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UNIVERSITY OF KANSAS

SCIENCE BULLETIN

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UNIVERSITY OF KANSAS PUBLICATIONS
University of Kansas Science Bulletin - Vol. XXXVII, Pt. I

October 15, 1955
Lawrence, Kansas

ANNOUNCEMENT

The University of Kansas Science Bulletin (continuation of the
Kansas University Quarterly) is issued in parts at irregular inter-
vals. Each volume contains from 300 to 700 pages of reading mat-
ter, with necessary illustrations. Exchanges with other institutions
and learned societies everywhere are sohcited. All exchanges should
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volumes of the University of Kansas Science Bulletin diflPers so
markedly from the dates bourne on the covers of the publication or
on the covers of the separata that it seems wise to oflFer a corrected
list showing the mailing dates. The editor has been unable to verify
mailing dates earlier than 1932. Separata were issued at the same
time as the whole volume.

Vol.

XX

Vol.

XXI

Vol.

XXII

Vol.

XXIII

Vol.

XXIV

Vol.

XXV

Vol.

XXVI

Vol.

XXVII,

Vol.

XXVIII,

Vol.

XXIX,

Vol.

XXX,

—October 1, 1932. Vol.

—November 27, 1934.
—November 15, 1935. Vol.
—August 15, 1936. Vol.

—February 16, 1938,
-July 10, 1939. Vol.

— No\'ember 27, 1940.

Pt. I— Dec. 30, 1941. Vol.

Pt. I— May 15, 1942.

Pt. II— Nov. 12, 1942.

Pt. I— July 15, 1943. Vol.

Pt. II— Oct. 15, 1943.

Pt. I— June 12, 1944.

Pt. II— June 15, 1945.

XXXI, Pt. I— May 1, 1946.
Pt. II— Nov. 1, 1947.
XXXII— Nov. 25, 1948.

XXXIII, Pt. I— April 20, 1949.
Pt. II— March 20, 1950.

XXXIV, Pt. I— Oct. 1, 1951.
Pt. II— Feb. 15, 1952.

XXXV, Pt. I— July 1, 1952.

Pt. II— Sept. 10, 1953.
Pt. Ill— Nov. 20, 1953.
XXXVI, Pt. I— June 1, 1954.
Pt. II— July 15, 1954.

Editor Edward H. Taylor

Editorial Board.

Edward H. Taylor, Chairman

Charles Michener

Paul Roofe

dwight j. mulford

worthie h. horr

Parke H. Woodard, Secretary

UNIVERSITY OF KANSAS

SCIENCE BULLETIN

DEVOTED TO

THE PUBLICATION OF THE RESULTS OF

RESEARCH BY MEMBERS OF THE

UNIVERSITY OF KANSAS

Volume XXXVII, Part I

University of Kansas Publications

Lawrence, October 15, 1955

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Contents of Volume XXXVII, Part I

PAGE

No.

1. The Effects of Internal Radiation from the Mothers Milk

upon the Spleen of the Young Albino Rat.

Vaid G. Roofe and Walter W. Hair, 3

2. A Study of the Effects of Protein Fats and Internal Irradia-

tion on the Bone Marrow of the Male Albino Rat.

Paul G. Roofe and Mdlard Spencer, 31

3. Taxonomy and Ecology of Kansas Hepaticae.

Ronald L. McGregor, 55

4. A Pollen Profile Study of the Muscotah Marsh.

W.H.Horr, 143

5. The Anatomy and Modes of Preservation of Genus Cardio-

carpiis spinatus Graham Elmer A. Roth, 151

6. The Distribution and Subspeciation of Ercbia epipsodea

Butler (Lepidoptera: Satyridae) Paid R. Ehrlich, 175

7. Some Taxonomic and Morphological Studies of Two Gen-

era of North American Dictyopharidae, ( Fulgoroidea,
Homoptera) Kathleen C. Doering, 195

8. Biological Studies and Taxonomic Notes in the Genus

Strongylogaster Dahlbom. (Hymenoptera: Tenthredin-
idae) Robert E. Beer, 223

9. Tromhicula giirneyi Ewing and Two New Related Chigger

Mites (Acarina, Trombiculidae ) . . . .Richard B. Looniis, 251

10. Comparison of the Effects of Administration of Propyl —

and lodothiouracil to Pregnant Guinea Pigs.

Roy R. Peterson, 269

11. A Taxonomic Revision of the Family Gelastocoridae (He-

miptera) Edward L. Todd, 277

12. Studies of the Agar-Plate Precipitin Test.

Charles A. Leone, A. Byron Leonard and Carlon Pryor, 477

13. Additions to the Known Herpetological Fauna of Costa

Rica with Comments on Other Species. No. II.

Edward H. Taylor, 499

MUS. COMP. ZOOL
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THE UNIVEESITY OP KANSAS

SCIENCE BULLETIN

Vol. XXXVII, Pt. I] October 15, 1955 [No. 1

The Effects of Internal Radiation from the Mothers Milk
upon the Spleen of the Young Albino Rat ^

BY

Paul G. Roofe and Walter W. Hair -

Abstract: Adult female albino rats were injected intraperitoneally with
approximately 30 micrograms of radium chloride. The females and their
litters were sacrificed at specific time intervals from four to fifty-two days
after birth. The spleens of the litter animals were examined quantitatively
en the basis of percentage composition for comparison of constituent cells.
A study of the peripheral blood was also made just preceding death. The
results of these studies have been stated and shown graphically. The following
are the most striking results:

1. The consumption of radium chloride through the milk of the mother by
the young rats causes changes in the splenic tissue of the rats. These
include loss of condensed splenic nodules, and a displacement of cell
function and nuclear organization.

2. The alpha irradiation from radium chloride caused considerable dis-
turbance in the relative quantities of the cellular constituents of the
spleen. These changes include:

( a ) An increase in the number of macrophages and an invasion by these

macrophages into the former splenic nodules;
{b) A breakdown and destruction of erythrocytes which in turn were

phagocytized by the reticuloendothelial cells;

(c) An increase in the amount of granulocytic myelopoiesis with a par-
ticular increase in eosinophile myelocytes;

(d) A decrease in the number of lymphocytes present, varies inversely
with the number of macrophages present;

(e) An increase in the number of megakaryocytes with heavy concen-
tration around the periphery of the organ.

