,  and  a  central  plasenta, 
to  which  the  seeds  are  attached. 

The  small  family  of  the  Phytolaccacea)  shows  great  variety  in  the 
plan  of  structure  of  the  flower,  including  many  apetalous  as  well 
as  polypetalous  forms.  Multiple  stamens  and  carpels  also  occur, 
and  Pax  (1892)  suggests  Phytolaccaceae  as  the  genetic  family  of 
the  order  CentrospermsD,  because  in  this  family  are  found  genera 
of  the  monochlamydeous  type  as  well  as  dichlamydeous  forms  in 
which  a  multiplication  of  stamens  and  carpels  has  occurred.  Family 
relations  of  Mollugo  may  be  illustrated  by  the  following  diagram, 
which  readily  shows  why  Gray  (1887)  first  placed  Mollugo  in  a 
suborder  of  the  Caryophyllacese  and  later  in  the  order  of  the  Che- 
nopodiales  (1908). 

The  Aizoaceae  resemble  Phytolaccaceae  in  their  apetalous  flowers 
and  in  the  anomaly  of  the  stem  structure  of  many  of  its  members. 
It  is  a  miscellaneous  group  of  herbs,  shrubs,  undershrubs  and  succu- 

FAMILY  RELATIONS  OP  MOLLUGO 


Chenopodiacese 


Aizoaceae 


fa.  Monochlamydeous 

Lb.  Stamens  in  one  series 

''a.  Monochlamydeous 

,.b.  Stamens,  3,  5,  or  o^> 


'a.   Dichlamydeous 
Portulaca- 


ceae  [b,   Antepetalous  stamens, 
3,  5,  or  increased. 


fia ,  Dichlamydeous 
b.  Starasns  8-10  in 
two  series. 


Payne:  Mollugo  Verticillata  L.  401 

lent  plants.  The  family  is  principally  tropical  and  is  found  in  great 
numbers  in  dry  places  and  the  warmer  parts  of  the  earth,  particu- 
larly in  South  America.  There  are  eighteen  genera  and  600  species. 
Four  hundred  of  these  belong  to  the  genus  Mesembryanthemum. 
The  familiar  ice  plant,  M.  crystallinum,  is  an  example  of  the  genus, 
and  thrives  in  California,  Australia  and  about  the  Cape.  Mollugo 
also  is  tropical,  but  it  has  invaded  the  northern  parts.  In  1892  it 
was  reported  from  central  Texas  by  Mrs.  Charles  Ward,  who  col- 
lected the  specimen  for  a  private  herbarium.  W.  A.  Wheeler,  in  the 
summer  of  1899,  collected  specimens  of  M.  verticillata  in  the  extreme 
southeastern  part  of  Minnesota,  and  the  specimens  on  which  the 
present  studies  were  made  were  collected  by  W.  C.  Stevens,  pro- 
fessor of  botany  of  Kansas  University,  near  Lawrence,  Kansas. 
Gray's  manual  (1887)  states  that  although  it  is  an  immigrant  from 
farther  south,  it  may  be  found  on  sandy  river  banks,  roadsides 
and  in  cultivated  ground  from  June  to  September.  The  inference 
is  that  it  is  rather  common.  Its  relative  unimportance  and  in- 
significance accounts,  perhaps,  for  the  lack  of  notation  in  the  litera- 
ture. 

HABITAT  AND  GENERAL  MORPHOLOGY 

M.  verticillata,  the  "carpet  weed,"  has  many  (12-18)  slender, 
jointed  branches  radiating  from  the  top  of  a  tapering  root.  The 
stems  are  prostrate  on  the  earth  and  form  a  i-ather  dense  mat  that 
measures  20  to  24  cm.  in  diameter.  The  flat  patches  of  greyish 
green  are  not  showy,  for  the  small  spatulate  whorls  of  leaves  and 
the  delicate  recurved  pedicels  bearing  single  flowers  are  inconspicu- 
ous. The  flowers  form  an  umbellike  cluster.  At  times  they  are  in 
the  axils  of  the  leaves,  but  more  often  five  or  six  flowers  are  aggre- 
gated on  one  side  of  the  stem  while  the  four  or  five  leaves  are 
crowded  in  a  semiwhorl  on  the  opposite  side.  Leaves  and  flowers 
together  suggest  the  umbel. 

The  stems  are  tumid  at  the  nodes,  the  swellings  decreasing  in  size 
at  nodes  farther  and  farther  removed  from  the  root.  The  main 
stems  are  3.5  mm.  in  diameter  at  the  point  of  origin  and  only  1.5 
to  2  mm.  in  cross  section  between  the  third  and  fourth  nodes.  They 
continue  to  decrease  in  diameter  to  the  growing  tip.  The  lenolh  of 
the  average  internode  is  25-35  mm. 

The  main  taproot  measures  4  mm.  in  diameter  at  the  point  of 
junction  with  the  stem. 

The  "carpet  weed"  can  grow  in  dry,  sandy  places  because  of  its 
eflficient  root  system  and  the  water-storing  tissue  in  the  leaves  and 


402  The  University  Science  Bulletin 

nodes.  The  flat  mat  conserves  the  moisture  in  the  soil  adjacent  to 
the  root  and  the  long  taproot  reaches  to  depths  of  18  to  20  cm. 
below  the  surface,  while  short  lateral  roots  coming  from  it  drain  a 
large  area  (PL  XLVIII,  fig.  6). 

The  small  hypogynous  flowers  are  borne  on  slender  pedicels  at 
the  nodes,  the  pedicels  being  as  long  or  slightly  longer  than  the 
mature  capsule,  which  may  measure  4  mm.  by  2  mm.  The  flowers 
are  regular  and  perfect,  but  incomplete,  having  a  whitish  perianth 
composed  of  five  small  curved  sepals,  three  short  stamens  with 
plump  anthers  alternating  with  the  cells  of  the  ovary,  and  a  short 
three-parted  style  over  a  large  three-chambered  ovary  (PI.  L,  figs. 

9,10). 

The  ovary,  when  mature,  is  crowded  with  large  anatropous  ovules 
arranged  on  axial  placentae.  Each  locule  contains  two  rows  of  seeds, 
one  on  either  side  of  a  partition  which  extends  from  the  wall  of  the 
ovary  in  the  center  of  each  locule  nearly  to  the  placenta.  There 
are  generally  five  or  six  seeds  in  a  row,  each  fully  developed  ovary 
containing,  therefore,  about  thirty  seeds.  The  seeds  are  kidney- 
shaped  and  a  reddish  brown.  The  brittle  testa  is  etched  in  delicate 
stria?  (PI.  XLVIII,  fig.  1)  and  within,  the  large  embryo  is  curved 
peripherally  about  the  fine,  starchy  endosperm.  The  epicotyl  oc- 
cupies the  upper  plump  portion  of  the  kidney-shaped  seed  (PI. 
XLVIII,  fig.  2). 

The  simple,  entire  spatulate  to  cuneiform  leaves  occur  in  whorls 
or  semiwhorls  at  every  node  of  the  stem.  There  are  four  to  eight 
leaves  in  a  whorl,  unequal  in  size,  even  in  a  single  whorl,  although 
the  variation  in  size  in  the  whorl  is  less  than  the  variation  in  leaf 
size  between  whorls.  The  leaves  that  compose  the  whorls  near  the 
point  of  origin  of  the  stem  measure  from  15  to  20  mm.  by  3  mm. 
(at  the  broadest  part  of  the  blade),  while  the  leaves  composing  the 
distal  whorls  measure  5  to  10  mm.  They  are  generally  pointed,  and 
a  few  tend  to  have  a  mucronate  apex  (PI.  XLVIII,  fig.  10). 

ANATOMY 

Several  methods  of  procedure  were  followed  in  preparing  the 
tissues  for  microscopic  examination.  Portions  of  the  specimens  had 
been  preserved  some  months  before  in  equal  parts  of  95  per  cent 
alcohol  and  glycerin  and  were  in  very  good  condition.  Before  em- 
bedding in  paraffin  small  pieces  of  stem,  root,  leaf,  flower  and 
pedicel  were  dehydrated  by  the  butyl-alcohol  method  (Zirkle,  1930), 
and  then  cut  in  cross  and  longitudinal  sections  varying  from  10  to 


Payne:  Mollugo  Verticillata  L.  403 

20  [X  in  thickness.  Some  sections  were  stained  in  Kernschwartz  and 
safranin,  others  in  gentian-violet  and  acid-fuchsin.  The  Kern- 
schwartz and  safranin  gave  good  contrast  and  made  dehcate  cell 
walls  stand  out  as  well  as  cytoplasmic  inclusions.  Other  sections 
were  simply  cleared  in  chloral  hydrate.  Others  still  were  treated 
with  various  acids  in  an  effort  to  determine  the  nature  of  the  crystal 
inclusions.  The  endosperm  material  was  tested  with  Sudan  III 
and  a  tincture  of  iodine.  Sudan  III  was  also  used  on  freshly 
mounted  cross  sections  of  root  and  stem.  The  amount  of  lignifica- 
tion,  as  determined  by  the  safranin  stain,  was  checked  by  the 
phloroglucin  test  (Stevens,  1924).  The  lignified  tissue  which  had 
been  immersed  in  alcohol  containing  a  trace  of  phloroglucin  turned 
a  deep  red  when  a  drop  of  hydrochloric  acid  was  added. 

Sections  for  photomicrographs  were  dehydrated  and  cleared  in 
xylol,  then  mounted  in  hyrax.  Exposures  varying  from  fourteen 
seconds  to  six  minutes  with  an  arc  light  illuminant  were  made  on 
hard-surface  sensitized  paper,  giving  a  negative  print.  A  few  dry- 
plate  negatives  were  developed  for  comparative  data. 

LEAF 

Leaves  cleared  in  chloral  hydrate  were  used  for  the  study  of  the 
epidermal  layers.  In  surface  view  most  of  the  upper  epidermal 
cells  are  relatively  large  and  irregular  with  pronounced  undulating 
margins.  The  length,  averaging  .082  mm.,  is  more  than  twice  the 
width  (PI.  XL VIII,  fig.  11).  Over  the  midrib  and  larger  veins  the 
epidermal  cells  are  elongated  and  rectangular  with  regular  margins 
(PI.  XLVIII,  fig.  12);  A  few  patches  of  small,  rounded  cells  can 
be  found  on  the  upper  surface  of  the  leaf,  while  cells  much  less  ir- 
regular in  contour  than  the  typical  undulating  epidermal  cells 
stretch  out  from  these  areas  forming  occasional  rosettes  in  the  epi- 
dermal pattern  (PI.  XLVIII,  fig.  14).  These  rosettes  suggest  the 
loci  of  trichomes,  but  trichomes  are  lacking  in  M.  verticillata.  Since, 
however,  so  many  related  forms  are  hairy  it  seems  probable  that 
this  peculiarity  is  a  vestigal  remnant  of  ancestral  forms. 

The  lower  epidermis  is  likewise  composed  of  sinuous  cells,  but 
the  undulation  of  the  walls  is  slightly  less  marked.  In  some  areas 
the  cells  approach  a  rough  rectangle  in  shape. 

Stomata  occur  on  both  upper  and  lower  surfaces.  A  stoma  is 
elongate-eliptical  in  shape  and  measures  about  0.018  mm.  in  length. 
Stomata  are  more  numerous  on  the  lower  than  on  the  upper  surface 
and  are  unevenly  distributed.    The  average  is  97.1  stomata  per  sq. 


404  The  University  Science  Bulletin 

mm.  for  the  upper  surface  and  117.9  stomata  per  sq.  mm.  on  the 
lower  surface  (PI.  XL VIII,  figs.  11  and  16). 

Stomata  occur  frequently  in  the  epidermis  of  the  stem  (PI.  XLIX, 
fig.  7). 

A  cross  section  of  the  leaf  (PI.  L,  fig.  12)  shows  a  much  thickened 
blade  with  the  region  over  the  midrib  depressed.  There  is  a  cor- 
responding protrusion  of  tissue  on  the  ventral  surface.  Cells  of  the 
simple  upper  epidermis  are  larger  in  cross  section  than  cells  of  the 
lower  epidermis,  the  former  being  about  two  times  longer  than  high 
and  nearly  oval  (PI.  XLVIII,  figs.  13,  15).  The  outer  wall  is  con- 
vex and  cutinized,  while  the  inner  wall  is  slightly  curved  and  very 
thin.    Upper  epidermal  cells  average  .031  mm.  in  radial  thickness. 

Cells  forming  the  lower  epidermis  have  about  0.6  the  diameter  of 
the  upper  epidermal  cells.  The  outer  wall  is  less  convex  and  not  as 
heavily  cutinized  as  in  the  upper  epidemiis. 

The  irregular  palisade-parenchyma  occurs  below  the  epidermis 
in  two  cell  layers.  The  cells  are  not  greatly  elongated,  some  of  them 
being  nearly  square.  In  the  lower  layer  many  of  the  palisade  forms 
pass  over  into  a  round  form.  Large  intercellular  spaces  intervene, 
particularly  in  the  upper  row.  Some  of  the  palisade  cells  are  typical 
— narrow  and  twice  as  long  as  neighboring  cells.  They  extend  the 
entire  breadth  of  the  palisade  parenchyma.  These  long  cells  are 
abundant  at  the  edge  of  the  leaf,  and  when  measured  in  the  surface 
view  average  .038  x  .033  mm.  Conspicuous  air  spaces  are  among 
the  palisade  cells. 

The  spongy  mesophyll  occupies  about  0.6  of  the  thickness  of  the 
entire  leaf.  It  is  seven  or  eight  layers  thick,  while  the  palisade 
parenchyma  occupies  only  about  .25  of  the  thickness.  The  border- 
parenchyma  cells  are  relatively  large,  sinuous,  thin-walled  cells,  and 
all  the  mesophyll  in  the  plane  of  the  veins  is  unusually  large.  The 
midrib  is  connected  to  both  surfaces  of  the  leaf  by  collenchyma 
tissue.  The  vascular  tissue,  forming  the  midrib,  is  slightly  concave 
above  and  convex  beneath,  the  phloem  flanking  the  xylem,  largely. 
There  is  a  tendency  for  it  to  completely  surround  the  xylem  elements. 

The  smaller  veins  are  completely  immersed  in  the  mesophyll  and 
also  have  a  sinuous  parenchyma  sheath.  A  cross  section  of  a  leaf 
shows  three  or  four  large  lateral  bundles  to  either  side  of  the  midrib 
(PI.  L,  fig.  12). 

When  a  cleared  leaf  is  brought  in  focus  the  most  striking  charac- 
teristic is  the  presence  of  innumerable  translucent,  simple  crystals 
and  tan-colored  compound  ones.    Masses  of  these  splotch  the  field 


Payne:  Mollugo  Verticillata  L.  405 

in  many  areas  and  obscure  the  underlying  tissue  (PI.  L,  fig.  11). 
Large  ones  are  scattered  at  random  just  under  the  epidermis,  but  a 
rapid  shift  of  focus  brings  many  more  from  lower  levels  into  view 
and  the  impression  is  gained  that  the  leaf  is  loaded  with  crystals. 
Ten  surface  counts  ranged  from  700  to  750  crystals  per  sq.  mm. 
Fields  can  be  selected,  however,  in  which  the  count  is  much  higher 
or  much  lower.  A  change  of  focus  to  lower  depths  of  the  tissue 
makes  an  accurate  estimate  of  frequency  impossible  due  to  the  fact 
that  crystals  crowd  and  overlap  from  one  local  plane  to  another 
(PI.  L,  fig.  8). 

The  crystals  are  of  two  forms  although  great  variation  occurs 
within  the  classes.  One  form  is  the  typical  spherical  rosette  and 
the  other  is  the  small,  colorless  octahedral  crystal.  Both  groups  are 
doubtlessly  forms  of  calcium  oxalate  (Ca  C2  O4  H2O),  for  they  dis- 
solve without  effervescence  when  immersed  in  a  2  per  cent  solution 
of  hydrochloric  acid.  They  go  into  solution  slowly,  however,  for 
after  immersion  for  thirty  or  forty  minutes  all  of  the  crystals  may 
not  have  disappeared  from  a  cross  section  of  a  leaf  .012  mm.  thick. 
The  very  small,  simple  crystals  and  the  rosette  forms  disappear  first; 
later  the  large  octahedron  type  (Rieder,  1899). 

The  habit  of  the  crystals  varies  greatly,  and  an  almost  endless 
number  of  combinations  of  simple  types  from  acute  to  obtuse  quad- 
ratoctahedra  occur.  Envelope  forms,  square  prisms  with  pyramidal 
ends,  truncated  pyramids,  elongated  octahedra  with  bevelled  edges 
or  truncated  angles  are  present.  Some  are  imperfectly  developed, 
but  most  of  them  can  easily  be  classified  as  octahedral  or  twinned 
octahedral  forms. 

Multiple  twinning  of  small  crystals  gives  rise  to  large  rosette 
forms  or  concretions.  Contact  twins  and  penetration  twins  occur 
frequently.  Twinned  structures  can  be  detected  by  means  of  the 
barbed  striae  which  meet  in  a  line  on  the  face  of  crystals  that  at 
first  seem  simple.  Discs  form  when  oxalate  of  lime  is  rapidly 
precipitated.  De  Bary  states  that  the  form  of  the  crystal  depends 
on  the  rapidity  with  which  it  is  deposited.  The  amount  of  water  of 
crystallization  also  varies  in  the  different  forms  (De  Bary,  1884). 

With  ordinary  illumination  many  of  the  very  small  crystals  seem 
irregular  or  indefinite  in  outline,  but  monochromatic  light  reveals 
sharp  edges  and  a  specific  contour.  When  studied  under  a  polariz- 
ing microscope  the  crystals  of  Mollugo  may  refract  one  of  several 
interference  colors  of  the  third  order;  blue,  green,  greenish-yellow 
or  reddish-violet  (Ford,  1922).    This  difference  in  the  birefringence 

27—3482 


406  The  University  Science  Bulletin 

is  due  to  the  size  or  thickness  of  the  crystal  and  its  orientation,  for 
the  greater  the  amount  of  double  refraction  the  higher  the  order  of 
interference  color.  The  refraction  of  the  colors  of  the  third  order 
supports  the  evidence  from  other  sources  that  the  crystals  are 
calcium  oxalate.  (Birefringence  of  calcium  oxalate  is  .163,  the  index 
of  refraction  for  one  ray  being  1.493,  and  for  the  other  1.656.) 

Longitudinal  and  cross  sections  of  the  leaves  .020  to  .040  mm. 
thick  show  that  crystals  of  both  groups  occur  plentifully  throughout 
the  spongy  tissue  and  in  the  palisade  cells.  Rows  of  simple  crystals 
fill  the  cells  that  border  the  veins  (PL  XLVIII,  fig.  9).  Small 
pyramidal  forms  occur  in  cells  of  Mollugo  more  distant  from  the 
veins,  and  a  few  of  the  upper  epidermal  cells  have  tiny  crystals  in 
the  vacuoles,  but  they  are  the  exception,  not  the  rule,  while  in 
Mesembryanthemum  lacerum  there  are  crystals  in  the  epidermal 
cells  in  large  numbers  (DeBary,  1884).  The  rosette  crystals  are 
massed  over  veins  or  in  areas  quite  far  removed  from  the  main 
veins.  If  one  rosette  crystal  occurs  in  a  cell  often  the  six  or  eight 
cells  adjacent  to  it  are  also  crystal  sacs.  The  crystals  are  always 
suspended  in  the  vacuoles  (PI.  XLVIII,  fig.  7). 

ANATOMY  OF  STEM 

Cross  sections  and  longitudinal  sections  of  stem  taken  from  parts 
adjacent  to  the  root,  at  nodes  and  at  internodes  were  examined. 
The  vascular  tissue  is  cylindrical  and  surrounds  a  relatively  large 
homogeneous  pith.  The  simple  epidermis  is  subtended  by  a  cortex 
which  varies  in  thickness  in  different  regions.  Some  of  the  inner 
cortical  cell  rows  are  heavily  lignified  and  sometimes  form  an 
undulating  fibrous  sheath  about  the  vascular  tissue,  sometimes  a 
broken  ring  of  bast  fibers  (PI.  XLIX,  figs.  5,  6,  7). 

The  pith  consists  of  several  longitudinal  rows  of  large,  thin-walled 
cells,  which  fill  the  center  of  the  pith  cylinder  and  are  surrounded 
peripherally  by  rows  of  smaller  thin-walled  cells.  The  parenchyma 
cells  are  very  small  at  the  periphery  of  the  pith,  measuring  in  cross 
section  .033  by  .041  mm.  in  diameter.  The  empty  central  pith  cells 
are  .146  mm.  by  .061  mm.  These,  by  far  the  largest  cells  found  in 
Mollugo,  in  a  longitudinal  section  appear  barrel-shaped.  The  thin 
walls  of  the  pith  cells  are  finely  pitted  and  small  triangular  in- 
terstices occur  among  them  (PI.  XLIX,  fig.  6).  Crystals  do  not 
occur  in  pith  cells. 

There  are  several  rows  of  protoxylem.  Next  to  the  pith  a  narrow 
vessel  .011  mm.  in  diameter,  with  a  heavy,  loose-coiled  spiral  occurs, 


Payne:  Mollugo  Verticillata  L.  407 

while  further  out  there  is  a  wider  spiral  vessel  (.015  mm.)  with  a 
finer  spiral  more  closely  coiled  (PI,  LI,  fig.  14).  Both  vessels  spiral 
from  left  to  right.  In  several  longitudinal  sections  the  inner  spiral 
was  already  distorted  through  excessive  elongation  of  the  branch. 
A  row  of  thin-walled  parenchymatous  cells  separate  the  spiral 
vessels. 

The  narrow  band  of  protoxylem  is  followed  sometimes  by  pitted 
tracheal  tubes,  sometimes  by  fiber  tracheids.  The  tracheal  tubes 
always  have  heavily  lignified  walls  and  are  scattered  irregularly 
among  the  thinner-walled  xylem  parenchyma  cells.  There  are  no 
medullary  rays.  The  large  pitted  tracheal  ducts  measure  .105  mm. 
in  diameter.  These  pitted  vessels  are  without  any  protoplasmic 
contents,  but  occasionally  crystals  and  crystal  sand  occurs.  Three 
large,  well-formed  rhombohedral  crystals  in  this  position  are  shown 
in  Plate  LI,  figure  15.  The  large  bordered-pitted  vessels  are  sur- 
rounded by  fibrous  tracheids  which  vary  greatly  in  size  and  form. 
Some  tracheids  measure  only  .010  mm.  in  diameter  and  are  many 
times  longer  than  wide  and  have  square  ends.  Others  are  .32  mm. 
in  diameter  and  have  the  end  walls  at  an  angle  of  40°.  The  wider 
tracheids  are  short  and  have  relatively  few  pits  in  the  lignified 
walls.  Xylem  parenchyma  is  not  abundant.  The  elements  are 
small,  oblong  and  thin-walled. 

In  cross  sections  of  larger  stems,  the  dense  cylindrical  arrange- 
ment of  tracheal  tubes  and  lignified  tracheids,  followed  by  a  cylinder 
of  less  compactly  disposed  tubes  and  fibrous  tracheids,  gives  the  im- 
pression of  annual  rings  (PI.  XLIX,  fig.  7).  This  pseudoannula- 
tion  is  characteristic  likewise  of  the  root. 

The  phloem  tissue  is  outside  the  xylem  cylinder  only  (PI.  XLIX, 
fig.  4).  A  cambial  layer  cannot  be  distinguished  in  mature  stems. 
The  arrangement  of  the  phloem  differs  in  cross  sections  taken  at 
the  nodes  and  between  the  nodes.  The  phloem  between  the  tumid 
regions  about  the  nodes  forms  a  complete  cylinder  consisting  of 
three  or  four  rows  of  sieve  tubes,  companion  cells  and  phloem 
parenchyma  surrounded  by  one  or  two  rows  of  heavier-walled  paren- 
chyma cells.  The  sieve  tubes  measure  only  .002  to  .003  mm.  in  di- 
ameter and  are  associated  with  companion  cells  that  are  even 
smaller.  The  tube  elements  are  about  .121  mm.  in  length  (PI.  LI, 
fig.  13). 

At  nodes  the  phloem  tissue  occurs  in  small  bundles  which  are 
separated  by  rays  of  parenchyma  cells,  like  extensions  from  the 
inner  cortex. 


408  The  University  Science  Bulletin 

The  cortex  is  variable  (PI.  XLIX,  figs.  1,  2,  4).  In  cross  sections 
taken  from  internodes  there  are  six  or  seven  layers  of  rounded  cells. 
The  subepidermal  layer  is  not  collenchymatous  and  no  endoder'mis, 
as  noted  above,  occurs,  but  an  undulating  bast-fiber  sheath  is  one 
or  two  rows  removed  from  the  phloem  cylinder.  The  walls  of  the 
bast  fibers  are  heavily  lignified,  in  some  instances  being  as  thick  as 
the  diameter  of  the  lumen  of  the  cell.  The  fibers  may  be  five  or 
six  deep  and  then  gradually  decrease  until  there  is  only  one.  A 
change  in  the  sheath  then  occurs,  for  the  next  cell  or  two  is  non- 
lignified  and  nonfibrous  (PI.  L,  fig.  6). 

Typical  cortical  cells  are  large,  rounded  and  active.  The  proto- 
plast contains  a  large  nucleus  and  abundant  chloroplasts  (PI.  LI, 
fig.  13).  Many  of  the  cortical  cells  are  crystal  sacs,  large  rosette 
crystals  forming  in  the  sap  cavities  (PI.  XLIX,  fig.  2.) .  These  cells 
have  thin  walls  and  measure  .039  mm.  in  diameter.  In  longitudinal 
sections  hypodermal  cells  are  only  slightly  elongated,  while  the 
deeper  cortical  cells  are  nearly  twice  as  long.  The  bast  fibers  are 
twenty  times  longer  than  wide.  A  few  small  pits  occur  in  the 
walls  (PI.  XLVIII,  fig.  8). 

In  cross  section,  epidermal  cells  bulge  slightly  and  are  also  cutin- 
ized  (PL  XLIX,  fig.  6).  In  surface  view  they  are  roughly  rec- 
tangular in  outline  and  measure  approximately  .113  by  .040  mm. 
Stomata  are  numerous  (PI.  LI,  fig.  16).  ' 

ROOT 

The  root  of  M.  verticillata  is  somewhat  anomalous  in  structure  in 
this  sense,  that  elements  of  the  single  cylinder  of  xylem  tissue  are  so 
distributed  that  they  give  the  impression  of  normal  annulation. 
There  are  three  pseudoannular  rings  of  xylem  in  cross  sections  of 
larger  roots  (PL  LI,  figs.  18,  19),  although  M.  verticillata  is  an 
annual. 

Adjacent  to  the  phloem  there  are  three  rows  of  tracheal  tubes, 
some  of  which  measure  .042  x  .146  mm.  in  diameter,  and  are  there- 
fore the  largest  elements  found  in  the  root.  The  walls  of  these  tubes 
are  thin  and  have  partially  or  completely  lignified  walls.  In  longi- 
tudinal sections,  the  rims  of  the  dissolved  cross  walls  are  remarkably 
distinct.  Groups  of  nonlignified  fiber-tracheids,  which  measure  only 
.007  to  .012  mm.  in  diameter,  are  clumped  about  or  between  the 
tracheal  tubes. 

A  cylinder,  one  to  two  layers  deep,  of  irregular,  small,  heavy- 
walled,  lignified  tracheids  follows  the  above-described  tissue.    These 


Payne:  Mollugo  Verticillata  L.  409 

cells  are  elongated  spindles,  and  it  is  their  arrangement  periodically, 
in  rings,  that  brings  about  the  pseudoannulation.  The  large,  rela- 
tively thin-walled  tracheal  tubes,  embedded  in  clumps  of  nonligi- 
fied  fiber-tracheids  follow.  In  a  cross  section  of  root  1.056  ram. 
in  diameter,  the  inner  or  central  "annual  ring"  measures  .273  mm. 
across  and  its  central  mass  as  well  as  the  three  or  four  peripheral 
rows  of  cells  are  the  small,  deeply-lignified  tracheids,  interspersed 
with  the  nonlignified  fiber-tracheids.  The  fiber-tracheids  have  dis- 
tinct cell  contents  and  are,  therefore,  living  cells.  In  a  sense  they 
take  the  place  of  xylem  parenchyma,  which  does  not  occur  in  the 
root  (PI.  LI,  fig.  18). 

In  macerated  tissue  the  nonlignified  fiber-tracheids  are  seen  to  be 
narrow  and  sharp  pointed.  Some  measure  .252  x  .011  mm.,  and  are 
therefore  23  times  longer  than  wide.  The  numerous  pits  in  the  large 
tracheal  tubes  are  heavily  bordered  and  arranged  in  relatively 
straight  rows.  Tracheal  tubes  are  more  numerous  than  any  other 
elements  in  the  xylem  tissue.  The  small,  pitted,  lignified  tracheids, 
measuring  .013  x  .038  mm.,  have  blunt  pointed  ends. 

Ciystals  also  occur  in  the  tracheal  tubes  of  the  root.  They  are 
large  and  simple,  but  very  irregularly  formed.  In  one  microscopic 
field  five  or  six  crystals  have  been  counted. 

In  the  same  cross  section  (1.056  mm.  in  diameter)  the  cortical 
tissue  measures  .189  mm.  in  width.  There  are  eleven  or  twelve  rows 
of  parenchyma,  the  outer  layers  being  only  slightly  compressed  or 
torn.  The  endodermis  is  not  definite,  but  the  two  or  more  inner 
rows  of  cells  have  rather  thickened  walls  and  measure  about  .041 
by  .018  mm.  in  diameter,  the  longer  axis  running  tangentially.  In 
longitudinal  sections,  all  cortical  cells  are  somewhat  elongated  and 
many  are  two  times  as  long  as  they  are  broad. 

As  is  the  case  in  the  phloem  of  the  stem,  the  sieve  tubes  and  com- 
panion cells  are  very  small  and  are  massed  in  small  clumps  at  reg- 
ular intervals.  Two  or  three  rows  of  thin-walled  phloem  paren- 
chyma cells  flank  these  clumps  exteriorly  and  complete  a  phloem 
cylinder  which  measures  .073  in  diameter  (PI.  LI,  fig.  19) . 


410  The  University  Science  Bulletin 

SUMMARY 

A  series  of  anomolies  of  growth  and  structure  is  to  be  found  in 
members  of  the  families  Chenopodiacese,  Aizoaceae,  Phytoloccacese, 
Nyctaginacese,  and  their  well-known  congeners.  Successive  rings 
of  distinct  vascular  bundles  appear  in  the  stems  and  even  in  the 
root  and  bring  about  secondary  thickening.  An  intermediate  or 
interfascicular  tissue  separates  the  zones  of  growth  (De  Bary,  1884) . 
Some  species  of  these  families,  however,  have  normal  growth,  for 
example  Rivina  brasilienses  and  R.  aurantiaca  among  the  Phyto- 
laccacese.  With  these  M.  verticillata  can  be  classed,  for  the  root  and 
stem,  while  they  show  some  peculiarities,  are  not  anomolous  in  the 
above  sense. 

Both  stem  and  root  of  Mollugo,  at  least  in  the  older  portions,  ap- 
pear to  have  annual  rings.  In  the  stem  the  amount  of  tissue,  as  well 
as  the  structure,  varies  between  nodes  and  internodes,  the  cortical 
parenchyma  being  nearly  twice  as  thick  at  the  nodes.  The  bast 
fibers  at  internodes  appear  as  a  broken  irregular  cylinder,  1  or  2 
cell  layers  in  thickness.  At  internodes  it  forms  a  more  prominent 
cylinder  with  an  undulating  margin  which  varies  from  1  to  6  cells 
in  thickness,  although  gaps  occur  here  and  there.  There  are  no 
medullary  rays.     Other  tissues  of  the  stem  are  relatively  normal. 

In  the  pseudoannual  rings  of  the  root,  the  zones  of  tracheal  tubes 
are  separated  from  each  other  by  rows  of  lignified  tracheids.  The 
units  which  form  the  tracheal  tubes  are  heavily  pitted,  and  sharp 
remnants  of  the  cross  walls  remain.  Between  the  tracheal  tubes 
are  groups  of  nonlignified  fiber-tracheids  which  remain  alive  and 
evidently  substitute  for  xylem  parenchyma. 

Simple  and  compound  crystals  of  calcium  oxalate  are  abundant 
in  the  mesophyll  tissue  of  the  leaf,  in  the  cortical  tissue  of  the  stem, 
and  are  found,  not  infrequently,  in  xylem  tubes  of  root  and  stem. 
Crystal  sand  occurs  in  a  few  of  the  epidermal  cells.  The  myriad 
minute  crystals  in  the  mesophyll  generally  have  definite  shape. 

Anatomical  modifications,  probably  held  in  common  with  more 
highly  specialized  xerophytic  ancestors,  are  found  in  the  thickened 
leaf  whose  middle  portion  is  occupied  by  water-storing  parenchyma 
cells,  and  in  the  vestigial  trichome-bases  which  occur  among  the 
typical  sinuous  epidermal  cells. 


Payne:  Mollugo  Verticillata  L.  411 


LITERATURE  LIST 

DeBary,  a.    1884.   Comparative  Anatomy  of  the  Vegetative   Organs  of  the 

Phanerogams  and  Ferns.    Clarendon  Press,  Oxford,  pp.  137,  567,  590. 
Ford,  W.  E.    1922.    Dana's  Textbook  of  Mineralogy.     John  Wiley  and  Sons, 

Inc.,  London,  pp.  133,  236. 
Gray,  A.   1887.   School  and  Field  Botany.    American  Book  Co.,  N.  Y.,  p.  199. 
1908.   New  Manual  of  Botany.     American  Book  Co.,  N.  Y.,  p.  24. 
International  Critical  Tables   of   Numerical  Data — Physics,   Chemistry 

AND  Technology.    Vol.  VII,  p.  24. 
Minnesota  Botanical  Studies.    1898.    Conway.    Macmillan  Co.,  N.  Y. 
Pax,  W.    1892.    Engler  and  Prantl.    Pflanzenfamilien,  III,  lb,  pp.  33,  39. 
Rendlb,   A.   B.    1925.   The   Classification   of   the  Flowering   Plants,  Vol.   11; 

Dicotyledons;  p.  8.    Cambridge  University  Press. 
Riedbr,  H.,  and   Moore,  F.   C.    1899.   Atlas  of  Urinary  Sediments.     Charles 

Griffin  &  Co.,  London,  pp.  13,  14,  17.    Plates  I  and  II. 
Stevens,  W.  C.    1924.    Plant  Anatomy.    P.  Blakiston's  Son  &  Co.,  Philadelphia, 

p.  334. 
Zirkle,  C.    1930.    The  Use  of  N-Butyl  Alcohol  in  Dehydrating  Woody  Tissue 

for  Paraffin  Embedding.    Science,  Vol.  LXXI,  No.  1830,  p.  103. 


412  The  University  Science  Bulletin 


PLATE  XLVm 

(Free-hand  and  Projection  Drawings  of  Mollugo  verticillata  L.) 

Fig.     1.   Seed  from  mature  ovary. 

Fig.    2.   Seed  with  testa  removed,  showing  contained  embryo  and  endosperm 

material. 
Fig.    3.    Old  flower,  carpels  and  central  placenta,  etc. 
Fig.    4.    Mature  flower,  sepals  reflexed  to  show  stamen. 
Fig.    5.   Epidermis  from  stem  (X  185). 
Fig.    6.    Mollugo  verticillata  L. 
Fig.    7.    Mesophyll  cells — crystal  sacs  (X  185). 
Fig.    8.   Sclerenchyma  cells  from  stem.    Macerated  tissue  (X185). 
Fig.    9.    Mesophyll  cells  adjacent  to  large  vein  (X160). 
Fig.  10.   Venation  of  leaf  tip  (X  140). 
Fig.  11.   Upper  epidermis  (X185). 
Fig.  12.   Epidermis  over  midrib  (X  185). 

Pig.  13.   Upper  epidermis,  cross  section,  upper  wall  cutinized  (X-160). 
Fig.  14.   Probable  vestigial  epidermal  base  of  trichome  (X  185). 
Fig.  15.    Lower  epidermis.    Cross  section  through  stoma  (X  160). 
Fig.  16.    Lower  epidemial  cells  (X  185). 


Payne:  Mollugo  Verticillata  L. 
PLATE  XLVIII 


438 


414  The  University  Science  Bulletin 


PLATE  XLIX 

(Photomicrographs    of   Stem) 

Fig.  1.    Cross  section  of  stem  through  region  of  node,  second  from  startmg 

point  (X22). 
Fig.  2.   Cross  section   of   stem  just  above  node.     Note  abundant  cortical 

tissue  (X22). 
Fig.  3.    Cross  section  through  node,  showing  traces  of  three  lateral  branches 

(X22). 
Fig.  4.    Cross  section  of  stem  from  4th  internode  (X39). 
P'iG.  5.    Cross   section    of   stem    at   node,    showing   broken   bast   fiber   ring 

(X105). 
Fig.  6.    Portion  of  Fig.  4  more  highly  magnified.    Bast  fibers  5  to  6  rows  deep 

(X95). 
Fig.  7.    Cross  section  through  large  stem  at  internode.    Pseudoannual  rings 

evident  (X  95). 


Payne:  Mollugo  Verticillata  L. 


415 


PLATE  XLIX 


416  The  University  Science  Bulletin 


PLATE  L 

(Pholoiiiicrographs  of  Flower,  Leaf,  e'.c.) 

Fig.  S.  Rosette  crystals  and  simple  rhombolicdral  crystals  in  tissue  of  leaf 
cleared  in  chloral  hydrate   (X  126). 

Fig.  9.  Longitudinal  section  through  flower.  Central  placenta,  short  style, 
seeds  in  carpels,  etc.  The  cuter  seed  coat  is  becoming  cutinized 
(X60). 

Fig.  10.  Cross  section  of  flower  showing  loci  and  arrangement  of  sepals, 
carpels,  placentae,  etc.  (,X  80). 

Fig.  11.  Upper  third  of  leaf  showing  crystals  in  the  cortical  tissue  (X  112). 
Cross  section  of  petiole. 

Fig.  12.  Cross  section  of  leaf.  Note  thickness  due  to  water-storage  par- 
enchyma, and  depression  over  midrib  (X  115). 


Payne:  Mollugo  Verticillata  L 


417 


PLATE  L 


418  The  University  Science  Bulletin 


PLATE  LI 

(Photomicrographs  of  Tissues  of  Stem,  Root,  etc.) 

Fig.  13.    Longitudinal   section   through   phloem  and  cortical   tissue.     8ieve 

tube  is  four  cells  removed  from  the  cortex  (X  372). 
Fig.  14.    Longitudinal  section  through  protoxylem  and  metaxylem  of  stem 

(internode,  X372). 
Fig.  15.   Longitudinal  section  of  stem  showing  crystal  sand  in  xylem  duct 

(X89). 
Fig.  16.   Stomata  in  epidermis  of  stem  (Xl^O). 
Fig.  17.   Bordered  pits  in  the  walls  of  tracheal  tube  and  adjacent  tracheids 

(X372). 
Fig.  18.    Cross   section    of   large     root    showing    pseudoannual     rings,    etc. 

(X130). 
Fig.  19.    Cross  section  of  root  showing  origin  of  secondary  rootlet  (X  130). 


Payne:    Molligo  Verticillata  L. 


419 


PLATE  LI 


THE  UNIVERSITY  OF  KANSAS 

SCIENCE  BULLETIN 

Vol.  XXL]  March,  1933.  [No.  13. 


Morphology  and  Anatomy  of  Lygodesmia  juncea 
(Pursh)  D.  Don. 

LIZA  SPANN 


Abstract:  Lygodesmia  juncea  (Pursh)  D.  Don.,  a  Compositse,  has  small 
leaves.  Stomata  occur  on  both  surfaces.  The  stomata  of  the  aboveground 
stem  are  almost  as  numerous  as  those  of  the  leaf.  The  aboveground  stem  has 
a  palisade  chlorenchyma  which  is  significant  in  photosynthesis. 

The  tissue  organization  of  the  aboveground  stem,  underground  stem  and 
root  are  discussed.  Laticiferous  tubes  traverse  the  length  of  the  plant  as  a 
continuous  anastomosing  system  occurring  in  the  phloem  and  pericycle.  An 
outstanding  characteristic  of  this  plant  is  that  the  leaf  trace  has  two  origins: 
(a)  the  part  which  forms  the  midrib  of  the  leaf  is  derived  from  a  bundle  of 
protoxylem  tubes  which  swing  sharply  out  and  pass  directly  into  the  leaf;  (6) 
the  part  which  forms  the  primary  veins  parallel  to  the  midrib  is  derived  from 
protoxylem  tubes  that  come  out  from  the  xylem  cylinder  considerably  below 
the  leaf  insertion. 

Resin  is  found  in  the  laticiferous  tubes,  parenchyma  cells  of  leaf,  cortex  of 
the  stem  and  tracheal  tubes  of  the  root.  Glucose  is  present  throughout,  but 
is  most  abundant  in  the  leaf  and  aboveground  stem.  A  glucoside,  lygodesmin. 
occurs  in  the  root,  stem  and  leaf.  Saccharose  is  present  only  in  the  above- 
ground  stem  and  leaf. 

Various  points  in  the  paper  are  illustrated  with  thirty-one  projection  draw- 
ings and  fourteen  photomicrographs. 


CLASSIFICATION  AND  DISTRIBUTION 

T  YGODESMIA  juncea  (Pursh)  D.  Don.  belongs  to  the  Chichorite 
■■-'  tribe  of  the  Compositse.  According  to  Gray  (1908)  Lygodesmia 
is  derived  from  two  Greek  words,  lugos,  a  pliant  twig,  and  desme, 
a  bundle,  from  the  fascicled  twiggy  or  rushlike  stems.  L.  juncea 
is  the  type  species  of  the  genus.  Britton  and  Brown  (1913)  relate 
that  about  six  species  are  natives  of  western  and  southern  North 
America,  but  they  describe  only  two  species,  L.  juncea  (Pursh)  D. 
Don.  and  L.  rostrata  Gray,  the  former  inhabiting  the  plains  from 
28—3482 

(421) 


422  The  University  Science  Bulletin 

Minnesota  to  Saskatchewan,  Wisconsin,  Missouri,  Nebraska  and 
Arizona  and  the  latter  occurring  on  plains  and  in  canyons  from 
South  Dakota  to  Saskatchewan,  Nebraska,  Kansas,  Colorado  and 
Wyoming. 

Coulter  and  Nelson  (1909)  give  four  species:  L  grandiflcra  T.  & 
G.,  L.  Spinosa  Nutt.  and  the  two  above-mentioned  sjiecies.  They 
say  that  L.  spinosa  is  distributed  on  gravelly  hills  and  plains  from 
eastern  Oregon  to  California,  Nevada  and  Idaho,  and  that  L.  grandi- 
flora  is  distributed  on  gravelly  hills  in  Wyoming  and  Colorado  to 
Utah. 

Chapman  (1897)  mentions  only  one  species,  L.  aphylla  D  C,  as 
occurring  in  the  southern  United  States,  specifically  on  dry,  sandy 
pine  barrens  in  Georgia  and  Florida. 

HABITAT  AND  GENERAL  MORPHOLOGY 

Lygodesmia  juncea  grows  in  sandy  soils  on  hills  that  are  exposed 
to  the  sun  and  on  sandy  plains.  The  particular  plant  upon  which 
this  work  is  based  was  collected  by  Prof.  W.  C.  Stevens  in  August 
in  northeastern  Kansas  on  a  bluff  near  the  Missouri  river. 

This  species  is  a  perennial.  The  stiff,  glabrous,  striate,  much- 
branched  stems  are  from  2  to  4  dm.  high.  The  diameter  of  the  main 
branches  at  their  bases  is  about  2.5  mm.  The  stem  gradually  tapers 
toward  the  apex,  the  average  diameter  there  being  0.5  mm.  The 
average  length  of  the  internodes  is  2  cm. 

The  leaves  are  rigid,  sessile  and  opposite.  The  lower  leaves  are 
acutely  linear-lanceolate  and  entire.  The  average  width  of  the 
basal  leaves  is  3.5  mm.  and  the  average  length  is  4.5  cm.  The  upper 
leaves  are  smaller,  averaging  0.5  mm.  in  width  at  the  base  and  2 
mm.  in  length,  or  they  may  be  reduced  to  subulate  scales. 

The  underground  stem  extends  to  a  considerable  depth  below  the 
surface.  Its  diameter  ranges  from  2  to  3  mm.  The  internodes  are 
about  2.5  cm.  in  length.  At  the  nodes  are  opposite,  scale-like  leaves 
which  average  4  mm.  in  width  at  the  base  and  7  mm.  in  length.  The 
scale  gradually  tapers,  forming  an  acute  apex. 

MATERIALS  AND  METHOD 

Several  methods  of  procedure  were  followed  in  preparing  the  tissue 
for  microscopic  examination.  The  specimen  had  been  preserved  in 
thirty  per  cent  alcohol.  Small  pieces  of  the  stem,  root  and  leaf  were 
dehydrated  and  embedded  in  paraffin  by  the  N-butyl  alcohol  method 
of  Zirkle  (1930)  and  then  cut  in  cross  and  longitudinal  sections 
varying  from  ten  to  forty  microns  in  thickness.  The  leaves  and 
some  of  the  stem  sections  were  cleared  in  chloral  hydrate. 


Spann:  Lygodesmia  Juncea  423 

Three  reagents  were  used  for  the  kinds  of  cell  walls:  Sudan  III 
for  cutinized  and  suberized  walls,  chloroiodide  of  zinc  for  cellulose 
walls,  and  phloroglucin  and  hydrochloric  acid  for  lignified  walls. 
For  cell  contents  Fehling's  solution  was  used  for  glucose,  and  sul- 
phuric acid  and  Fehling's  solution  for  glucosides.  Czapek  (1897) 
and  Hoffmeister's  (1898)  invertase  method  was  used  for  saccharose. 
To  distinguish  between  fats  and  resins  the  saponification  test  of 
Moenikes  (1924)  was  run  and  several  solvents,  such  as  alcohol  and 
carbon  disulphide,  were  used. 

Sections  for  photomicrographs  were  dehydrated  and  cleared  in 
toluol,  and  then  mounted  in  hyrax.  Exposures  varying  from  fifteen 
seconds  to  three  minutes  with  an  arc  light  illuminant  were  made  on 
hard-surfaced  sensitized  paper,  giving  a  negative  print.  The  draw- 
ings were  made  with  an  arc-light  projectoscope. 

ABOVEGROUND  STEM 

The  stem  has  two  kinds  of  epidei-mal  cells.  Those  over  the  bast 
fibers  are  vertically  elongated  and  rectangular  with  regular  margins 
and  have  no  stomata  (PL  LII,  fig.  1),  while  those  over  the  chlorcn- 
chyma  are  relatively  large  with  irregular  margins  and  have  stomata. 
The  outer  cell  wall  of  the  epidermis,  as  seen  in  cross  section,  is 
slightly  convex,  and  the  inner  wall  is  flat.  The  radial  diameters  of 
the  cells  average  0.02  mm.  and  the  tangential  diameters  vary  from 
0.025  mm.  to  0.05  mm.,  the  shortest  cells  being  in  front  of  the  bast 
fibers.  The  epidermis  has  a  thin  cuticle  and  the  outer  cell  walls  are 
cutinized  about  half  across.  These  walls  are  0.006  mm.  thick,  but 
the  inner  and  radial  walls  are  thinner  and  are  cellulose  throughout. 
The  average  number  of  stomata  per  sq.  mm.  over  the  entire  surface 
of  the  stem  is  thirty.  In  areas  between  the  bast  fibers  the  average 
is  forty-five  per  sq.  mm. 

The  parenchyma  of  the  cortex  is  in  the  form  of  palisade  chloren- 
chyma  (PI.  LII,  fig.  15),  three  to  four  cell  layers  thick,  the  diameters 
of  the  cells  averaging  0.016  mm.  tangcntially  and  0.039  radially. 
The  walls  are  cellulose  and  very  thin.  Along  with  chloroplasts  are 
droplets  of  resin.  Bundles  of  bast  fibers  alternate  with  patches  of 
chlorenchyma  of  the  cortex  (Pi.  LIV,  fig.  2).  The  bast  fibers  are 
lignified  and  are  0.007  mm.  thick  and  from  2  to  2.5  mm.  long  (PI. 
LII,  fig.  3). 

Beneath  the  chlorenchyma  of  the  cortex  is  a  definite  endodermis 
(PI.  LV,  figs.  3  and  4,  and  PI.  LIV,  fig.  2)  whose  cells  have  radial 
diameters  of  0.017  to  0.025  mm.  and  tangential  diameters  averaging 


424  The  University  Science  Bulletin 

0.04  mm.     The  walls  are  cellulose  and  are  0.002  mm.  thick.     The 
endodermis  contains  droplets  of  resin  (PL  LII,  fig.  19). 

Bast  fibers  also  occur  in  the  pericycle  just  beneath  the  bundle  of 
bast  fibers  in  the  cortex  (PI.  LIV,  fig.  2).  These  are  thicker  walled 
than  those  of  the  cortex. 

According  to  Col  (1904)  the  secreting  system  is  found  existing 
in  three  distinct  forms  in  the  Compositae:  (1)  anastomosing  latex 
tubules,  (2)  secreting  canals,  and  (3)  cells  secreting  latex.  The 
form  existing  in  this  stem  is  of  the  first  type.  The  tubes  form  a 
much-branched  anastomosing  system  throughout  the  phloem  and 
pericycle  (PI.  LIII,  fig.  9)  and  contain  a  resinous  secretion. 

The  vascular  tissue  is  in  the  form  of  a  cylinder  surrounding  a 
relatively  large  pith.  The  phloem  elements  are  very  minute  with 
correspondingly  thin  walls.  The  parenchyma  cells  are  isodiametric 
in  cross  section,  having  an  average  diameter  of  0.013  mm.  The 
sieve  tubes  give  a  distinct  protein  reaction. 

The  xylem  cylinder  is  composed  of  two  distinct  parts  (PI.  LIV, 
fig.  2) :  that  part  which  contains  the  protoxylem  points  and  is  rela- 
tively broad  radially,  having  protoxylem,  metaxylem  and  second- 
ary xylem,  is  made  up  of  tracheal  tubes,  xylem  parenchyma,  and 
fiber-tracheids ;  and  that  part  which  is  radially  narrow,  embracing 
metaxylem  and  secondary  xylem  only,  is  made  up  of  xylem  paren- 
chyma cells  (PL  LIII,  fig.  3)  which  are  0.08  mm.  long  and  0.02  mm. 
wide  in  longitudinal  section,  and  fiber-tracheids  which  are  0.2  mm. 
long  and  0.01  mm.  wide.  The  tracheal  tubes  have  thick,  pitted- 
lignified  walls  (PL  LIV,  fig.  3)  and  have  an  average  diameter  of 
0.058  mm.    The  protoxylem  tubes  average  0.016  mm.  in  diameter. 

The  leaf  traces  have  two  origins.  Those  vascular  elements  which 
form  the  midrib  of  the  leaf  come  from  the  primary  xylem  and  are 
composed  of  protoxylem  only  (PL  LIII,  fig.  2).  These  bundles  of 
protoxylem  pass  up  the  stem  and  make  an  abrupt  turn  out  into  the 
leaf,  while  those  which  enter  the  leaf  on  either  side  of  the  midrib 
pass  out  from  the  xylem  considerably  below  the  leaf  insertion, 
travel  up  the  stem  just  under  the  phloem,  then,  passing  out  through 
the  phloem  and  pericycle,  ascend  beneath  the  endodermis  (PL  LIII, 
fig.  5)  and  finally  pass  out  into  the  sessile  leaf  and  continue  on 
either  side  of  the  midrib  about  two-thirds  the  length  of  the  leaf.  In 
Plate  LIII,  figure  1  the  relation  of  the  two  origins  of  the  leaf  traces 
are  represented. 

The  spiral  tubes  of  the  traces  on  either  side  of  the  midrib  have 
an  average  diameter  of  0.011  mm.    They  are  found  in  cross  sections 


Spann:  Lygodesmia  Juncea  425 

of  stems  just  beneath  the  phloem,  in  the  phloem  and  pericycle,  and 
just  beneath  the  endodermis,  singly  or  in  bundles  of  from  two  to 
four  strands. 

The  parenchyma  cells  of  the  pith  are  smaller  near  the  periphery 
and  larger  near  the  center.  The  cell  walls  are  slightly  lignified. 
The  cells  are  isodiametric  in  cross  section,  and  their  diameters  vary 
from  0.09  to  0.16  mm.  This  part  of  the  stem  contains  glucose, 
saccharose  and  a  glucoside  which  we  have  called  lygodesmin. 

The  stem  is  infested  by  an  insect  larva  which  causes  a  globose 
gall  varying  from  5  mm.  to  8  mm.  in  diameter.  The  galls  are  in 
groups  of  from  two  to  ten  on  one  side  of  the  stem.  The  epidermal 
cells  (PI.  LIV,  fig.  4)  of  the  gall  are  similar  to  those  of  the  normal 
stem,  but  smaller,  and  the  outer  cutinized  walls  thinner.  Beneath 
the  epidermis  of  the  gall  is  a  layer  of  chlorenchyma  containing 
chloroplasts  and  resin.  This  layer  is  thinner  than  the  corresponding 
layer  of  the  stem  and  contains  no  bast  fibers.  Just  beneath  the 
rather  indistinct  endodermis  in  the  parenchyma  of  the  pericycle 
there  is  a  system  of  anastomosing  latex  tubes  filled  with  resinous 
material.  These  tubes  seem  to  be  scattered  promiscuously  through 
the  gall,  following  the  vascular  system  until  they  reach  the  tissue 
surrounding  the  insect  larva.  Here  they  form  a  network  around 
the  entire  cavity.  The  vascular  system  in  cross  section  is  arranged 
in  bundles  near  the  periphery,  w^iich  run  in  various  directions 
through  the  stele.  The  bulk  of  the  gall  has  been  formed  by  the 
proliferation  of  the  parenchyma  cells  of  the  stele,  which  are  smaller 
than  the  corresponding  cells  of  the  stem.  The  cell  walls  of  the  outer 
layer  of  pith  cells  are  lignified,  but  the  inner  layers  of  cells  have 
cellulose  walls.  The  inner  cells  are  broken  down,  forming  a  cavity 
in  which  the  insect  resides.  Kiister  (1911),  the  eminent  authority 
on  galls,  does  not  mention  this  one. 

LEAF 
The  leaf  is  parallel  veined  (PI.  LII,  fig.  11)  with  anastomoses 
between  the  veins.  In  surface  view  the  upper  and  lower  epidermal 
cells  of  the  leaf  vary  in  size  and  shape,  but  have  almost  even  margins 
(PL  LII,  figs.  7  and  9).  The  epidermal  cells  over  the  midrib  and 
veins  are  elongated,  rectangular  cells  with  regular  boundaries  (PI. 
LIV,  figs.  7  and  8).  In  cross  section  the  cells  of  the  upper  and 
lower  epidermis  are  about  the  same  size,  averaging  0.03  mm.  in  tan- 
gential diameter  and  0.026  mm.  in  radial  diameter.  The  outer  walls 
of  both  are  partly  cutinized,  but  those  of  the  lower  epidermis  are 


426  The  University  Science  Bulletin 

slightly  thicker  than  those  of  the  upper  (PL  LII,  figs.  13  and  18). 
The  radial  walls  are  thinner  than  the  outer  walls  and  the  inner  walls 
are  the  thinnest. 

Stomata  occur  on  both  surfaces  of  the  leaf.  The  openings  are 
elongate-elliptical  in  outline  and  measure  0.02  mm.  in  length.  The 
stomata  are  only  slightly  more  numerous  on  the  lower  epidermis 
than  on  the  upper,  averaging  53  per  sq.  mm.  on  the  lower  and  51 
per  sq.  mm.  on  the  upper. 

Just  beneath  the  lower  epidermis  is  one  row  of  interrupted  iso- 
diametric  collenchyma  cells.  These  cells  also  occur  at  the  margins 
of  the  leaf  and  are  especially  prominent  beneath  the  midrib  and 
primary  veins  (PI.  LV,  fig.  1).  The  palisade  cells  are  on  each  side 
of  the  leaf  in  two-cell  layers.  The  cells  are  about  three  times  as 
long  as  broad  and,  in  addition  to  the  chloroplasts,  contain  frequent 
droplets  of  resin. 

The  parenchyma  sheath  of  the  veins  contains  resin,  and  besides 
functioning  in  connecting  the  vascular  system  with  the  other  tissues 
it  has  a  considerable  water-storage  capacity. 

The  laticiferous  tubes  (PI.  LIII,  fig.  7)  follow  the  veins  outside  of 
the  parenchyma  sheath  and  form  a  network  between  the  veins. 
DeBary  (1884)  states  that  in  the  ribs  of  leaves  the  main  trunks 
of  latex  tubes  lie  chiefly  in  the  tissue  bordering  the  phloem  portions  of 
the  vascular  bundles,  following  the  longitudinal  course  of  the  latter, 
and  as  seen  in  cross  section  are  scattered  without  strict  regularity 
among  the  surrounding  parenchyma.  This  is  correct  for  this  leaf. 
As  in  the  stem,  the  tubes  are  filled  with  resinous  material.  Glucose, 
a  glucoside,  an  abundance  of  saccharose,  and  resin  are  found  in  the 
leaf. 

UNDERGROUND  STEM 

The  outer  cell  walls  of  the  epidermis  are  cutinized  and  thinner 
than  the  corresponding  cell  walls  of  the  aboveground  stem.  In 
cross  section  the  radial  and  tangential  diameters  of  these  cells 
measure  about  the  same,  being  about  0.02  mm.  There  is  one  row 
of  collenchyma  cells  just  beneath  the  epidermis  (PI.  LII,  fig.  17), 
which  is  the  only  strengthening  tissue  of  the  cortex. 

The  cortex,  excluding  the  collenchyma,  is  made  up  of  about  nine 
layers  of  parenchyma  cells  with  lignified  walls  (seen  in  cross  section 
in  PI.  LIV,  fig.  1,  and  in  longitudinal  section  PI.  LV,  fig.  5).  The 
diameters  of  the  cells  vary  from  0.033  to  0.05  mm.,  the  smaller  ones 
being  near  the  periphery.  Following  this  are  three  layers  of  cork 
cells  (PI.  LV,  fig.  5)  which  have  originated  from  the  cells  of  the 


Spann:  Lygodesmia  Juncea  427 

pericycle.  The  tangential  diameter  of  these  cells  is  0.03  mm.  and 
the  radial  diameter  is  0.012  mm.    The  cell  walls  are  suberized. 

The  parenchyma  cells  of  the  pericycle  have  thin  cellulose  walls. 
The  latex  tubes  (PI.  LIII,  fig.  8)  form  a  richly  anastomosing  system 
extending  out  into  the  pericycle,  but  are  more  numerous  in  the 
phloem.  The  phloem  is  otherwise  composed  of  phloem  parenchyma 
and  sieve  tubes. 

The  xylem  tissues  form  a  continuous  cylinder  around  the  pith  (PI. 
LIV,  fig.  1).  The  cylinder  is  composed  of  two  parts,  as  in  the  above- 
ground  stem:  metaxylem  parenchyma  cells  and  fiber  tracheids 
alternating  with  groups  of  metaxylem  subtended  by  the  protoxylem. 
The  parenchyma  cells  of  the  pith  in  cross  section  are  large,  iso- 
diametric  cells  with  lignified  walls. 

The  anomalous  protoxylem  tubes  are  also  found  in  the  under- 
ground stem,  coming  out  from  the  xylem,  traveling  up  the  stem  just 
outside  the  xylem  cylinder,  crossing  through  the  phloem  and  peri- 
cycle, and  up  the  stem  under  the  cork  cells. 

The  glucoside,  lygodesmin,  is  abundant  in  the  parenchyma  cells 
of  the  cortex.  Glucose  is  not  as  abundant  as  in  the  leaves  and 
aboveground  stem,  and  saccharose  is  not  present. 

ROOT 

In  the  root  section  shown  in  Plate  LIV,  figure  5,  cork  cells  have 
been  formed  and  have  cut  off  all  the  cells  outside  the  pericycle.  The 
cell  walls  of  this  tissue  are  thin  and  suberized.  In  cross  section  the 
cells  have  a  radial  diameter  of  0.02  mm.  and  a  tangential  diameter 
of  0.05  mm. 

Beneath  the  cork  cells  are  parenchyma  cells  of  the  pericycle  with 
thin,  cellulose  walls.  The  latex  tubes  form  a  network  (PL  LII,  fig. 
4)  extending  out  into  this  layer  of  cells  and  into  the  phloem,  being 
more  numerous  in  the  phloem.  According  to  Col  (1904)  the  laticif- 
erous  tubes  in  the  Compositse  are  always  situated  in  the  pericycle  of 
the  stem,  in  the  primary  phloem  of  the  root,  and  in  the  secondary 
phloem  of  both  these  organs.  This  plant  is  an  exception  in  that  the 
laticiferous  tubes  occur  in  the  pericycle  and  phloem  of  both  the  root 
and  stem.  Mocnikcs'  (1924)  statement  that  in  all  the  compositse 
no  mucilage  is  laid  down  in  the  secretion  passages  is  not  contradicted 
by  this  plant. 

The  xylem  tissues  of  the  root  are  divided  into  wedges  by  rays 
which  vary  from  one  to  three  cells  across  tangentially  (PL  LIV,  fig. 
5).    The  ray  cells  (PL  LIII,  fig.  6j  are  about  twice  as  long  radially  as 


428  The  University  Science  Bulletin 

tangentially.  They  have  thin  cellulose  walls.  The  diameters  of 
the  tracheal  tubes  are  larger  than  those  of  the  stem  (PI.  LII,  figs. 
5  and  6) ,  varying  from  0.033  mm.  to  0.083  mm.  Some  of  the  tracheal 
tubes  next  to  the  phloem  are  filled  with  resinous  material  (PI.  LII, 
fig.  16)  similar  to  that  in  the  latex  tubes.  The  protoxylem  elements 
have  a  rather  compact  spiral  and  are  about  the  same  size  as  those 
of  the  stem.  Between  the  tracheal  tubes  throughout  the  xylem  are 
parenchyma  cells  and  fiber  tracheids.  The  anomalous  protoxylem 
tubes  are  found  occasionally  in  the  root  in  the  pericycle  just  beneath 
the  cork  cells.  Glucose  and  a  glucoside  are  present  in  the  root,  but 
no  saccharose  was  found. 

SUMMARY 

Lygodesmia  juncea  (Pursh)  D.  Don.,  a  member  of  the  Composite, 
is  a  rush-like  perennial  about  4  dm.  high.  There  are  six  species  of 
this  genus  native  of  the  United  States. 

The  upper  and  lower  epidermis  of  the  leaf  and  the  epidermis  of 
the  aboveground  stem  are  very  much  alike,  having  alternating  longi- 
tudinal rows  of  irregular  cells  with  stomata  and  of  rectangular  cells 
without  stomata,  the  latter  overlying  and  underlying  the  main  veins. 
Again,  the  parenchyma  cells  of  the  cortex  of  the  stem  are  similar 
to  the  parenchyma  cells  of  the  leaf  in  character  of  walls,  shape, 
chloroplasts,  and  resin  content. 

The  stem  above  ground  is  the  only  part  of  the  plant  that  has  bast 
fibers,  which  occur  in  the  cortex  and  pericycle.  The  stem  is  infested 
by  an  insect  larva  which  causes  a  globose  gall. 

Laticiferous  tubes  traverse  the  entire  body  of  the  plant  as  a  con- 
tinuous anastomosing  system  occurring  in  the  phloem  and  pericycle. 
The  tubes  are  filled  with  resinous  material. 

As  to  cell  contents,  resin  is  the  most  abundant  material,  since  it 
is  found  in  the  laticiferous  tubes  of  all  organs  of  the  plant,  in  the 
parenchyma  cells  of  the  leaf,  the  cortex  of  the  stem,  and  the  tracheal 
tubes  of  the  root.  Glucose  is  present  in  all  the  organs,  but  is  most 
abundant  in  the  leaf  and  in  the  aboveground  stem.  A  glucoside, 
lygodesmin,  occurs  in  all  of  the  organs.  Saccharose  is  present  only 
in  the  aboveground  stem  and  in  the  leaf,  being  most  abundant  in 
the  leaf.    The  phloem  contains  protein. 

The  outstanding  characteristic  of  this  plant  is  that  the  leaf  trace 
has  two  origins:  (1)  a  part  which  forms  the  midrib  is  derived  from 
a  bundle  which  separates  from  the  vascular  system  of  the  stem  and 
swings  abruptly  outward  into  the  leaf,  and  (2)  a  part  which  forms 


Spann:  Lygodesmia  Juncea  429 

the  primary  veins  parallel  to  the  midrib,  the  tracheal  elements  of 
these  veins  being  protoxylem  tubes  which  separate  from  the  vascular 
cylinder  of  the  stem  considerably  below  the  leaf  insertion,  pass  up 
the  stem  outside  the  xylem  cylinder,  cross  through  the  phloem  and 
pericycle,  ascend  the  stem  beneath  the  endodermis  for  some  distance, 
and  then  pass  out  into  the  leaf. 

In  conclusion,  the  writer  wishes  to  express  her  appreciation  to 
Prof.  W.  C.  Stevens  for  his  suggestions,  helpful  criticisms  and  ad- 
vice during  the  entire  progress  of  this  investigation. 


430  The  University  Science  Bulletin 


PLATE  LIT 

Projection  Drawings  of  Lygodesmia  juncca  (Pursh)   D.  Don. 

Fig.  1.    Epidermis  of  the  aboveground  stem  (X140). 

Fig.  2.   Cross  section  of  the  pith  cells  of  the  aboveground  stem  (X  140). 

Fig.  3.   Bast  fiber  from  the  aboveground  stem  (X40). 

Fig.  4.   Laticiferous  tubes  of  the  root  (X40). 

Fig.  5.    Elements  from  metaxylem  and  protoxylem  of  the  root  (X  140). 

Fig.  6.  Elements  from  metaxjdem  and  protoxylem  of  the  aboveground 
stem  (X  140). 

Fig.  7.   Upper  epidermis  of  the  leaf  (X  140). 

Fig.  8.  Longitudinal  section  of  the  pith  cells  of  the  aboveground  stem 
(X140). 

Fig.  9.   Lower  epidermis  of  the  leaf  (X  140). 

Fig.  10.   Tip  of  the  leaf  (X8). 

Fig.  1L   Base  of  the  leaf,  showing  parallel  veins  (X8). 

Fig.  12.   Cross  section  of  the  cork  cells  of  the  root  (X  140)- 

Fig.  13.  Cross  section  of  the  upper  epidermis  of  the  leaf,  showing  a  stoma 
(X140). 

Fig.  14.   Cross  section  of  the  metaxylem  tracheal  tubes  of  the  root  (X  140). 

Fig.  15.  Cross  section  of  the  parenchyma  cells  of  the  cortex  of  the  above- 
ground  stem  (X  185). 

Fig.  16.  Semidiagrammatic  drawing  from  the  cross  section  of  the  root, 
showing  some  xylem  tubes  filled  with  resin  (X  17). 

Fig.  17.  Cross  section  of  the  epidermis  and  collenchyma  of  the  underground 
stem  (X  185). 

Fig.  18.  Cross  section  of  the  lower  epidermis  of  a  leaf,  showing  a  stoma 
(X140). 

Fig.  19.  Cross  section  of  the  endodermis  of  the  aboveground  stem,  showing 
resin  in  the  cells  (X  185). 

Fig.  20.  Cross  section  of  the  parenchyma  cells  of  the  cortex  of  the  under- 
ground stem  (X  185). 

Fig.  21.   Cross  section  of  cork  cells  of  the  underground  stem  (X  140). 

Fig.  22.  Cross  section  of  a  bundle  of  bast  fibers  from  the  aboveground  stem 
(X  140). 


Spann:  Lygodesmia  Juncea 


431 


PLATE  LII 


W^l 


432  The  University  Science  Bulliotin 


PLATE  LIIl 

Projection  Drawings 

Fig.  1.  Semidiagrammatic  drawing  of  a  cross  section  of  the  aboveground 
stem,  showing  a  leaf  gap,  midrib  trace,  and  protoxylem  tubes  passing  out  of 
the  pericycle  to  enter  veins  parallel  to  the  midrib  (X8). 

Fig.  2.  Semidiagrammatic  drawing  of  the  longitudinal  section  of  the  above- 
ground  stem  showing  a  leaf  and  branch  gap,  a  midrib  trace,  and  protoxylem 
tube  in  the  pericycle  (X  16). 

Fig.  3.  Longitudinal  section  of  xylem  parenchyma  cells  of  the  aboveground 
stem  (X  140). 

Fig.  4.  Longitudinal  section  of  the  pith  cells  of  the  underground  stem 
(X185). 

Fig.  5.  Semidiagrammatic  drawing  of  the  longitudinal  section  of  the  above- 
ground  stem  showing  a  protoxylem  tube  which  contributes  to  a  vein  parallel 
to  the  midrib,  passing  through  the  xylem,  the  phloem,  and  the  pericycle,  and 
up  the  stem  under  the  endodermis  (X  120). 

Fig.  6.   Cross  section  of  the  ray  cells  of  the  root  (X  185). 

Fig.  7.    Laticiferous  tubes  of  the  leaf  (X40). 

Fig.  8.    Laticiferous  tubes  of  the  underground  stem  (X40). 

Fig.  9.  Laticiferous  tubes  of  the  aboveground  stem  (X40).  In  the  semi- 
diagrammatic drawings  1,  2  and  5  the  xylem  is  shaded  with  parallel  lines,  the 
phloem  and  pericycle  are  stippled,  midrib  trace  is  cross-hatched,  and  the 
protoxylem  tubes  are  solid  black. 


Spann:  Lygodesmia  Juncea 
PLATE  LIII 


433 


434  The  University  Science  Bulletin 


PLATE  LIV 

Photomicrographs  of  Root  and  Stem  Sections  and  Epidermis 

Fig.  1.   Cross  section  of  the  underground  stem  (X30). 
Fig.  2.   Cross  section  of  the  aboveground  stem  (X30). 
Fig.  3.   Longitudinal  section  of  the  aboveground  stem,  showing  the  pith, 
protoxylem,  and  metaxylem  (X  105). 
Fig.  4.    Cross  section  of  a  gall  (X30). 
Fig.  5.   Cross  section  of  root  (X25). 
Fig.  6.   Epidermis  of  the  aboveground  stem  (X30). 
Fig.  7.    Lower  epidermis  of  the  leaf  (X30). 
Fig.  8.   Upper  epidermis  of  the  leaf  (X30). 


Spann:  Lygodesmia  Juncea 
PLATE  LIV 


435 


«€P 


Jl?^-. 


436  The  University  Science  Bulletin 


PLATE  LV 

Photomicrographs  of  Leaf  and  Stem  Sections 

Fig.  1.   Cross  section  of  a  leaf  (X30). 

Fig.  2.  Cross  section  of  a  leaf,  showing  a  stoma  in  the  lower  epidermis 
(X  105). 

Fig.  3.  Longitudinal  section  of  aboveground  stem,  showing  the  protoxylem 
tubes  just  beneath  the  endodermis  (X  105). 

Fig.  4.  Longitudinal  section  of  the  aboveground  stem,  showing  a  stoma, 
chlorenchyma,  endodermis,  one  or  two  rows  of  pericycle  cells,  and  protoxylem 
tubes  just  inside  the  phloem  (X  105). 

Fig.  5.   Longitudinal  section  of  underground  stem  (X30). 

Fig.  6.  Longitudinal  section  of  the  underground  stem,  showing  the  phloem 
and  metaxylem  (X  105). 


Spann:  Lygodesmia  Juncea 


437 


PLATE  LV 


•t^i^^':'H¥ 


A 

i     tlflin'i 

111  Mp  ill 


29-3482 


438  The  University  Science  Bulletin 


BIBLIOGRAPHY 

1.  Britton  and  Brown.    1913.    An  Illustrated  Flora  of  the  Northern  United 

States,  Canada,  and  the  British  Possessions.     Charles  Scribner's  Sons, 
N.  Y.,  Vol.  Ill,  p.  322. 

2.  Chapman,  A.     1897.     Flora  of  the  Southern  United  States.     Cambridge 

Botanical  Supply  Co.,  Cambridge,  Mass.,  p.  275. 

3.  CoL.  A.    1904.    Recherches  sur  I'appareil  secreteur  interne  des  Composees. 

Journal  de  Botanique.    18e  Annee.    No.  5,  p.  153. 

4.  Coulter  and  Nelson.     1909.    New  Manual  of  Rocky  Mountain  Botany. 

American  Book  Co.,  N.  Y.,  p.  591. 

5.  CzAPEK.  F.     1897.     Uber  die  Leitungswege  der  organischen  Baustoffe  im 

Pflanzenkbrper.    Sitzgsber.    Wiener  Akad.,  CVI,  1,  Sep.,  S.  14. 

6.  De  Bary,  A.     1884.     Comparative  Anatomy  of  the  Vegetative  Organs  of 

the  Phanerogams  and  Ferns.     Oxford,  at  the  Clarendon  Press,  p.  432. 