3. The alpha radiation from radium chloride caused peripheral blood
damage which includes:

(a) A lowering of the white blood count below that of "normal."

(b) An increase in the number of normoblasts present in the peripheral
blood.

1. (Work done under contract with the Office of Naval Research. N6 ori 220 T. O. 1)

2. Department of Anatomy, University of Kansas, Lawrence, Kansas.

(3)

4 The University Science Bulletin

4. The effect of the radiation on the weight of the animals was dramatic,
showing a distinct loss in weight gain after the twentieth day when the
rats were weaned.

The effect of small repeated doses of an alpha emitting substance have a
(luantitatively observable effect on the cellular constituents of the spleen of
young albino rats. This deleterious effect seems to be reversible with the
return to normal by various cell constituents in accordance with their radio-
sensitivity. It is apparent that an attempt is being made by the animal to
compensate for cellular damage throughout its body and a fraction of this
compensation is observable in the spleen.

TABLE OF CONTENTS

PAGE

Introduction 5

Methods and Materials 6

Results and Discussion 7

Reticulo-Endothelial Cells 8

Myeloid Cells 9

Lymphoid Cells 10

Other Cell Types 11

Peripheral Blood 12

Body Weight 14

Radium Retention 14

Summary and Conclusion 15

Literature Cited 16

Tables and Graphs 18

INTRODUCTION

Relatively little experimentation has been done involving the
study of radiation damage due to emission of radiation from an in-
ternal source. The literature shows considerable variance in the
work so reported. This is to be expected however, because the
number of variables are many and are difficult to control. In the
study with which this paper deals, a concerted effort was made to
control as many of these variables as possible.

The purpose of this study was, first, to observe the effects of
small repetitive internal doses of radium chloride on the general
adaptation and metabolic processes of the young albino rat; sec-
ondly, to observe the cellular changes in the spleens of these rats,
due to radiation damage; thirdly, to observe the effects of the ra-
dium on the peripheral blood stream of the rats; and fourthly, to
observe how much of the radium is retained in the animal at vari-
ous intervals. There have been many observations concerning the
cell changes in all of the hemopoietic organs but few quantitative
measurements have been made of the cellular damage. There also

Effect of Radiated Milk on Spleen 5

is a noticeable lack of normal quantitative measurements of hemo-
poietic activity in the literature, especially for young animals.

Radium chloride is an alpha emitter and has a penetration
range of 25 microns, therefore the area subjected to radiation
can be limited to the tissues composing and adjacent (25m) to
the alimentary tract and blood stream, since the radium salt was
administered orally. Most of the damage done by alpha particle
emission occurred within the first few hours when the radium
remained in the circulating blood stream; however the half life
of radium is 1,750 years, thus the fixed radium will continue to
emit alpha particles for some time (Martland, 1929). The alpha
particles from radium are ejected with an initial velocity of
1 20 to 1/12 that of light. If a nucleus of a lighter atom is
struck squarely a high velocity hydrogen atom may be given o£F.
This in turn strikes another atom. Such activity is a possible
mechanism of action of the destructive force of an alpha emitter
(Martland, 1929).

An instance was observed in which several radium dial painters
became ill and died, presumably from radium ingested when they
pointed the tips of their brushes with their tongues. The insol-
uble sulphates of the radium were fixed in the colloids and de-
posited in the reticulo-endothelial system of the bone marrow,
spleen and liver (Martland, 1929). Thomas and Brunner (1933),
after Barker, suggests that radium, in the animal system, may fol-
low the course of calcium. Oral administration of radium sulphate
showed a decrease in the mature lymphocytes in the spleen and
replacement by stem cells which subsequently degenerated. The
sinuses were wide and empty and a myeloid metaplasia was evident
(Rosenthal, Maurice, and Grace, 1936). Elimination of the radium
in the first three or four days takes place mainly through the
feces (Seil, Viol, and Gordon, 1915). However, approximately
thirty percent of the administered radium is retained in the bone
matrix (Hoecker and Roofe, 1949). As long as there is radium
in the system of the animal, some radium and radon will be in
the blood and thus the blood cells are constantly being bombarded
by alpha particles. Human beings receiving as much as 890 micro-
grams or more of radium, showed a significant retention of radium
after two years but dosages below this amount showed no reten-
tion or only traces (Barker and Schlundt, 1930).

This problem arose out of an interest in the possibility of trans-
mission of radium through the mammary gland of the female rat.
We observed that by allowing the young to nurse after the RaClj

6 The University Science Bulletin

injection of the mother, repeated low dosages of radium could be
be given internally to the young in this manner. Thus both immedi-
ate and long range effects could be observed without subjecting the
young rats to excessive handling.

METHODS AND MATERIALS

The animals used were female Sprague-Dawley albino rats. The
rats were pregnant when received and bore their litters several days
after arrival. Twenty rats were used, ten of which were control and
ten of which were experimental. Upon birth of the litter all but six
young were removed. On the second and third day after the birth
of a litter in the experimental group, 15 micrograms * of radium
chloride was injected intraperitoneally into the adult female so that
each adult female of this group received a total of 30 micrograms.
Likewise on the second and third day after the birth of a litter in the
control group, the adult female received 1.5cc of physiological saline
(.9%) intraperitoneally. The young were allowed to nurse at will
and a minimum of handling was involved. All litters were weaned
on the twentieth day after birth. On the fourth, eighth, twelfth, six-
teenth, twentieth, twenty-fourth, thirty-first, thirty-eighth, forty-
fifth,** and fifty-second day after birth, one of the adult females
and her litter were sacrificed, both in the control and experimental
groups. The adult female was sacrificed by striking her at the base
of the skull with a blunt instrument. The young were sacrificed by
using ether. Latta and Ehlers (1931) report that ether anesthesia
produced no changes in rats outside physiological limits. Before the
animals were sacrificed each one was weighed and counts were
made on the peripheral blood. Also an attempt was made to obtain
blood directly from the heart by puncturing through the thorax with
a needle. After the preliminary tests were made by standard labora-
tory procedures and the animal was sacrificed, the spleens were
rapidly removed and placed in zenker-formol fixative, embedded in
paraffin and sectioned at eight microns. The sections were stained
with hematoxylin-eosin stain and hematoxylin-eosin-azure stain. Af-
ter removal of the hemopoietic tissues from the animal, it was sealed
in a jar for thirty days to allow the radium-radon equilibrium to be
established (Cameron and Viol, 1915). Each litter was sealed sepa-
rately in order to measure radium retention. The radiation counts
were made with a "64-Scaler" using standard procedure.

" Due to some absorption of the radium on the side of the vial, exactly 15 micrograms
was not given but an approximate thereof.

"" The 4.5-day experimental animals were sacrificed on the forty-second day because it
was becoming obvious that they would not live until the forty-fifth day. Thus it was of
particular interest that they be used in the study.

Effect of Radiated Milk on Spleen 7

Two animals were picked at random from the litter of six and dif-
ferential cell counts of the spleen were made using a modified Chalk-
ley ( 1942 ) procedure. Five hundred cells were counted in each
tissue and the results of these counts were converted to percentage.
Oil immersion was used for the cell counts with a magnification of
950 diameters. The counts were made without regard to the type of
field since in most of the experimental group no distinction could be
made between red pulp and white pulp. Not all of the intermediate
cell types could be definitely distinguished, consequently several
larger categories were counted which included intermediate cell
stages. The following is a brief description of each category:

( 1 ) Reticular Cell — Is an elongated cell (8 to 14 microns ) with a very
clear cytoplasm. The nucleus is practically indistinguishable except
in outline. The width of the cell ranges from four to eight microns.

(2) Macrophage — Is a cell of the size of a large lymphocyte, however it
contains fragments of eosinophiHc staining debris. Any chromatin, if
present, is also fragmented. The cytoplasm stains a light blue.

(3) Neutrophilic Myelocyte — Like the eosinophile myelocyte, the neutro-
phile myelocyte category includes slight variations just before or after
the myelocyte stage. The cell contains neutrophilic staining (wine
colored) granules. The nucleus is large and undivided. The cell
ranges from 10 to 14 microns in diameter.

(4) Eosinophile Myelocyte — All cells of the granulocyte series were imma-
ture. Consequently the cells in this category include shght variations
just before and after the myelocyte stage. The cell contains large
eosinophilic staining granules. The nucleus is large and undivided.
The cell ranges from 10 to 14 microns in diameter.

(5) Large Lymphocyte — Contains a dark staining nucleus which generally
fills most of the cell. Frequently the chromatin was in the process of
fragmentation. The cytoplasm stains a light blue. The cell ranges
from 10 to 12 microns in diameter. Earlier members of the lympho-
cytic series could not be definitely detennined so they were included
in the category of large lymphocytes.

(6) Small Lymphocyte — Contains a small dark staining nucleus which com-
pletely fills the cell. The cell ranges from six to eight microns in
diameter.

(7) Megakaryocyte — Is a very large cell containing a polymorphous
nucleus. It usually stains lightly and has irregular shapes. The cell
ranges from 40 to 120 microns in diameter.

Plasma cells, monocytes, hemocytoblasts, and basophiles were found in such
low frequency that they were considered insignificant, consequently they were
not included in the results. The erythroid series is explained later in the paper.

RESULTS AND DISCUSSION

The general gross appearance of the spleens from the control and
experimental groups did not differ markedly. This is to be expected
since the dosage received by the young animals was less than the

8 The University Science Bulletin

lethal range. Dunlop (1942) and Kolodny, (1925) report that the
spleen undergoes brief early swelling and then shrinks rapidly. In
some cases the spleen enlarged after massive exposure, according to
(Thomas and Brunner, 1933). The younger experimental animals
showed a definite contrast to the controls of the same period. They
exhibited no nodules or a few with light staining cells. There was
an obvious increase in eosinophile myelocytes in the experimental
group which were found in clusters. Only a few were seen in the
controls. The megakaryocytes were apparently increased in num-
ber in the experimental group and they were seen more frequently
toward the pheriphery of the organ. The actual cell counts show
a much greater contrast in cell types than is apparent upon inspec-
tion at low power under the microscope.

The following are the results and a discussion of a comparative
study of the various cell types.

Reticulo-Endothelial Cells

The reticulo-endothelial cells include those which are phagocytic
in nature. These are reticular cells and macrophages.

Reticular Cells — Because the reticular cells stain lightly they are
frequently unobserved and are covered by fragments of red cells
and macrophages. The experimental group showed a slight increase
in reticular cells as compared to the control group (see Graph V).
They were generally seen grouped together forming a reticular
framework for the organ. In some cases they were seen to be part
of epithelioid islands as described by ( Bloom, 1948 ) . These "free"
reticular cells apparently possessed the property of being able to
phagocytose debris. The "fixed" reticular cells became more con-
densed in the experimental group, which is similar to the work
reported by (Brown, 1954). Hsii and Ma (1940) observed these
phagocytes being transformed into erythrocytes, however no such
transformation was seen here.

Macrophages — The macrophages were seen very soon to invade
the splenic nodules of the experimental animal and were observed
in abundance from the fourth day to completion of the experiment.
In the control grovip the percent of macrophages was highly erratic,
varying from 32% in the four-day animals to 71.8% in the 52-day
animals. This is possibly a reflection of the cell maturation process,
inasmuch as there was a corresponding decrease in lymphocytes.
The experimental group also showed erratic variation but was
generally higher than the control (see Graph XIII). The 42-

Effect of Radiated Milk on Spleen 9

day animals, in particular, showed a high macrophage count, which
is particularly interesting since these animals had to be sacrificed
three days earlier than planned because they were obviously not
going to survive. The macrophages contained nuclear and cellular
debris, a great deal of which appeared to be remnants of erythro-
cytes. The macrophages composed the greater part of the epithe-
lioid islands mentioned above and seemed to be most prevalent
when the animal was having its most severe reaction to the radia-
tion. The 52-day animals showed some return of the normal
constituents of the splenic nodules and a reduction in the number
of macrophages. As can be seen by the accompanying graphs
( see Graph X and XI ) , the number of macrophages varies inversely
with the number of large lymphocytes and this inverse ratio be-
tween the two is almost constant. It is reasonable to assume from
this that the latter is the precursor of the former. Thomas and
Brunner (1933) report gross loss of Malpighian corpuscles and
microscopically a great decrease in the prominence of these struc-
tures. Rosenthal (1936) describes the increase in macrophages as
a hyperplasia of the stem cells. Latta and Ehlers (1931) report
an increase in macrophages which are laden with pigment. Bloom
( 1948 ) however, declares no reticulo-endothelial hyperplasia. Thus
it is seen that there is certainly no agreement on how the debris is
phagocytized or how these reticulo-endothelial cells are formed.
This investigation tends to affirm that lymphocytes are precursors
of macrophages.