7.  Gray,  A.     1908.     New  Manual  of  Botany.     American  Book  Co.,  N.  Y., 

p.  868. 

8.  KxJSTER,  E.  1911.  Die  Gallen  Der  Pflanzen.  Verlag  Von  S.  Hirzel,  Leipzig. 

9.  Moenikes,  a.     1924.     Zur  Frage  der  Harzbildung  bei  den  Umbelliferen, 

Compositen   und   Araliaceenwurzeln.     Botanisches   Archiv.   Band.     V, 
p.  91. 

10.  HoFFMEiSTEE,  C.     1898.    Uber  den  mikrochemischen  Nachweis  von  Rohr- 

zucker  in  pflanzlichen  Geweben.    Jahrb.  f.  Wiss.  Bot.,  XXXI,  S.  688. 

11.  Stevens,  W.  C.    1924.    Plant  Anatomy.    P.  Blakiston's  Son  &  Co.,  Phila- 

delphia, pp.  324,  328,  342. 

12.  ZiRKLB,  C.     1930.     The  use  of  N-Butyl  Alcohol  in  Dehydrating  Woody 

Tissue  for  Paraffin  Embedding.    Science,  Vol.  LXXI,  No.  1830,  p.  103. 


THE  UNIVERSITY  OF  KANSAS 

SCIENCE  BULLETIN 

Vol.  XXI.]  March,  1933.  I  No.  14. 


Comparative  Anatomy  of  Mentzelia  oligosperma  and 
M.  decapetala 

(Contribution  from  the  Botanical  Laboratories  of  the  University  of  Kansas) 
W.  E.  BOOTH 


Abstract:  The  gross  morphology  of  the  two  species  and  detailed  account 
of  their  microscopic  anatomy  are  given.  Microchemical  tests  are  reported 
showing  the  kinds  of  cell  walls  and  cell  contents.  The  principal  anatomical 
differences  between  the  two  species  are  pointed  out.  The  two  species  are  shown 
to  be  alike  in  the  origin  of  the  phellogen  of  stem  and  root,  growth  in  diameter 
of  pericycle  of  root  and  stem,  silicified  trichomes,  predominance  of  wood 
fibers  in  the  xylem,  and  in  fatty  oils  and  proteins  in  the  endosperm  of  seeds. 
The  species  are  shown  to  differ  in  stored  corbohydrates — starch  in  oligosperma, 
glucosides  in  decapetala — in  the  presence  of  palisade  cells  in  the  cotyledons  of 
oligosperma  and  their  absence  in  decapetala,  in  the  smaller  average  of  cell 
size  in  oligosperma,  and  in  unlike  character  of  stone  cells  of  the  seed  testas. 
The  paper  is  illustrated  by  plates  embracing  thirty-two  drawings,  fifteen 
photomicrogiaphs  and  two  habit  photographs. 


GEOGRAPHICAL  DISTRIBUTION 

Mentzelia  oligosperma  Nutt,  Mentzelia  decapetala   (Pursh)  Urban  and  Gelg. 

'\/TENTZELIA  is  a  member  of  a  comparatively  small  family, 
-*•'-*•  Loasacese,  which  is  characterized  particularly  by  stinging 
or  barbed  hairs,  and  a  flower  structure  similar  to  that  of  the  cacti. 
This  genus  is  the  only  representative  of  its  family  found  in  this 
region  (4),  except  for  species  formed  by  a  division  of  the  above 
species  (5). 

Mentzelia  was  named  by  Linnaeus  in  1753  for  Mentzel,  a  German 
botanist  (I ) .  Only  two  species  of  Mentzelia  are  known  to  be  in  our 
range  (Kansas),  but  twelve  species  are  described  for  the  Central 
Rocky  Mountain  region  (2),  and  thirty-four  in  that  part  of  the 
United  States  north  of  New  Mexico  (3). 

(439) 


440  The  University  Science  Bulletin 

Collections  of  M.  oligosperma  were  made  by  Prof.  W.  C.  Stevens 
and  by  the  author  from  a  limestone  bluff  (Botany  Bluff)  located 
five  and  one-half  miles  northeast  of  Lawrence.  The  presence  of  this 
plant  is  quite  uncommon  except  on  limestone  banks  or  talus  slopes, 
where  it  may  be  found  in  some  abundance.  The  specimen  of  M. 
decapetala  used  in  this  study  was  collected  during  August  by  Pro- 
fessor Stevens  in  Barber  county,  Kansas.  Its  habitat  was  that  of 
a  gently  rolling  xeric  prairie. 

The  distribution  of  M.  oligosperma  is  primarily  that  of  the  great 
plains  region.  Throughout  this  region  it  may  truly  be  classified 
as  a  xerophyte.  In  the  eastern  extremity  it  inhabits  rocky  banks, 
growing  side  by  side  with  typical  mesophytic  plants,  but  its  habit 
of  growth  clearly  indicates  that  of  a  low  water  requirement. 

The  range  of  M.  decapetala  is  similar  to  that  of  M.  oligosperma. 
It  does  not  extend  as  far  east,  however,  and  is  found  as  far  north  as 
Saskatchewan.  Although  the  distribution  of  M.  decapetala  has  been 
described  as  extending  east  as  far  as  Iowa  (4),  no  specimens  are 
recorded  in  the  herbarium  of  the  University  of  Kansas  from  the 
eastern  part  of  this  state.  Its  presence  is  quite  evident  in  the 
western  part  of  Kansas,  and  has  been  found  as  far  east  as  the  central 
part  of  the  state. 

GENERAL  MORPHOLOGY 

M.  oligosperma  (stick  leaf)  grows  from  3  to  9  decimeters  in 
height  and  is  highly  branched.  The  type  of  branching  which  is 
quite  characteristic  of  this  plant  is  sometimes  described  as  dichoto- 
mous  (2).  Upon  examination,  however,  it  is  quite  evident  that  the 
main  axes,  3  to  5  dm.  in  height,  possesses  a  racemose  monopolial 
system  of  branching.  The  further  growth  in  height  of  this  shoot 
of  the  first  order  is  arrested  by  the  formation  of  a  single  terminal 
inflorescence.  Just  below  this  terminal  inflorescence,  three  lateral 
branches  of  equal  strength  usually  arise.  These  three  lateral 
branches  in  turn  divide  repeatedly,  presenting  an  appearance  very 
much  like  a  cymose  monopodial  inflorescence.  The  shoots  of  the 
second  order,  however,  do  not  terminate  in  inflorescens,  as  it  at 
first  appears,  but  rather  give  rise  to  lateral  shoots,  of  the  third 
order,  which  are  of  nearly  equal  strength  with  fruit  in  the  axes,  the 
shoots  of  the  second  order  proceeding  in  an  oblique  direction.  There- 
fore, those  parts  of  the  shoot  formed  by  the  triple  branching  of  the 
main  axis  also  present  a  true  monopodial  racemose  branching.  It 
is  very  common  to  find  shoots  of  the  fifteenth  order  represented  on  a 
plant  measuring  about  6  dm.  in  height. 


Booth:  Two  Species  of  Mentzelia  441 

The  oval-shaped  leaves  of  M.  oligosperma  are  comparatively 
small,  with  coarsely  dentate  margins.  The  lower  leaves  are  short 
petioled  and  narrowed  at  the  base.  The  upper  sessile  leaves  are 
somewhat  broader  and  have  a  truncate  base.  The  flowers  are  com- 
posed of  five  yellow  petals,  five  sepals,  many  stamens,  and  an 
inferior  one-celled  ovary,  which  develops  into  a  linear  capsule. 
Capsules  collected  in  this  locality  showed  the  number  of  seeds  per 
capsule  not  to  exceed  three;  however,  plants  have  been  reported  with 
capsules  containing  nine  seeds. 

The  mode  of  dehiscence,  which  is  often  described  as  occurring  at 
the  summit,  is  somewhat  different  from  the  common  type  of  opening 
by  pores.  The  tissue  which  completes  the  capsule  wall  at  the  sum- 
mit is  very  different  in  nature  from  that  below  it  and  upon  maturity 
may  fall  out,  leaving  an  opening  through  which  seeds  might  escape. 
However,  tissue  at  the  summit  of  the  capsule  does  not  usually  fall 
out  easily,  and  this  is  of  value  in  dispersal  of  the  seeds  into  other 
localities.  The  capsule,  because  of  an  abcission  layer,  is  easily  re- 
moved from  the  plant  and  may  be  carried  on  the  coats  of  animals. 
Later,  due  to  freezing  and  thawing  or  to  seed  germination,  this  tissue 
is  loosened  and  falls  out. 

The  root  system  is  unextensive  and  much  reduced,  except  for  a 
short,  fleshy,  primary  root.  Both  primary  and  secondary  roots  are 
often  distorted,  because  of  the  rocky  soil  in  which  they  grow,  but  in 
general  they  extend  themselves  parallel  with  the  surface  of  the 
ground  in  the  uphill  direction. 

M.  decapetala  is  larger  than  M.  oligospema  in  its  parts,  but 
seldom  grows  to  a  height  of  more  than  six  decimeters.  The  branches, 
which  arise  from  a  woody  root,  are  sparsely  branched  and  often 
terminate  in  a  flower.  The  arrangement  of  leaves  on  the  stem  is  in 
spiral  fashion,  with  an  approximate  two-fifths  phyllotaxis.  The 
lower  leaves  are  short  petioled  and  are  somewhat  lanceolate  in 
shape.  The  upper  leaves  are  sessile,  and  as  they  near  the  apex  of 
the  shoot  they  are  broader  and  not  so  long.  Hypsophylls  are  quite 
numerous  below  the  flowers  and  are  very  leaflike  in  form.  In  a 
developmental  stage  they  compare  very  closely  to  the  same  stage  of 
the  foliage  leaf  formation.  The  deeply  cleft  margin  found  in  both 
hypsophylls  and  foliage  leaves  remains  deeply  cleft  in  the  hypso- 
phylls, while  the  margin  of  the  foliage  leaves  are  widely  and  coarsely 
dentate. 

Much  larger  and  more  conspicuous  flowers  than  obtain  in  M. 
oligosperma  are  characteristic    of   M.    decapetala.    These   flowers 


442  The  University  Science  Bulletin 

regularly  open  more  or  less  widely  in  the  evening,  and  are  commonly 
from  six  to  eight  centimeters  broad.  They  have  five  sepals,  ten 
petals,  and  many  stamens.  The  ovary  is  inferior  and  develops  into 
a  many-seeded  capsule,  which  dehisces  at  the  summit. 

The  main  root  which  extends  nearly  parallel  to  the  surface  of  the 
ground  in  only  one  direction  is  common,  as  is  also  true  of  M.  oligos- 
perma.  This  root  system  is  probably  more  extensive  than  that  of 
M.  oligosperma,  but  in  form  is  very  similar. 

MATERIAL  AND  METHODS 

Specimens  upon  which  this  study  was  based  were  collected  during 
August  and  September.  In  order  to  prevent  excessive  wilting  the 
plants  were  wrapped  in  moist  sphagnum,  and,  as  soon  as  they 
reached  the  laboratory,  they  were  preserved  in  thirty  per  cent 
alcohol. 

Microchemical  tests  were  made  from  material  preserved  in  thirty 
per  cent  alcohol.  All  parts  to  be  sectioned  for  photographing  and 
for  finer  study  of  structure  were  first  desilicified  by  full-strength 
hydrofluoric  acid  for  a  week;  then  washed,  put  through  a  gradual 
series  of  butyl  alcohol  and  embedded  in  paraffin.  The  sections  to 
be  photomicrographed  were  mounted  in  hyrax,  and  negative  prints 
made  on  number  thirty-one,  hard,  single-weight  photographic  paper. 

All  drawings  were  made  by  the  aid  of  a  projectoscope  from  both 
sectioned  and  macerated  materials. 

Standard  microchemical  tests  (6)  were  used,  with  few  exceptions, 
in  the  determination  of  plant  products  present.  The  Czapek  in- 
version test  (7)  was  used  in  testing  for  glucocides,  and  the  presence 
of  hemicellulose  was  demonstrated  by  dissolving  it  in  a  three  per 
cent  solution  of  potassium  hydroxide  (9). 

COMPARATIVE  ANATOMY 

Within  the  genus  Mentzelia  very  little  anatomical  research  has 
been  published  prior  to  the  time  of  this  study.  Investigations  have 
been  carried  on  by  Gilg  in  reference  to  types  of  epidermal  hairs 
present  in  the  family  Loasacese,  and  by  Solereder  (8) ,  who  also  de- 
scribes epidermal  hairs  and  vascular  bundles  in  M.  albescens  and 
M.  pumila. 

Mentzelia  oligosperma 

Stem.  The  epidermis,  along  with  several  rows  of  cortical  cells, 
break  away  from  the  stem  proper  after  the  formation  of  a  phellogen 
by  the  pericycle.     This  layer  of  dead  tissue  sheaths  the  stem  and 


Booth:  Two  Species  of  Mentzelia  443 

presents  a  grey,  shaggy  appearance,  which  is  very  characteristic 
(PI.  LVIII,  fig.  2).  The  epidermal  cells  are  in  face  view  somewhat 
rectangular  in  shape,  but  in  cross  section  appear  as  if  crushed. 
The  cell  walls  are  relatively  thin,  the  outer  being  but  slightly  thicker 
than  the  inner  walls.  Unicellular  epidermal  hairs  of  three  general 
types  are  regularly  present  in  the  epidermis,  averaging  seven  hairs 
per  square  millimeter.  About  43  per  cent  of  the  number  present  are 
long,  reflex-barbed  hairs  (PI.  LVI,  fig.  1).  The  barbs  are  usually  in 
definite  rows,  and  at  the  apex  are  five  reflexed  barbs  which  in  face 
view  appear  star-shaped.  Forty-three  per  cent  are  short  hairs, 
usually  without  barbs  except  at  the  apex  (PL  I,  fig.  3).  The  re- 
maining 14  per  cent  are  long-pointed  hairs  with  barbs  which  point 
outward  and  toward  the  apex  (PI.  LVI,  fig.  2).  These  hairs  are 
often  quite  long,  measuring  from  .7  to  1.0  millimeter  in  length.  The 
cell  walls  of  the  three  types  of  hairs  represented  are  of  strongly 
silicificd  cellulose.  Around  the  base  of  the  long-type  hairs  a  single 
ring  of  subsidiary  cells  is  commonly  present,  and  these  cells  are  also 
often  silicified  (PI.  LVI,  fig.  5;  PI.  LVIII,  fig.  8). 

The  epidermis  and  underlying  cortical  cells,  which  are  not  entirely 
sloughed  off,  undoubtedly  serve  as  a  protection.  In  addition  to  this 
protection  a  well-formed  periderm  is  present,  which  develops  im- 
mediately on  the  inner  side  of  the  pericyclic  fibers.  Specimens  col- 
lected early  in  September  had  a  phellem  consisting  of  not  less  than 
two  cells  in  thickness  even  on  the  youngest  branches.  The  phel- 
logen,  at  the  time  of  collection,  had  ceased  activity,  and  all  cells  of 
the  phellem  were  equally  well  suberized. 

The  pericycle,  which  is  disrupted  by  the  formation  of  a  phello- 
derm,  is  composed  of  cells  of  three  types:  parenchyma,  pericyclic 
fibers,  and  stone  cells  (PI.  LVII,  fig.  7).  The  pericyclic  fibers, 
which  are  situated  at  the  extreme  outer  edge  of  the  pericycle,  are, 
upon  formation  of  a  phellogen,  separated  from  the  rest  of  the  peri- 
cycle. On  mature  stems  occasional  fibers  may  still  be  seen  clinging 
to  the  first-formed  cells  of  the  phellem.  Longitudinal  rows  of  stone 
cells  occur  at  irregular  intervals,  usually  two  or  three  rows  of  cells 
being  grouped  together.  Both  pericyclic  fibers  and  stone  cells  are 
highly  lignified. 

A  narrow  band  of  phloem,  which  is  somewhat  crushed  in  places, 
surrounds  the  cylinder  of  xylem.  The  sieve  tubes  are  quite  promi- 
nent in  the  uncrushed  areas,  but  sieve  plates  seemingly  are  few  in 
number. 

The  secondary  xj^lem   can   be   distinguished   from   the   primary 


-i44  The  University  Science  Bulletin 

xylem  by  the  fact  that  the  cells  of  the  secondary  xylem  are  very 
regularly  formed,  which  is  not  so  true  of  the  primary  xylem;  also 
the  cells  are  not  so  large  in  diameter  in  the  secondary  xylem. 
Tracheal  vessels  are  comparatively  few  in  number  and  have  an  ir- 
regular distribution  throughout  the  xylem.  They  are  represented 
by  spiral  and  scalariform  types  of  secondary  thickening  in  the  pri- 
mary xylem,  and  by  reticulate  and  reticulate-pitted  in  the  secondary 
xylem.  The  bulk  of  the  wood  is  composed  of  wood  fibers  and 
tracheids  (PI.  LVII,  figs.  4  and  6),  no  parenchyma  being  present. 
The  comparative  numbers  of  fibers  and  tracheids  present  seem  to 
be  slightly  in  favor  of  tracheids  in  both  primary  and  secondary 
xylem;  however,  the  tracheids  are  very  fiberlike  in  form  (PI.  LVII, 
fig.  4),  especially  in  the  secondary  xylem. 

Parenchyma  cells,  occupying  the  central  position  in  the  stem,  are 
large  cells  with  diameters  essentially  equal.  The  cell  walls  are  com- 
paratively thin,  but  are  well  lignified  throughout  the  pith. 

Food  products  present  in  the  stem  during  late  summer  are  starch 
and  protein.  The  starch,  although  not  in  any  great  abundance,  is 
stored  in  the  pith.  Protein  is  present  in  the  phloem  in  sufficient 
quantities  to  give  a  good  protein  test. 

Root.  In  the  early  ontogeny  of  the  roots  a  phellogen  is  formed 
by  cells  of  the  pericycle  just  beneath  the  endoderrais.  The  epi- 
dermis, cortex  and  endodermis  are  ruptured  and,  due  to  decay  and 
mechanical  strain,  are  soon  no  longer  visible.  The  cork  tissue  forms 
a  thick,  smooth  layer,  broken  only  by  occasional  lenticels.  The  cell 
walls  are  rigid  and  well  suberized. 

The  pericycle,  remaining  inside  of  the  periderm,  as  seen  in  trans- 
verse sections,  is  an  unbroken  ring  of  tissue  with  minute  intercellular 
spaces.  Its  component  cells  are  largely  parenchymatous  in  nature, 
with  scattered  groups  of  stone  cells  formed  in  longitudinal  series 
(PI.  LVII,  fig.  8).  The  primary  cells  of  the  pericycle  retain  their 
ability  to  initiate  new  cells,  and  so  in  the  mature,  fleshy  root,  meas- 
uring about  ]5  mm.  in  diameter,  it  is  quite  common  to  find  the  peri- 
cycle 1.4  mm.  in  width.  The  cell  walls  of  the  pericyclic  parenchyma 
are  of  cellulose,  while  the  stone  cells  have  heavily  lignified  walls. 

The  central  part  of  the  root  is  composed  largely  of  parenchyma- 
storage  cells,  among  which  are  scattered  tracheal  vessels,  tracheids, 
and  wood  fibers.  The  fibers  and  conducting  elements  are  few  in  num- 
ber, especially  are  there  few  fibers.  The  conducting  and  strength- 
ening elements  of  the  xylem  are  in  small  radial  strands.  The  water- 
conducting  elements,  although  few  in  number,  are  larger  in  diameter 


Booth:  Two  Species  of  Mentzelia  445 

than  those  of  the  stem  (PL  LVII,  fig.  3) ,  and  because  of  the  low 
water  requirement  of  the  shoot  they  are  able  to  carry  an  ample 
supply. 

Parenchyma  cells  of  the  xylem  are  highly  functional  as  storage 
organs  for  starch.  Starch  is  also  stored  in  the  pericyclic  paren- 
chyma, but  in  decidedly  smaller  amounts. 

Leaf.  The  normal  epidermal  cells  are  probably  rectangular  in 
face  view,  but  because  of  the  frequent  irregularities  caused  by  epi- 
dermal hairs  and  stomata,  there  are  many  which  differ  from  this 
type.  The  number  of  hairs  on  the  upper  surface  of  the  leaf  averages 
about  40  per  square  mm.  and  on  the  lower  surface  about  44,  On  the 
upper  surface  45  per  cent  of  the  epidermal  hairs  are  long,  reflex- 
barbed  hairs,  52  per  cent  are  short  hairs  with  reflex  barbs  at  the 
apex  only,  and  2  per  cent  are  long,  pointed  hairs,  as  described  in 
connection  with  the  stem  of  this  plant.  The  proportion  of  hair 
types  for  the  under  surface  of  the  leaf  is  somewhat  the  same,  there 
being  slightly  fewer  large,  reflex-barbed  hairs,  and  more  long, 
pointed  hairs. 

The  cell  walls  of  the  epidermal  hairs  and  subsidiary  cells,  as  was 
also  true  of  the  stem,  are  composed  of  strongly  silicified  cellulose. 
Calcium  carbonate  as  a  lining  of  the  large  types  of  epidermal  hairs, 
and  of  the  subsidiary  cells  in  less  amount,  is  very  common.  This 
lining  many  times  is  quite  thick  in  the  basal  region  of  the  hair,  being 
often  thicker  than  the  cell  wall,  while  the  lumen  of  the  narrower 
part,  from  just  above  the  base  to  the  apex,  is  entirely  filled  with 
calcium  carbonate.  The  epidermal  hairs  are  covered  with  the  thin 
cuticle  which  is  present  on  the  regular  epidermal  cells. 

About  16  stomata  per  square  mm.  are  present  on  the  upper  surface 
of  the  leaf,  and  23  on  the  lower.  These  stomata,  with  raised  guard 
cells,  are  of  the  common  type.  Although  this  type  of  stoma  is  most 
common  among  the  mesophytes,  in  M.  oligosperma  it  is  well  pro- 
tected by  the  dense  covering  of  epidermal  hairs. 

The  mesophyll  of  the  leaf  consists  of  a  single  layer  of  palisade 
parenchyma  occupying  approximately  the  upper  half  of  the  leaf, 
and  a  loosely  joined,  spongy  parenchyma.  A  small  amount  of 
starch  may  be  found  in  the  palisade  parenchyma,  and  a  larger 
amount  in  the  guard  cells  and  spongy  parenchyma. 

Vascular  bundles  forming  the  veins  are  not  prominent,  those 
present  having  as  their  main  conducting  tissue  small  groups  of 
spiral  elements.  The  proportion  of  phloem  and  strengthening  tissue 
is  small.    The  midrib,  although  quite  outstanding,  has  also  but  little 


446  The  University  Science  Bulletin 

vascular  tissue  (PI.  LIX,  fig.  3).  The  strengthening  tissue  of  the 
midrib,  in  addition  to  the  water-conducting  tissue,  is  composed  of  a 
small  group  of  fibers  just  beneath  the  vascular  bundle,  and  a  few 
collenchyma  cells  at  the  lower  and  upper  sides. 

Fruit.  The  epidermis  of  the  capsule  is  very  similar  to  that  of  the 
foliage  leaf,  with  the  exception  of  absence  of  stomata.  Chloroplasts 
are  present  in  the  outermost  parenchyma  cells  of  the  mesophyll,  and 
a  small  amount  of  starch  is  found  throughout  the  entire  tissue. 
Near  the  center  of  the  mesophyll  vascular  strands  are  present  at 
regular  intervals.  These  strands,  about  twelve  to  fourteen  in  num- 
ber, are  made  up  of  conductive  tissue  and  very  thick-walled  fibers. 
They  enter  the  mesophyll  at  the  base  of  the  capsule  and  continue, 
without  branching,  to  the  summit. 

At  maturity  the  seeds  have  testas  composed  of  very  characteristic 
stone  cells,  which  in  face  view  appear  very  ordinary  (PI.  LVI,  fig. 
11),  but  in  cross  section  are  quite  unlike  most  stone  cells  (PI.  LVI, 
fig.  12).  They  are  composed  of  a  primary  wall,  a  secondary  wall 
of  cellulose  with  which  is  associated  hemicellulose,  and  a  tertiary 
wall  of  cellulose.  The  secondary  wall  has  every  appearance  of 
formation  in  both  centrifugal  and  centripetal  direction.  Although 
developmental  studies  were  not  made  of  these  cells,  it  seems  justifi- 
able to  conclude  that  points  projecting  from  regions  of  maximum 
thickness  are  formed  in  a  centrifugal  direction,  while  an  even  layer 
of  nearly  equal  thickness  formed  in  a  centripetal  direction  is  present 
throughout  the  entire  cell.  The  tertiary  wall  lining  the  lumen  of 
the  cell  is  of  cellulose  and  has  thickenings  in  the  form  of  longitudinal 
bars.  The  secondary  wall  in  particular  has  an  abundance  of  calcium 
carbonate  in  a  noncrystalline  form. 

Just  beneath  the  testa  is  the  tegmen,  consisting  of  a  thin  layer  of 
crushed  cells  (PI.  LVI,  fig.  8-B).  These  cells  have  cellulose  walls, 
and  in  many  sections  are  difficult  to  distinguish  from  a  similar  layer 
just  beneath,  which  is  a  remnant  of  the  nucellus.  A  well-developed 
endosperm  is  present,  which  is  composed  of  rounded  parenchyma 
cells.  The  cell  walls  are  of  cellulose,  and  intercellular  spaces  are 
large  and  numerous.  This  endosperm  is  especially  rich  in  stored 
foods  in  the  form  of  fatty  oils  and  aleurone.  Some  seeds,  when 
sectioned,  show  the  presence  of  homogeneous  masses  of  oil  and  pro- 
tein around  the  embryo,  or  in  breaks  in  the  endosperm  and  between 
loosely  joined  cells  of  the  endosperm  (PI.  LVI,  fig.  8-E).  It  is  very 
likely  that  this  material  has  been  turned  loose  by  the  breaking 
down  of  endosperm  cells. 


Booth:  Two  Species  of  Mentzelia  447 

The  embryo  is  composed  of  relatively  large  plano-convex  cotyle- 
dons, and  a  short,  radicle-hypocotylar  region.  A  differentiation  of 
tissue  is  shown  in  the  radicle  and  hypocotyl  by  a  definite  epidermis, 
isodiametric  parenchyma  cells,  and  by  procambium  strands.  The 
cotyledons  also  show  the  presence  of  an  epidermis,  procambium 
strands,  isodiametric  parenchyma,  and  palisade  parenchyma  (PI. 
LVIII,  fig.  2).  The  palisade  layer,  as  was  also  true  of  the  foliage 
leaf,  is  one  cell  in  width. 

Reserve  food  is  stored  in  all  parts  of  the  embryo.  This  stored 
food  is  very  similar  to  that  of  the  endosperm,  there  being  large 
quantities  of  both  fatty  oil  and  aleurone  present. 

Mentzelia  deoapetala 

Stem. — The  epidermis  of  the  stem  of  M.  decapetala  is  composed 
of  irregular  cells  which  often  have  undulated  radial  walls,  the  outer 
wall  being  slightly  thicker  than  the  inner,  and  in  face  view  ap- 
pearing as  if  furrowed,  as  will  be  explained  in  more  detail  in  rela- 
tion to  leaves,  where  this  feature  is  more  pronounced. 

The  functional  duration  of  the  epidermis  is  usually  that  of  a  single 
season,  after  which  time  a  phellogen  is  formed  by  the  pericyclic 
parenchyma  just  beneath  the  pericyclic  fibers.  On  the  oldest  stems 
the  pericyclic  fibers  are  still  visible  clinging  to  the  first-formed  cells 
of  the  phellem.  Also  patches  of  cortex  and  epidermis  still  cling 
to  the  stem,  but  are  not  present  in  sufficient  quantities  to  affect  the 
appearance  of  the  stem. 

The  cortex  of  a  one-year-old  stem  is  composed  of  a  narrow  band 
of  large,  nearly  isodiametric,  cells.  At  a  slightly  older  stage  the  cell 
walls  become  undulated,  a  character  of  the  cell  wall  very  probably 
caused  by  the  formation  of  the  phelloderm  and  possibly  one  or  two 
layers  of  phellem  when  the  stem  is  quite  young,  prohibiting  the 
passage  into  the  cortex  of  water  and  food  substances,  so  the  cells 
of  the  cortex  and  epidermis  gradually  lose  their  turgidity  and  dimin- 
ish in  size  to  such  an  extent  that  their  thin  cell  walls  fall  into  folds 
(PI.  LIX,  fig.  1). 

The  pericycle  in  the  young  stem  is  composed  of  thin-walled  paren- 
chyma cells  and  pericyclic  fibers.  These  fibers  are  unusually  large 
cells,  commonly  being  about  3  mm.  in  length  (PI.  LVII,  fig.  15). 
The  cell  walls  are  only  moderately  thick  and  but  slightly  lignified. 
Compound  crystals  of  calcium  oxalate  are  occasionally  present  in  the 
pericycle  parenchyma  and  in  the  pith  (see  below) . 

The  phloem  is  represented  by  a  narrow  band  of  very  small  cells, 


448  The  University  Science  Bulletin 

the  sieve  tubes  measuring  not  more  than  5  microns  in  diameter. 
Although  sieve  tubes  and  companion  cells  may  definitely  be  seen, 
the  presence  of  sieve  plates  is  not  apparent. 

Tracheal  vessels  are  quite  numerous  in  both  primary  and  sec- 
ondary xylem;  they  are,  however,  very  small  in  diameter.  The 
primary  xylem  is  represented  by  vessels  with  spiral,  scalariform 
and  reticulate  type  of  secondary  thickenings,  while  the  secondary 
xylem  has  only  pitted-type  vessels.  Tracheids  and  fibers  are  both 
present  in  great  abundance  in  the  secondary  xylem.  Pith-rays  ex- 
tend for  a  short  distance  into  the  primary  xylem,  but  never  extend 
into  the  secondary  xylem. 

The  cells  of  the  pith  lack  uniformity  in  size.  The  larger  and  the 
smaller  cells,  however,  are  usually  grouped  together  (PI.  LIX,  fig.  7). 
This  grouping  may  not  always  be  seen  in  a  transverse  section,  be- 
cause a  longitudinal  grouping  is  most  common.  Calcium  oxalate 
crystals  are  not  uncommon  in  the  pith,  but  when  present  the  com- 
pound crystals  are  usually  small. 

Reserve  food  in  the  form  of  glucosides  is  present  in  large  quantities 
throughout  the  stems,  but  is  present  in  larger  quantities  in  the  peri- 
cycle. 

Root.  Epidermal,  cortical  tissues  and  endodermis  are  early  in  the 
development  of  the  root  sloughed  off,  because  of  the  formation  of  a 
pericyclic  periderm.  The  phellogen  takes  its  origin  from  the  outer 
pericyclic  parenchyma  cells  and  is  functional  only  for  a  short  period, 
then  its  cells  mature  into  cork  cells.  The  cork  cells  on  the  old  roots 
are  usually  badly  crushed,  but  are  highly  suberized,  so  serve  well  as 
a  protection. 

The  pericycle  of  the  root  increases  in  width  until  in  a  mature 
root  measuring  about  25  mm.  in  diameter  the  pericycle  may  com- 
monly be  as  much  as  2.5  mm.  in  width.  The  bulk  of  this  region  is 
made  up  of  parenchymatous  cells  varying  in  shape.  Groups  of  stone 
cells  are  scattered  irregularly  among  the  typical  parenchyma  cells, 
but  are  confined  mostly  to  the  inner  half  of  the  pericycle.  These 
stone  cells  are  usually  in  groups  of  two  or  three,  as  seen  in  trans- 
verse section,  forming  long  radial  strands.  Compound  crystals  of 
calcium  oxalate  are  very  abundant  in  the  pericycle.  Usually  they 
are  situated  in  small  parenchyma  cells  measuring  about  .024  mm. 
in  diameter. 

In  structure  the  phloem  in  the  root  corresponds  very  closely  with 
that  of  the  stem,  with  possible  exception  of  being  even  reduced  more 


Booth:  Two  Species  of  Mentzelia  449 

in  extent.     Sufficient  protein  is  present  in  scattered  groups  of  cells 
in  the  phloem  to  give  a  protein  test. 

Tracheal  vessels  are  quite  numerous  in  both  primary  and  second- 
ary xylem.  Conducting  elements  in  the  primary  and  secondary 
xylem  are  represented  by  vessels  with  scalariform  and  reticulate 
secondary  thickenings  as  the  dominant  cell  type.  Few  tracheids  and 
fiber  tracheids  are  present  in  the  primary  xylem  and  only  occasional 
spiral  element,  but  both  tracheids  and  wood  fibers  are  quite  abun- 
dant in  the  secondary  xylem.  Xylem  rays,  usually  about  four 
cells  in  width  in  the  secondary  xylem  and  much  wider  in  the  primary 
xylem,  extend  from  near  the  center  of  the  root  to  the  pericycle. 
Compound  crystals  of  calcium  oxalate  are  present  in  scattered  cells 
of  the  xylem  rays,  but  are  by  far  less  abundant  than  in  the  pericycle. 

The  reserve  food,  which  is  in  the  form  of  glucosides,  is  present  in 
the  stele  of  the  root,  but  the  amount  present  is  smaller  than  that  of 
the  stem. 

Leaf.  The  epidennis  of  the  leaf  is  composed  of  comparatively 
small  cells  with  thick  cell  walls.  The  regular  epidermal  cells  and 
subsidiary  cells  of  the  hairs  are  covered  with  a  cuticle  which  is 
formed  in  ridges  which  may  be  continuous  from  one  cell  to  another. 
These  ridges  are  most  commonly  from  two  to  three  microns  in 
height,  but  it  is  not  uncommon  for  them  to  be  as  high  as  four  or 
five  microns.  Their  composition  is  purely  that  of  cutin,  which  may 
be  completely  removed  by  boiling  in  a  3  per  cent  alcoholic  solution 
of  potassium  hydroxide. 

Heavy  silicified  epidermal  hairs  are  quite  abundant  on  both  upper 
and  lower  leaf  surfaces,  there  being  about  eight  hairs  per  square  mm. 
of  leaf  surface.  About  75  per  cent  of  the  hairs  present  are  conical 
in  shape,  with  blunt  barbs  (PI.  LVI,  fig.  4),  and  25  per  cent  are  short 
hairs  with  reflexed  barbs  at  the  apex  (PI.  LVI,  fig.  3).  The  inner 
wall  of  the  conical  hairs  and  of  the  subsidiary  cells  are  roughened 
and  are  made  to  seem  more  irregular  by  the  presence  of  small 
patches  of  calcium  oxalate. 

The  mesophyll  of  the  leaf  is  composed  largely  of  palisade  par- 
enchyma. A  single  layer  of  these  cells  is  present  beneath  both  upper 
and  lower  epidermal  surfaces,  leaving  only  a  relatively  small  area 
through  the  center  of  the  leaf  for  spongy  parenchyma  and  vascular 
tissue.  Frequently  the  cell  walls  of  palisade  parenchyma  in  contact 
with  the  enlarged  base  of  an  epidermal  hair  become  silicified.  When 
this  is  reduced  to  ash  a  skeleton  of  silica  remains,  appearing  as  a 


450  The  University  Science  Bulletin 

meshwork  below  the  bulbous  base  of  the  epidermal  hair  (PI.  LVIII, 

fig.  4). 

The  midrib,  although  not  prominent,  contains  considerarble  con- 
ductive tissue.  At  the  base  of  the  leaf,  and  proceeding  into  the  leaf 
for  often  8  mm.,  are  three  separate  vascular  bundles,  which  then 
merge  to  form  a  single  bundle.  The  strengthening  tissue  consists  of 
xylem,  and  collenchyma  at  both  upper  and  lower  surfaces  of  the 
midrib  near  the  base  of  the  leaf,  but  it  is  not  formed  near  the  tip. 

Fruit.  The  epidermis  of  the  capsule  is  very  similar  to  that  of  the 
stem,  with  exception  of  there  being  hardly  so  many  epidermal  hairs 
per  surface  area.  The  mesophyll  is  composed  largely  of  irregular 
parenchyma  cells  with  large  intercellular  air  spaces.  Compound 
crystals  of  calcium  oxalate  are  very  abundant  in  this  tissue,  but  are 
localized  largely  in  the  innermost  cells,  and  in  particular  around  the 
seeds.  Mechanical  strength  is  furnished  by  thick-walled  fibers  as- 
sociated with  numerous  radially  arranged  vascular  strands.  The 
strands  are  frequently  connected  by  branching  vascular  elements, 
fibers  and  thick-walled  parenchyma. 

The  testa  and  wing  of  the  seed  are  composed  of  characteristic 
stone  cells  with  thick  cellulose  walls  (PI.  LVI,  fig.  13).  These  cells 
are  quite  irregular  in  outline  as  is  also  the  case  of  their  homologues 
in  M.  oligosperma.  The  tegmen  is  composed  of  a  single  layer  of 
crushed  cells  which  are  difficult  to  distinguish  from  a  similar  layered 
remnant  of  the  nucellus.  Endosperm  is  present  as  a  comparatively 
thick  tissue  surrounding  the  embryo.  It  is  especially  rich  in  stored 
protein  and  fatty  oils.  In  the  mature  seed  empty  cells  of  the  endo- 
sperm adjoining  the  embryo  are  quite  common,  the  food  from  these 
cells  having  undoubtedly  been  used  in  the  formation  of  the  embryo. 

The  embryo  is  straight  and  has  a  prominent  radical-hypocotylar 
region.  The  component  tissues  of  the  embryo  are  the  epidermal,  the 
parenchymatous  cells  and  the  procambium  strands.  The  mesophyll 
of  the  cotyledons  is  composed  entirely  of  small,  nearly  isodiametric, 
cells,  there  being  no  differentiation  into  palisade  and  spongy  paren- 
chyma. The  amount  and  types  of  food  stored  in  the  embryo  are 
practically  the  same  as  of  the  endosperm. 

The  presence  of  glucosides  in  the  mesocarp  of  the  fruit  is  common, 
but  is  hardly  as  prominent  as  in  the  root,  stem,  and  leaves. 


Booth:  Two  Species  of  Mentzelia  451 

SUMMARY 

Principal  anatomical  similarities  and  differences  between  Ment- 
zelia oligosperma  and  M.  decapetala: 

POINTS   OF    SIMILARITY 

1.  Formation  of  phellogen  by  the  pericycle  in  root  and  stem. 

2.  Component  cell  types  of  the  pericycle  in  root  and  stem. 

3.  Growth  of  pericycle  by  cell  multiplication  in  roots. 

4.  Presence  of  silicij&ed  subsidiary  cells  and  epidermal  hairs. 

5.  Kinds  of  food  stored  in  embryo  and  endosperm — fatty  oils  and 

proteins. 

6.  Relatively  large  amounts  of  wood  fibers  in  the  xylem. 

7.  A  low  water  requirement  is  indicated  in  the  stems  of  M.  oligo- 

sperma by  the  relatively  small  number  of  water-conducting 
elements,  and  in  M.  decapetala  by  only  a  slightly  larger  num- 
ber of  water-conducting  elements  which  are  smaller  in  diame- 
ter than  those  of  M.  oligosperma. 

POINTS   OF   DIFFERENCE 

1.  Food  stored  in  stem,  root,  and  leaf — starch  in  M.  oligosperma 

and  glucosides  in  M.  decapetala. 

2.  Differentiation  of  cell  types  in  cotyledons — palisade  cells  present 

in  M.  oligosperma  and  absent  in  M.  decapetala. 

3.  Type  of  stone  cell  making  up  the  testa. 

4.  Size  of  component  cells  throughout  the  plant — smaller  in  M. 

oligosperma  than  in  M.  decapetala. 

5.  Epidermis  and  cuticle — undulate  radial  walls  and  ridged  cuticle 

in  M.  decapetala  without  counterpart  in  M.  oligosperma. 

6.  Presence  of  calcium  oxalate  in  stone  cells  and  epidermal  hairs  of 

M.  oligosperma,  and  calcium  carbonate  in  epidermal  hairs  and 
as  compound  crystals  widely  distributed  throughout  the  paren- 
chymatous tissues  of  M.  decapetala. 

7.  Epidermal  and  cortical  tissues  of  the  stem  of  M.  decapetala  are 

partly  sloughed  off,  only  patches  remaining  attached  to  the 
cork.  The  epidermis  and  cortex  of  M.  oligosperma  remains 
more  or  less  intact  as  a  loose  sheath  around  the  stem. 

8.  Stems  of  M.  decapetala,  because  of  their  having  a  longer  growth 

period,  reach  a  larger  diameter  than  do  the  annual  stems  of 
M.  oligosperma. 
There  are  many  anatomical  differences  between  the  two  species 


452 


The  University  Science  Bulletin 


investigated,  but  points  of  similarity  are  sufficiently  striking  to 
indicate  a  close  relationship. 

The  writer  wishes  to  acknowledge  the  assistance  received  from 
Prof.  W.  C.  Stevens,  who  suggested  this  problem,  supplied  material, 
and  offered  many  valuable  suggestions  throughout  the  course  of 
investigation  and  in  preparation  of  the  manuscript.         - 


Cell  Measurements  from  Tissues  Represented  in  Stem  and  Root 


Cell  type  from  stem. 

Mentzelia  i 
diameter 

Hnoxperma, 
in  microns. 

Mentzelia 
diameter 

decapetala, 
n  microns. 

Long. 

Tang. 

Radial. 

Wall. 

Long. 

Tang. 

Radial. 

Wall. 

Epidermis 

22 

20 

44 

3.2 

30 

23 

3,2.8 

Parenchyma  of  cortex 

52 

23 

15 

2.0 

100 

28 

10 

1.3 

920 

52 

48 
46 

24 
32 

5.0 
2  2 

3,100 
30 

18 
20 

20 
36 

8.0 

Pericyclic  parenchyma.  .  . 

3.0 

Pericyclic  stone  cell 

131 

43 

39 

12  4 

12 
6± 

8 
6± 

.8 
.8 

8 
5± 

8 
5± 

.8 

.8 

Vessel  segment 

320 

41 

58 

5.2 

20 

32 

2.2 

580 
611 
120 

24 

20 

100 

28 

20 

100 

5.0 
6.0 
3.0 

590 

630 

60 

14 

10 

52-30 

16 
16 
80 

2.0 

Wood  fibre 

2.0 

Parenchyma  of  pith 

1.3 

Phellem 

240 

40 

IS 

3  2 

78 

52 

22 

Cell  type  from  root. 
Phellem 

96 

60 

24 

3.0 

102 

60 

15 

2.0 

Pericyclic  parenchyma.  .  .  . 

100 

100-4 

40-32 

2.5 

140 

140-30 

52-32 

1.3 

Pericyclic  stone  cells 

71 

25 

25 

9.2 

120 

40 

36 

10.0 

12 

7— 
76 

12 
36 

1.0 
1.0 
2.0 

40  ± 

8 

6± 

12  ± 

8 

6=*= 
10  ± 

.8 

.8 

Parenchyma  of  xylem .... 

78 

1.3 

Tracheal  vessel 

240 
300 

160 
20 

144 
20 

6.0 
4.0 

40 
600 

53 
32 

58 
30 

6.0 

Tracheid 

3.5 

Wood  fibre 

381 

20 

20 

4.5 

654 

23 

23 

3.4 

Booth:  Two  Species  of  Mentzelia  453 


BIBLIOGRAPHY 

1.  Clements  and  Clements.     1928.    Rocky  Mountain  Flowers.     Wilson  Co., 

New  York. 

2.  Coulter  and  Nelson.     1909.     New  Manual  of  Botany  of  Central  Rocky 

Mountains.    Amer.  Book  Co.,  New  York,  pp.  324-325. 

3.  Keller,  A.  A.     1900.    Catalogue  of  North  American  Plants  North  of  New 

Mexico.    Uni.  of  Min. 

4.  Robinson  and  Fernald.     1908.     Gray's  New  Manual   of  Botany,   Amer. 

Book  Co.,  New  York,  p.  588. 

5.  Rydberg,  p.  a.     1917.     Flora  of  Rocky  Mountains  and  Adjacent  Plains, 

pp.  169-175. 

6.  Stevens,  W.  C.     1924.    Plant  Anatomy.    Blakiston's  Son  &  Co.,  Philadel- 

phia. 

7.  Schneider-Zimmerman.     1922.     Die    Botanische    Mikrotechnik.     Gustav 

Fisher,  Jena,  p.  259. 

8.  SoLEREiDER,   H.     1908.     Systematic   Anatomy   of  the   Dicotyledons.     Engl. 

Trans.    Carendon  Press,  Oxford,  1 :  378-381. 

9.  Tunman,  O.    1913.    Pflanzenmikrochemie.    Berlin,  1931. 


;«)-;M,s2 


454  The  University  Science  Bulletin 


I 


PLATE  I.VI 

Fig.  1.  Reflex-barbed  epidermal  hair  from  the  leaf  of  M.  oligosperma. 
X  73.3. 

Fig.  2.  Long-pointed  epidermal  hair  from  the  leaf  of  M.  oligosperma. 
X  73.3. 

Fig.  3.  Short  epidermal  hair  which  is  present  on  either  M.  oligosperma  oi- 
M.  decapetala.    X  73.3. 

P^G.  4.    Short-pointed  epidermal  hair  from  leaf  of  M.  decapetala.    X  73.3. 

Fig.  5.  Face  view  of  the  subsidiary  cells  of  an  epidermal  hair  from  the  leaf 
of  M.  oligosperma.    X  73.3. 

Fig.  6.  Face  view  of  the  subsidiary  cells  of  an  epidermal  hair  from  the  leaf 
of  M.  decapetala.    X  73.3. 

Fig.  7.  Outline  drawing  from  a  transverse  section  of  a  seed  of  M.  decapetala. 
X  40.  A,  testa;  B,  tegmen;  C.  perisperm;  E,  oily-proteinaceous  material  from 
the  endosperm;  f,  cotyledon. 

Fig.  8.  Outline  drawing  from  a  transverse  section  of  a  seed  of  M.  olig- 
osperma. X  40.  A,  testa;  B,  tegman;  C,  perisperm;  E,  endosperm;  F, 
cotyledon. 

Fig.  9.  Outline  drawing  from  face  view  of  cells  composing  the  outer  seed 
coat  of  M.  oligosperma.    X  75. 

Fig.  10.  Outline  drawing  from  face  view  of  cells  composing  the  outer  seed 
coat  of  M.  decapetala.    X  75. 

Fig.  11.  Transverse  section  of  stone  cells  from  testa  of  M.  oligosperma. 
X  150. 

Fig.  12.   Face  view  of  a  stone  cell  from  the  testa  of  M.  oligosperma.   X  150. 

Fig.  13.  Transverse  section  of  stone  cells  from  testa  of  M.  decapetala. 
X150. 

Fig.  14.   Seed  of  M.  decapetala.    X  3.3. 

Fig.  15.   Seed  of  M.  oligosperma.    X  3.3. 


Booth:  Two  Species  of  Mentzelia 
PLATE  LVI 


455 


45H  The  University  Science  Bulletin 


PLATE  LVII 

Fig.  1.  Part  of  tracheal  elements  from  primary  xylem  of  stem  of  M. 
oligosperma.    X  183. 

Fig.  2.   Vessel  segment  from  stem  of  M.  oligosperma.    X  183. 

Fig.  3.   Vessel  segment  from  root  of  M.  oligosperma.    X  183. 

Fig.  4.   Tracheid  from  stem  of  M.  oligosperma.    X  183. 

Fig.  5.   Wood  fiber  from  root  of  M.  oligosperma.    X  183. 

Fig.  6.   Wood  fiber  from  stem  of  M.  oligosperma.    X  183. 

Fig.  7.   Stone  cell  from  stem  of  M.  oligosperma.    X  183. 

Fig.  8.   Stone  cell  from  root  of  M.  oligosperma.    X  183. 

Fig.  9.  Part  of  tracheal  elements  from  primary  xylem  of  stem  of  M. 
decapetala.    X  183. 

Fig.  10.   Vessel  segment  from  stem  of  M.  decapetala.    X  183. 

Fig.  11.    Wood  fiber  from  root  of  M.  decapetala.    X  183. 

Fig.  12.   Wood  fiber  from  stem  of  M.  decapetala.    X  183. 

Fig.  13.  Tracheid  from  stem  of  M.  decapetala.    X  183. 

Fig.  14.   Part  of  a  pericyclic  fiber  from  stem  of  M.  decapetala.    X  183. 

Fig.  15.  Pericycle  fiber  from  stem  of  M.  decapetala.  Half  of  cell  is  shown. 
X40. 

Fig.  16.   Vessel  segment  from  root  of  M.  decapetala.    X  183. 

Fig.  17.    Stone  cell  from  pericycle  of  root  of  M.  decapetala.    X  183. 


Booth:  Two  Species  of  Mentzelia 
PLATE  LVII 


45' 


458  The  University  Science  Bulletin 


PLATE  LVIII 

Fj(;.  1.    A  part  of  a  young  shoot  of  M.  oligosperma. 

Fig.  2.    Longitudinal  section  from  the  seed  of  M.  oligosperma.    X  37.5. 

Fig.  3.  M.  oligosperma  showing  a  part  of  the  fleshy  root  system  and  the 
shaggy  characteristic  of  the  stems.    X  37.5. 

Fig.  4.   Silicified  subsidiaiy  cells  of  M.  decapetala. 

Fig.  5.   Transverse  section  from  the  seed  of  M.  decapetala.    X  37.5. 

Fig.  6.   Leaf  margin  of  M.  decapetala.    X  65. 

Fig.  7.  Longitudinal  section  from  the  seed  of  M.  decapetala  showing  but 
little  differentiation  of  tissue.     X  37.5. 

Fig.  8.  Ash  from  M.  oligosperma  showing  the  silicified  basal  cells  of  an 
epidermal  hair. 

Fig.  9.  Transverse  section  from  the  seed  of  M.  oligosperma  through  hypo- 
cotylar  region.    X  37.5. 

Fig.  10.    M.  decapetala  in  native  habitat. 

Fig.  1L   Leaf  margin  of  M.  oligosperma.    X  65. 


Booth:  Two  Species  of  Mentzelia 


459 


PLATE  LVIII 


460  The  University  Science  Bulletin 


PLATE  LIX 

Fig.  1.   Sector  from  transverse  section  of  stem  of  M.  decapetala.    X  65. 

Fig.  2.   Sector  from  transverse  section  of  stem  of  M.  oligosperma.    X  65. 

Fig.  3.   Transverse  section  of  midrib  of  M.  decapetala.    X  65. 

Fig.  4.   Transverse  section  of  midrib  of  M.  oligosperma.    X  65. 

Fig.  5.  Longitudinal  section  of  pith  cells  from  the  stem  of  M.  decapetala 
showing  the  irregularity  in  size  and  shape  of  the  cells.    X  200. 

Fig.  6.  Longitudinal  section  of  pith  cells  from  the  stem  of  M.  oligosperma 
showing  that  there  is  not  a  great  irregularity  in  shape  of  the  component 
cells.    X  200. 


Booth:    Two  Species  of  Mentzelia 


461 


PLATE  LIX 


THE  UNIVERSITY  OP  KANSAS 

SCIENCE  BULLETIN 

Vol.  XXL]  March,  1933.  [No.  15. 


Some  Studies  on  the  Transference  of  Ions 
in  Anhydrous  Acetic  Acid  Solution 

ARTHUR  W.  DAVIDSON  and  VERNON  HOLM,' 
Department  of  Chemistry,  University  of  Kansas 


Abstract:  Transfei-ence  measurements  were  made  by  the  Hittorf  analytical 
method  on  sodium  and  ammonium  acetates  in  acetic  acid  solution,  as  well  as 
on  sodium  zinc  acetate  and  on  ammonium  cupric  acetate  solutions.  In  the 
case  of  the  alkali  acetates  the  apparent  transport  numbers  of  the  cations  were 
found  to  be  surprisingly  small,  and  to  decrease  with  increasing  dilution,  be- 
coming practically  zero  in  the  most  dilute  solutions  studied  (about  1  mole  per 
cent,  or  0.15  molal).  It  is  suggested  that  these  anomalous  values  may  be  due 
to  the  presence  of  complex  ions.  In  the  case  of  the  double  acetate  solutions 
it  had  been  expected  that  the  less  positive  metal  might  be  found  to  migrate 
toward  the  anode,  but  no  such  result  was  obtained. 


INTRODUCTION 

AMPHOTERISM  IN  WATER,  AMMONIA  AND  ACETIC  ACID 
SOLUTIONS 

IT  IS  a  familiar  fact  that  zinc  hydroxide  dissolves  in  aqueous 
solutions  of  sodium  hydroxide,  and  that  at  very  high  concentra- 
tions of  alkali  a  compound  called  sodium  zincate  may  be  isolated 
from  such  solutions.  The  formula  for  this  compound  which  appears 
to  be  best  substantiated  is  that  given  by  Goudriaan,^  Na^ZnOs  . 
4H2O.  It  is  obvious  that  this  compound  might  equally  well  be 
formulated  as  ZnfOH)^  .  2NaOH  .  2H2O,  and  might  be  regarded 
as  an  addition  compound  of  the  two  hydroxides.  Analogous  com- 
pounds are  formed  when  strongly  alkaline  solutions  react  with  any 

1.  This  paper  was  constructed  from  a  thesis  submitted  by  Vernon  Hohn  in  partial  fulfill- 
ment of  the  requirements  for  the  degree  of  Doctor  of  Philosophy  at  the  University  of  Kansas, 
June,  1932. 

2.  Goudriaan,  Rec.  trav.  chim.,  39,  505  (1920). 

(463) 


464  The  University  Science  Bulletin 

of  the  so-called  amphoteric  hydroxides,  such  as  those  of  aluminum, 
lead  and  tin.  Further,  this  type  of  reaction  is  not  peculiar  to  aque- 
ous solutions.  It  has  been  shown,  mainly  by  Franklin  and  his  co- 
workers,^ to  be  of  even  more  common  occurrence  in  liquid  ammonia ; 
a  large  number  of  amides  (bases  of  the  ammonia  system)  react 
with  potassium  amide  to  form  addition  compounds.  Further  evi- 
dence of  the  general  nature  of  this  phenomenon  has  been  found 
from  the  study  of  solutions  in  acetic  acid.  In  a  recent  paper  from 
this  laboratory^  it  was  showTi  that  some  of  the  properties  of  solutions 
of  zinc  and  sodium  acetates  in  acetic  acid  are  closely  parallel  to 
those  of  the  corresponding  aqueous  system;  the  solubility  of  zinc 
acetate  increases  with  increasing  concentration  of  sodium  acetate 
until  the  compound  Zn  (0211302)2  •  2NaC2H302  .  4HC2H3O2  appears 
as  solid  phase.  Similarly,  cupric  acetate  dissolves  in  potassium  or 
ammonium  acetate  solutions,  and  in  the  latter  case  a  compound  of 
the  formula  Cu(C2H302)2  •  4NH,C2H302  .  4HC2H3O2  has  been  iso- 
lated.^ If  an  amphoteric  base  be  regarded  as  one  which  is  capable 
of  combining  with  stronger  bases  to  give  addition  compounds,  all  of 
the  cases  cited  may  obviously  be  regarded  as  instances  of  ampho- 
terism. 

In  the  case  of  aqueous  solutions,  compounds  of  the  type  under 
consideration  are  commonly  regarded  as  salts,  and  are  assumed  to 
dissociate  so  as  to  yield  anions  containing  the  less  positive  metal. 
The  name  "metallate"  has  been  suggested  ^  as  a  general  term  for  all 
such  salts.  In  general,  the  evidence  for  this  sort  of  dissociation  is 
far  from  conclusive.  However,  in  the  case  of  sodium  zincate,  at 
least,  it  has  been  shown  that  during  electrolysis  the  concentration 
of  zinc  increases  in  the  neighborhood  of  the  anode,'^  which  is  in 
accord  with  the  hypothesis  that  the  zincate  ion,  HZnOo",  is  present 
in  such  solutions.  In  the  ammonia  system  the  existence  of  metallate 
ions  was  merely  taken  for  granted  until  very  recently,  when  Detrick 
and  Fernelius,  in  an  unpublished  paper,^  reported  that,  in  the  elec- 
trolysis of  liquid  ammonia  solutions  of  potassium  ammonoaluminate, 
A1(NH2)3  •  KNH2  .  NH3  or  AI(NH2)2NHK  .  2NH3,  potassium  am- 
monoplumbite,  PbNH  .  KNHg  or  PbNK  .  NH3,  and  potassium  am- 
monocuprite,  CUNH2  .  2KNH2  .  NII3  or  CuNK^  .  3NH3,  the  less 

3.  For  a  summary  see  Bergstrom  and  Fernelius,  Cham.  Reviews,  12,  51  (1933). 

4.  Davidson  and  McAllister,  J.  Am.  Chem.  Soc,  52,  519  (1930). 

5.  Davidson  and  Griswold,  ibid.,  53,  1311   (1931). 

6.  Fernelius  and  Bergstrom,  J.  Phys.  Chem.,  35,  741   (1931). 

7.  Kremann,  Z.  anorg.  Chem.,  33,  87   (1902). 

8.  Presented  at  the  Indianapolis  meeting  of  the  American  Chemical  Society,  April,  1931. 


Davidson  and  Holm:   Transference  of  Ions  465 

positive  metal  was  almost  quantitatively  deposited  on  the  anode,  in 
every  case,  in  the  form  of  a  nitrogen  compound.  The  authors  regard 
this  as  evidence  that  these  compounds  are  true  metallates  and  not 
merely  double  amides. 

Because  of  the  marked  analogy  between  the  sodium  zinc  acetate 
and  the  ammonium  cupric  acetate  mentioned  above,  and  corre- 
sponding compounds  in  the  water  and  ammonia  systems,  the  question 
naturally  arises  whether  or  not  the  former  compounds  are  true 
metallates ;  or,  in  other  words,  whether  the  amphoterism  of  zinc  and 
cupric  acetates  in  acetic  acid  includes  the  property  of  forming 
metallate  ions  in  the  presence  of  excess  of  acetate  ion,  or  whether 
it  is  limited  to  their  tendency  to  form  addition  compounds  with 
stronger  bases.  It  was  in  the  attempt  to  throw  additional  light 
on  this  question  that  the  present  work  was  undertaken. 

METHOD 

General  Plan.  It  was  hoped  that  some  indication  of  the  ionic 
species  present  in  such  acetic  acid  solutions  as  have  just  been  dis- 
cussed might  be  obtained  by  means  of  a  study  of  their  behavior 
during  electrolysis.  The  method  chosen  was  that  first  used  by 
Hittorf  ^  in  the  determination  of  transference  numbers — that  is,  the 
measurement  of  the  changes  in  concentration,  during  electrolysis,  of 
the  solution  surrounding  one  or  both  of  the  electrodes.  It  was 
originally  planned  to  employ  this  method  merely  to  detennine  the 
direction  of  migration  of  the  less  positive  metal,  but  the  results  of 
preliminary  experiments  indicated  such  marked  anomalies  of  trans- 
ference that  it  appeared  desirable  first  to  determine  quantitatively 
the  migration  ratios  in  some  simpler  systems,  especially  since  no 
such  experiments  on  acetic  acid  solutions  have  hitherto  been  re- 
ported.^" 

Preliminary  Experiments.  In  the  earliest  experiments  it  was 
found  that,  due  to  the  high  resistance  of  the  acetic  acid  solutions 
to  be  studied,  an  electromotive  force  of  110  volts  was  insufficient 
to  bring  about  appreciable  concentration  changes  in  a  reasonable 
period  of  time.  Hence  a  transformer  and  rectifier  capable  of  giving 
600  volts  were  used  in  the  subsequent  work.    This  device,  as  well 

9.  (a)  Hittorf,  "Ueber  die  Wanderungen  der  lonen  wahrend  der  Elektrolyse,"  Ostwald's 
Klassiker  der  exacten  Wissenschaften,  vol.  21,  23,  Leipzig,  1903-4.  See,  also,  (b)  Drucker, 
"Ostwald-Luther  Hand  und  Hilfsbuch  zur  Ausfiihrung  physiko-chemischer  Messungen,"  Leip- 
zig,  1931,  pp.  688-96. 

10.  Transference  numbers  in  acetic  acid  solutions  have,  however,  been  determined  by  the 
moving  boundary  method  by  Longsworth  (Thesis,  University  of  Kansas,  1928),  who  concluded 
that  the  transference  numbers  of  sodium  and  potassiiun  acetates  changed  little  with  concen- 
tration, and  approached  a  value  of  0.5  in  dilute  solutions. 


466  The  University  Science  Bulletin 

as  the  cell  used  for  the  quantitative  experiments,  will  be  described 
later. 

When  a  solution  of  sodium  and  zinc  acetates  was  electrolyzed,  at 
a  low  current  density,  between  platinum  electrodes,  hydrogen  was 
evolved  at  the  cathode,  but  no  gas  bubbles  whatever  could  be  ob- 
served at  the  anode.  It  appeared,  therefore,  that  it  would  be  pref- 
erable to  study  concentration  changes  around  the  anode,  where 
there  would  be  no  disturbance  due  to  evolution  of  gas,  and  some 
preliminary  trials  were  made  with  this  end  in  view.  In  one  case  a 
white  deposit,  shown  by  analysis  to  be  zinc  acetate,  was  obtained 
on  the  anode.  This  result,  which  suggested  the  presence  of  an 
anion  containing  zinc,  could  not  be  duplicated,  however.  Further 
experience  showed  that  it  would  be  difficult  to  determine  the  exact 
nature  of  the  anode  reaction  occurring  in  these  solutions,  or,  for  that 
matter,  even  in  solutions  of  alkali  acetates  electrolyzed  under  these 
conditions.^  ^  It  was  accordingly  decided  to  set  up  a  cell  with  a 
reversible  and  definitely  known  anode  reaction.  After  some  un- 
successful experiments  with  zinc  amalgam,  a  mercury  anode  was 
tried  and  proved  to  be  satisfactory.  On  passage  of  the  current,  a 
white  deposit  appeared  above  the  mercury;  the  decrease  in  weight 
of  the  metal  agreed  with  that  calculated,  on  the  basis  of  oxidation 
to  the  mercurous  state,  from  the  quantity  of  electricity.  The  de- 
posit was  found  to  yield  mercurous  ion,  but  no  mercuric,  and  must 
therefore  have  been  mercurous  acetate;  this  compound  was  so  in- 
soluble that  no  perceptible  precipitate  with  hydrogen  sulfide  was 
obtained,  in  any  case,  in  the  supernatant  solution.  No  evolution  of 
gas  was  detectable  with  this  electrode  at  current  densities  up  to 
0.8  ampere  per  square  decimeter. 

Apparatus.  The  cell  ^^  used  in  the  quantitative  experiments  was 
of  the  Loeb-Nernst  type,^^  and  is  pictured  in  figure  1.  Mercury  in 
the  bottom  of  the  longer  leg  served  as  anode,  connection  to  which 
was  made  by  means  of  a  platinum  wire  sealed  through  a  glass  tube. 
This  tube,  filled  with  mercury,  was  held  in  place  by  a  rubber  stopper, 
which  carried  also  a  tube  filled  with  calcium  chloride,  in  order  that 
moisture  might  be  excluded.  The  cathode  was  a  piece  of  platinmn 
foil,  to  which  connection  was  made  by  means  of  a  long  platinum 
wire  of  fine  gauge.     In  order  to  minimize  the  stirring  effect  of  gas 

11.  Hopfgartiier  (Monatsh.,  32,  523  [1911])  studied  the  electrolysis  of  sodium  acetate 
between  platinum  electrodes  and  found  that  carbon  dioxide,  ethane,  oxygen,  methyl  acetate 
and  other  substances  were  formed  at  the  anode,  the  relative  quantities  varying  with  current 
density,  concentration  and  temperature. 

12.  The  writers  are  indebted  to  Mr.  Jesse  Stareck  for  the  construction  of  this  cell. 

13.  Ref.    9b,   p.    689. 


Davidson  and  Holm:  Transference  of  Ions 


467 


liberated  at  the  cathode,  a  funnel-shaped  deflector  was  ]ilaced  with 
its  wider  end  over  the  cathode,  while  the  smaller  end  passed  through 
the  stopper  of  the  cathode  compartment  and  was  connected  to  a 
second  drving  tube.     A  small  hole  in  the  deflector  tube  above  the 


Fig.   1.     Cell   used   for  transference   measurements. 


468  The  University  Science  Bulletin 

surface  of  the  liquid  allowed  atmospheric  pressure  to  be  maintained 
in  this  part  of  the  cell.  The  cathode  lead  wire  passed  out  through 
the  end  of  the  deflector  and  under  the  rubber  tubing  which  made 
connection  with  the  drying  tube,  as  shown  in  the  diagram. 

The  source  of  current  was  a  Majestic  B  battery  eliminator  which 
had  been  rebuilt  to  deliver  a  maximum  of  about  600  volts  when 
connected  in  the  110-volt  alternating-current  circuit;  a  Kenetron 
tube  (type  280)  served  as  rectifier,  and  a  rheostat,  connected 
across  the  alternating-current  input,  made  it  possible  to  adjust  the 
voltage.  The  secondaiy  (direct-current)  circuit  contained  the  elec- 
trolytic cell  described  above,  a  standard  copper  eoulometer  and  a 
Weston  three-range  ammeter. 

Preparation  of  Materials.  The  anhydrous  acetic  acid  was  pre- 
pared by  the  method  of  Kendall  and  Gross  j^'*  the  product  used  in 
this  work  had  a  freezing  point  of  16.5-6°.  The  sodium  acetate  was 
dehydrated  for  several  days  at  150°  and  then  kept  in  stoppered 
bottles  in  a  desiccator.  Its  purity  was  verified  by  checking  the 
freezing  points  of  several  solutions  of  the  salt  in  acetic  acid  against 
the  data  of  Kendall  and  Adler.^^  Anhydrous  zinc  acetate  was  pre- 
pared as  described  in  a  previous  paper  from  this  laboratory  ;^  analy- 
sis for  zinc  by  potentiometric  titration  with  potassium  ferrocyanide 
solution  gave  35.70  per  cent,  as  compared  with  the  calculated  35.65 
per  cent.  Ammonium  acetate  was  obtained  by  passing  anhydrous 
ammonia  over  acetic  acid,  as  described  in  the  same  paper;  it 
was  found  that  either  the  normal  salt  NH^C.HaOo  or  the  solvate 
NH^CoHgO,  .  HC2H3O2  might  be  obtained  by  this  method,  but  since 
the  concentration  of  all  ammonium  acetate  solutions  was  determined 
by  analysis,  the  relative  amounts  of  the  components  ammonia  and 
acetic  acid  in  the  salt  used  for  preparing  them  was  of  no  consequence, 
as  long  as  no  other  substances  were  present.  Cupric  acetate  was 
prepared  as  described  in  another  paper  already  referred  to;^  analysis 
for  copper  by  the  electrolytic  method  gave  34.96  per  cent,  as  com- 
pared with  the  calculated  35  per  cent. 

An.\lysis  of  the  Solutions.  Solutions  of  sodium  acetate  were 
analyzed  by  adding  a  slight  excess  of  sulfuric  acid,  evaporating 
slowly  to  dryness,  heating  gently  until  the  excess  sulfuric  acid  had 
been  driven  off  and  then  strongly  to  constant  weight  of  sodium  sul- 
fate. In  the  case  of  solutions  containing  both  sodium  and  zinc,  the 
zinc  was  determined  by  potentiometric  titration  with  potassium 
ferrocyanide  solution.     Attempts  to  determine  both  metals  in  the 


14.  Kendall  and  Gross,  J.  Am.  Cliem.  Soc,  43,  1426  (1921). 

15.  Kendall  and  Adler,  ibid.,  43,  1470   (1921). 


Davidson  and  Holm:   Transference  of  Ions  409 

same  sample  failed  to  yield  results  as  consistent  as  desired,  and 
since  a  high  degree  of  accuracy  in  the  analyses  was  essential  if  the 
transport  figures  were  to  have  any  significance,  this  plan  was  not 
followed.  The  analyses  for  cupric  acetate  were  made  by  the  iodi- 
metric  method,  and  those  for  ammonium  acetate  by  adding  an  excess 
of  sodium  hydroxide  solution  and  distilling  the  liberated  ammonia 
into  standard  hydrochloric  acid.  In  almost  every  case  duplicate 
analyses  were  made,  which  agreed  to  well  within  the  probable  error 
of  the  Hittorf  method. 

Measurement  of  Transference.  Transference  studies  were 
made  on  acetic  acid  solutions  of  sodium  and  ammonium  acetates 
over  as  wide  a  range  of  concentrations  as  possible,  namely  from 
about  ]  to  7  mole  per  cent  of  solute  (0.15  to  1.25  molal).  More 
dilute  solutions  could  not  be  used  because  of  their  extremely  low  con- 
ductivity, while  at  high  concentrations  the  relative  change  during 
electrolysis  would  be  so  small  as  to  impair  the  accuracy  of  the  re- 
sults. A  study  was  made  also  of  the  transference  of  zinc  ion  in  solu- 
tions containing  both  sodium  and  zinc  acetates,  and  of  cupric  and 
ammonium  ions  in  solutions  of  both  of  these  acetates.  (In  these 
cases  deposits  of  zinc  and  of  copper,  respectively,  appeared  on  the 
cathodes  during  electrolysis,  while  no  trace  of  zinc  or  of  cupric 
acetate  could  be  detected  in  the  anode  deposit.) 

The  procedure  in  making  a  run  was  as  follows.  About  thirty 
grams  of  redistilled  mercury  were  accurately  weighed,  and  trans- 
ferred to  the  longer  leg  of  the  electrolytic  cell,  which  was  to  serve 
as  the  anode  compartment.  The  solution  to  be  electrolyzed  was 
then  introduced,  reaching  a  level  slightly  above  that  of  the  connect- 
ing tube.  The  stoppers,  fitted  as  described  above,  were  put  in  place 
and  the  electrical  circuit  was  completed.  The  current  was  adjusted 
to  the  desired  value  of  a  few  milliamperes  and  electrolysis  continued 
until  0.05  to  0.15  gram  of  copper  had  been  deposited  on  the  cathode 
of  the  coulometer;  the  time  reciuired  for  this  varied  from  six  to 
twenty-four  hours.  At  the  end  of  the  electrolysis  this  cathode  was 
immediately  washed,  dried  and  weighed.  The  stopcock  in  the  cath- 
ode leg  was  opened  and  the  solution  from  this  compartment  and  the 
connecting  tube  slowly  withdrawn.  Then  the  total  remaining  con- 
tents of  the  anode  leg  were  transferred  to  a  weighed  stoppered 
flask.  After  it  had  been  weighed  again,  this  was  set  aside  to  allow 
the  mercurous  acetate  to  settle  out  completely.  Portions  of  the  solu- 
tion were  then  decanted  into  weighing  bottles  for  analysis. 

As  has  been  stated  previously,  at  the  beginning  of  this  work  it  was 

31—3482 


470 


The  University  Science  Bulletin 


not  intended  to  determine  more  than  the  direction  of  transference 
of  the  metallic  constituents;  the  data  obtained,  however,  made  it 
possible  to  calculate  the  apparent  transport  numbers  of  the  cations 
in  the  pure  sodium  and  ammonium  acetate  solutions,  and  the  frac- 
tion of  the  current  apparently  carried  by  the  less  positive  metal 
in  the  mixed  acetates,  although  the  highest  degree  of  accuracy  in 
these  numerical  values  was  not  striven  for.  The  temperature  was 
that  of  the  laboratory,  about  25°C.  No  thermostat  was  used,  nor 
was  the  middle  portion  analyzed  to  insure  constancy  of  concentra- 
tion in  this  region.  It  is  believed,  however,  that  the  anode  portion 
taken  for  analysis  (usually  about  30  grams,  or  about  two-fifths  of 
the  whole  solution)  included  all  the  changes  caused  by  transference 
and  electrode  reactions  at  the  anode.  The  changes  in  density  were 
very  slight.  The  transport  numbers  given  are  probably  correct  to 
±0.05.  It  should  be  mentioned  further  that  no  attempt  was  made 
to  correct  the  apparent  transport  numbers  for  possible  solvation  of 
the  ions,  as  this  would  have  involved  a  tremendous  increase  in  ex- 
perimental difficulties  without  commensurate  increase  in  the  sig- 
nificance of  the  values  obtained. 

RESULTS 

Space  does  not  permit  the  tabulation  of  all  the  data  obtained. 
The  results  are  summarized  in  the  following  tables. 


TABLE  1. — Transference  data 

for  sodium  acetate 

Concentration  of 
original  solution. 

Trial 

Concentra- 
tion of 
anode 
solution 

Copper 
deposited  in 

Transference 

of  sodium 

acetate 

Apparent 

transport 

Mole 

Molal- 

Weight 

number. 

after 
electrolysis, 

coulometer, 
grams. 

from 
anode, 

number 
of  cation. 

per 
cent. 

ity. 

per 
cent. 

weight 
per  cent. 

grams. 