Myeloid Cells

Erythroid cells, neutrophiles, eosinophiles and basophiles are
considered in this section. In the normal spleen these cells are
found in the red pulp but in the irradiated animals their location
was inconstant.

Erythroid Cells — Comparative counts of the erythroid series
were not attempted here since the animals were subjected to
trauma just previous to being sacrificed in an attempt to obtain
heart blood directly for a comparative study with the peripheral
blood. Such trauma would cause a dilatation and contraction of
the splenic sinuses and thus render a count of the erythroid series
inaccurate. This hypothesis was verified upon low power in-
spection of many of the slides.

Neutrophiles — In both the neutrophile and eosinophile series,
only the myelocyte stage was observed. In no instance was a
mature granulocyte seen. Whether these cells mature to the

10 The University Science Bulletin

myelocyte stage and then defer their further maturation until they
get into the circulation is an item for speculation but one about
which no conclusive statements can be made here. In the adult
ablino rat there is no myelopoiesis in the spleen and the small
amount seen here in the control group is attributed to the fact that
these are young developing rats. Both the neutrophiles and
eosinophiles approach zero toward the fifty-second day. The
neutrophile myelocytes in the experimental group follow the con-
trol group very closely until the twenty-fourth day, at which
time they remain somewhat elevated above the control group. On
the twentieth day the animals were weaned and thus the source of
radiation from the mothers milk was terminated. Thus the experi-
mental group is shown in Graph VII dropping below the control
about four days after weaning. As will be seen below however,
this same thing was not manifested by the eosinophiles. Neverthe-
less the experimental group does show an increase in neutrophilic
myelocytes. Thomas and Brunner (1933), (Latta and Ehlers,
1931) and (Rosenthal, 1936) report extra-medullary myelopoiesis
in the spleen. Bloom ( 1948 ) and ( Sabin and Doan, 1932 ) , how-
ever, report comparatively little increase in ectopic myelopoiesis.

Eosinopliiles — The eosinophile myelocytes of the experimental
animals were consistently more numerous than those of the control
animals (see Graph VIII). They were also many times more
numerous than the neutrophile myelocytes. They were found in
clusters in almost all cases. The 52-day experimental group
showed a sharp rise in eosinophile myelocytes. Minot and Spur-
ling (1924) reported an increase in eosinophiles after repeated
short treatments with X-rays.

Basophiles — The basophiles were in such low incidence that it was
thought that no significant data would be reflected in a comparative
count.

Lymphoid Cells

Lymphoid cells include large, medium and small lymphocytes,
along with monocytes which have a dubious origin but which are
placed in this category arbitrarily. The large and medium lympho-
cytes have been grouped under one heading.

Large Lymphocijtes— These cells, as was mentioned previously,
varied in an inverse proportion with the macrophages. Like the
macrophages, their percentage of frequency was highly erratic
both in the experimental and control group. Their incidence was
generally lower in the experimental group than in the control

Effect of Radiated Milk ox Spleen 11

group, and especially, the 42-day experimental animals showed a
very low percentage of lymphocytes ( see Graph XII ) . The lympho-
cytes of the control group were concentrated in the center of the
splenic nodules but those of the experimental group showed no
specific location. In the 52-day experimental animals some return
of the lymphocytes to the splenic nodules was observed. In the
experimental group many of the lymphocytes exhibited a pycnotic
nucleus and many were in the process of shattering. Bloom ( 1948 )
and (Thomas, 1933) reported similar results. The latter also
reported immature lymphocytes present in greater numbers than
normal.

Spurling and Lawrence ( 1925 ) have shown that leucocytes are
not affected directly, in the blood stream, within an hour and a
quarter after irradiation, but most of the degenerated forms are
due to an indirect eflFect.

Small Lijjnphocijtes — The number of small lymphocytes did not
change significantly in either the control or the experimental group,
except in the very young animals (see Graph VI). They were,
in fact, relatively infrequent. Warren, MacMillan and Dixon
(1950) using a beta emitter on mice, report a decrease in the
number of small lymphocytes.

Monocytes — It was considered impractical to attempt to distin-
guish between large lymphocytes and monocytes, so a discussion of
this group will not be included in this paper.

Other Cell Types

Megakaryocytes — The number of megakaryocytes in the experi-
mental group from the eighth day to sacrifice were quite signifi-
cantly higher than those of the control group. The control group
by the thirty-eighth day showed practically no incidence, while the
experimental group showed a variable but high incidence of mega-
karyocytes (see Graph IX). Bloom (1948) reports degeneration
and disappearance of the megakaryocytes. Thomas and Brunner
(1933) report an increase in megakaryocytes, in some cases, being
very numerous.

Kolodny (1925) after experimental deep roentgen irradiation,
found an increase in blood coagulability and postulated that due
to destruction of splenic follicles, thrombokinetie and thrombo-
plastic substances were released. While attempting to get peri-
pheral blood from the tail of the rats in our experimental group,
an increase in coagulability was also noticed. Since the theory for
origin of platelets has found considerable proof recently (Russell

12 The University Science Bulletin

and Roofe, 1954) the increase in megakaryocytes could account for
this increase in coagulability; however, most of the platelets are fur-
nished by the megakaryocytes of the bone marrow. The platelet
count is extremely variable and does not correspond to the changes
in megakaryocytes in the spleen. Dunlop ( 1942 ) reported a short-
ening of coagulation time but does not attempt an explanation.

Plasma Cells — Thomas and Brunner (1933) found many lymph-
ocytes converted to plasma cells, which are common reactive cells
in certain chronic diseases. None were found however in either the
control or experimental groups here.

Peripheral Blood

A study of the peripheral blood was made on each animal before
it was sacrificed. This study includes a white cell count, red cell
count, platelet count and a differential count.