0.99 

0.167 

1.34 

1 
2 

1.36 
1.35 

0.02C3 
0.0580 

—0.004 
—0.003 

—0.08 
—0.02 

3 

1.34 

0.0170 

0 

0 

4 

1.34 

0.0176 

0 

0 

M 

—0.03 

2.76 

0.473 

3.74 

5 

3.69 

0.0592 

0.017 

0.11 

5.06 

0.888 

6.80 

6 

7 

6.62 
6.68 

0.1279 
0.0716 

0.066 
0.048 

0.20 
C.26 
0.23 

7.02 

1.257 

9.35 

8 

9.25 

0.0755 

0.040 

0.21 

Davidson  and  Holm:   Transference  of  Ions  471 

TABLE  2. — Transference  data  for  ammonium  acetate 


Concentration  of 
original  solution. 

Trial 
number. 

Concentra- 
tion of 
anode 
solution 
after 
electrolysis, 
weight 
per  cent. 

Copper 
deposited  in 
coulometer, 

grams. 

Transference 
of  ammonium 

acetate 
from 

anode, 

grams. 

Apparent 

Mole 
per 
cent. 

Molal- 
ity. 

Weight 
per 
cent. 

transport 
number 
of  cation. 

0.84 

0.140 
0.312 
0.556 

1.248 

1.07 
2.35 
4.10 

8.78 

1 

2 

Mean 

3 

4 

Mean 

5 

6 

Mean 

7 

8 

Mean 

1.06 
1.04 

0.0789 
0.1058 

0.002 
0.010 

0.01 
0.04 
0  03 

1.84 

2.28 
2.33 

0.1533 
0.1076 

0.025 
0.008 

0.07 
0.03 
0.05 

3.23 

4.00 
4.02 

0.0701 
0.0844 

0.032 
0.027 

0.19 
0.13 
0.16 

6.97 

8.68 
8.62 

0.0752 
0.1175 

0.035 
0.061 

0.19 
0.21 
0.20 

TABLE  3.- 

-Transference  data  for  sodium  acetate- 

zinc  acetate 

Concentration  of  original  solution. 

Mole 
ratio 
zinc 
acetate: 
total 
solute. 

Copper 
deposited  in 
coulometer, 

grams. 

Transference 
of  zinc 
acetate 
from 
anode, 
grams. 

Sodium  acetate. 

Zinc  acetate. 

Fraction 

of  current 

carried 

Mole 
per^cent. 

Molal- 
ity. 

Mole 
per  cent. 

Molal- 
ity. 

by  zinc. 

3.35 
6.66 

0.579 
1.192 

0.27 
0.26 

0.047 
0.047 

0.074 
0.038 

0.0797 
0.0997 

0.046 
0.034 

0.20 
0.12 

472  The  University  Science  Bulletin 

TABLE  4. — Transference  data  for  ammonium  acetate -cupric  acetate 


Concentration  of 

original  solution. 

Fraction 

Mole 

ratio 

cupric 

acetate: 

Trial 
No. 

Copper 
depos- 
ited in 
coulom- 

Transference  from 
anode,  grams. 

of  current 

Ammonium 
acetate. 

Cupric  acetate. 

cairied 
by  cation. 

Mole 
per 
cent. 

Molal- 
ity. 

Mole 
per 
cent. 

Molal- 
ity. 

total 
solute. 

eter, 
grams. 

NH4C2H3O2 

CuCCzHsOo)^ 

NH4+ 

Cu+^- 

0.81 

0.136 

0.214 

0.036 

0.209 

1 

0.1342 

0.C15 

0.089 

0.04 

0.23 

2 

0.1119 

0.011 

0.083 

0.04 

0.26 

3 

Mean 

0.1379 

0.088 

0.04 

0.22 

0.23 

2.68 

0.460 

0.300 

0.052 

0.101 

4 

0.1438 

0.015 

C.099 

0.04 

0.24 

5 

0.2339 

0.028 

0.116 

0.05 

C.17 

6 

C.1345 

0.020 

0.077 

0.06 

0.20 

Mean 

0.05 

0.20 

8.30 

1 .  .555 

0,440 

0.081 

0.049 

7 
8 

9 
Mean 

C.1818 
C.2112 
0.0693 

0.060 

0.073 
0.072 

0.14 

0.12 
0.13 

0.14 
0.12 

0.020 

0.13 

10.83 

2.030 

0.292 

0.055 

0.026 

10 

0.0976 

0.088 

0.024 

0.37 

0.08 

11 

0.0956 

0.100 

0.022 

0.43 

0.08 

Mean 

0.40 

0.08 

22  30 

4.802 

0.346 

0.075 

0.015 

12 

13 

\'ean 

0.1203 
0.1194 

0.006 
0.013 

0  02 

0  04 

0.03 

Davidson  and  Holm:  Transference  of  Ions 


473 


DISCUSSION 

SODIUM  ACETATE  AND  AMMONIUM  ACETATE  SOLUTIONS 

The  variation  of  the  transport  number  of  the  cation  in  these  solu- 
tions with  concentration  is  shown  graphically  in  figures  2  and  3. 
The  rapid  increase  of  the  apparent  transport  number,  and  its  ap- 
proach to  a  normal  value,  with  increasing  concentration,  is  quite 
different  from  the  behavior  of  most  salts  in  aqueous  solution,  where 
migration  ratios  do  not  vary  greatly  with  concentration.  The  fact 
that  in  the  most  dilute  solution  of  sodium  acetate  the  transport  num- 
ber apparently  has  a  slightly  negative  value  is  particularly  anom- 


o 
<_) 

O 

a: 
en 


cc 
o 

a 

2 
< 


0.4 
0.3 
0.2 
0.1 
0.0 


o 

o 

y 

^ 

/ 

/ 

/ 

o 
< 


(X 
aJ 


ct: 
o 


0.4 


0.3 


0.2 


0.0 


-0.1 


o    . 

^^ 

/o 

7 

MOLL   PLR  CLNT.  OF 
SODIUM    ACLTATL 

Fig.  2.     Apparent  transport  number  of  cation  in 
sodium  acetate  solutions. 


MOLE.    PER   CLNT.   OF 
AMMONIUM    ACLTATE. 

Apparent  transport  number  of  cation  in 
ammonium  acetate  solutions. 


alous.  In  the  case  of  aqueous  solutions  marked  abnormalities  of  this 
sort  are  ordinarily  attributed  to  the  formation  of  complex  ions,  as, 
for  instance,  in  the  case  of  cadmium  iodide,  where  ions  such  as 
Cdlj-  or  Cdl^--  are  assumed  to  be  present.  It  is  possible  that  similar 
phenomena  may  occur  in  acetic  acid  solutions.  Thus,  in  the  case  of 
sodium  acetate,  if  we  were  to  suppose  that  the  dissociation  followed 
the  course  shown  by  the  equation — 

2NaC2H302  ^  Na-  +  NaCC^H^O^),- 
and,  further,  that  the  two  ions  here  shown  had  nearly  the  same 
mobilities,  we  could  thereby  account  for  the  fact  that  there  appears 
to  be  no  migration  of  sodium  from  the  anode  in  dilute  solutions.  It 
would  be  difiBcult,  however,  to  explain  why  such  an  abnormality 
should  be  greater  in  dilute  than  in  concentrated  solutions. 


474  The  University  Science  Bulletin 

If  some  knowledge  of  the  molecular  species  present  in  an  acetic 
acid  solution  of  a  typical  electrolyte  like  sodium  acetate  (which  is 
the  one  for  which  the  most  data  are  available)  could  be  gained  from 
a  study  of  other  properties  of  such  a  solution,  light  might  be  thrown 
also  on  the  anomalous  transference.  It  might  be  expected,  for  in- 
stance, that  a  study  of  freezing-point  data  would  give  some  clue  as 
to  the  molecular  state  of  the  solute.  In  the  following  table  the  freez- 
ing points  of  sodium  acetate  solutions,  from  the  data  of  Kendall  and 
Adler^^  and  of  Webb,^^  are  compared  with  those  calculated^'^  for  a 
perfect  undissociated  solute  in  acetic  acid. 


TABLE  5.- 

—Freezing  points  of  acetic  acid  solutions 

Mole  per  cent  solute ....        0 

Ideal  solute,   °C 16.60 

Sodium  acetate,   °C.  ...   16.60 

0.3           0.5              1                2               3 
16.42        16.30       16.00       15.40       14.79 
16.45       16.36        16.07        15.45       14.77 

4 
14.17 
14.10 

6 
13.55 
13.47 

Evidently,  despite  the  dissociation  of  sodium  acetate  into  ions,  the 
freezing-point  depressions  throughout  this  range  are  almost  the 
same  as  the  ideal  values;  for  ammonium  acetate  the  corresponding 
depressions  are  even  smaller.  While  it  is  apparent  that  we  have  here 
a  state  of  affairs  quite  different  from  that  existing  in  aqueous  solu- 
tions, the  correct  interpretation  of  the  data  is  by  no  means  obvious, 
since,  due  to  the  very  low  dielectric  constant  of  the  solvent,  large 
interionic  forces  are  present,  and  these  may  be  mainly  responsible 
for  the  deviation  from  the  behavior  of  an  ideal  ionized  solute. 
Widely  divergent  views  have  been  expressed  as  to  the  molecular 
state  of  sodium  acetate  in  these  solutions.  Thus  Walden^^  appar- 
ently regarded  this  compound  as  being  only  slightly  dissociated  into 
ions  and  probably  associated  to  some  extent  into  polymeric  mole- 
cules. Webb,^^  on  the  other  hand,  assuming  the  electrolyte  to  be 
practically  completely  dissociated,  accounted  for  the  low  freezing- 
point  depressions  in  terms  of  interionic  forces  as  calculated  accord- 
ing to  the  Debye-Hiickel  theory.    A  viewpoint  intermediate  between 


16.  Webb,  J.  Am.  Ghem.  Soc,  48,  2263  (1926) 

AH 

17.  From  the  equation  In  Ni  = 


R      \T„,     T/  i2r,„/Ai/ .  InNi 

where  T  is  the  freezing  point  of  the  solution,  Tm  that  of  pure  acetic  acid,  289.7°, 
AH  the  molal  heat  of  fusion  of  acetic  acid,  2,780  calories  (de  Visser,  Rec.  trav. 
chim.,  12,  101  [1893]),  and  Ni  the  mole  fraction  of  acetic  acid;  R  has  its  usual 
significance.     On  substituting  the  numerical  values  and  changing  to  common 

289.7 

logarithms,  this  equation  takes  the  form  T  = . 

1—0.4771  log  Ni 

18.    Walden,     "Molekulargrossen     von     Elektrolyten     in     nichtwasserigen     Losungsmitteln," 
Dresden  and  Leipzig,  1923,  p.  132-5. 


Davidson  and  Holm:   Transference  of  Ions  475 

these  extremes  was  adopted  by  Hall  and  Werner/^  who  concluded, 
on  the  basis  of  electrometric  titration  and  dilution  curves,  that  al- 
though sodium  acetate  is  to  be  classed  among  the  "stronger"  bases 
in  acetic  acid,  yet  it  cannot  be  regarded  as  being  completely  dis- 
sociated in  the  usual  sense,  since  its  basicity  changes  with  dilution 
as  though  the  electrolyte  obeyed  the  dilution  law. 

It  is,  of  course,  possible,  by  thermodynamic  treatment  which  does 
not  require  any  hypothesis  as  to  the  actual  molecular  state  of  the 
solute,^"  to  calculate  the  activity  coefficient  of  the  electrolyte  from 
the  freezing-point  data.  This  having  apparently  not  been  done 
hitherto,  we  have  made  the  calculation  by  the  method  of  RandalF^ 
for  several  concentrations,  with  the  results  shown  in  Table  6. 


TABLE  6.- 

— Activity  coeiBcient  of  sodium  acetate  in  acetic  acid 

0.001          0.01            0.1 

1.0 

0.366          0.121          0.032 

0.010 

Our  knowledge  of  acetic  acid  solutions  is,  in  our  opinion,  as  yet 
inadequate  to  provide  a  satisfactory  explanation  for  these  veiy  low 
activity  coefficients,  or  even  to  permit  a  certain  decision  among  the 
conflicting  viewpoints  as  to  the  molecular  state  of  this  electrolyte. 
Little  additional  light  is  thrown  upon  the  question  by  the  available 
conductivity  data  for  sodium  acetate,^^  which  show  a  minimum 
value  of  the  equivalent  conductance  at  a  concentration  of  about 
0.05  M.  It  is  evident  that  any  attempt  to  calculate  the  degree  of 
dissociation  from  conductivity  data  by  the  classical  method  of 
Arrhenius  would  be  quite  futile.  On  the  other  hand,  it  appears  that 
interionic  attraction  theories  alone,  based  upon  the  assumption  of 
practically  complete  ionization,  might  be  equally  inadequate  to  deal 
with  this  case,  since  some  sort  of  ionic  association  almost  certainly 
occurs.  The  very  recent  work  of  Fuoss  and  Kraus^^  seems  to  indi- 
cate, however,  that  a  better  understanding  of  the  state  of  electrolytes 
in  solvents  of  low  dielectric  constant  may  eventually  be  attained  by 
means  of  a  further  study  of  conductance  data  for  dilute  solutions  by 
a  method  derived  from  both  viewpoints. 

19.  Hall  and  Werner,  J.  Am.  Cliem.  Soc,  50,  2367  (1928);  see,  also,  Hall,  Cheni.  Re- 
views, 8,  208  (1931). 

20.  See  Randall  and  Allen,  J.  Am.  Chem.  Soc.,  52,  1814  (1930). 

21.  Randall,  ibid.,  48,  2512  (1926).  The  value  of  A  was  taken  as  3.604,  and 
the  Debye-Hiickel  limiting  law,  in  the  form  logy  =  —  25  mi,  was  used  as  an 
aid  in  determining  the  limiting  value  of  jlm\  at  infinite  dilution. 

22.  Hopfgartner,  Sitz.  Akad.  Wiss.  Wien,  120,  II,  1024  (1911);  Longsworth,  Thesis, 
University  of  Kansas,  1928. 

23.  Fuoss  and  Kraus,  J.  Am.  Chem.  Soc,  55,  1919,  2387  (1933). 


476  The  University  Science  Bulletin 

It  should  be  mentioned  also  that  Freed  and  Kasper,"-*  on  the  basis 
of  magneto-chemical  studies,  question  the  validity  of  the  transport 
number  as  a  criterion  of  the  formation  of  complex  ions,  believing 
that  anomalous  transference  may  eventually  be  accounted  for  on  the 
basis  of  a  hypothesis  of  ionic  clusters.  For  the  present  we  can  say 
only  that  although  the  abnormal  transference  numbers  of  sodium 
and  ammonium  acetates  rnay  be  due  to  the  presence  of  complex  ions, 
we  have  no  positive  evidence  that  this  is  the  case. 

SODIUM-ZINC  AND  AMMONIUM-CUPRIC  ACETAl'E  SOLUTIONS 

Because  of  the  amphoteric  properties  exhibited  by  zinc  and  cupric 
acetates  in  acetic  acid,  it  was  expected  that  some  indication  might 
be  found  of  the  migration  of  zinc  or  copper  toward  the  anode.  This 
appeared  particularly  likely  in  the  presence  of  a  large  excess  of  ace- 
tate ion  (that  is,  in  solutions  containing  high  concentrations  of 
sodium  or  ammonium  acetate) ,  in  view  of  the  frequent  occurrence  of 
such  reversed  transference  in  aqueous  solutions  under  similar  con- 
ditions.25  However,  the  data  of  Tables  3  and  4  show  a  decrease  in 
zinc  or  cupric  acetate  in  the  anode  compartment  in  every  case,  and 
hence  give  no  evidence  of  reverse  migration.  The  determination  of 
the  conventional  transport  numbers  for  zinc  or  copper  in  the  corre- 
sponding acetates,  under  these  conditions,  would  have  required  a 
knowledge  of  the  distribution  of  the  current  between  the  two  bases 
present  together  in  each  solution,  and  even  an  approximate  calcula- 
tion of  such  distribution  was  impossible.  Thus,  in  a  solution  of 
sodium  and  zinc  acetates,  the  fraction  of  the  current  carried  by  the 
zinc  acetate  would  depend  both  on  the  relative  concentrations  and  on 
the  relative  conductivities  of  the  two  bases ;  but  no  conductivity  data 
for  zinc  acetate  are  available,  and  in  fact  this  compound  is  so  slightly 
soluble  that  solutions  of  the  concentration  here  used,  but  containing 
no  sodium  acetate,  could  not  possibly  be  prepared.  However,  the 
data  sufficed,  in  every  case,  for  the  calculation  of  the  fraction  of  the 
total  current  carried  by  the  less  positive  cation. 

In  figure  4  the  fractions  apparently  transported  by  ammonium  and 
by  copper  in  the  mixed  acetate  solutions  are  plotted  against  the  con- 
centration of  ammonium  acetate.  It  is  evident  that  the  transference 
of  copper  towards  the  cathode  decreases  with  increasing  concentra- 
tion of  ammonium  acetate,  but  this  result  would  necessarily  follow 


24.  Freed  and  Kasper,  ibid.,  .52,  2G32   (1930). 

25.  McBain    and    Van   Rysselberge,   J.    Am.    Chem.    Soc,    .50,    3009    (1928),   and    52,    2336 
(1930). 


Davidson  and  Holm:  Transference  of  Ions 


477 


from  the  increasing  fraction  of  current  carried  by  the  hitter  base. 
In  fact,  both  zinc  and  copper,  in  the  mixed  solutions,  appear  to  carry 
a  far  larger  share  of  the  total  quantity  of  electricity  than  would  be 
expected  from  their  relative  concentrations.  These  experiments, 
then,  give  no  evidence  of  the  presence  in  the  solutions  examined  of 
complex  anions  containing  either  copper  or  zinc.  In  view,  however, 
of  the  highly  anomalous  nature  of  the  conductance  phenomena  even 
in  solutions  of  single  salts  in  this  solvent  it  can  scarcely  be  concluded 
that  the  existence  of  such  metallate  ions  is  definitely  disproved,  and 
the  nature  of  amphoterism  in  acetic  acid  solutions  remains  a  sub- 
ject for  further  study. 


^ 


< 
o 

u-I 
CK 

ID 


06 
0.5 
0.4 
03 
02 


2  01 


0.0 


i 

COPF 
AMMONI 

UM» 

1 

^^ 

/ 

« 

— 

o^ — — 

^~~'~— 



0 

5  10 

lOLL  PtR  CtNT.  OF 


15  20 

AMMONIUM  ACtTATL 


Fig.  4.     Transference  of  copper  and  of  animonium  in  ammonium  cupric  acetate  solutions  con- 
taining 0.3  mole  per  cent  of  cupric  acetate. 


THE  UNIVERSITY  OF  KANSAS 

SCIENCE  BULLETIN 

Vol.  XXL]  March,  1933.  [No.  16. 


The  Effects  of  a  Minimal  Lethal  Dose  of  X-Rays  upon 
Chick  Embryos 

H.  ERNEST  CROW,  Friends  University,  Wichita,  Kan. 


Absthact  :  Chick  embryos  of  28,  48,  60,  75,  96  and  120  hours  of  incubation 
were  exposed  to  unfiltered  X-rays  with  the  current  set  at  22  a.  and  90  v.  for  15 
minutes  at  a  distance  of  18  cm.  This  do.sage  was  found  to  be  the  mmimal 
amount  that  would  cause  death  of  the  chicks  in  15-20  hours.  The  different 
tissues  and  organs  of  the  chicks  were  carefully  studied  in  serial  sections  at 
periods  from  20  minutes  to  20  hours  after  raying.  Effects  became  observable 
in  about  two  hours  and  became  more  pronounced  with  the  passage  of  time. 
All  the  tissues  of  the  body  were  affected,  but  the  greater  modifications  occurred 
in  the  blood  and  nervous  systems.  Separation  and  fraying  of  cell  layers,  break- 
ing of  limiting  membranes  and  capillary  walls,  accompanied  by  the  accumula- 
tion of  disorganized  cellular  debris  within  the  brain  and  other  cavities,  and 
extensive  hemorrhages,  were  common,  general  effects.  In  dividing  cells  the 
principal  modifications  were  inhibited  mitoses,  clumped  chromosomes,  unequal 
divisions,  pycnosis  and  fragmentation  of  chromosomes.  In  other  cells  nuclear 
and  cytoplasmic  substances  were  separated  into  large,  irregular  granules  and 
clear  vacuoles,  cell  walls  were  broken,  cytoplasm  oozed  from  the  ends  of  cells 
and  nuclei  became  shrunken.  The  characteristic  affinity  of  chromatin  for  Iron 
Hsematoxylin  stain  was  much  disturbed  after  a  few  hours,  a  faded  gray  or  al- 
most no  color  being  common. 


INTRODUCTION 

IN  the  three  and  one-half  decades  since  the  discovery  by  Rontgen 
of  the  rays  which  bear  his  name,  or  popularly  called  X-rays, 
much  interest  has  grown  around  the  use  of  X-rays  in  the  treatment 
of  disease.  Only  a  very  limited  number  of  workers  have  used  X-rays 
in  experimental  research  to  determine  the  effects  produced  upon  the 
cells  rayed.  The  greater  number  of  these  have  confined  their  at- 
tention to  tissues  grown  in  vitro.  The  embryo  of  the  chick  is  a 
favorite  experimental  animal,  yet  only  a  few  workers  have  used  it 

(479) 


480  The  University  Science  Bulletin 

for  the  study  of  the  effects  of  X-rays  upon  the  Hving  animal.  It 
was  with  a  desire  to  investigate  the  effects  of  X-rays  upon  the  tissues 
of  the  young  chick  embryo  that  this  work  was  undertaken. 

The  experiments  herein  recorded  have  for  their  object: 

To  investigate  the  effects  of  single  exposures  of  unfiltered  X-rays 
of  constant  dosage  upon  chick  embryos  of  different  ages. 

To  discover,  if  possible,  what  tissue,  if  any,  is  more  afTected  than 
others. 

To  learn  what  cellular  changes  take  place  as  a  result  of  the 
X-rays. 

ACKNOWLEDGMENTS 

To  Prof.  H.  H.  Lane,  head  of  the  Department  of  Zoology,  Uni- 
versity of  Kansas,  under  whose  direction  the  present  work  was 
done,  for  his  kindly  interest  and  helpful  suggestions,  the  author 
wishes  to  express  his  sincere  gratitude.  Also,  he  wishes  to  thank 
Prof.  C.  V.  Kent,  professor  of  physics  in  the  ITniversity  of  Kansas, 
for  his  help  in  the  use  of  the  X-ray  apparatus. 

HISTORICAL 

Rolleston  (75)  has  summarized  in  a  recent  paper  the  growth  in 
interest  in  X-ray  effects  on  living  tissues  as  well  as  showing  their 
harmful  effects  upon  man. 

Experimental  study  of  effects  of  X-rays  on  living  cells  was  be- 
gun by  Perthe  (66),  who  exposed  the  eggs  of  Ascaris  ryiegalocephala 
to  X-rays  and  found  delayed  development  of  body  and  the  pro- 
duction of  individuals  abnormal  in  some  parts  of  the  body.  In 
1904  Oilman  and  Baetjer  (35)  exposed  hen  eggs  to  X-rays  for 
short  periods  on  successive  days  and  noted,  at  first,  a  slight  ac- 
celeration of  development,  followed  by  production  of  abnormali- 
ties, deformities,  retardation  of  eyes  and  adherence  of  membanes 
to  the  body  of  the  embryo. 

Hartman  (41)  in  1920  exposed  frog  larvae  to  X-rays  and  found 
reduction  in  number  of  leucocytes;  while  retardation  of  growth 
and  reduction  of  cell  number  in  lymphoid  tissue  in  rats  and  mice 
when  exposed  to  X-rays  were  observed  by  Cramer,  Drew  and 
Mottram  (27),  in  1921.  Jolly  (49),  1924,  confirmed  their  results 
in  experiments  on  guinea  pigs  where  a  difference  in  susceptibility 
to  X-rays  was  noted  between  bone  and  lymphoid  tissues;  and  in 
1927  Brambell,  Parkes,  and,  Fielding  (18)  caused  sterility  in  the 
adult  mouse  by  raying  its  ovary,  and  in  the  young  by  exposure 
before  birth. 


Crow:  Effects  of  X-Rays  ox  Chick  Embryos  481 

Wright  and  Bulmar  (93)  in  1929  found  rabbits  and  cats  were 
killed  by  X-rays,  but  the  former  requiring  twice  the  time  of  ex- 
posure for  the  effect  to  be  lethal. 

Eggs  of  frog,  rayed  by  Ancel  and  Vintemberger  (5)  1924,  did 
not  show  effects  immediately,  but  death  occurred  in  a  few  hours 
after  gastrulation  began.  The  same  authors  (14) ,  1928,  found  a 
given  dosage  affected  frog  eggs  to  a  greater  extent  when  employed 
as  a  single  dose  than  when  spread  over  two  or  more  exposures. 

Specific  effects  upon  cells  were  reported  by  Ancel  and  Vintem- 
berger (14),  1928,  on  frog  eggs.  It  was  found  that  the  time  of 
least  susceptibility  was  when  the  egg  was  in  a  state  of  nuclear 
repose.  Vintemberger,  1928,  by  careful  and  ingenious  experimen- 
tation on  frog  eggs  proved  that  the  nuclei  are  very  susceptible  to 
X-rays  and  the  cytoplasm  very  little,  if  at  all,  affected;  this  con- 
firmed the  results  secured  by  Albert  and  Politzer  (3),  1924,  on 
Ambystoma  larvae  where  the  X-rays  affected  the  arrangement, 
splitting  and  distribution  of  chromosomes.  Chick  embryos  have 
proven  good  materials  for  experimental  work.  Ancel  and  Vintem- 
berger (8),  1924,  in  their  work  on  chick  embryos  found  support 
for  the  prevalent  idea  that  cells  are  more  sensitive  to  X-rays  when 
in  mitosis.  Heim  (42),  1927,  raying  eggs  before  they  were  laid  and 
embryos  at  various  ages,  found  in  all  cases  growth  retarded  and 
marked  eye  and  brain  defects,  the  effects  varying  inversely  with 
the  age.  In  1922  and  1926  Colwell,  Gladstone  and  Wakely  (24- 
25),  by  subjecting  chick  embryos  of  various  ages  to  daily  doses  of 
X-rays,  produced  marked  degeneration  of  all  tissues,  but  especially 
epithelial,  nerve  and  blood  tissues.  By  single  doses  of  different 
intensity  Strange  ways  and  Fell  (80)  produced  similar  degenerative 
effects  to  those  produced  when  heavy  doses  were  used,  but  there 
was  a  partial  recovery  from  lighter  doses.  Yamamoto  (94)  found 
inhibition,  retardation  or  modification  of  whole  or  parts  of  organs, 
and  death  in  most  cases  resulted  when  chick  embryos  were  exposed 
to  sufficient  dosage  of  X-rays.  In  his  report  only  gross  exteraal 
modifications  are  described. 


482  The  University  Science  Bulletin 

MATERIALS  USED 

The  chick  embiyos  used  are  in  seven  series,  i.  e.,  rayed  at  seven 
different  ages,  as  follows: 

Series  I rayed  at      0  hours  of  incubation 

Series  I rayed  at  100  hours  of  incubation 

Series  I rayed  at  110  hours  of  incubation 

Series  I rayed  at  140  hours  of  incubation 

Series  I rayed  at  160  hours  of  incubation 

Series  II rayed  at     28  hours  of  incubation 

Series  III rayed  at     48  hours  of  incubation 

Series  IV rayed  at     60  hours  of  incubation 

Series  V rayed  at     75  hours  of  incubation 

Series  VI rayed  at     96  hours  of  incubation 

Series  VII rayed  at  120  hours  of  incubation 

With  the  exception  of  Series  I,  in  which  time  of  exposure  was 
variable,  all  were  exposed  to  the  X-rays  for  the  same  length  of 
time,  i.  e.,  fifteen  minutes.  This  is  the  time  that,  for  a  given  dis- 
tance from  the  target,  and  at  a  certain  dosage,  was  the  minimal 
lethal  dose. 

TECHNIQUE  OF  IRRADIATION 

The  apparatus  employed  was  one  of  the  coils  fitted  with  a 
Coolidge  tube  used  in  the  physics  department  of  the  University 
of  Kansas.  The  current  was  set  at  22  a.  and  90  v.  The  eggs  were 
placed  at  a  distance  of  18  cm.  from  the  Tungsten  target  during 
time  of  raying  without  any  filter.  All  the  time  the  eggs  were  out 
of  the  incubator  they  were  kept  well  wrapped  in  layers  of  warm 
cotton  in  order  to  avoid,  as  far  as  possible,  any  change  of  tem- 
perature. After  the  period  of  raying  the  eggs  were  returned  at 
once  to  the  incubator,  where  a  temperature  of  39°  C.  was  main- 
tained. 

HISTOLOGICAL  PREPARATION 

After  their  respective  periods  of  incubation  the  eggs  were  opened, 
a  hole  an  inch  in  diameter  being  made  on  the  side  toward  the 
larger  end,  and  part  of  the  albumen  pulled  out  and  cut  off.  Careful 
observations  were  made  of  the  living  embryos,  noting  any  redden- 
ing on  the  body  of  the  blastodermic  disc ;  attention  was  paid  to  the 
heartbeat  and  to  any  other  features  that  might  appear  noteworthy. 
Then,  by  means  of  a  capillary  pipette,  several  cubic  centimeters 
of  Bouin's  fixative  were  injected  beneath  the  body,  thereby  lifting 
it  slightly  from  the  yolk.  A  generous  amount  of  the  solution  was 
also  poured  over  the  surface  of  the  embryo.     The  fixing  solution 


Crow:  Effects  of  X-Ravs  ox  Chick  Embryos  483 

was  replaced  as  needed,  for  it  was  believed  best  partially  to  fix 
the  embryo  in  situ  in  order  to  minimize  the  disturbance  in  the  dis- 
tribution of  the  blood.  In  this  way  the  chicks  were  fixed  for  a 
period  of  ten  to  twenty  minutes.  Then  the  embryo  and  much  of 
the  blastodermic  disc  were  cut  around  and  loosened  from  the  re- 
mainder of  the  egg,  and  carefully  floated  off  and  transferred  di- 
rectly to  a  dish  of  the  fixing  solution.  The  embryos  were  allowed 
to  remain  in  this  solution  from  2  to  18  hours  according  to  size; 
they  were  then  washed  for  a  few  minutes  in  distilled  water  to  re- 
move the  excess  fixative  and,  as  rapidly  as  safety  would  permit, 
carried  through  ascending  grades  of  alcohol  to  70  per  cent,  where 
they  were  held,  with  frequent  changes,  until  the  alcohol  was  no 
longer  colored.  In  the  same  strength  of  alcohol  they  were  pre- 
served. Subsequently  those  which  were  to  be  sectioned  were  passed 
through  ascending  grades  of  alcohols  to  absolute.  They  were  then 
cleared  by  gradual  addition  of  cedar  oil  to  the  alcohol,  until  the 
alcohol  was  completely  replaced  by  the  oil.  After  passing  through 
two  changes  of  oil  the  embryos  were  placed  in  a  paraffin  bath  for 
embedding.  After  embedding  they  were  cut  in  serial  sections  seven 
or  ten  micra  in  thickness,  and  subsequently  stained  in  (a)  haema- 
toxylin  and  counterstained  with  eosin,  or  (6)  iron  hsematoxylin 
and  counterstained  with  eosin  or  acid  fuchsin. 

After  mounting  in  Canada  balsam  all  of  the  sections  were  care- 
fully examined  with  a  Leitz  binocular  microscope,  using  16  X 
oculars  and  45  X  objective.  The  finer  cellular  details  of  all  tissues 
were  studied  with  16  X  oculars  and  95  X  achromatic  oil-immersion 
objective. 


484 


The  University  Science  Bulletin 


SERIES  I 

In  series  I  are  included  all  of  the  results  of  the  preliminary  ex- 
periments performed  to  determine  what  is  the  minimal  lethal  dose. 
All  the  chicks  were  exposed  to  the  same  voltage,  i.  e.,  22  a.,  90  v.,  the 
distance  and  time  being  changed. 

The  experiments  were  grouped  under  A,  B,  C  and  D,  as  follows: 

SERIES  I 


Number 

of 

individuals. 


Distance 
target. 


Length 
of  expos- 
ure. 


Age  at 
time  of 
raying. 


Age 
opened. 


Results. 


GROUP  A 


4 

4 

90v.,  22a. 
90 v.,  22a. 
90v.,  22a. 
90v.,  22a. 
90v.,  22a. 
90v.,  22a. 
90v.,  22a. 

40  cm. 
40  cm. 
40  cm. 
40  cm. 
40  cm. 
40  cm. 
40  cm. 

15  min. 
30  min. 
40  min. 
60  min. 
60  min. 
90  min. 
90  min. 

0 
0 
0 
0 
0 
0 
0 

120  hrs. 
120  hrs. 
1.50  hrs. 
150  hrs. 
240  hrs. 
288  hrs. 
384  hrs. 

Normal  development. 
Normal  development. 

4 

Normal  development. 

2   

Normal  development. 

2 

Normal  development. 

2 

Normal  development. 

2 

Normal  development. 

GROUP  B 


4 

90v.,  22a. 

18  cm. 

10  min. 

0 

30  hrs. 

Normal  development. 

4 

90v„  22a. 

18  cm. 

15  min. 

0 

30  hrs. 

Almost  normal  develo.i- 
ment — dead. 

4 

90v.,  22a. 

18  cm. 

20  min. 

0 

30  hrs. 

Developed  very  little — ■ 
dead. 

4 

90 v.,  22a. 

18  cm. 

25  min. 

0 

30  hrs. 

.■Mmost  no  development 
—dead. 

GROUP  C 


2   

90v. 
90v. 
90v. 

22a. 
22a. 
22a. 

18  cm. 
18  cm. 
18  cm. 

20  min. 
20  min. 
20  min. 

100  hrs. 
100  hrs. 
100  hrs. 

100  ^  hrs. 
102  hrs. 
117  hrs. 

Living. 

2 

2 

Living. 
Dead. 

GROUP  D 


8 

90v.,  22a. 
90v.,  22a. 
90v.,  22a. 
90 v.,  22a. 
90v.,  22a. 
90v.,  22a. 
90v.,  22a. 

18  cm. 
18  cm. 
18  cm. 
18  cm. 
18  cm. 
18  cm. 
18  cm. 

75  min. 
60  min. 
45  min. 
30  min. 
30  min. 
25  min. 
20  min. 

110  hrs. 
140  hrs. 
140  hrs. 
140  hrs. 
160  hrs. 
160  hrs. 
160  hrs. 

135  hrs. 
143  hrs. 
143  hrs. 
143  hrs. 
167  hrs. 
167  hrs. 
167  hrs. 

Dead  about  112  hours. 

3 

Dead. 

3 

3 

Dead. 

Two  dead,  one  living. 

3 

Dead. 

.3 

Two  dead,  one  living. 

.3 

Two  dead,  one  living. 

CONCLUSIONS 


Lethal  effects  do  not  appear  even  after  long  exposure  at  a  distance 
of  40  cm. 

Growth  is  not  retarded  or  accelerated  by  exposure  of  as  long  as 
90  minutes  at  a  distance  of  40  cm. 


Crow:  Effects  of  X-Rays  on  Chick  Embryos  485 

Growth  is  stopped  and  death  occurs  within  a  few  hours  after  ex- 
posure of  20  minutes  or  more  at  a  distance  of  18  cm. 

Death  of  embryo  does  not  occur  for  about  20  hours  after  exposure 
of  15  minutes  at  18  cm. 

Exposure  of  10  minutes  at  18  cm.  produced  no  effects. 

SERIES  II  TO  VH 

In  the  following  pages  each  stage  has  been  given  two  numbers, 
separated  by  a  hyphen,  by  which  it  will  be  known  in  referring  to  it 
either  in  discussion  or  in  explanation  of  the  plates.  The  first  num- 
ber gives  the  age  at  which  the  chick  was  exposed  to  the  X-rays; 
the  second  figure  gives  the  age  of  the  chick  at  which  it  was  killed 
and  studied.  Thus  28-34  means  the  chick  was  28  hours  old  when 
rayed  and  34  hours  old  when  killed. 

In  the  series  following,  all  the  embryos  which  were  exposed  to  the 
X-rays  received  the  same  dosage.  The  current  was  set  at  90  v.  and 
22  a. ;  the  time  of  exposure  was  15  minutes,  with  the  eggs  placed  18 
centimeters  from  the  target. 

The  following  tables  present  in  condensed  form  the  effects  ob- 
served in  the  chick  embryos  included  in  series  II  to  VII.  No  record 
is  given  of  the  conditions  found  in  the  control  specimens.  Without 
exception  they  all  showed  normal  conditions  in  growth,  mitosis, 
chromosome  arrangement  and  staining  qualities. 