White Blood Cell Count — The white cell count of the control and
experimental group remained approximately the same until the
twelfth day, at which time the experimental group showed a definite
decrease in white cells, (see Graph I). The decrement becomes
greater toward termination of the study. This decrease does not
correlate with the white cells in the spleen. Dunlop (1942) after
Liechti, reported an incomplete summation of divided doses, and
less blood damage resulting than if a total dose had been given at
a single time (after Gloor and Suppinger). He also states that
young subjects are more susceptible to radiation than normal adults
( after Heim and Steton ) . From this we can assume that the summa-
tion of dosages tends to produce less effect than had the dose been
given at a single time; however, due to the youth of the animals more
peripheral blood damage should be apparent than an adult would
show. Murphy and Nakahara ( 1920 ) report small doses of X-rays
stimulate lymphocyte production. However, (Mottram and Russ,
1921) claim that this lymphocytosis was a compensatory reaction
due to an initial lymphopenia immediately following irradiation.
Lawrence ( 1948 ) reports direct irradiation of hemopietic tissue pro-
duces a leukopenia but radiation in other parts of the body does not
have a similar effect. Hsii and Ma (1940) report lymphopenia and
polymorphonuclear leukopenia as constant findings.

In normal adult albino rats, Downey (1938) reports an average
leucocyte count of 11,000 with a range from 5,800 to 29,800. The
normal animals in our work from four to fifty-two days in age,
showed a range in leucocyte count of 2,025 to 25,650 (see table I).

Effect of Radiated Milk on Spleen 13

The leucocyte count started low (4,400) in the four-day animals
and rose steadily but showed no definite leveling off stage within
the 52-day period ( see Graph I ) .

Kindred ( 1937, 1940 ) reports most of the lymphocytes of normal
young animals (15 to 20 days old) are small lymphocytes whereas
the lymphocytes of older animals ( 80 days old ) are mostly medium
lymphocytes. The normal animals in our study show a very small
percentage of small lymphocytes in the 15- to 20-day-old animals,
with most of the lymphocytes being large ones (see table II).
As the age of the animal increases this predomination of large
lymphocytes remains.

Red Blood Cell Count — The red cell count in the experimental
group remained approximately the same as the control group
throughout (see Graph II). The experimental animals showed a
slight erythropenia but followed the same contour as the control
group. From this work and the observations of other workers, the
theory that the erythrocytes are more resistant to radiation than the
lymphocytes or granulocytes, seems to be substantiated. Hsii and
Ma (1940) report the erythrocytes to be definitely affected and
report an increase in erythropoiesis in the spleen. As was stated
above, in the experimental group it was impossible to accurately
determine the amount of erythropoiesis because of the engulfing
of an abnormal number of erythrocytes by macrophages. Kolodny
(1925), (Hsii and Ma, 1940) also report similar observations.
Brown ( 1954 ) reports an abnormal number of normoblasts in
the peripheral blood. Our control group also showed normoblasts
in the peripheral blood but less abundantly than in the experi-
mental group and they did not remain as long (see Graph IV).

Downey (1938) reports for the adult albino rat an average red
cell count of 6,180,000 with a range from 5,800,000 to 11,200,000.
Our control animals had a range of 2,760,000 to 9,375,000, showing
a steady climb from the four-day animals to the 52-day animals
(see Graph II). The red cell count showed a tendency to level off
at approximately 6,000,000 by the twenty-fourth day, however
later this count varied. The normoblasts of the control group were
as high as 21 per 100 leucocytes but had practically disappeared by
the twenty-fourth day (see Graph IV), Those of the experimental
group however rose as high as 63 per 100 leucocytes and were in
evidence as long as the forty-second day.

Platelet Count — The platelet count of both the control and ex-
perimental group was highly variable (see Graph III). Justifica-

14 The University Science Bulletin

tion for the variability in platelet count in these animals does
not seem to be obvious. The rise and fall of platelets does not
correlate with the megakaryocyte curve or the change in coagu-
lation time.

Downey (1938) reports a range in platelets from 121,000 to
460,000 for the adult albino rat. Some give an average as high
as 1,275,000 (after Downey). The range in our control animals
showed a total variation from 369,000 to 1,075,000, from a four-
day to a 52-day period (see Table I ).

Differential Count — The differential count of the experimental
group is similar to that of the control group throughout the 52-day
period ( see table I ) . The only difference seen is a slight prevalence
of immature cells in the experimental group. This is possibly a
reflection of the radiation damage on the centers of hemopoiesis.

Body Weight

The weight of the experimental animals held very close to the
weight of the control group until the twentieth day, at which time
the animals were weaned. From this time on the control animals
continued to gain steadily, whereas the experimental animals
remained about the same weight or gained very slightly ( see Graph
XV). Rosenthal (1936) reports a market loss in weight and patho-
logical fractures. Also by oral administration of radium sulfate,
gingivitis, alopecia, abrasion of teeth and necrosis of bone were
demonstrated by Rosenthal. We have similar observations, with
a tendency of the bones to fracture.

Radium Retention

The percentage of radium consumed by the young rats is im-
possible to ascertain since the actual amount of radium being ob-
tained through the milk of the mother is unknown and the frequency
which they nursed was uncontrollable. The number of micro-
grams retained by the whole litter of six was measured however.
The amount of radium retained by the litter decreases steadily
from the eight-day animals to the 52-day animals, with two notable
exceptions (see Graph XIV). The explanation for these incon-
sistencies is unknown. Seil, Viol and Gordon (1915) report that
60% to 98% of the radium taken into the human system is eliminated
principally in the feces (90%) and urine (10%). When taken by
mouth 65% to 75% is eliminated in the first four days and by the tenth
day the daily elimination rate is less than 1%. Thereafter there is a
gradual diminution in its removal so that after several years the

Effect of Radiated Milk on Spleen 15

rate of elimination is .002% to .005% per day. Since the mothers of
our animals were given approximately 30 micrograms intraperi-
toneally and the young were receiving their source of internal radia-
tion from the milk of the mother, it is seen that a number of possible
exits for the radium is available. Yet the retention data of the
young animals shows a total variation of 5.4 micrograms to 12.8
micrograms retained by the litter (see Graph XIV). Thus the
litter retained from Vg to more than Vs of the total amount of radium
chloride received by the mother, showing that a relatively high
percentage of the total amount of radium chloride injected was
passing thru the milk to the young. Hoecker and Roofe (1949)
report a retention of about 30% in the animal and most of this is
retained in the bone ash; however the radium is eliminated from
the blood stream in a very few minutes. The above data reported
by Seil, Viol and Gordon did not include data on lactating females.
Seil, Viol and Gordon (1915), after Ketron and Bellingham-Smith,
report practically no radium being eliminated through the skin,
which is contradictory to the results of our study if the mammary
gland is considered a modified sebaceus gland. A number of
methods have been attempted to increase the rate of excretion of
radium with some success (Thomas and Brunner, 1933) but all
seem to have unwanted side reactions.