Series  II 
In  series  II  the  control  and  rayed  embryos  have  been  carried,  side 
by  side,  through  all  the  steps  from  the  time  the  eggs  were  placed 
in  the  incubator  until  the  sections  were  mounted,  except  for  the 
short  time  when  the  latter  were  being  exposed  to  the  X-rays.  Dur- 
ing that  short  period  the  control  specimens  were  taken  from  the 
incubator  and  placed  beside  the  X-ray  apparatus,  yet  always  com- 
pletely shielded  from  the  X-rays.  The  rayed  specimens  have  shown, 
without  exception,  modifications  and  abnormalities,  which  have  in- 
creased in  extent  and  magnitude  with  the  length  of  time  after  raying. 
The  control  specimens  have  shown  normal  conditions  in  every 
tissue.  It  appears,  therefore,  that  whatever  modifications  have  oc- 
curred in  the  rayed  embryos,  have  been  caused  only  by  the  X-rays. 


-3482 


486 


The  University  Science  Bulletin 


SERIES  II 


Age  at  time 
of  raying  and 
observation, 

Ectoderm. 

Neural  tube 
and  optic 

Entoderm. 

Mesoderm. 

Blood. 

hours. 

vesicles. 

28-30 

Few  mitoses.. 

Few  mitoses; 

No  apparent 

No  visible 

Few   mitoses. 

normal  ap- 

change in 

change;  few 

pearance. 

cells;  few  mi- 
toses. 

mitoses. 

28-34 

Rare  mitoses; 

Rare  mitoses ; 

Rare  mitoses; 

Rare  mitoses; 

Some  clump- 

stain is  poor. 

cytoplasm 

stain  is  dim; 

some  chromo- 

ing of  chromo- 

more granular. 

greater  granu- 

somes 

somes    in    the 

lation. 

clumped; 
more     granu- 
lation. 

rare     mitoses ; 
some    granula- 
tion. 

2»-45 

Much    sepa- 

Debris within 

Lining  of 

Cytoplasm 

Karyosomes 

ration  of  lay- 

brain vesicle; 

foregut     bro- 

often shrunken 

prominent ; 

ers;  cells  bro- 

cells stain 

ken  and 

about   nuclei, 

stain  dim; 

ken  and 

dimly;  chromo- 

frayed;    some 

granular,  large 

granules  large; 

frayed;    large 

somes     indis- 

debris  within 

vacuoles,   ap- 

many cells 

irregular  gran- 

tinct; some 

cavity;  few 

parent    disso- 

broken;     very 

ules  and  clear 

granulation 

large  granules 

lution  of 

rare      mitoses; 

spaces;    chro- 

and   vacuola- 

in  cytoplasm; 

chromosomes 

all     abnormal; 

mosomes     in- 

tion. 

chromosomes 

in  dividing 

chromosomes 

distinct. 

indistinct  and 
apparently  dis- 
solved in  cyto- 
plasm in  early 
mitoses;  some 
nucleishrunken. 

cells. 

partly  dis- 
solved ;    two 
cells   with   tri- 
polar  spindles. 

CONCLUSION 

Raying  effects  are  most  marked  in  nerve  tissue. 

Cytoplasm  becomes  more  granular  and  nuclei  less  distinct  after 
raying.  Mitoses  are  not  stopped  suddenly,  but,  if  in  division,  a  cell 
goes  on  and  completes  it.  Probably  no  cell  begins  and  completes 
division  after  raying. 


Crow:  Effects  of  X-Rays  on  Chick  Embryos  487 

Series  III 
In  series  III  the  embryos  which  have  been  used  as  controls  have 
been  kept  under  the  same  conditions  as  the  rayed  specimens,  save 
only  that  they  were  not  exposed  to  the  X-rays.  Even  during  the 
short  time  when  the  rayed  specimens  were  being  exposed  to  the 
X-ray,  the  controls  were  beside  the  X-ray  apparatus,  but  fully 
shielded  from  the  rays.  Through  fixation,  dehydration,  clearing, 
embedding,  staining,  mounting,  etc.,  the  control  and  the  rayed  em- 
bryos have  been  carried  under  as  near  identical  conditions  as  it  is 
possible  to  secure.  If  faulty  technique  had  been  the  cause  of  modifi- 
cations and  degeneration  in  one  set  of  embiyos  it  should  have  shown 
similar  changes  in  the  other  group.  In  all  of  the  irradiated  embryos 
these  modifications,  as  given  in  the  preceding  pages,  have  appeared, 
while  in  not  a  single  control  specimen  have  any  than  normal  con- 
ditions been  found.  In  view  of  all  the  evidence  only  one  conclusion 
could  be  made,  namely:  the  X-rays  have  been  the  cause  of  the  modi- 
fications occurring  in  the  irradiated  embryos. 

CONCLUSION 

The  effects  of  the  X-rays  are  shown  by  the  nervous  tissue  in  which 
there  occur:  separation  of  limiting  membranes,  breaking  of  neural 
wall  in  places,  and  partial  filling  of  canal  with  disorganized  cellular 
debris. 

The  number  of  cells  in  mitoses  decreases  rapidly,  none  occurring 
after  about  six  hours.  A  few  show  chromosomes  somewhat  clumped. 
The  cytoplasm  and  nuclei  show  granules  which  appear  to  increase  in 
size  with  degeneration  of  the  cells. 

Fraying  of  epithelium  occurs  in  many  places. 

Debris  accumulates  within  cavities  or  tubes  of  the  body. 

Tissues  stain  less  well. 


488 


The  University  Science  Bulletin 


1 

>> 

03 
g 

Few  cells  di- 
viding; slight 
clumping  in  few 
cells. 

Staining     not 
good  color;  large 
granules  in  cyto- 
plasm; very  few 
mitoses. 

Cell  walls 
poorly    defined, 
some  broken;  no 
cells  dividing. 

o 
o 

5 

Cytoplasm  con- 
tains large  gran- 
ules;    cell     wall 
broken  in  many 
cells;  some   nu- 
clei fragmented; 
few  mitoses;   in 
some  cells  chro- 
mosomes are 
clumped. 

Karyosomes 
prominent;  stain 
not  sharp;   rare 
mitoses;      some 
with    condensed 
chromosomes;  a 
few  cells  broken. 

Much   granu- 
lation; poor  dif- 
ferentiation; few 
broken  cells; 
very     rare     mi- 
toses; some  chro- 
mosomes    frag- 
mented. 

E 

o 
a) 

Mitoses      are 
infrequent;  kary- 
osomes     promi- 
nent; large  gran- 
ules. 

Very  few  cells 
in  division 
stages;  some 
clumping  of 
chromosomes; 
stain  is  not 
sharp. 

Some  shrink- 
ing of  cytoplasm; 
mesothelium 
separated  and 
broken  in  some 
places;    absence 
of  mitoses. 

£ 
o 

Few    mitoses 
with  some 
clumping  of 
chromosomes; 
some      fibrinoid 
material  in 
pharynx. 

Infrequent 
mitoses;  some 
chromosome 
condensation; 
debris   in   phar- 
ynx; stain  poor. 

Much    debris 
in  cavity,  some 
fraying  of  inner 
layer;    vacuoles 
prominent  in 
some    cells;    no 
mitoses;  gray 
stain. 

0) 

> 

>> 
t~, 
o 

3 
< 

Few    mitoses 
with    some    con- 
densation of 
chromosomes; 
stain    not    defi- 
nite;  some   ma- 
terial in  the 
vesicle. 

Rare  mitoses; 
some      chromo- 
some condensa- 
tion; stain  gray; 
debris     in     the 
cavity. 

Mucn  material 
in   vesicle;    wall 
broken;  stain 
poor  with  little 
differentiation; 
few  cells  divid- 
ing  show   chro- 
mosomes  poorly 
defined. 

i 

[o 

> 

.2 
a 
O 

Mitoses     rare 
with  some  con- 
densation of 
chromosomes; 
some   fibrin-like 
material   within 
vesicle. 

Mitoses  rare; 
some  clumping 
of  chromosomes; 
debris  in  vesicle; 
stain,     gray    in 
color. 

Much    debris 
in   vesicle;  poor 
differentiation 
of    stain ;    gran- 
ules large;  very 
rare  mitoses; 
compact  chromo- 
some     arrange- 
ment. 

3 
3 

Rare   mitoses 
not  clear-cut; 
granulation  and 
vacuolation  dis« 
organized  mate- 
rial     in      brain 
cavity. 

Dividing  cells 
swollen,    a    few 
bursted  with 
contents  passing 
into  cavity; 
other  debris 
within  brain 
cavity. 

Canal    nearly 
filled    witli    dis- 
organized cellu- 
lar mass;  limit- 
ing    membrane 
broken  and 
frayed;  stain 
very  poor;  rare 
mitoses. 

E 

Very  few  cells 
dividing;     chro- 
mosomes abnor- 
mal;   cytoplasm 
granular  and 
vacuolated. 

No  increase  in 
size;   separation 
in  many  places; 
granulation;  few 
mitoses  with 
condensed  chro- 
mosomes. 

Much  separa- 
tion; large  gran- 
ules and  vacu- 
oles;  walls  dim; 
rare  mitoses. 

Age  at 

time  of 

raying  and 

observation, 

hours. 

in 
CO 

1 

i 

•>* 

tf3 
K 

00 

Crow:  Effects  of  X-R.\ys  on  Chick  Embryos 


489 


E 
.2 

1 
ci 

"o 

O 

ity     for     stain; 
granules     large; 
no  mitoses. 

a 

ID 

■a 

3 

eration  of  cells; 
all  appear  faded 
gray;     no     cells 
dividing. 

o 

s 

Large  cyto- 
plasmic gran- 
ules; walls  bro- 
ken; nuclear  ma- 
terial condensed 
upon  wall;  cells 
shrunken  and 
fissured;  no  mi- 
toses; poor  stain. 

Much  frag- 
mentation    and 
breaking  of  walls; 
karyosomes 
fragmented;    no 
cells      dividing; 
poor  staining; 
differentiation. 

s 

(U 

o 

Cytoplasm 
shrunken  and 
granular;    myo- 
cuele  filled  with 
material;     walls 
leaking;  no  cells 
dividing. 

Much  degen- 
eration; walls 
leaking  and 
forming     debris 
in  cavities;  cyto- 
plasm granular; 
gray    stain;    no 
cells  in  division. 

E 

1 
a 

Pharynx  cells 
broken  and 
frayed;  debris  in 
cavity;  large  ir- 
regular granules; 
no  cells  dividing 
cells;  stain 
poorly. 

Much    debris 
in    gut    cavity; 
wall  broken  and 
frayed  in  many 
places;  much  de- 
generation;    no 
mitoses. 

o 

3 

Cellular  debris 
in  vesicle;  bro- 
ken walls;  large 
granules  and  vac- 
uoles; karyo- 
somes irregular; 
no  cells  dividing. 

Limiting  mem- 
brane frayed 
with  cells  form- 
ing    debris     in 
vesicle;  large 
vacuoles  and 
granules;    some 
karyosomes   are 
fragmented;   no 
cells  dividing. 

> 
.2 

0. 

O 

Vesicle    filled 
with    cells    and 
granules;  exuda- 
tions from  walls; 
no  mitoses. 

Development 
inhibited;  small 
vesicle  filled 
with  debris; 
walls  broken 
and   frayed;   no 
cells      dividing; 
stain  poor. 

6 

3 

3 
ai 

Canal  almost 
filled   with  bro- 
ken cells;  mem- 
brane broken 
and   frayed;    no 
cells      dividing; 
large      granules 
resemble  broken 
chromosomes. 

Much    debris 
in  canal;  wall  of 
brain  broken;  no 
cells      dividing; 
stain   is    a    dull 
gray;  much  dis- 
organization. 

E 

a) 

o 

K 

Much  separa- 
tion and  degen- 
eration; stain 
dim;  some   cells 
broken;  gran- 
ules large;  karyo- 
somes frag- 
mented; no  cells 
dividing. 

Layers    sepa- 
rated and  frayed; 
almost  no  stain; 
cells  almost 
structureless 
with    few    large 
granules;  no 
cells  in  division. 

Age  at 

time  of 

raying  and 

observation, 

hours. 

c 

01 

■* 

490  The  University  Science  Bulletin 

Series  IV 

The  irradiated  embryos  of  this  series  have  had  controls  checked 
against  them  at  every  point  from  beginning  of  incubation  to  the 
mounting  finally  of  the  sections.  Only  in  the  item  of  irradiation 
has  there  been  a  difference  in  treatment.  Any  changes  occurring 
in  specimens  which  have  been  irradiated  and  not  appearing  in 
the  controls  could  be  interpreted  as  having  been  caused  by  the 
X-rays.  In  the  report  on  the  embryos  constituting  series  IV,  given 
in  the  following  pages,  there  is  given  a  recital  of  the  changes  and 
modifications  which  were  found  in  the  irradiated  specimens.  With 
very  great  similarity  in  the  extent  of  the  effects,  the  type  of  modi- 
fication and  the  time  of  their  appearance,  these  changes  have  ap- 
peared in  all  the  individuals  which  were  exposed  to  X-rays.  Not 
one  of  the  controls  showed  any  of  the  marked  modifications  so 
characteristic  of  the  irradiated  specimens,  but  all  were  normal  in 
every  way.  The  X-rays  have  been,  without  doubt,  the  cause  of 
all  of  the  destructive  changes  which  have  appeared  in  the  embryos 
irradiated. 

conclusion 

Exposure  to  X-rays  has  caused  the  accumulation  of  much  dis- 
organized material  within  the  cavity  of  the  brain  and  cord,  the 
optic  vesicles,  pharynx  and  coelom. 

There  is  a  rapid  decrease  in  number  of  cells  dividing,  only  a  few 
showing  after  six  hours.  Slight  condensation  of  chromosomes  into 
a  clump  shows  in  a  few  cells. 

The  granules  in  the  nuclei  and  cytoplasm  have  become  more 
prominent. 

The  cells  do  not  retain  the  stain  well. 


Crow:  Effects  of  X-Rays  ox  Chick  Embryos 


491 


Pi 

as 


.s 

greater  granula- 
tion; no  cells  di- 
viding. 

Large      gran- 
ules;    rare     mi- 
toses; individual 
chromosomes 
cannot  be  distin- 
guisned  because 
of  compact 
shape. 

o 

S 

Pronounced 
granulation;  al- 
most no  mitoses; 
chromosomes 
condensed;  a 
few    nuclei    are 
fragmented. 

Prominent 
granules  in  cyto- 
plasm; nuclei 
shrunken,  wrin- 
kled    and     fis- 
sured;  rare   mi- 
toses; stain  less 
sharp. 

£ 

0) 

■a 
o 

o 

Slight  leaking 
of      mesothelial 
membrane;  very 
few  cells  divid- 
ing. 

A  little  fray- 
ing of  walls,  and 
some    debris    in 
the  coelom;  few 
cells  show   nor- 
mal chromosome 
arrangement. 

E 

1 

c 

Normal      ap- 
pearance in 
most  cells;  slight 
condensation  of 
chromosomes  in 
the     few     cells 
dividing. 

Pharyngeal 
wall   broken    in 
few  places;  cells 
passing  into  cav- 
ity;    cytoplasm 
granular;  few 
cells  dividing. 

>2 

a) 
> 

p 

3 

«j3 

Cell  walls  in- 
distinct; no  mi- 
totic figures. 

Cells  indis- 
tinct; a  very  few 
cells  in  division; 
some  with 
clumped     chro- 
mosomes;   vesi- 
cles   filled    with 
fibrinoid    mate- 
rial. 

> 
.2 

0, 

O 

Some  material 
between    coats; 
not   many   cells 
dividing;  karyo- 
somes   resemble 
condensed  meta- 
phases. 

Distinct  separa- 
tion    of     coats 
with  space  filled 
with     disorgan- 
ized    cells     and 
fibrin-like     sub- 
stance; no  cells 
dividing;     cyto- 
plasm oozing 
from  inner  end 
of  lens  cells. 

6 

XI 
3 

"S 

S 

Little  change; 
limiting      mem- 
brane broken  in 
few  places;  little 
fibrinoid    mate- 
rial   in    cavity, 
mitoses  less  than 
normal  number. 

Inner  limiting 
membrane   bro- 
ken   in    places, 
and  cells  passed 
into  brain  cavity; 
mitoses  few; 
chromosomes  in 
few  slightly 
clumped;    stain 
not  well  re- 
tained. 

E 
o 

Some  separa- 
tion   of    layers; 
increase  in  vac- 
uoles and  gran- 
ules; mitoses 
normal  in  num- 
ber and  arrange- 
ment. 

More  separa- 
tion   of    layers; 
stain     not     re- 
tained well ;  vac- 
uoles   common, 
few  cells  divid- 
ing, chromosomes 
not  definite. 

Age  at 

time  of 

raying  and 

observation, 

hours. 

1     : 

• 

to 
o 

ID 

492 


The  University  Science  Bulletin 


o 


i 

(O 

>> 

>. 
a 

Some  leaking 
of  walls;  debris 
inducts,    indefi- 
nite  cell   walls; 
granules  promi- 
nent  in    nuclei, 
and  irregular  in 
shape;     no    mi- 
toses; loss  of  af- 
finity for  stain. 

o 

s 

ib    1^' °  .  2  s  ..  t  g  J  . 

S.Sccg^Sg£2g.SSJ 

s 

1 

(V 

.2       ?  J  ..  i^-3  ^ 
1  g  g|,|  8  S  S 

^  1 1  ^  i  1 2- 1 

.Q  i:  .2  .S  B  .5  2 

o 
o 

"S 

3    i    O    b  ~          -b' 

.-        -^    q    &    c  .5 
1 1  1   c  "S  .^   "   § 

^  1  -S   g  1  1   §   o 

> 

o 

a;    S    03    5     .     C  .5 

■C  "u  .s   c   £  c   M 

i 

O 

3 
3 

■g  c  ==  c  ^  >>  •-  -2  S  g  -«  -3  o  >. 

3    O  •-    =    g  9  .2  •£    §    "^   iJ    E    S  -TS 

1 

■o  >>      .2  .= 

c  S  >?  S  - 
2    £    g  "3  "3 

Age  at 

time  of 

raying  and 

observation, 

hoirs. 

1 

i      § 

Crow:  Effects  of  X-Rays  ox  Chick  Embryos  493 

Series  V 

In  all  the  steps  of  preparation  of  the  embryos  for  microscopic 
study,  control  and  irradiated  specimens  have  been  subjected  to  as 
nearly  the  same  conditions  as  possible,  except  in  the  matter  of 
irradiation.  Under  these  conditions  all  the  control  specimens  have 
shown  normal  development  and  cell  structure,  while  the  irradiated 
embryos  have  in  every  case  shown  modifications  and  degeneration. 
In  all  control  specimens  mitoses,  with  clearly  defined  chromosomes, 
have  been  found  abundant,  but  in  irradiated  embryos  the  number 
of  mitoses  decreases  rapidly  till  after  six  hours  none  may  occur. 
In  the  controls  the  chromosomes  occur  regular  in  arrangement  and 
definite  in  outline,  but  in  the  irradiated  specimens  clumping  of 
chromosomes  occurs  frequently,  and  in  a  few  cells  the  chromosomes 
are  so  disorganized  as  to  be  little  more  than  a  granular  mass  of 
irregular  shape.  In  the  control  embryos  the  cytoplasm  and  nuclei 
rarely  contain  prominent  granules;  but  in  a  few  hours  after  ex- 
posure to  X-rays  embryos  show  very  large  granules  in  many  cells. 
The  separation  of  epithelial  membranes  from  underlying  mesoderm 
or  the  breaking  of  these  membranes  does  not  occur  in  control  speci- 
mens. Cellular  exudations,  which  occur  so  prominently  in  the 
later  stages  projecting  into  the  cavities  of  the  brain,  cord,  gut  and 
urinary  ducts,  and  into  the  coelom  and  the  vesicles  of  the  sense 
organs,  are  not  found  at  any  place  in  the  control  specimens. 
Hemorrhages  occur  in  embryos  a  few  hours  after  irradiation.  These 
become  greater  with  the  passage  of  time,  until  in  the  75-94  speci- 
mens all  except  one  individual,  which  was  little  affected  by  the 
X-rays,  showed  widespread  areas  involved.  In  the  controls  not 
one  showed  any  hemorrhages  whatever.  The  dark-blue  color  and 
clear  differentiation  characteristic  of  tissues  stained  with  hsema- 
toxylin  were  found  in  every  control  specimen,  while  in  those  ir- 
radiated the  characteristic  color,  after  a  few  hours,  did  not  appear; 
in  fact,  the  affinity  for  all  stains  employed  became  less  with  the 
passage  of  time.  Since  great  changes  have  occurred  in  the  struc- 
ture of  the  cells,  as  shown  by  the  granules,  vacuoles  and  fragmenta- 
tion, it  seems  reasonable  to  suppose  that  the  chemical  constitution 
of  the  cells  may  have  been  altered,  and,  if  so,  then  the  change  in 
staining  reaction  may  be  this  expected  result. 

Inasmuch  as  all  the  irradiated  specimens  have  shown  destruc- 
tive modifications  which  have  become  greater  with  the  passage  of 
time  after  irradiation,  and  since  in  not  one  of  the  control  speci- 


494  The  University  Science  Bulletin 

mens  did  any  of  the  modifications  occur,  proves,  without  reason- 
able doubt,  that  the  X-rays  have  caused  the  destructive  changes. 

CONCLUSION 

Few  mitoses  are  found  four  or  more  hours  after  irradiation. 

There  is  much  material  in  the  neural  canal,  in  the  vesicles  of  the 
eye,  ccelom,  and  lumina  of  digestive  and  urinary  systems.  This 
material  varies  in  amount  with  the  length  of  time  after  raying. 

The  effects  seem  to  be  general  rather  than  local  and  do  not  make 
their  appearance  for  some  time  after  irradiation. 

All  the  effects  of  raying  appear  to  be  destructive,  i.  e.,  no  accel- 
eration occurs,  and,  given  enough  time,  all  tissues  show  degenera- 
tion. This  consists  of  separation  of  epithelial  membranes,  appear- 
ance of  granules,  and  vacuoles  in  the  cytoplasm,  loss  of  affinity  for 
stains,  so  that  the  nuclei  and  cytoplasm  are  poorly  differentiated, 
and  in  the  cells  dividing  often  a  condensation  of  chromosomes 
occurs. 

There  appears  to  be  a  difference  in  degree  of  susceptibility,  one 
specimen  of  75-79  showing  a  few  mitoses,  others  none. 

In  75-94  stage  one  chick,  which  was  much  older  than  the  others, 
showed  little  effects  of  raying,  having  recovered  enough  that  many 
cells  were  in  division. 


Crow:  Effects  of  X-Rays  on  Chick  Embryos 


495 


E 

o 

1 

s 
c 

5 

From      many 
cells,  cytoplasm 
has  escaped  into 
lumen  of  duct  or 
tubule;  a  small 
number  of  cells 
dividing      these 
often  bulging 
into  lumen. 

c 

75-78;  a  few  tu- 
bules broken; 
one   chick  only 
showed    mitotic 
stages;  slight 
clumping  of 
chromosomes 
appeared. 

i 

o 

5 

Many  small 
cells;  large  cells 
often  have  frag- 
mented   nuclei; 
large      granules 
abundant;  few 
pells  dividing. 

Hemorrhages 
in  many  places; 
granules      large 
and  irregular, 
often  condensed 
upon    cell   wall ; 
many  cells  bro- 
ken and   others 
with  fragmented 
nuclei;  rare  cell 
divisions;  a  few 
appear  as  chro- 
mosomes melted 
together. 

E 

O 

1 

Not  good  dif- 
ferentiation; 
karyosomes 
fragmented    in 
some  cells;  ami- 
tosis  in  few  cells; 
not  many  cells 
dividing. 

Much  like 

75-78     material 
in  ccelom;  clouds 
of    cytopla,»m 
passing  from 
cells;     cells     in 
head  mesen- 
chyme shrunken. 

s 

o 

2 
c 

Little     effect 
on  cell  appear- 
ance; clouds  of 
cytoplasm    ooz- 
ing from  cells  of 
hind  gut;  a  few 
cells  in  division 
apparently  nor- 
mal. 

Gut  in  most 
places  filled  with 
material;      wall 
broken  in  places; 
liver  antl  blood 
cells  mixed;  few 
cells  in  division, 
none  normal; 
some  fragment- 
ing of  chromo- 
somes. 

o 
> 

>> 

o 

•5 

3 

< 

Some    break- 
ing of  inner  mem- 
brane where 
cells   are   added 
to  debris  in  ves- 
icle;   few    cells 
dividing. 

Inner      mem- 
brane broken; 
cytoplasm     and 
nuclei  oozed 
into  vesicle; 
stain  poorly  dif- 
ferentiated; 
large     granules; 
rare  mitoses; 
some    clumping 
of  chromosomes. 

Optic  vesicles. 

Small  space 
between      coats 
partly  filled  with 
debris;  lens  also 
has  material  in 
vesicle;  some  in- 
crease in  granu- 
lation; few  cells 
dividing. 

Coats  widely 
separated      and 
space  filled  with 
disorganized 
material;  cell 
walls  indistinct; 
lens  vesicle  filled 
with  debris; 
long  cells 
shrunken;  chro- 
mosomes in  the 
rare  mitoses 
clumped. 

6 

Lack  of  sharp- 
ness    in     cells; 
small  amount  of 
material  in  cav- 
ity;    clouds     of 
cytoplaem  pro- 
ject from    cells 
into  canal;   few 
mitoses  with  lit- 
tle sharpness. 

Spaces  in 
brain  wall  from 
which  cells  have 
passed  into 
canal;  cell 
shapes  indefi- 
nite;   stain    not 
sharp;  few  cells 
dividing  in  one 
specimen,    none 
in  others. 

Ectoderm. 

A  little  sepa- 
ration of  layers; 
some  fragmenta- 
tion   of    a    few 
karyosomes; 
stain  not  sharp; 
few  mitoses; 
some  chromo- 
some groups; 
compact  spin- 
dles invisible. 

Much  like 
75-78;  few  large 
granules;  no 
cells  dividing; 
one  specimen,  a 
few  mitoses. 

Age  at 

time  of 

raying  and 

observation, 

hours. 

: 
_: 

00 

496 


The  University  Science  Bulletin 


E 
1 

c3 
3 

Tubules    bro- 
ken in  many 
places,    and    lu- 
mina  filled  with 
blood  and  other 
cells;     stain     is 
poor;  no  nutosea 
except     in     one 
chick,    and    but 
few  there. 

1 
O 

s 

Great  hemor- 
rhages; stain  not 
uniform;      dim; 
many  cells  irreg- 
ular; some 
shrunken,  others 
with  fragmented 
or  no  nuclei;  ap- 
pear as  if  nuclei 
were     dissolved 
in  cytoplasm; 
one  specimen 
had     few     cells 
dividing,  all 
others  none; 
chromosomes 
showed  con- 
densed arrange- 
ment. 

s 

O 

1 

Stain  dim ; 
cells   in   mesen- 
chyme shrunken 
and  irregular; 
mesothelum 
often  broken 
and  flayed;  de- 
bris   in    coelom, 
one  chick 
showed    a    few 
cells      dividing; 
all     small     and 
compact,  others 
none. 

s 

1 

3 

Pharynx  walls 
broken;  cells  in 
cavity  debris; 
poor  differentia- 
tion;   nuclei    of 
numy  cells 
shrunken  and 
angular;  large 
granules;  cells  of 
liver  and  blood 
mingling;  no  mi- 
toses, except  in 
one      specimen, 
where  some  cells 
appear  to  be  di- 
viding by   ami- 
tosis. 

> 

o 

■5 

3 

Wall  folded 
and  broken;  ves- 
icle   filled;    nu- 
clei irregular; 
karyosomes 
much  faded;  no 
cells  dividing. 

_a3 

a> 
> 

a 

Great  hemor- 
rhages     around 
eye  causing  sep- 
aration of  coats 
and     filling     of 
vesicles;  wall 
broken  fre- 
quently; large 
granules 
grouped  around 
cell  wall;  shrink- 
age in  lens  cells; 
no  cells  dividing 
except     in     one 
specimen. 

a; 

.a 

3 
3 

z 

Great  hemor- 
rhages in  area 
surrounding 
brain  have  bro- 
ken    wall,     and 
material  in  cav- 
ity contains 
blood  cells;  wall 
is  folded  and  sep- 
arated in  places; 
spinal  canal 
filled;    no    cells 
dividing;      very 
dim  gray  stain; 
one    chick    had 
more     the     ap- 
pearance of 
those    of    75-78 
stage,     it     was 
10-15  hours 
older. 

1 

Much  separa- 
tion of  layers;  a 
few  appear 
swollen   as   if   a 
prophase     had 
been  aborted:  in 
such,  no  karyo- 
somes     appear; 
large      granules 
and  vacuoles;  no 
mitoses;  one 
specimen  sev- 
eral cells  divid- 
ing. 

Age  at 

time  of 

raying  and 

observation, 

hours. 

1      : 

Oi 

Crow:  Effects  of  X-Rays  ox  Chick  Embryos  497 

Series  VI 

In  this  as  well  as  in  all  the  other  series,  irradiated  specimens  and 
controls  have,  in  their  preparation  for  microscopic  study,  been 
carried  as  near  side  by  side  as  possible.  In  the  long  staining  process 
not  only  have  both  received  exactly  the  same  treatments,  but  often 
a  slide  on  which  are  sections  of  an  irradiated  embryo,  and  one  on 
which  are  sections  of  a  control,  have  been  carried  through  the  dif- 
ferent staining,  washing  and  dehydrating  jars,  back  to  back.  In 
this  way,  if  effects  due  to  faulty  technique  appear  in  one  they  would 
appear  also  in  the  other.  In  the  irradiated  specimens,  except  those 
in  the  96-20-minutes  stage,  which  were  killed  before  effects  became 
visible,  there  have  occurred  such  modifications  as  have  been  reported 
at  length  in  the  preceding  pages.  Not  one  irradiated  specimen 
killed  two  hours  or  more  after  irradiation  was  normal,  while  all  the 
control  specimens  were  normal  in  every  respect.  It  is  very  evident 
that  the  many  degenerative  changes  that  occur  in  irradiated  embryos 
are  due  wholly  to  the  action  of  X-rays. 

The  effects  of  X-rays  on  chick  embryos  of  different  ages,  as  re- 
ported in  this  paper,  have  always  appeared  as  degenerative  modifi- 
cations. These  have  grown  in  magnitude  with  the  passage  of  time 
after  irradiation.  In  the  96-20-minutes  stage  an  attempt  has  been 
made  to  discover  how  soon  effects  of  irradiation  appear.  In  this 
stage  the  embryos  were  killed  twenty  minutes  after  exposure  to  the 
X-rays.  These  showed  such  minor  changes  that  it  could  not  be 
determined  whether  they  were  caused  by  the  X-rays  or  were  merely 
individual  variations.  Two  hours  after  irradiation  modifications 
became  apparent  and  grew  greater  each  hour. 

It  appears  that  cells  are  affected  at  the  time  of  irradiation,  but 
there  elapses  a  period  in  which  no  effects  are  visible,  but  which  be- 
come manifest  at  the  end  of  that  time.  There  is  doubtless  produced 
within  the  cellular  substance  by  irradiation  chemical  changes  which 
cause  serious  disturbance  of  the  metabolic  balance,  the  result  of 
which  may  be  seen  in  the  formation  of  granules,  vacuoles  and 
chromosome  clumps  and  fragmentation  of  nuclei  and  the  breaking 
up  of  whole  cells. 

CONCLUSION 

The  effects  of  raying  are  general  rather  than  local,  and  degenera- 
tive changes  occur  in  all  tissue,  but  they  become  visible  first  in  the 
nerve  tissue  and  sense  organs. 

Separation  of  retinal  coats,  limiting  membranes  of  the  brain  and 


498  The  University  Science  Bulletin 

cord,  the  mesothelium  and  epithelium,  occurs.  All  of  these  occur 
over  greater  areas  with  the  passage  of  time  after  irradiation. 

Cellular  differentiation  becomes  less  with  time,  the  cytoplasm 
and  nuclei  becoming  more  granular  and  vacuolated. 

Mitoses  have  not  been  affected  in  twenty  minutes,  but  are  reduced 
in  number  in  two  hours,  and  in  four  hours  no  new  figures  have 
occurred.  A  few  of  the  figures  have  their  chromosomes  condensed 
into  a  clump. 

Degenerative  material  accumulates  in  the  neural  canal,  vesicles 
of  the  sense  organs,  alimentary  canal,  urinary  lumina  and  coelom. 

Great  hemorrhages  occur  after  a  few  hours,  probably  causing 
much  of  the  debris  within  body  cavities  and  distortion  of  walls  and 
membranes. 


Crow:  Effects  of  X-Rays  on  Chick  Embryos 


499 


CQ 

H-l 


1 

a 

Little    fibrin- 
like material  in 
ducts;   few  mi- 
toses. 

Cytoplasm 
more     granular 
and  dimly 
stained;  tubules 
have    filled    lu- 
mina;   few    mi- 
toses. 

o 

s 

Fewer  cells  in 
division  than 
control. 

Veins  and 
capillories 
gorged;     small- 
sized  cells;  stain 
poorly;  many 
large  granules 
in    other    cells; 
few  mitoses; 
metaphase 
plates  condensed. 

1 

4) 

No  effect  ap- 
parent; number 
of  mitoses 
smaller  than 
control. 

Some  loss  of 
affinity  for  stain; 
mesothelium 
lifted   and   bro- 
ken; material  in 
coelom;  cyto- 
plasm granular; 
nuclei  of  many 
cells    shrunken; 
few  mitoses. 

a 

(1) 

13 

2. 
a 
H 

Scant  amount 
of  fibrin-like 
si'bstance  in 
lumina   of   lung 
buds  and  oesoph- 
agus and  cells  in 
pharynx;  mitoses 
about  one-half 
normal  number. 

Debris  in  cav- 
ity in  most 
places;    number 
of    mitoses    de- 
creased; some 
chromosome 
plates  compact. 

Auditory  vesicles. 

Slight  separa- 
tion    of     inner 
limiting     mem- 
brane. 

Stain  not 
clear;     chromo- 
somes not   well 
separated  or 
distinct    in    the 
small  number  of 
mitoses. 

J 
> 

a 
O 

No  changes 
in  structure; 
small  number  of 
mitosos. 

Slight  separa- 
tion   of    coats; 
from    lens    cells 
cytoplasm  oozes 
into  vesicle ;  slight 
condensation  of 
chromosomes  in 
the  few  mitoses. 

Neural  tube. 

Apparently 
normal;  number 
of  mitoses,  one- 
half  control; 
may  be  normal 
number. 

Outer     limit- 
ing    membrane 
separated  in  few 
places ;  some  de- 
bris within 
brain;    few    mi- 
toses and  slight 
clumping  of 
chromosomes. 

Ectoderm. 

No  gross 
ohanges;     num- 
ber   of    mitoses 
about     one-half 
the    number    in 
control  speci- 
men. 

Separation  of 
layers  in  few 
places;  some 
cells    shrunken; 
few  mitoses. 

Age  at 

time  of 

raying  and 

observation, 

hours. 

1     • 
o 

(N 

500 


The  University  Science  Bulletin 


o 


Urinary  system. 

Much  ^f  ducts 
and  tubules 
show  cellular  de- 
bris in   lumina; 
granules  large 
and     gray     col- 
ored; rare  cell  di- 
visions; chro- 
mosomes in 
dense  masses. 

Much   degen- 
eration and  de- 
struction; lu- 
mina filled; 
dull,  gray  stain, 
allantois  affected 
little;    no    cells 
dividing. 

Blood. 

Hemorrhages 
common;  stain 
gray;     granules 
common;  many 
karyosomes 
fragmented;  in- 
dividual chro- 
mosomes cannot 
be    followed    in 
the  few  cells  di- 
viding. 

Great  destruc- 
tion and  hemor- 
rhages; cells 
broken  or  frag- 
mented, and  cy- 
toplasm   oozing 
out ;    very    dim 
stain;     granules 
condensed  on 
cell  walls. 

Mesoderm. 

Cells  indis- 
tinct; stain, 
dark  gray ;  mem- 
branes often 
broken,  with 
cells    contribut- 
ing to  debris  in 
coelom;  very  few 
cells  dividing. 

Cytoplasm 
shrunken    upon 
nuclei;  stain, 
dull  gray ;  meso- 
thelium  broken, 
and    much    ma- 
terial in  ocelom; 
no  cells  in  divi- 
sion states. 

Entoderm. 

Walls  broken 
in    few    places; 
much  loss  of 
cells  into  lu- 
mina;  liver   tu- 
bules broken 
and  mixed  with 
blood;   few   mi- 
totic cells  have 
heavy     spindles 
and  clumped 
chromosomes. 