Summary and Conclusion

Adult female albino rats were injected intraperitoneally with ap-
proximately 30 micrograms of radium chloride. The females and
their litters were sacrificed at specific time intervals from 4 to 52 days
after birth. The spleens of the litter animals were examined quan-
titatively on the basis of percentage composition for comparison of
constituent cells. A study of the peripheral blood was also made just
preceding death. The results of these studies have been stated and
shown graphically. The following are the most striking results:

1. The consumption of radium chloride through the milk of the mother by
the young rats causes changes in the splenic tissue of the rats. These in-
clude loss of condensed splenic nodules, and a displacement of cell func-
tion and nuclear organization.

2. The alpha irradiation from radium chloride caused considerable disturb-
ance in the relative quantities of the cellular constituents of the spleen.
These changes include:

( a ) An increase in the number of macrophages and an invasion by these

macrophages into the fonner splenic nodules;
(h) A breakdown and destruction of er>'throc>tes which in turn were

phagocytized by the reticulo-endothelial cells;

16 The University Science Bulletin

(c) An increase in the amount of granulocytic myelopoiesis wath a par-
ticular increase in eosinophile myelocytes;

(d) A decrease in the number of lymphocytes present, varies inversely
with the number of macrophages present;

(e) An increase in the number of megakaryocytes with heavy concentra-
tion around the periphery of the organ.

3. The alpha radiation from radium chloride caused peripheral blood dam-
age which includes:

(a) A lowering of the white blood count below that of "normal."

(b) An increase in the number of normoblasts present in the peripheral
blood.

4. The effect of the radiation on the weight of the animals was dramatic,
showing a distinct loss in weight gain after the twentieth day when the
rats were weaned.

The effect of small repeated doses of an alpha emitting substance
have a quantitatively observable effect on the cellular constituents of
the spleen of young albino rats. This deleterious effect seems to be
reversible with the return to normal by various cell constituents in
accordance with their radiosensitivity. It is apparent that an at-
tempt is being made by the animal to compensate for cellular dam-
age throughout its body and a fraction of this compensation is ob-
servable in the spleen.

LITERATURE CITED

Barker, Howard and Herman Schluntdt

1930. The Detection, Estimation and Elimination of Radium in Living
Persons Given Radium Chloride Internally. H American Journal
of Roentgenology and Radium Therapy, vol. 24, pp. 418-423.
Bloom, W.

1948. Histopathology of Irradiation from External and Internal Sources.
National Nuclear Energy Series, Division IV, vol. 22 I, New York,
McGraw Hill Book Company, Inc., pp. 21-22, 243-347.
Brown, J. W.

1954. The Effect of Protein Deficiency on the Submaxillary Lymph
Nodes and Spleens of Albino Rats, University of Kansas Science
Bulletin, vol. 36, pp. 41-54.
Cameron, W. H. and C. H. Viol

1915. Classification of Various Methods Employed in the Internal Admin-
istration of Radium Emanation and Radium Salts. Radium, vol.
4, pp. 66-77.
Chalkley, H. W.

1942. Method for the Quantitative Morphologic Analysis of Tissues.
Journal of National Cancer Institute, vol. 4, pp. 47-53.
Downey, Hal

1938. Handbook of Hematology, vol. II, Hoeber, Inc., New York, N. Y.
pp. 813-820.

Effect of Radiated Milk on Spleen 17

DUNLAP, C. E.

1942. Effects of Radiation on the Blood and the Hemopoietic Tissues

Including the Spleen, Thymus and Lymph Nodes. Archives of

Pathology, vol. 34, pp. 562-608.
Hsu, C. L. and W. C. Ma

1940. Direct and Indirect Effects of Roentgen Radiation on the Blood

Forming Organs of Rats. American Journal of Cancer, vol. 39,

pp. 319-333.
Kindred, J. E.

1938. A Quantitative Study of the Lymphoid Organs of the Albino Rat.

American Journal of Anatomy, vol. 62, pp. 453-473.
1940. A Quantitative Study of the Hemopoietic Organs of Young Albino

Rats. American Journal of Anatomy, vol. 67, pp. 99-149.

KOLODNY, A.

1925. Tissue Changes After Experimental Deep Roentgen Irradiation.
American Journal of Pathology, vol. 1, pp. 285-293.
Latta, J. S. and O. C. Ehlers

1931. Effects on Blood and Hemopoietic Organs of Repeated Short Ex-
posures to X-Ray. American Journal of Anatomy, vol. 47,
pp. 447-474.
Lawt^ence, J. S., M. C. Valentine and A. H. Dowdy

1948. The EfiFect of Radiation on Hemopoiesis. Is There an Indirect
Effect? Blood, vol. 3, pp. 593-611.

Martland, H. S.

1929. Occupational Poisoning in Manufacture of Luminous Watch Dials.
Journal of American Medical Association, vol. 92, pp. 552-554.
Minot, G. R. and R. G. Spurling

1924. The Effect on the Blood of Irradiation, Especially Short Wave
Length Roentgen-Ray Therapy. American Journal of Medical
Sciences, vol. 168,^pp. 215-241.
MoTTRAM, J. C. and S. Russ

1921. Lymphopenia Following Exposures of Rats to Soft X-Rays and
B-Rays of Radium. Journal of Experimental Medicine, vol. 34,
p. 271,
Murphy, J. and W. Nakahara

1920. Studies on X-Ray Effects. Effects of Small Doses of X-Rays of
Low Penetration on the Lymphoid Tissue of Mice. Journal of
Experimental Medicine, vol. 31, pp. 13-18.
Hoecker, F. E., and P. G. Roofe

1949. Structural Differences in Bone Matrix Associated with Metabolized
Radium. Radiology, vol. 52, pp. 856-864.

Rosenthal, Maurice and J. E. Grace

1936. Bone Marrow and Lymph Node Changes in Rabbits, Produced by
Oral Administration of Radium Sulfate. American Journal of
Medical Science, vol. 191: 607-.
Russell, M. and P. G. Roofe

An Experimental Approach to the Origin of Blood Platelets.