Walls    folded 
and  fissured 
where  cells  have 
passed   into   lu- 
mina; blood  and 
liver   cells  min- 
gle; granules 
large;  nuclei 
fragmented; 
some    areas  al- 
most without 
color;  no  mitoses. 

Auditory  vesicles. 

Thin  wall 
folded  and  bro- 
ken;   debris    in 
vesicle;  stain 
dim,    very    few 
mitoses,  chro- 
mosomes indefi- 
nite. 

Some  strip- 
ping of  wall; 
cellular      debris 
in  vesicle;  large 
granules  con- 
densed upon  cell 
wall;  stain,  dark 
gray;  no  mitotic 
figures;     a     few 
nuclei  and  kary- 
osomes frag- 
mented. 

Optic  vesicles. 

Greater  sepa- 
ration   of    coats 
than  96-98; 
blood  cells  have 
broken  coats 
and  partly  filled 
space    between; 
stain  not  sharp; 
lens  cavity 
filled;    rare    mi- 
toses. 

Growth 
stopped  and 
much  degenera- 
tion; walls  bro- 
ken and  folded; 
much   debris  in 
vesicles ;      more 
granules;  nuclei 
indistinct  and 
some  shrunken; 
no  mitoses. 

Neural  tube. 

Distinct    mo- 
rain     of     disor- 
ganized material 
within  brain 
cavity;  cyto- 
plasm slightly 
granular,  and 
staining  dimly ; 
rare     cell    divi- 
sions; some 
sliow  dense  mass 
instead  of  chro- 
mosomes. 

Much   degen- 
eration; brain 
cavity  almost 
filled    with    de- 
bris; brain  wall 
broken  fre- 
quently; very 
poor,  dim  stain; 
many     granules 
in  nuclei;  karyo- 
somes frag- 
mented in  some ; 
no  mitosPS. 

Ectoderm. 

Greater  sepa- 
ration of  layers; 
shrinkage  of  cy- 
toplasm;   many 
granules  and 
vacuoles;  rare 
mitoses;  chro- 
mosomes bro- 
ken. 

Much  separa- 
tion    of     layers 
with  material  in 
space;  poor  dif- 
ferentiation ; 
karyosomes 
often  frag- 
mented; no  mi- 
toses. 

Age  at 

time  of 

raying  and 

observation, 

hours. 

96-100 

'XI 

Crow:  Effects  of  X-Rays  on  Chick  Embryos  501 

Series  VII 

The  control  and  the  irradiated  embryos,  which  have  been  studied 
in  this  series,  have  been  handled  with  great  care  and  kept  under  as 
near  identical  conditions  as  possible  from  the  time  the  eggs  were 
placed  in  the  incubator  until  the  sections  were  stained  and  mounted, 
differing  only  in  one  particular — irradiation.  Those  embryos  which 
have  been  exposed  to  X-rays  have  shown  destructive  changes  in 
every  tissue,  all  of  which  have  been  described  in  the  preceding  pages. 
Very  similar  changes  have  been  found  in  all  the  irradiated  in- 
dividuals of  the  other  series,  the  degree  of  destruction  in  each  series 
corresponding  to  that  in  other  series  at  a  given  time  after  irradiation. 

In  contrast  with  the  irradiated  specimens,  which  have  shown  the 
destructive  effects  already  described,  the  controls  have,  without 
exception,  shown  normal  development  and  normal  cellular  structure. 

With  degenerative  changes  occurring  in  the  irradiated  embryos, 
and  not  one  of  the  controls  showing  any  of  these  modifications,  only 
one  conclusion  can  be  made,  namely:  the  changes  in  the  irradiated 
specimens  are  the  effects  of  exposure  to  X-rays. 

CONCLUSION 

Degenerative  effects  show  mostly  in  the  neural  tissue  and  sense 
organs.  All  other  tissues  are  affected  somewhat  later  or  less 
promptly. 

Great  amounts  of  material  occur  within  all  cavities  after  a  few 
hours. 

After  a  few  hours  hemorrhages  are  present  in  many  of  the 
mesenchymal  areas  of  the  body  and  have  probably  caused  separation 
of  the  epithelial  membranes. 

All  the  effects  of  raying  appear  to  be  destructive.  No  acceleration 
of  growth  or  activity  occurs  in  any  tissue. 

No  division  of  cells  occurs  four  hours  after  the  embryos  have 
been  rayed.  Some  of  the  chromosome  arrangements  are  very  com- 
pact or  clumped. 

The  allantois  is  affected  least  of  all  tissues,  probably  because  of 
its  very  thinness  and  conseciuent  ease  with  which  gaseous  exchange 
takes  place.  This  is  in  keeping  with  results  obtained  by  Strange- 
ways  and  Fell  in  1927,  who  found  lethal  effects  were  due  to  absence 
of  gas  exchange. 


33—3482 


502 


The  University  Science  Bulletin 


> 

I— I 

w 

OQ 


s 

!  i 

03 
3 

Greater  vacu- 
olation  of  some 
cells;  fewer  mi- 
toses   and    less 
sharpness    than 
control. 

Some    break- 
ing of  walls,  and 
filling  ot  tubules 
and  ducts;  gran- 
ules  and    vacu- 
oles common; 
stain  faded  gray. 

1 
1 
1 

§ 

5 

Small  areas  of 
head  gorged 
with  blood; 
stain  is  not 
sharp   in    color; 
mitoses  less 
than  control. 

Great  hemor- 
rhages; poor  dif- 
ferentiation and 
uniformity; 
many    cells    ir- 
regular  or  bro- 
ken;   cells    with 
two  and  no  nu- 
clei occur;  kary- 
osoines  are  fre- 
quently frag- 
mented; rare 
mitoses;  distinct 
chromosome 
clumDs. 

£ 

0^                    1 

1        1 
1        j 

Stain  is  dim, 
and  cells  poorly 
diffeientiated; 
mitoses  one-half 
number  in  con- 
trol. 

Hemorrhages 
have     distorted 
niosenchyme    of 
head;  some 
shrinkage  of 
cells;     stain     is 
dark  gray;  very 
few  mitoses; 
small  cells  show 
condensed  chro- 
mosomes. 

E 

1 

No  structural 
change,  stain 
not    sharp;    nu- 
clei slightly 
granular;    chro- 
mosomes in  few 
cells  clearly 
clumped. 

Stomach   and 
hind  gut  almost 
filled    with    de- 
bris ;    wall    bro- 
ken   in    several 
places;  granules 
largo     in     liver 
cells)     the     few 
cells  in  division 
sliow    clumping 
of  chromosomes; 
stain  is  dim. 

>            1 

o 
^             1 

3                1 

■*"                1 

Slight  fraying 
of  inner  limiting 
membrane;    mi- 
toses almost 
normal  number. 

Surface  of 
wall  roughened; 
stain,  dull  gray ; 
no  mitoses; 
some  cells  have 
granules  of  ap- 
pearance of 
fragmented 
chromosomes. 

'i 
> 

a 

O 

A  little  mate- 
rial is  oozing 
through  inner 
limiting     mem- 
brane;  granules 
more  prominent; 
mitoses  com- 
mon; some  cells 
show  condensa- 
tion of  chromo- 
somes. 

Coats     some- 
wha  t   separated 
and  fissured 
with  loss  of  cells 
into  vesicle; 
stain  gray; 
many  nuclei  an- 
gular and 
shrunken;     rare 
mitoses;      chro- 
mosomes 
clumped. 

Neural  tube. 

Separation  of 
limiting     mem- 
branes    in     few 
places;  fibrinoid 
mateiial    abun- 
dant    in     brain 
cavity;    mitoses 
near  number  in 
control;    a    few 
show  distinct 
clumping  of 
chromosomes. 

Limiting  mem- 
branes widely 
broken  and  cells 
occupy  brain 
cavity;  no  cells 
dividing;     cyto- 
plasm   contains 
many  large 
granules  ar- 
ranged about 
walls;  stain  gray 

Ectoderm. 

Slight  separa- 
tions   of    mem- 
branes; stain 
less  definite  and 
sharp;  few  cells 
in  division; 
chromosomes  in 
some    in     com- 
pact tangle. 

Sepal  ations  in 
many  places; 
stain    dim    and 
gray;     granules 
large;  karyo-  , 
somes  frag- 
mented   in    few 
cells;     no     cells 
dividing. 

Age  at 

time  of 

raying  and 

observation, 

hours. 

CO 

6 

i 

Crow:  Effects  of  X-Rays  on  Chick  Embryos 


503 


o 


h- < 

Pi 

03 


s 

i 

v-- 

3 

^ 

c 

>. 

T3 

I- 

>1 

t 

"3 

"2 
3 

1 

•a 

'3 

03 

o 
2 

> 

03 

i 

0 

i 

E 

_3 

"c 

"o 

E 

3 

> 

M 

5 

0 

g 

3 

M 

m 

p 

_m" 

■53 

-a 

3 

0 

£ 

s 

03 

^ 

■5 

^ 

3. 

"3 

■S 

■§ 

a 

03 

o 
o 

M 

1 

"3 

3 
C 

"3 

c 

'i 

3 
3 

o 

u 

1 

-2 
"3 

3 
c3 

1 

-p 

3 

"2 

08 

i 
03 

"5. 

3 

1 
0    b' 

s 

1 

3 

1 

-0 

.S 

>1 

3 
03 

3 

— 

"3 

_ 

3   .0 

3 

■3 

5 

S 

^ 

o 

0 

5 

i 

■s 

c3 

C3 

j: 

^ 

2 

1 

o 

3 

3 

"3 

3 
03 
O 

E 

io 

'3 

3 

„ 

0 

o 

O 

■& 

"3 

03 

3 

E 

[3 

3 
£ 

bi 

|3 

03 

"g 

> 

o 

3 

E 

tc 

o 

O 

O 

> 

c; 

13 

. 

^ 

J! 

>. 

K 

^ 

^ 

^ 

_> 

1 

a; 

1 

d 

s 

T) 

J3 

2. 

o 

O 

3 

1 

x 

^ 

2 

3 
03 

3 

a 

"a 

3 

bi 

o 

M 

0 

3 

3 

"3 

X 

s 

u 

c 

Ml 

s 

-g 

"T" 

-a 

^ 

tJ 

'o 

> 

S 

"3 

£ 

3 

E 

3 

c 

'3 

CC 

m 

o 

>, 

^ 

0 

o 

w 

t^ 

0 

_£ 

c 

o 

o 

03 

^ 

"3 

^ 

_>> 

'e 

-3 

T3 

■^ 

^C 

a 

C 

— 

0 

< 

03 

•- 

S 

■3 

3 

_o 

O 

_aj 

03 

s 

13 

"c 

"3 

£ 
p 

C 

3 

4) 

0; 

a 

c 

03 

1 

> 

C 
o3 

'S 

3 

a 

3 

b 
>> 

^- 

u 

3 

1 

C3 

> 

"S 

O 

03 

.E 

00 

0 
3 

~ 

"0 

a. 
O 

O 

O 

1 
"S 

^ 

'o 

o 

3 

1 

S 

0 

3 

0 
3 

01 
M 

-2 

a; 

-n 

c 

i 

a 

o 

3 
"3 

3 

3 
o3 

— 

1 

0 

3 

J2 

03 

3 

^ 

-d 

3 

« 

o 

s 

4) 

S      3 

'3 

0 

1.= 

3 
3 

03 

3 

0) 

E 

1. 
§    S 

^  E 

"3 

03 

0) 

1 

T3 

5 

3 

"p. 

"o 

3 

> 

E 

03 

a 

u 

'3 

-^ 

u 

0 

0 
a 

0 

03 

0) 

x:  -a 
e  .2 

bi 

.3 

03 

■a 

3 
03 

jl. 

, 

Lh 

^_ 

^ 

m 

i 

1 

a 

a; 

-o 
c 

oJ 

0 

a 

3 

o 

3 
0 

o 
'c 
E 

c3 

3 

■3 

3 
0 

1 

o 

3 

.2 

_0 

T. 

i 
J 

-a 

^ 

_2 

2 

K 

^ 

C3 

Is 

'o3 

3 
03 

3 

"3 

'> 

3 

3 

u 

03 

0 

0; 

03 

01 

bC  73 

=" 

3 

.-^'S 

o 

^  c  s 

0)   S   M 

>  3 

„• 

^ 

1^ 

"7 

aiXi 

S 

u 

o 

:j 

504  The  University  Science  Bulletin 

DISCUSSION 

In  series  II  to  VII  of  this  paper  all  factors,  i.  e.,  dosage,  time  and 
distance,  have  been  kept  constant.  A  better  comparison  could,  in 
this  way,  be  obtained.  The  results  have  been  in  accord  with  expecta- 
tions. Whether  a  certain  dosage  given  all  at  one  exposure  or  broken 
into  two  or  more  fractional  parts  and  administered  on  successive 
days,  has  the  same  effect  has  been  made  the  subject  of  experimenta- 
tion by  Oilman  and  Baetjer  (35),  who  found  exposures  of  ten  min- 
utes per  day  for  four  days  produced  abnormalities,  deformities  and 
retardations  in  chick  embryos.  Ancel  and  Vintemberger  (14)  ob- 
served that  frog  eggs  exposed  to  X-rays  showed  less  degree  of  effect 
for  a  given  dose  when  it  was  spread  over  two  or  more  applications 
than  when  employed  as  one  dose;  but  in  an  earlier  paper,  in  1925, 
Ancel  and  Vintemberger  (12)  had  declared  that  the  destructive  ef- 
fects of  raying  on  the  chick  blastodemi  were  not  modified  by  divid- 
ing the  dose  into  fractional  parts.  Colwell,  Gladstone  and  Wakely 
(25)  caused  profound  degeneration  in  tissue  of  the  chick  by  doses 
on  three  and  five  successive  days  and  Strangeways  and  Oakley  (79) 
found  similar  cellular  changes  when  a  single  dose  of  soft  rays  was 
employed  upon  chick  choroid  growing  in  vitro,  which  was  confirmed 
by  Strangeways  and  Fell  (80)  on  living  chick  embiyos  one  day  and 
six  days  old.  Yamamoto  (94)  made  an  excellent  test  of  this  idea 
when  he  exposed  chick  embryos  of  similar  age  to  single  doses  and 
to  fractional  doses.  His  conclusions  show  a  greater  number  of  deaths 
occurred  in  the  group  exposed  to  one  dose. 

The  eflfect  of  irradiation  on  the  chromosomes  was  recorded  for 
Ascaris  megalocephala  by  P.  Hertwig  (43)  as  a  great  disturbance  of 
the  chromosome  arrangement,  bearing  little  resemblance  to  that  of 
normal  cell  division. 

Gatenby  and  Wigoder  (34)  found  pycnosis,  similar  to  the  "pyk- 
nosen  and  pseudo-amitosen"  of  Politzer  (68),  giant  cell  formation, 
necrosis  and  other  abnormalities.  Colwell,  Gladstone  and  Wakely 
(25)  observed  condensation  of  chromosomes  into  distinct  clumps, 
while  Strangeways  and  Fell  (80)  found  pluripolar  spindles  and 
fragmentation  of  chromosomes. 

In  an  earlier  paper,  1923,  Strangeways  and  Oakley  (79)  ob- 
served on  chick  tissues  growing  in  vitro,  when  exposed  to  soft  X- 
rays,  very  distinct  condensation  of  chromosomes  into  clumps, 
which,  because  of  this  massing,  could  not  divide,  but  instead  broke 
up  and  disintegrated.  In  other  cells  attempting  to  divide,  most  of 
the  chromosomes  reached  the  anaphase  or  telophase,  leaving  the 


Crow:  Effects  of  X-Rays  on  Chick  Embryos  505 

others  only  partially  split,  and  thus  were  made  to  lag  behind.  In 
still  others,  when  hit  just  before  entering  the  prophase,  the  break- 
ing down  of  the  chromosome  complex  did  not  take  place  till  the 
telophase,  or  "daughter-cell,"  stage. 

The  findings  in  the  present  series  of  experiments  are  very  much 
in  harmony  with  those  of  the  workers  just  cited.  There  have  oc- 
curred metaphases  in  many  of  the  embryos  in  which  the  chromo- 
somes are  condensed  into  quite  distinct  clumps  (figs.  31,  75,  76, 
77).  A  few  cells  show  a  condition  where  the  chromatin  is  as  one 
heavy  cord  formed  into  a  much-tangled  knot  (figs.  26,  29,  34). 
There  were  found  a  small  number  of  cells  with  tripolar  spindles 
(figs.  10,  11)  in  one  specimen,  and  in  rare  cases  in  a  few  other 
individuals.  The  cells  having  two,  three  or  four  nuclei  of  unequal 
size  may  be  the  result  of  such  abnormal  spindle  formation  (figs. 
8,  36).  A  number  of  cells  in  different  embryos  show  such  unusual 
chromosome  arrangements  as  to  bear  little  resemblance  to  those 
in  regular  mitosis  (figs.  16,  19,  28,  34).  Figures  19  and  28  resemble 
the  "pseudo-amitosen"  of  Politzer. 

In  a  small  number  of  cells  in  a  few  of  the  embryos  the  chromo- 
somes are  so  widely  separated  and  disarranged  as  to  suggest  a  dis- 
persal while  in  the  prophase  (figs.  13,  16,  38,  63). 

In  all  embryos,  four  hours  after  being  rayed,  cells  occur  that 
have  nuclei  in  various  stages  of  degeneration  (figs.  5,  9,  24,  49,  50, 
55,  56,  57,  58,  59,  68,  69,  70,  71,  72,  73).  In  many  of  these  the  size 
and  number  of  granules  suggest  dispersal  and  partial  fragmentation 
of  the  chromosomes. 

The  work  of  Strange  ways  and  Fell  (80)  upon  the  chick  embryos 
of  20-25  hours,  6  days  and  17  days  is  similar  to  that  reported  in 
this  paper.  Using  embryos  of  wider  age  difference  has  not  modified 
conditions  materially.  Two  dosages  were  used  by  these  workers, 
the  one  about  double  the  strength  of  the  other.  With  the  heavier 
dose  changes  occurred  somewhat  more  quickly  and  with  greater 
destruction.  The  embryos,  after  exposure,  were  incubated  for 
periods  varying  from  none  to  six  days.  The  20-25  hour  embryos, 
observed  immediately  after  raying,  showed  no  microscopic  changes, 
but  sections  showed  a  great  reduction  of  mitotic  figures.  At  80 
minutes,  with  lighter  dose,  mitoses  were  rare;  with  heavier  dose, 
absent.  The  general  conditions  of  the  tissues  under  similar  ex- 
posures were,  respectively,  slight  cellular  degeneration  and  greater 
cellular  degeneration. 

After  24  hours  a  partial  recovery  seemed  to  have  taken  place,  for 


506  The  University  Science  Bulletin 

mitosis  became  active  again,  though  the  tissues  showed  abnormal 
mitoses,  reduced  growth,  and  degeneration.  Some  of  the  abnormal 
mitoses  showed  pluripolar  spindles  and  others  fragmentation  of 
chromosomes.  Nuclear  and  cellular  disintegrations  were  present 
and  the  mesenchyme  had  become  edematous.  In  embryos  exposed 
at  six  days  and  examined  immediately  there  was  a  striking  reduc- 
tion of  mitoses,  some  cells  showed  slight  degenerative  effects  and 
the  surface  of  the  body  was  somewhat  reddened.  In  26  minutes 
sections  showed  degeneration  in  many  cells,  mitoses  rare,  and  large 
numbers  of  cells  in  the  delicate  vascular  walls  breaking  down.  In 
52  minutes  dark  splotches  occurred  in  most  specimens,  which,  on 
being  sectioned,  showed  great  extravasation  of  blood.  There  were 
no  mitoses  observed.  After  80  minutes  much  crimson  color  of 
body  showed.  An  enonnous  number  of  degenerated  cells  occurred 
in  all  tissues,  especially  in  the  retina  and  central  nervous  system. 
The  color  of  body  was  due  to  the  engorgement  of  the  superficial 
vessels,  the  walls  of  which  had,  in  many  places,  broken  down  and 
liberated  blood  into  the  surrounding  tissues.  Mitoses  were  very 
rare  or  absent.  After  24  hours  no  sign  of  recovery  could  be  noted. 
There  was  some  hemolysis,  which  reddened  the  amniotic  fluid.  Most 
cells  were  degenerating  and  many  broken  dow^n.  There  were  no 
mitoses. 

The  results  shown  in  this  paper  parallel  very  closely  those  given 
above  as  reported  by  Strangeways  and  Fell  (80).  The  series  of 
chicks  used  were  of  ages  with  shorter  time  intervals  between,  e.  g., 
28,  48,  60,  75,  96  and  120  hours.  The  intervals  after  exposure  were 
somewhat  more  variable  than  those  of  Strangeways  and  Fell  (80). 
The  degenerative  effects  were  not  so  quickly  shown  nor  were  they 
so  striking,  there  being  no  great  reduction  in  number  of  mitoses  ob- 
served after  two  hours  and  no  complete  disappearance  of  them  be- 
fore six  hours  in  some  individuals.  In  no  case  was  there  any  re- 
coveiy  or  recurrence  of  mitosis,  but,  rather,  degeneration  became 
greater  with  time,  with  very  great  disorganization  of  much  of  the 
nervous  tissue  before  20  hours.  There  was  not^only  accumulation 
of  disorganized  material  within  the  retina  and  central  nervous  sys- 
tem as  reported  by  Strangeways  and  Fell  (80),  but  within  the  tubes 
and  cavities  in  other  parts  of  the  body  as  well,  yet  this  did  not  ap- 
pear in  great  quantity  for  four  or  six  hours.  There  was  extravasa- 
tion of  blood  in  embryos  after  two  hours,  and  ever  increasing  with 
time  after  raying  until  in  some  specimens  brain  and  retinal  walls 
were  folded  and  most  of  the  mesenchymal  spaces  gorged.    For  the 


Crow:  Effects  of  X-Rays  on  Chick  Embryos  507 

pluripolar  spindles,  fragmentation  of  chromosomes  and  other  abnor- 
mal mitoses  of  Strangeways  and  Fell  (20)  there  were  found  tripolar 
spindles  (figs.  10,  11),  fragmented  (figs.  13,  38)  and  dispersed 
chromosomes  (figs.  13,  38)  and  in  others  condensed  in  a  clump 
(figs.  31,  75,  76,  77),  the  last  not  being  found  by  other  workers. 

Though  different  in  method  of  exposure,  the  chicks  being  exposed 
to  X-rays  on  successive  days,  yet  the  results  obtained  by  Colwell, 
Gladstone,  and  AVakely  were  very  similar  to  those  reported  in  the 
present  paper.  In  their  paper  of  1922,  series  I  (24),  the  conclusion 
was  reached  that  of  those  specimens  which  did  develop  after  raying, 
all  the  organs  and  tissues  which  should  develop  during  the  period 
•examined  were  differentiated,  but  were  smaller  in  size.  In  series  II 
and  III  (25)  the  same  authors  conclude  that  the  effect  is  always  de- 
structive and  is  exerted  chiefly  on  the  ectoderm,  neural  tube  and  its 
associated  structures,  such  as  the  retina  and  optic  vesicle  the  epithe- 
lium of  glandular  structures,  as  the  liver,  or  the  mesonephros,  and 
upon  the  red  blood  corpuscles.  The  effects  consist  in  the  detach- 
ment of  the  epithelium;  diminution  of  the  transparency  of  the 
cytoplasm,  or  the  formation  in  the  cytoplasm  of  minute  granules  or 
vacuoles;  variation  in  degree  of  distinctness  with  which  the  chromo- 
somes may  be  distinguished  in  the  dividing  nuclei,  and  in  the  stain- 
ing reactions  of  the  nuclei  and  cell  bodies.  The  effect  on  the  blood 
corpuscles  appears  also  to  be  destructive.  In  a  few  cases  a  fibrinous 
clot  appears  in  the  heart  or  larger  vessels.  In  their  specimens  re- 
ceiving the  full  dosage,  hemorrhages  were  frequent  in  the  cavity  of 
central  nervous  system  and,  to  a  less  degree,  other  cavities  of  the 
body  were  partly  filled  with  fibrinoid  and  cellular  material.  The 
cells  in  division  after  irradiation  show  chromosomes  which  are  con- 
densed into  a  clump.  There  appears  to  be  no  reduction  in  number 
of  mitoses  nor  is  there  any  stimulation. 

The  findings  in  the  present  paper  present  results  similar  to  those 
reported  by  Colwell,  Gladstone  and  Wakeley  (25).  There  is  less 
separation  of  the  ectoderm  and  very  little  of  the  epitrichium.  The 
effect  upon  the  neural  tissue  and  sense  organs  is  as  great  as  observed 
by  these  workers.  In  the  more  advanced  stages  after  raying  the 
results  are  probably  much  greater  than  they  report.  In  only  one 
situation  do  they  report  breaking  of  ''internal  limiting  membrane 
lining  the  central  canal  of  the  cord  and  the  canal  partially  filled  with 
fibrinous-like  deposit  containing  a  group  of  small  oval  cells  derived 
from  the  neuro-syncytium,  forming  the  wall  of  the  injured  tube." 
Most  specimens  described  in  the  present  paper  show  exudations  from 


508  The  University  Science  Bulletin 

brain  cells  and  often  much  debris  within  its  cavity  accompanied 
by  definite  spaces  or  fissures  from  which  cells  or  cell-parts  have 
come.  The  effect  on  the  coats  of  the  eye  is  much  the  same  as  that 
observed  on  the  neural  tissue.  There  is  generally  separation  of  the 
two  coats  within  two  hours,  the  distance  increasing  with  time  after 
raying.  In  nearly  every  case  this  space  is  filled  with  fibrinous-like 
material  to  which  disorganized  cells  are  added  in  a  few  hours,  the 
total  mass  becoming  so  great  as  to  cause  folding  of  retinal  coats. 
Both  coats  may  be  broken  or  fissured  which  may  admit  blood  cells 
to  the  mass  within  the  space.  Within  the  lens  vesicle  disorganized 
material  occurs  after  a  few  hours.  Colwell,  Gladstone  and  Wakeley 
(25)  report  no  widespread  modifications  in  the  eye,  reporting  only 
on  the  distribution  of  pigment  granules  and  staining  qualities  of  the 
pigment  cells.  They  report  the  effect  of  the  X-rays  on  the  entoderm 
and  its  derivatives  as  being  very  slight,  while  the  present  paper  re- 
ports degenerative  effects,  in  all  advanced  stages.  They  report  little 
or  no  effect  in  other  tissues  except  blood. 

It  is  in  comparison  of  the  blood  and  mitoses  that  the  present  paper 
agrees  most  closely  with  these  workers.  In  both  extensive  hem- 
orrhages are  reported  and  in  both  destructive  effects  are  noted. 
These  effects  are  shown  by  both  in  the  breaking  up  of  cytoplasm 
(figs.  1,  2,  15),  the  formation  of  granules,  the  fragmentation  of 
karyosomes  (figs.  39,  42,  46),  and  occasionally  the  fragmentation 
of  the  whole  cell  (figs.  24,  70,  72).  Both  report  cells  without  nuclei. 
(Figs.  6,  7.) 

The  appearance  of  the  cells  which  are  in  division  is  reported  much 
the  same  in  both  papers.  Both  report  variation  in  degree  of  dis- 
tinctness with  which  chromosomes  may  be  distinguished  in  dividing 
nuclei.  Both  report  the  condensation  of  the  chromosomes  into 
clumps  (figs.  31,  75,  76,  77)  in  which  the  individual  elements  could 
not  be  followed.  It  appears,  then,  that  with  lighter  doses  given  at 
daily  intervals  cell  division  and  cellular  structure  may  be  modified, 
but  widespread  destruction  is  not  produced,  while  with  single,  heavy 
dosage,  as  reported  in  the  present  paper,  modifications,  similar  to 
those  reported  by  the  workers  above,  appear,  after  a  few  hours,  to 
be  followed  by  even  greater  degeneration  in  later  stages. 

Several  workers  have  employed  Radium  and  others  ultra-violet 
light  in  their  experiments  on  living  tissues.  The  effects  of  both  of 
these  are  very  similar  to  those  produced  by  exposures  to  X-rays. 

One  very  striking  modification  of  living  cells  produced  by  X-rays, 
radium  and  ultra-violet  light  is  the  suppression  of  mitosis.     In  all 


Crow:  Effects  of  X-Rays  on  Chick  Embryos  509 

the  embryos  on  which  report  is  made  in  this  paper,  with  the  excep- 
tion of  one  individual,  whose  susceptibility  to  the  dosage  of  X-rays 
employed  must  have  been  lower  than  all  the  others,  a  very  definite 
reduction  in  number  of  mitoses  occurred  in  two  hours  with,  in  al- 
most all  cases,  complete  disappearance  of  all  cell  division  in  six 
hours.  The  individual  variation  that  occurred  was  not  great  enough 
to  be  a  matter  of  record.  No  reduction  in  number  or  other  change 
could  be  observed  in  embryos  twenty  minutes  after  raying,  but 
Strangeways  and  Fell  (80)  found  a  very  rapid  fall  in  thirty  min- 
utes. After  the  abrupt  fall  a  further  reduction  continued  much 
more  gradually  for  three  hours  when  there  began  a  rise  in  number 
which  continued  to  increase  till  the  original  number  had  been  sur- 
passed. Canti  and  Spear  (23)  recorded  a  similar  sharp  reduction, 
followed  by  a  gradual  rise  which  sometimes  was  equal  to  the  fall  for 
cells  growing  in  vitro  and  exposed  to  gamma  radiation.  Grasnick 
(36)  found  the  nuclei,  especially  those  in  mitosis,  more  affected  by 
gamma  rays  while  the  cytoplasm  was  influenced  more  by  the  beta 
rays  of  radium. 

Canti  (21)  demonstrated  the  effect  of  radium  upon  chick  embryo 
tissues  growing  in  vitro  by  making  slow  pictures  of  them.  In  this 
unique  method  are  shown  cells,  which  moving,  cease  mobility,  be- 
come round  and  form  bubbles  in  their  process  of  breaking  down. 
Mitosis  is  stopped  in  20  minutes.  Two  daughter  cells  moving  apart 
are  stopped. 

Canti  and  Donaldson  (20)  found  similar  stopping  of  mitoses  by 
exposure  to  radium,  the  choroid  tissue  of  chick  embryos  growing 
in  vitro.  In  this  it  was  observed  that  the  more  intense  the  irradi- 
ation the  sooner  was  mitosis  stopped. 

Reiss  (73)  compares  the  action  of  X-rays  and  radium  on  the 
nucleus  and  cytoplasm  in  "Pour  nos  experiences,  le  rapport  des 
sensibilites  du  cytoplasme  et  du  noyau  serait  de  18  :  1  pour  le 
radium,  et  de  3.8  :  1  pour  les  rayons  X." 

That  ultra-violet  light  acts  similarly  to  X-ray  and  radium  is 
shown  by  Earl  (31)  who  observed  its  action  upon  blood  and  fibro- 
blasts. The  erythrocytes  became  swollen  and  lost  their  hemoglobin; 
while  white  blood  cells  were  fixed  and  coagulated.  Fibroblasts  from 
chick  embryos  were  observed  to  swell,  develop  vacuoles  and  coagu- 
late. 

Hinricks  (45,  46)  has  done  extensive  experimenting,  using  ultra- 
violet light  upon  chick  embryos,  and  finds  abnormalities  and  wide- 
spread modifications. 


510  The  University  Science  Bulletin 

Short  exposures  of  tissue  cultures  of  fibroblasts  to  ultra-violet 
light  was  recorded  by  Kler  (54)  as  having  very  little  effect;  to  pro- 
duce a  lethal  effect  heavier  doses  being  necessary.  In  a  similar  treat- 
ment of  chick  heart  fragments  after  an  exposure  of  15  seconds  to  2 
minutes  a  stimulation  of  contraction  was  observed  by  Kler  (55)  who 
found  for  the  intensity  of  light  employed  2-3  hours  exposure  was 
necessary  to  be  lethal.  Watrin  (89)  exposed  tadpoles  in  water  to 
X-rays  and  to  ultra-violet  light,  finding  the  former  had  no  effect, 
but  during  exposure  to  the  latter  the  tadpoles  moved  rapidly  and 
after  3  doses  at  5-day  intervals  they  lost  weight  and  pigment,  and 
died  in  5  days. 

GENERAL  CONCLUSIONS 

The  effects  of  irradiation  with  X-rays  appear  as  destructive 
changes  in  all  tissues,  varying  directly  with  the  lengih  of  time  after 
exposure. 

These  effects  for  the  most  part  are  general  rather  than  local. 

Effects  make  their  appearance  at  about  the  same  time  after  ir- 
radiation in  all  series. 

Mitoses  are  not  stopped  suddenly,  but  a  gradual  diminution  takes 
place  until,  after  4-6  hours,  few  or  none  occur.  Some  mitotic  figures 
show  chromosomes  somewhat  condensed  into  a  clump  so  that  in- 
dividual elements  cannot  be  discerned. 

No  acceleration  of  growth  can  be  found. 

The  greatest  effects  are  shown  in  the  nerve  tissue,  sense  organs, 
epithelium  and  blood  cells. 

Pronounced  effects  are  shown  in:  Accumulation  of  disorganized 
material  in  vesicles,  canals  and  cavities  of  body;  detachment  of 
epithelial  membranes,  limiting  membranes,  and  separation  of  retinal 
walls;  great  hemorrhages  into  loose  mesenchyme;  appearance  of 
granules  in  the  cytoplasm ;  decrease  in  differentiation  of  nuclear 
structure. 

BIBLIOGRAPHY 

1.  A.nLER,  Karl.     1930.     Die  biologische  Wirkung  dei\  kurzwelligen  Strahlen 

auf  den  Stoffwechsel  der  Zelle.     Strahlentherapie  36,  1  :  1-31. 

2.  Alberti,  W.,  and  Politzer,  G.     1923.     Uber  den  Einfluss  der  Rontgen- 

strahlen  auf  die  Zellteilung.     Arch.  f.  Mikr.  Anat.     100  :  83. 

3.  Alberti,  W.,  and  Politzer,   G.     1924.     Uber  den  Einfluss  der  Rontgen- 

strahlen  auf  die   Zellteilung.     II.     Mitteilung.     Arch.  f.  Mikr.   Anat. 
103  :  284. 

4.  Amato,  a.     1911.     Uber  die  Wirkung  der  Rontgenstrahlen  auf  in  karyo- 

kinese  begriffene  Zellen.     Zeitschr.  f.  Rontgenk.     13.     (Quoted  by  Al- 
berti and  Politzer.     [2]). 

5.  Ancel,  p.,  and  Vintemkerger,  P.     1924.     Action  acceleratrice  des  rayons 

X  sur  de  developpement  embryonnaire.     Comp.  rend.  Soc.  biol.  91  : 
606-609. 


Crow:  Effects  of  X-Rays  ox  Chick  Embryos  511 

6.  Ancel,  p.,  and  Vixtemberger,  P.     1924.    De  action  des  rayons  X  sur  la 

segmentation  et  la  gastnilation  chez  Rana  fusca.     Comp.  rend.  Soc. 
biol.  91  :  1267-70. 

7.  AxcEL,  p.,  and  Vixtemberger,  P.     1924.    La  sensibilite  d'une  cellule  aux 

rayons  X,  est-elle  plus  grande  au  cours  de  la  mito.se  que  pendant  le 
repos?    Comp.  rend.  Soc.  biol.  91  :  1271-73. 

8.  Ancel,  p.,  and  Vixtemberger,  P.     1925.     Influence  de  Tactivite  cellulaire 

sur  la  manifestation  des  lesions  produites  dans  le  blastoderme  de  I'oeuf 
de  poule  par  les  ra5'ons  X.    Comp.  rend.  Soc.  biol.  91  :  1425-26. 

9.  AxcEL,  P.,  and  Vixtemberger,  P.     1925.     Etude  de  Taction  des  rayons 

X  sur  le  developpement  embiyonnaire.    Arch,  de  biol.  35  :  1-43. 

10.  Axc-el,   p.,  and  Vixtemberger,   P.     1925.     Sur  revolution   comparee   de 

I'embn'on  et  du  blastoderme  dans  les  oeufs  soumis  a  Taction  des  rayons 
X.     Comp.  rend.  Soc.  biol.  92  :  172-4. 

11.  Axcel,  p.,  and  Vixtemberger,  P.     1925.    Sur  la  radiosensibilite  cellulaire. 

Comp.  rend.  Soc.  biol.  92  :  517-26. 

12.  AxcEL,  P.,  and  "\'ixtemberger,  P.    1925.    Sur  la  radiosensibilite  des  cellules 

en  carj'ocinese.    Comp.  rend.  Soc.  biol.  92  :  986-8. 

13.  AxcEL,  P.,  and  Vixtemberger,  P.     1925.    Comparison  entre  les  effets  des 

ravons  X  et  ceux  du  vieillissement  sur  Toeuf  de  poule.     Comp.  rend. 
Soc.  biol.  92  :  1401-3. 