Unpublished manuscript

18 The University Science Bulletin

Sabin, F. R., C. a. Doan and C. E. Forkner

1932. The Production of Osteogenic Sarcomata and the Effects on
Lymph Nodes and Bone Marrow of Intravenous Injections of
Radium Chloride and Mesothorium in Rabbits. Journal of Experi-
mental Medicine, vol. 56, p. 267.

Seil, H. a., C. H. Viol and M. A. Gordon

1915. Elimination of Soluble Radium Salts Taken Intravenously and Per
Os. Radium, vol. 5, pp. 40-44.
Spurling, Roy G. and J. S. Lawrence

1925. Direct Effect of Radium Irradiation on Leucocytes. American
Journal of Medical Sciences, vol. 169, p. 157.
Thomas, H. E. and F. H. Brunner

1933. Clironic Radium Poisoning in Rats. American Journal of Roent-
genology, vol. 29, p. 641.

Warren, Shields, J. C. MacMillan and F. J. Dixon

1950. Effects of Internal Radiation of Mice with P32: Spleen, Lymph
Nodes, Thymus, Bone and Bone Marrow. Radiology, vol. 55,
pp. 375-388.

TABLES AND GRAPHS

TABLE I
Peripheral Blood Data

Explanation:

The column headed "Date" refers to the number of days between birth of
the litter and the time of sacrifice.

The column headed "Type" gives reference to either experimental or control
animals.

The succeeding columns are self explanatory and give both the average and
the range. The range is the actual raw data of the two animals used in each
age group. The red counts, white counts and platelet counts are based on cells
per mm-^. The differential and normoblast counts are based on 100 counted
leukocytes.

Those spaces which are blank under "range" indicate that there was no range,
either due to inability to get the count, or due to the absence of that cell type.

Those which are blank under "average" indicate that it was impossible to
get a count.

The weights of the 45- and 52-day control animals were not taken.

* The 45-day experimental animals were sacrificed on the forty-second day
because it was becoming obvious that they would not live until the forty-fifth
day.

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Effect of Radiated Milk on Spleen 21

TABLE II
Spleen Differential Count
Explanation:

The column headed "Date" refers to the number of days between birth of
the litter and the time of sacrifice.

The column headed "Type" gives reference to either experimental or
control animals.

The succeeding columns are self explanatory and give both average and
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indicate that there was no range, either due to inability to get the count, or
due to absence of that cell type.

* The 45-day experimental animals were sacrificed on the forty-second day
because it was becoming obvious that they would not live until the forty-fifth
day.

22

The University Science Bulletin

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Effect of Radiated Milk on Spleen 23

GRAPH I
Peripheral Blood
White Cell Count
Explanation:

"Days" refers to the number of days after birth of the litter.
The ordinate represents the leukocytes in thousand per nim^.
The solid line represents the control group and the broken Une represents
the experimental group.

GRAPH H
Peripheral Blood
Red Cell Count
Explanation:

"Days" refers to the number of days after birth of the litter.
The ordinate represents the erythrocytes in million per mm^.
The solid line represents the control group and the broken line the experi-
mental group.

GRAPH HI
Peripheral Blood
Platelet Count
Explanation:

"Days" refers to the number of days after birth of the litter.
The ordinate represents the platelets in hundred-thousand per mni-*^.
The solid line represents the control group and the broken line the experi-
mental group.

GRAPH IV
Peripheral Blood
Normoblasts
Explanation:

"Days" refers to the number of days after birth of the litter.
The ordinate represents the nonnoblasts per 100 leukocytes.
The soHd line represents the control group and the broken line the experimen-
tal group.

GRAPH V
Spleen-Differential Coltnt ,
Reticular Cells
Explanation:

"Days" refers to the number of days after birth of the litter.
The ordinate represents the percentage of reticular cells.

The solid line represents the control group and the broken line the experimen-
tal group.

24 The University Science Bulletin

GRAPH VI

Spleen-Differential Count

Small Lymphocytes

Explanation:

For explanation of abscissa see above.

The ordinate represents the percentage of small lymphocytes.
The sohd hne represents the control group and the broken line the experimen-
tal group.

GRAPH VII

Spleen-Differential Count

Neutrophile Myelocytes
Explanation:

"Days" refers to the number of days after birth of the litter.

The ordinate represents the percentage of neutrophile myelocytes.

The solid line represents the control group and the broken line the experimen-
tal group.

Effect of Radiated Milk on Spleen

25

GRAPHS I to VII

24 32 40 48 St

OATS

\ 6

Z

o

'6 24 32 40 4« S8

OAyS

70

\

U 4

is:

•

GRAPH VI

6

4
2

0

— -^v^

-^O^C:-^

24 32 40 4«

OAVS

I 8

16 24 32

DArs

40 4S se

26 The University Science Bulletin

GRAPH VIII
Spleen-Differential Count
Eosinophile Myelocytes
Exphmotion:

For explanation of abscissa see above.

The ordinate represents the percentage of eosinophile myelocytes.
The solid line represents the control group and the broken line the experimen-
tal group.

GRAPH IX

Spleen-Differential Count

Megakaryocytes
Explanatian:

"Days" refers to the number of days after birth of the litter.
The ordinate represents the percentage of megakaryocytes.
The solid line represents the control group and the broken line the experimen-
tal group.

GRAPH X

Spleen-Differential Count

Large Lymphocytes and Macrophages (control)
Explanation:

"Days" refers to the number of days after birth of the litter.

The ordinate represents the percentage of large lymphocytes and macro-
phages.

The solid line represents the macrophages of the control group and the
broken line represents the large lymphocytes of the control group.

GRAPH XI

Spleen-Differential Count

Large Lyinphocytes and Macrophages (experimental)
Explanation:

"Days" refers to the number of days after the birth of the litter.

The ordinate represents the percentage of large lymphocytes and macro-
pages.

The solid line represents the macrophages of the experimental group and the
broken line represents the large lymphocytes of the experimental group.

GRAPH XII

Spleen-Differential Count

Large Lymphocytes
Explanation:

"Days" refers to the number of days after the birth of the litter.
The ordinate represents the percentage of large lymphocytes.
The solid line represents the control group and the broken line the experi-
mental group.

Effect of Radiated Milk on Spleen

27

GRAPHS VIII to XII

Y

V

0 8 16 24 32 40 ia i6

DAYS

L, *0

IS 24 32 40 48

0*TS

16 34 32 40 48 S«

0*TS

28 The University Science Bulletin

GRAPH XIII
Spleen-Differential Count

Macrophages
Explanation:

"Days" refers to the number of days after the birth of the litter.