14.  AxcEL,  p.,  and  Vixtemberger,  P.    1928.    Sur  Teffet  du  fractionnement  des 

deses  de  ravons  X  et  Thvpersensibilite  des  cellules  en  mito.se.    Comp. 
rend.    Soc.  biol.  99  :  852-4. 

15.  Baldavix.  Wm.    1920.    Effects  Produced  by  X-rays  on  Frog  Ova.    Science. 

52:229-30. 

16.  Br.\mbell,  F.  W.  R..  Parkes,  A.  S.,  and  Fieldixg  U.    1927.    Changes  in  the 

Ovary  of  the  Mou.se  Following  Exposure  to  X-ravs.    Part  I,  Irradiation 
at  three  weeks  old.    Proc.  Roy.  Soc.  lOlB  :  706,  29-56,  3  pis. 

17.  Brambell,  F.  W.  R.,  Parkes,  A.  S..  and  Fieldixg  U.    1927.    Changes  in  the 

Ovary  of  the  Mouse  Following  Exposure  to  X-ravs.    Part  II,  Irradiation 
at  or  before  birth.    Proc  Roy.  Soc.  lOlB  :  707,  95-114,  2  pis. 

18.  Brambell,  F.  W.  R.,  and  Parkes,  A.  S.    1928.    Changes  in  Ovary  of  Mouse 

Following  X-rays.     Part  IV,  Corpus  lutem  in  the  sterilized  ovarj^  and 
some  concluding  experiments.    Proc.  Roy.  Soc.  102B  :  719,  383-396. 

19.  Burrows,  M.  T.,  Jorst.\d,  L.  H.,  and  Erxst,  E.  C.     1927.    The  Action  of 

X-rays  on  the  Organism.    Proc.  Soc.  Exp.  Biol,  and  Med.  24  :  311-12. 

20.  Caxti,  R.  G.,  and  Doxaldsox,  M.    1926.    The  Effects  of  Radium  on  Mi- 

tosis in  Vitro.    Proc.  Roy.  Soc.  lOOB  :  705,  413-419. 

21.  Canti,  H.     1928.     Cinematograph  Demonstration  of  Living  Tissue  Cells 

Growing  in  Vitro.    Archiv.  f.  exper.  Zellf.  B6. 

22.  C.AXTi,  R.  G.,  and  Spear,  F.  G.     1927.     Effect  of  Gamma  Irradiation  on 

Cell  Division  in  Tissue  Culture  in  Vitro.    Proc.  Roy.  Soc.  102B  :  92-101. 

23.  Caxti,  R.  G..  and  Spe.ar,  F.  G.     1929.    The  Effect  of  Gamma  Irradiation 

on  Cell  Di-i.-ision  in  Tissue  Culture  in  Vitro.    Part  II.  Proc.  Roj-.  Soc. 
lOoB  :  93-98. 

24.  CoL^VELL,  H.  A.,  Russ,  S..  and  Wakely,  C.  P.  G.    1924.    Radium,  X-rays 

and  the  Living  Cell.    Introduction  to  the  Study  of  X-rays  and  Radium. 
G.  Bell  and  Sons.     (London.) 

25.  CoLAAELL,  H.  A.,  Gladstoxe,  R.  G.,  and  Wakely,  C.  P.  G.     1922.    Action 

of  Repeated  Doses  of  X-rays  Upon   the   Developing   Chick   Embiyo. 
Jour.  Anat.    57  :  1-11. 

26.  CoLWELL,  H.  A.,  Gladstoxe,  R.  C...  and  Wakely,  C.  P.  G.    1926.    Action 

of  X-ravs  Upon  the  Developing  Chick  Embrvo.    Series  2  and  3.    Jour. 
Anat.  60  :  207-228. 

27.  Cramer.  W..  Drem-,  A.  H.,  and  Mottham.  J.  C.    1921.    Similarity  of  Effects 

Produced  by  Absence  of  Vitamins  and  bv  Exposure  to  X-ravs.    Lancet. 
1  :  963-4. 


512  The  University  Science  Bulletin 

28.  Crowther,  J.  A.     1924.     Some  considerations  Relative  to  the  Action  of 

X-rays  on  Tissue  Cells.    Proc.  Roy.  Soc.  96B  :  207-11. 

29.  Dawson,  Alden  B.    1927.    Histological  Study  of  the  response  of  intestinal 

Mucosa  of  Dog  to  Irradiation  With  Special  Reference  to  Giant-cell 
Formation.    Jour.  Exp.  Zoology  46,.  4  :  467-491,  2  pis. 

30.  DoAN,  C.  A.,  Cunningham,  R.  S.,  and  Sabin,  F.  R.    1925.    Experimental 

Studies  on  the  Origin  and  Maturation  of  Avian  and  Mammalian  Red 
Blood-cells.  Cont.  to  Embryol.,  Carnegie  Inst.  Wash.,  No.  83,  Vol. 
16  :  163-226. 

31.  Earl,  W.  R.     1928.    Studies  Upon  the  Effects  of  Light  on  Erythrocytes 

in  Vitro.    Jour.  Exp.  Med.  48  :  669-82,  683-94. 

32.  Gassxjl,  R.     1927.    Uber  Rontgensthahlenwirkung  auf  lebende  Gewebe  in 

vitro.    Strahlentherapie  27  :  545-560.     19  bildem. 

33.  Gassul,  R.    1928.    Uber  Rontgenstrahlenwirkung  auf  lebende  Gewebezellen 

in  vitro.     (Explantationversuche)  Archiv.  f.  exper.  Zellf.  B6. 

34.  Gatenby,  J.  B.,  and  Wigoder,  Sylvlv.    1929.    The  Effect  of  X-radiation  on 

the  Spermatogenesis  of  the  Guinea  Pig.  Proc.  Roy.  Soc.  104B  :  731,  351- 
371. 

35.  GiLMAN,  P.  K.,  and  Baetjbr,  F.  G.     1904.     Some  Effects  of  the  Rontgen 

Rays  on  the  Development  of  Embryos.    Am.  Joiu-.  Phys.  X  :  222. 

36.  Grasnick.    W.      1917.     Die    Wirkung    der   Radiumstrahlen    auf   tierische 

Gewebe.    Arch.  f.  Mikr.  Anat.  90  :  1-35. 

37.  Hance,  R.  T.     1925.     Fixation  of  Avian  Chromosomes.     Anat,  Record, 

31  :  87-92. 

38.  Hance,  R.  T.     1926.     A  Comparison  of  Mitosis  in  Chick  Tissue  Culture 

and  in  Sectioned  Embryos.    Biol.  Bull.    50  :  155-59. 

39.  Hance,  R.  T.    1926.    The  Chromosomes  of  the  Chick  Soma.    Biol.  Bull., 

51  :  443-449. 

40.  Hance,  R.  T.     1928.    Hereditary  Constitution  and  X-rays.    Sci.  Monthly 

27,  3  :  264-226. 

41.  Hartmann,   a.     1920.     Uber  die    Einwirkung   von    Rontgenstrahlen   auf 

Amphibienlarv-en.  Einwirkung  geringer  Strahlendosen  auf  das  Blut 
imd  das  blutbildende  Gewebe  von  Rana  temporaria  Larven.  Arch.  f. 
Entwicklungsmech,  47  :  131-209. 

42.  Heim,  Konrad.    1927.    Biologische  Rontgenwirkungen  verfolgt  beim  Huhn 

vom  Ei  bis  zum  Organexplantat.    Strahlentherapie,  27,  4  :  694-710. 

43.  Hertwig,  p.     1911.     Durch  Radiumbestrahlung  hervorgerufene  Verander- 

ungen  in  den  Kemteilungsfiguren  von  Ascaris  megalocephala.  Arch, 
f.  Mikr.  Anat.  B77. 

44.  Hertwig,  O.     1907.     Handbuch  der  vergleichenden  und  experimentellen 

Entwickelungslehre  der  Wirbeltiere.  (Edited  by  Hertwig,  written  by 
numerous  collaborators).    Fischer,  Jena. 

45.  Hinrichs,  Marie  A.     1926.     Modification  of  Development  in  Chick  Em- 

biyos  Induced  bv  Ultra-violet  Radiation.  Proc.  Soc.  Exp.  Biol,  and 
Med.  23.  8  :  786-788. 

46.  Hinrichs,  Marie  A.     1927.     Modification  of  Development  on  the  Basis 

of  Differential  Susceptibility  to  Radiation.  IV.  Chick  Embryos  and 
Ultra-violet  Radiation.    Jour.  Exp.  Zoology,  47  :  309-342. 

47.  Hoffman,  V.    1922.    Uber  Erregung  und  Liihmung  tierischer  Zellen  durch 

Rontgenstrahlen.  I.  Untei-suchungen  an  Froscheirn  und  Larven. 
Strahlentherapie,  13  :  277-98. 

48.  Holthl'SEN,  H.    1921 .    Beitriige  zur  Biologie  der  Strahlenwirkung.    Pfliiger. 

Arch.  f.  Physiologie  187B^  1-24. 

49.  Jolly,  J.     1924.    I.    Action  des  rayons  X  sur  les  cellules  modifications  de 

la  radiosensibilite  par  ligature  des  connecions  vasculaires.  II.  Sensi- 
bilite  comparee  des  differents  organes  Ivmphoides  aux  ravons  X.  Comp. 
rend.  Soc.  hiol.  91  :  351-6. 


Crow:    Effects  of  X-Rays  on  Chick  Embryos  513 

50.  Jolly,  J.    1924.    Mode  d'action  des  rayons  X  sur  les  cellules.    Irradiations 

d'orgaues  isoles.    Comp.  rend  Soc.  biol.  91  :  79. 

51.  K.ARCZAG,  L.,  Farklas,  G.  v.,  and  Gyorgyi,  G.    1927.    Uber  die  biologische 

Indifferenz  der  Rontgenstrahlen  gegeniiber  Kiinstlichen  Gewebskul- 
turen.    Arch.  f.  exp.  Zellforsch  4,  a  :  206-11. 

52.  Keibei,,  F.,  and  Abraham,  K.    1900.    Normentafeln  zur  Entwickelungsge- 

schichte  des  Huhnes  (Gallus  domesticus).    Fischer,  Jena,  132  pp,  3  pi. 

53.  KEiLLicx)TT,  W.  E.    1913.    Outlines  of  Chordate  Development.    Holt,  N.  Y., 

471  pp. 

54.  Kler,  S.     1925.    Action  de  la  lumiere  ultra-violette  sur  les  cultures  tissu- 

laires  in  vitro.    Comp.  rend.  Soc.  biol.  93  :  1389-91. 

55.  Kler,  S.     1925.     Action  de  la  lumiere  ultra-violette  sur  les  fragments  de 

coeur  contractiles  in  vitro.    Comp.  rend.  Soc.  biol.  93  :  1391-94. 

56.  liEE,  A.  B.     1921.    The  Microtomist's  Vade-mecum.    8th  ed.     Blakiston's 

Son  and  Co.    Phila.  594  pp. 

57.  Lacassagne,  A.,  and  Gricouraff,  G.    1927.    Au  sujet  de  Taction  direct  ou 

indirecte  des  rayons  X.  Recherches  sur  les  tissus  lymphoides  en  survie. 
Comp.  rend.  Soc.  biol.  96  :  862. 

58.  Lillie,  F.  R.    1919.    Development  of  the  Chick.    Holt,  New  York,  XI,  472 

pp. 

59.  Markovits,  J.    1928.    Strahlenwirkung  auf  die  Zellteilung.    Archiv.  f.  ex- 

per.  Zellf.  B6. 

60.  Mandeleeff,  p.     1926.     Actions  des  Irradiations  sur  I'organisme  des  co- 

bayes.    Comp.  rend.  Soc.  biol.  94  :  1274. 

61.  MoppETT,  W.    1929.    The  Differential  Action  of  X-rays  on  Tissue  Growth 

and  Vitality.    Proc.  Roy.  Soc.  105B  :  402-21. 

62.  Nemenow,   M.  Y.     1926.     A   Contribution   to   the  Biological   Action   of 

Roentgen  Rays.     British  Jour.  Radiol.  31,  313  :  302-9. 

63.  Partces,  a.  S.,  and  Brambell,  F.  W.  R.    1927.    The  Effect  of  X-rays  Steri- 

lization on  the  Development  of  the  Accessary  Organs  of  Reproduction 
in  the  Mouse.  Parts  I  and  II.  Jour.  Roy.  Micro.  Soc.  47,  2  :  97-106, 
3  pi. 

64.  Patten,  B.   M.     1929.     The   Early   Embryology   of  the   Chick.     3d  ed. 

Blakiston's  Son  and  Co.    Phila.  XIII,  228  pp. 

65.  Payne,  F.     1913.     A  Study  of  the  Effect  of  Radium  Upon  the  Eggs  of 

Ascaris  Megalocephala  Univalens  Arch.  f.  Entw-Mech.  d.  Organ  B.  36, 
H.  3,  287-293.    Plates  19-21. 

66.  Perthes.     1904.     ^'ersuche   iiber   den   Einfluss  der  Rontgenstrahlen  und 

Radiumstrahlen  auf  die  Zelltielung.  Deutsche  Medizin.  Wochenschr. 
30  :  632,  668.    (Quoted  by  Payne,  F.  [65].) 

67.  Petersen,  W.  F.,  and  Saelhof,  C.  C.     1921.     Organ  Stimulation  by  the 

Roentgen  Ray.    J.  Amer.  Med.  Assoc.  76  :  718-20. 

68.  Politzer,  G.     1928.     Uber  die  specifische  Wirkimg  der  Rontgenstrahlen, 

Strahlentherapie,  B.  27. 

69.  Politzer,  G.     1929.    Uber  den  Einfluss  der  Rontgenstrahlen  auf  die  Zell- 

teilung.   Natur  u.  Museum  59  :  12,  583-88. 

70.  Regand,  C,  Lacassagne,  A.,  and  Javin.    1925.    Lesions  microscopiques  de- 

terminces  par  les  rayons  X  dan  I'embiyon  de  Poulet.  Comp.  rend.  Soc. 
biol.  93  :  1587-89. 

71.  Regand,  C,  and  Lacassagne,  A.    1925.    I>a  caryocinese  est  un  moment  de 

moindre  resistance  de  la  cellule  aux  radiations.  Comp.  rend.  Soc.  biol. 
92  :  123-5. 

72.  Regand,   C,  and  Lacassagne,   A.     1927.     Effets  histophysiologiques  des 

rayons  de  Roentgen  et  de  Becquerel-Curie  sur  les  tissus  normaux  des 
animaux  superieurs.    Radiol  and  Radiotherapie  I,  1  :  1-128. 


514  The  University  Science  Bulletin 

73.  Reiss,  p.  1925.  Sur  Taction  comparee  des  rayons  X  et  du  radium  siir  le 
noyau  et  le  cytoplasme.    Comp.  rend.  Soc.  biol.  92  :  1403-06. 

74  RoFFO  A.  H.  1927.  Culture  in  vitro  de  la  rate  soumise  aux  rayons  Roent- 
gen.   Comp  rend.  Soc.  biol.  97  :  1034-36. 

75.  RoLLESTON,  Humphry.     1930.     Critical  Review.     The  harmful  effects  of 

IiTadiation  (X-rays  and  radium.)    The  Quarterly  Jour,  of  Med.  93  :  101- 
131. 

76.  Rusa,  S.    1920.    Biological  Action  of  Radiations,  J.  Rontgen  Soc.  16  :  134- 

41. 

77.  Shultz,  Audrey  Flitch.     1922.    The  Mitotic  Index  of  the  Chick.     Proc. 

Okla.  Acad.  Science,  New  Series.    247  No.  18,  45. 
78    Spe.\r,  F.  G.     1930.    The  delaved  Effect  of  Radium  on  Tissue  Culture  in 
Vitro.    Proc.  Roy.  Soc.  106B  :  44-49. 

79.  Strangew.^ys,  T.  S.  P.,  and  O.akley,  H.  E.  H.     1923.     The  Immediate 

Changes  Observed  in  Tissue  Cells  After  Exj^osure  to  Soft  X-rays  While 
Growing  in  Vitro.    Proc.  Roy.  Soc.  95B  :  373-81. 

80.  Strangew.^ys,  T.  S.  P.,  and  Fell,  H.  B.    1927.    A  Study  of  the  Direct  and 

Indirect  Action  of  X-rays  on  Tissues  of  EmbiTonic  Fowl.    Proc.  Roy. 
Soc.     102B  :  713,  9-28. 

81.  Strangeways,  T.  S.  P.,  and  Hopwood,  F.  L.    1926.    Effects  of  X-rayg  Upon 

Mitotic   Cell   Division   in   Tissue   Cultures   in   Vitro.     Proc.   Roy.  Soc. 
lOOB  :  703,  283-293. 

82.  Taylor,  H.  D.,  Witherbee.  W.  D.,  and  Murphy,  J.  B.     1919.    Studies  on 

X-ray   Effects.     I.     Destructive   action   on   blood   cells.     Jour.   Exper. 
Med!  29  :  53-73,  75-82,  83-87,  89-96. 

83.  TsuzuKi.  Masao.     1926.     Experimental  Studies  on  the  Biological  Action 

of   Hard   Roentgen   Ravs.     Amer.   Jour.   Roentgenol.     16,   2  :  134-150. 
8  figs. 

84.  Vintemberger,  P.  1928.    Sur  des  variations  de  la  radiosensibilite  eu  cours 

des    premiere    mitoses    de    segmentation    dans    I'oeuf    de    Rana    fusca. 
Comp.  rend.  Soc.  biol.  98  :  532-5. 

85.  Vintemberger,  P.     1928.    Sur  le  resultat  de  I'application  d'une  dose  forte 

de  ravons  X  a  ime  region  de  la  cellule  ne  renferment  par  le  noyau. 
Comp\  rend.  Soc.  biol.  99  :  1965-7. 

86.  Vintembbrgee,  P.     1928.     Sur  les  effets  d'applications  de  rayons  X  lo- 

calisees  soit   au  protoplasm,  soit  a   la  region  nucleaire   de  la  cellule. 
Comp.  rend.  Soc.  biol.  99  :  1968-71. 

87.  Vintemberger,  P.     1930.     E'tude   experimentale  sur  la   mitose  envisogie 

comme    facteur   de    radiosensibilite.    Archiv.    d'    Anat    d'Hi.stologie    et 
d'Embiyologie  XII  :  229-466,  6  pis.,  9  figs.,  10  tables. 

88.  Warren,  S.  L.,  and  Whipple,,  G.  H.     1922.     Roentgen-ray   Intoxication 

III.     Speed  of  utolysis  of  various  body  tissues  after  lethal  X-ray  ex- 
posures.  Jour.  Exp.  Med.  35  :  213. 

89.  Watiun,  J.     1926.     Actions   Comparatives  des  rayons  X   et  des  rayons 

ultra-violet    sur    les    developpements    de    larvas   de    Rana    temporaria. 
Comp.  rend.  Soc.  biol.  95  :  189. 

90.  Weysse,  a.  W..  and  Burgess,  W.  S.     1906.     Histogenesis  of  the  Retina. 

Am.  Naturalist  4C  :  611-638. 

91.  Willis,  D.  A.,  and  Bachem,  A.     1928.    Die  Wirkung  der  Rontgenstrahlen 

auf  die  Niere.     Strahlentherapie,  B.  27. 

92.  Wilson,  F.  B.    1925.    The  Cell  in  Development  and  Heredity.    Macmillan, 

N.  Y.,  3d  ed.,  IV-1232  pp. 

93.  Wright,  S.,  and  Bulman,  H.  A.     1929.     Phvsiological  Action  of  X-rays. 

J.   Physiology.  68  :  IV-V. 

94.  Yamamoto,  T.     1927.     Effects  of  Roentgen  Rays  on  the  Development  of 

the  Embrv-o  of  the  Hen.     Japanese  Jour.  Obstetr.  and  Gvnecol.     10, 
3  :2-16,  I'pl. 


EXPLANATION  OF  PLATES 


All  the  figures  in  plates  LX  to  LXIII  are  drawings  made  in  all 
cases  with  the  aid  of  a  camera  lucida  using  a  Leitz  16  X  ocular  and 
95  X  oil  immersion  objective.  All  drawings  were  made  at  a  magnifi- 
cation of  2,800  diameters,  and  reduced  to  one-half  in  reproduction. 


(515) 


516  The  University  Science  Bulletin 


PLATE  LX 

Fig.  1.    28-45.  Ectoderm  showing  vacuoles  and  fraying  of  outer  wall. 

Fig.  2.    28-45.  Entoderm  from  open  gut  showing  fraying  of  outer  walls. 

Fig.  3.    28-45.  A  mesodermal  cell,  very  dense  nucleus  and  many  prominent, 
dark  granules. 

Fig.  4.    28-45.  A  mesenchymal  cell  with  vacuolated  nucleus. 

A  group  of  blood  cells  from  28-45: 

Fig.  5.   A  large  blood  cell  undergoing  cytoplasmic  fragmentation. 

Figs.  6  and  7.   Degenerating  cells  without  nuclei. 

Fig.  8.   Cells  with  two  nuclei. 

Fig.  9.  Blood  cells,  one  of  which  has  no  nucleus. 

All  are  degenerated  as  shown  by  the  granules  within. 
Figs.  10  and  11.    Cells  with  tripolar  nuclei. 
Fig.  12.    A  giant  blood  cell. 


Crow:    Effects  of  X-Rays  on  Chick  Embryos  517 

PLATE  LX 


518  The  University  Science  Bulletin 


PLATE  LXI 

Fig.  13.   48-60.    Entodermal  cell  with  dispersed  chromosomes. 

Fig.  14.  48-54.  Cells  composing  the  urinary  tubule  showing  numerous 
granules  in  the  cytoplasm. 

Fig.  15.  48-62.  Cells  from  the  auditoiy  vesicle  showing  the  frayed  cytoplasm 
and  fragmented  karyosomes. 

Fig.  16.   48-54.    A  blood  cell  with  disarranged  chromosomes. 

Fig.  17.  48-54.  Blood  cell  showing  condensation  of  the  chromosomes  into 
a  clump. 

Fig.  18.  48-54.  Blood  cell  with  prominent  granules  and  broken  cellular 
wall. 

Fig.  19.  48-54.  Blood  cell  with  clumped  chromosomes  somewhat  disarranged 
and  degenerating. 

Fig.  20.  48-60.    Blood  cells  with  broken  cell  walls  and  oozing  cytoplasm. 

Figs.  21  and  22.  48-60.  Mesenchymal  cells  with  somewhat  shrunken  cyto- 
plasm. 

Figs.  23  and  24.  48-60.  Blood  cells  undergoing  degeneration  and  fragmen- 
tation. 


Crow:    Effects  of  X-Rays  on  Chick  Embryos  519 

PLATE  LXI 


i^'f^^^ 


520  The  University  Science  Bulletin 


PLATE  LXII 

Fig.  25.  60-64.  A  cell  from  the  auditory  vesicle  showing  slight  clumping  of 
chromosomes. 

Fig.  26.  60-64.  Cell  from  auditory  vesicle  with  chromatin  condensed  into 
a  t-angled  knot.    Pycnosis. 

Figs.  29  and  34.  60-64.  Cell  from  mid-gut  showing  a  condensation  of  the 
chromosomes. 

Fig.  30.  60-64.  Cell  from  the  auditory  vesicle  showing  much  vacuolated 
nucleus. 

Figs.  27,  31,  32,  33.    60-64.    Blood  cells  with  shrunken  nuclei. 

Fig.  28.  60-64.  Blood  cell  with  chromosomes  disarranged  suggesting  pseudo- 
amitosis. 

Fig.  36.   60-64.   Blood  cell  with  four  nuclei. 

Fig.  37.  60-64.  Mesodermal  cell  with  clumped  chromosomes  and  granular 
cytoplasm. 

Fig.  38.  60-64.  Mesodermal  cell  with  dispersed  and  fragmented  chromo- 
somes. 

Figs.  39,  40,  41,  42,  43,  44,  45.  60-66.  Blood  cells  with  prominent  granules 
and  more  or  less  shrunken  and  degenerating. 

Fig.  46.  60-66.  Cells  from  the  urinary  tubule  showing  granular  cytoplasm 
and  fragmented  karyosomes. 


Ceow:    Effects  of  X-Rays  on  Chick  Embryos  521 

PLATE  LXII 


44 


46 


522  The  University  Science  Bulletin 


PLATE  LXIII 

Figs.  47,  48.    75-94.   Blood  cells  shrunken  and  degenerating. 

Fig.  49.    75-94.    Blood  cell  almost  clear. 

Fig.  50.   75-94.   Blood  cell  with  fragmented  nucleus. 

Fig.  51.   75-94.   Blood  cell  which  is  stained  very  darkly. 

I"^GS.  52,  53.   75-78.    Mesodermal  cells  whose  nuclei  are  dividing  by  amitosis. 

Fig.  54.   75-78.     Mesodermal  cell  with  fragmented  nucleus. 

Figs.  55,  56,  57,  58,  59.  75-M.  Blood  cells  with  fragmented  nuclei  and 
granular  cytoplasm. 

Figs.  61,  62.    75-78.    Small  blood  cells. 

Fig.  63.  75-78.  Elongated  cell  from  the  neural  tube,  showing  dispersed 
chromosomes. 

Figs.  64,  65,  66,  67,  74.  96-114.  Blood  cells  much  degenerated  and  almost 
clear. 

Figs.  68,  69,  70,  71,  72,  73.  96-114.  Blood  cells  which  are  degenerating  and 
breaking  up.  Pseudopodium-like  projections  extend  from  the  cytoplasm  of 
several  cells. 

Figs.  75.  76.  120-123.  Cells  from  the  brain  showing  clumping  of  the  chro- 
mosomes. 

Fig.  77.     120-123.    Liver  cells  showing  chromosome  clump. 

Fig.  78.    120-140.    Shrunken  Pigment  cells  from  the  retina. 


Crow:   Effects  of  X-Rays  on  Chick  Embryos  523 

PLATE  LXIII 


n 


15-3482 


Publications  of  the  University  of  Kansas 


Recently  adopted  postal  charges  are  1  cent  for  each  two  ounces  in  the  United 
States  and   possessions,    and    1%    cents   to   all    foreign   countries.      In   transmitting 
postage  for  mailing,   find  proper  amount  of  postage   for  your  zone  by  weight  in- 
dicated. 
Volume  KANSAS  university  quarterly 

I No.    1,   weight,    12    ounces.      Nos.    2,    3,    supply   exhausted.      No.    4, 

weight,  12  ounces. 
11 Nos.  1,  2,  3,  4,  supply  exhausted. 

Ill Nos.    1,    2,    supply   exhausted.      No.    3,    weight,    16    ounces.      No.    4, 

weight,  12  ounces. 

IV No.  1,  weight,  9  ounces.     No.  2,  weight  12  ounces.     Nos.  3,  4,  weight 

each,  8  ounces. 

V No.  1,  weight,  8  ounces.     No.  2,  weight,  6  ounces.     Vol.  V  oonsista  of 

only  two  numbers. 
VI,  A... Nos.   1,  2,  3,  4,  supply  exhausted. 
VI,  B...Nq.   1,  weight,  8  ounces.     No.   2,  weight,   12  ounces.     No.   3,  weight, 

8  ounces.     No.   4,  weight,  12  ounces. 
VII,  A... Nos.  1,  2,  3,  4,  supply  exhausted. 

VII,  B...Nos.    1-2,    weight,    12    ounces.      No.    3,    weight,    8    ounces.      No.    4, 

weight,  16  ounces. 
VIII,  A... No.   1,  weight,   9  ounces.     No.   2,  weight,   10   ounces.     No.   3,  weight, 
12  ounces.     No.  4,  weight,  12  ounces. 

VIII,  B...No.  1,  weight,  8  ounces.     Publication  of  Series  B  was  suspended  with 
this  number. 
IX No.  1,  weight,  10  ounces.     No.   2,  weight,  10  ounces.     No.  3,  weight, 

9  ounces.     No.  4,  weight,  12  ounces. 
X Nos.  1,  2,  3,  4,  weight  each,  12  ounces. 

Volume  SCIENCE  bulletin 

I Nos.    1-4,    weight,    8    ounces.      Nos.    5-9,    weight,    12    ounces.      Nos. 

10-12,  weight,  6  ounces. 

II Nos.    1-3,   weight,    20   ounces.      Nos.    4-9,   weight,    11    ounces.      Nos. 

10-15,  weight,  20  ounces. 

Ill Nos.  1-6,  weight,  33  ounces.     Nos.  7-10,  weight,  25  ounces. 

IV Nos.   1-6,  weight,  33  ounces.     Nos.  7-20,  weight,  28  ounces. 

V Nos.  1-11,  weight,  33  oxmces.     Nos.  12-21,  weight,  27  ounces. 

VI No.  1,  weight,  27  ounces.     Nos.  2-7,  weight,  19  ounces. 

VII Nos.  1-17,  we'''-+    50  ounces. 

VIII Nos.  1-10,  wBi  ,    ounces. 

IX Nos.  1-21,  weigi.        ^  ounces. 

X Nos.  1-15,  weight,  i7  ounces. 

XI No.  1,  weight,  20  ounces. 

XII Nos.  1-2,  weight,  19  ounces. 

XIII Pt.  I,  Nos.  1-9,  weight,  12  ounces.     Pt.  II,  Nos.  10-15,  weight,   10 

ounces. 

XIV Nos.  1-21,  weight,  34  ounces. 

XV Nos.  1-6,  weight,  18  ounces. 

XVI Nos.  16,  weight,  14  ounces. 

XVII Pt,  I,  No.  1,  weight,  18  ounces.     Pt.  II,  Nos.  2,  3,  4,  5,  6,  7,  weight, 

8  ounces. 

XVIII Nos.  1-13,  weight,  38  ounces. 

XIX Pt.   I,   Nos.    1-7,    weight,    6    ounces.      Pt.    II,   Nos.    8-14,   weight,    16 

ounces. 

XX Pt.  I,  Nos.   1-6,  weight,   11  ounces.     Pt.  II,  Nos.  7-21,  weight,   15 

ounces. 

The  Kansas  University  Quarterly  and  the  Science  Bulletin  will  be  sent  in  ex- 
change for  other  publications  of  like  character,  or  will  be  sent  on  receipt  of  the 
amount  of  postage  according  to  weight  mentioned  above,  or  may  be  sent  by  ex- 
press, charges  collect.  Separates  of  all  articles  in  the  Science  Bulletin  not  out  of 
print  are  available.  Applications  should  be  made  to  Science  Bulletin,  Library  of 
the  University  of  Kansas. 


BULLETINS   OP    DEPARTMENT    OF    ENTOMOLOGY 

'Two  Grain  Insects."     V.  L.  Kellogg  (with  F.  H.  Snow). 

'Common  Injurious  Insects  of  Kansas."     V.  L.  Kellogg. 

'The  Horn  Fly  of  Cattle."     V.  L.  Kellogg  (with  F.  H.  Snow). 

'The  More  Destructive  Grasshoppers  of  Kansas."     Hunter  and  Snow. 

'Scale  Insects  Injurious  to  Orchards."     S.  J.  Hunter. 

'Alfalfa,  Grasshoppers,  Bees;    Their  Relationships."     S.  J.  Hunter. 

'The  Honey  Bee  and  Its  Food  Plants  in  Kansas."     S.  J.  Hunter. 

'The  Green  Bug  and  Its  Natural  Enemies."     S.  J.  Hunter.  _^ 

'Report  of  Results  of  University  Research  Commission  on  Horse  Plague. 

S.  J.  Hunter,  A.  L.  Skoog,  W.  K.  Trimble,  N.   P.  Sherwood. 
"Orchard  Problems  and  How  to  Solve  Them."     H.  B.  Hungerford. 
"Studies  in  Kansas  Insects."     Bulletin  11. 

1.  Grasshoppers;    Melanopli  of  Kansas.     P.  W.  Claassen. 

2.  Grasshoppers;   CEdipodinse  of  Kansas.     R.  H.  Beamer. 

3.  Dragonflies  of  Kansas.     C.  H.  Kennedy. 

4.  Scale  Insects  Injurious  to  Fruit  and  Shade  Trees.     P.  B.  Lawson. 

5.  Spring  Cankerworm  and  Its  Control.     W.   H.   Wellhouse. 
"Insect  Pests  About  the  House."     H.  B.  Hungerford. 

Application  should  be  made  to  the  State  Entomologist,  University  of  Kansas. 


li 


STATE  GEOLOGICAL  SURVEY  OF  KANSAS 

I,  1896 General  Stratigraphy  of  Eastern  Kansas;   exhausted. 

II,  1897 General  Geology  of  Western  Kansas;   exhausted. 

III,  1898 Special  Report'on  Coal;    exhausted. 

IV,  1898 Upper  Cretaceous  Paleontology;    exhausted. 

V,  1899 Gypsum  and   Gypsum  Cement  Plasters. 

VI,  1900 Carboniferous  Invertebrates,   Cretaceous   Fishes;    exhausted. 

VIl'   1902 Special  Report  on  Mineral  Waters;    exhausted. 

VIII,  1904 Special  Report  on  Lead  and  Zinc. 

IX,   1909 Special  Report  on  Well  Waters  in  Kansas;    exhausted. 

Bulletin    1,   1913 Special  Report  on  Well  Waters  in  Kansas. 

Bulletin    2,  1915.  ..  .Crystalline  Rocks   in  Kansas;    exhausted. 

Bulletin    3,  1917 Oil  and  Gas  Resources  of  Kansas;    exhausted. 

Bulletin    4,  1918.  .  .  .En^aronmient  of  Camp  Funston. 

Bulletin    5,  1918 Elk  City  Gas  Field. 

Bulletin    6,  1918 Oil  and  Gas  Resources  of  Kansas. 

1920 Part  1.    General  Geology  of  Oil  and  Gas. 

1920 Part  2.    Geology  of  Kansas. 

1920.... Part  5.    Allen  and  Neosho  Counties. 
1921.... Part  6.    Wilson  and  Montgomery  Counties. 
1927.  . .  .Part  7.    Anderson  County. 

Bulletin    7,  1921 Geology  of  El  Dorado  Oil  and  Gas  Field. 

Bulletin    8,  1921 Economic  Geology  of  the  Arkansas  C'tv  District. 

Bulletin    9,   1924 Geology   and    Invertebrate    1  logy    of    the    Comanchean 

and  "Dakota"  Formr     -'  ,        Kansas. 
Bulletin  10,  1925.  ..  .Geology   of   Russell   Counr,       *  .asas,    with    Special    Reference 

to  Oil  and  Gas  Resoun      ;    exhausted.  •  ,   -d  r 

Bulletin  11,   1926 Geologic  Investigation  in  Western  Kansas,  with   Special  Ref- 
erence to  Oil  and  Gas  Possibilities;    exhausted. 

Bulletin  12,   1929 Geology  of  Cowley  County. 

Bulletin  13,  1927 ...  .Underground   Resources  of  Kansas;    exhausted. 
Bulletin  14,   1928.  ..  .Volcanic  Ash  Resources  of  Kansas. 

Bulletin  15,  1930 Geology   of  Cloud  and  Republic  Counties. 

Bulletin  16,   1930 Geology  of  Mitchell  and  Osborne  Counties. 

Bulletin  17,  1930 Fauna  of  the  Drum  Limestone  of  Kansas   and  Western 

Missouri. 
Bulletin  18,  1932 Geology  of  Wallace  County,  Kansas. 

MINERAL  RESOURCES  OF   KANSAS 

Report  for  1897,  1898,  1900-'01,  1902;   exhausted. 

Report  for  1899,  1903;   exhausted.  '     •     ,00-7 

Mineral  Resources  Circular  1.    Oil  and  Gas  Resources  of  Kansas  m  19^7. 

Circular  3,  1931.    Diatomaceous  Marl  from  Western  Kansas,  a  Possible  Source  ot 

Hydraulic  Lime.  ,     .     ^       j.      -v 

Circular  4,  1931.    Mineral  Resources  of  Wyandotte  County,  Kansas. 

CONTRIBUTIONS    TO   PALEONTOLOGY 

Grasses  and  Other  Plants  from  the  Tertiary  Rocks  of  Kansas  and  Colorado. 
Pliocene  Diatoms  of  Wallace  County,  Kansas. 

Publications  of  the  State  Geological  Survey  are  distributed  from  the  office  of 
the  State  Geologist,  Lawrence,  Kan.  Apply  to  this  office  for  list  of  mailing 
charges. 


i. 


I|!|{!i||||||!l|{|||l{||l||{l{||{i|ji{i  i||{ll|>|||l|  {IPIIIP 


3  2044  093 


361    939 


Date  Due 


FtlB-^fS- 


LLlAL 


APiH~^-i6Q0