The ordinate represents the percentage of macrophages.

The solid line represents the control group and the broken line the experi-
mental group.

GRAPH XIV
Radium Retention

Explanation:

"Days" refers to the number of days after birth of the litter.

The ordinate represents the number of micrograms of radium retained. The
great deviations in the general trend of the curve is unexplainable. There is
a possibility that there was a leak in the jars in w^hich the animals vv'ere sealed.

GRAPH XV
Body Weight

Explanation:

"Days" refers to the number of days after birth of the litter.

The ordinate represents the weight of the animals in grams.

The solid hne represents the control group and the broken line represents tlie
experimental group. The line represented by long and short dashes indicates
the probable trend of the 45- and 52-day control animals, since that data was
not obtained.

Effect of Radiated Milk on Spleen

29

GRAPHS XIII to XV

THE UNIVERSITY OP KANSAS

SCIENCE BULLETIN

Vol. XXXVII, Pt. I] October 15, 1955 [No. 2

A Study of the Effects of Protein Fast and Internal

Irradiation on the Bone Marrow of the

Male Albino Rat ^

BY

Paul G. Roofe and Millard Spenceb 2

Abstract: Two groups of experimental animals were used in this experi-
ment. The animals in the first group were subjected to protein fast and the ani-
mals in the second group were injected intraperitoneally with radium chloride.
Differential counts were made by a random selection procedure on the bone
marrow taken from these animals. Toluidine blue was used as the stain to de-
termine the effect the protein fast and irradiation had on the nucleic acid con-
tent of the tissue.

1. Cellular basophilia and nucleic acid content is not noticeably affected by
protein fast and radiation under the conditions described in this experiment.

2. Total bone marrow basophiha and nucleic acid content is reduced during
protein fast due chiefly to reduction in cellularity.

3. Fatty displacement of tissue occurs in bone marrow during protein fast.

4. Mitotic activity is decreased by both protein fast and radiation.

5. Changes which take place in the bone marrow during protein fast are
overcome when animals are returned to normal protein diets.

6. The myeloid, erythroid ratio increases during protein fast and following
irradiation.

7. In the bone marrow erythropoietic elements are the most easily affected
by protein fast and internal irradiation, and granulocytopoietic elements are
more susceptible than are the megakaryocytes.

CONTENTS

PAGE

Introduction 32

Materials and Methods 32

Results 34

Weight Results 34

Histological Results 35

Geiger Count Results 37

General and Histochemical Observations 38

Discussion 38

Summary 41

Bibliography 41

1. Sponsored by the Office of Naval Research under contract N6 ori 220 T. O. 1.

2. Department of Anatomy, University of Kansas, Lawrence, Kansas.

(31)

32 The University Science Bulletin

INTRODUCTION

It is well known that protein is one of the most important dietary
constituents, and that adequate amounts of protein are essential
for normal nutrition and body metabolism ( Orten and Orten, 1943;
Wright, 1946). It is also an accepted fact that the protein supplied
in the diet must contain suitable quantities of the essential amino
acids to meet the body needs for protein biosynthesis (Benditt,
1947). Recently experimental evidence has shown that the nucleic
acids play an important role in protein synthesis, but the exact
nature of the nucleic acid action is still unknown. In a review,
Caspersson (1947) has suggested that the nucleic acids are essen-
tial for such self-reproducing units as the gene, and that they are
abundant in cells in which rapid protein synthesis is taking place
either for the production of gene-proteins, or in the production of
the main bulk of cytoplasmic proteins for growth or secretion.

Originally, nucleic acids were thought to be localized solely in
the cell nucleus. It is now an established fact that two kinds of
nucleic acids exist — ribo and desoxyribonucleic acids — and that
RNA is localized in the cytoplasm with DNA localized in the cell
nucleus (Davidson, 1950). The nucleic acids have been identified
and localized by basic staining, enzyme digestion techniques, the
Feulgen reaction, and ultraviolet absorption. Caspersson (1947)
has attempted to use the latter technique as a means of quanti-
tatively determining nucleic acid content of tissues. In a review,
Roofe and Lesher (1951) have pointed out that many of the
results obtained by the absorption method are quantitatively in-
valid, but the qualitative information is valuable.

Since the beginning of the atomic era investigators have become
curious as to the effect the products of atomic fission will have on
living tissue. A great deal of time and effort has been and is being
devoted to determine some of the effects. This study was made to
obtain more data on the histological changes in the bone marrow
which occur during protein fast and after internal irradiation.
A qualitative histochemical procedure was also employed to deter-
mine what effect these conditions would have upon the nucleic
acid content of the bone marrow.

MATERIALS AND METHODS

Male Sprague-Dawley rats served as the experimental animals.
Those rats used in the radiation experiment were mature animals
while those used in the protein experiment were fifty days old
when placed on their respective diets. For the duration of the

Irradiation on Bone Marrow 33

experiment all animals were confined in individual cages in a room
where the temperature was maintained at approximately 70 F.

In the protein deficiency experiment the following dietary groups
were formed:

Group A — Purina fox cliow diet

Group B — Synthetic protein diet

Group C — Synthetic 0.0% protein diet

Group R — 0.0% protein for 35 days and fox chow from days 35-56.

Animals in Group C served as experimental animals. Those in
Groups A and B served as control animals, and the animals in Group
R served as experimental recovery animals. The constituents of the
diets for Groups B and C are shown in Table I. All animals were
maintained on their respective diets until a 14-day interval had
elapsed, when representatives from each group were sacrificed.
Others were then sacrificed at one week intervals. Animals referred
to as R (recovery) were placed on the 0.0% casein diet on the same
date as those in Group C. This feeding was continued for 35 days
and at that time they were placed on the Purina fox chow diet.
After a two week interval, animals in Group IR were sacrificed as
were those in Group 2R the following week.

In the radiation experiment, 14 animals were used: Seven experi-
mental and seven control. These animals were designated as Ral-
Ra7 and N1-N7 respectively. All animals were fed Purina fox chow.
Group Ra animals were injected intraperitoneally with 10 micro-
grams of radium chloride. Geiger counts were made on these ani-
mals on the injection date and at one-week intervals for the duration
of the experiment. The data obtained from these counts is shown in
Graph VIII. Animal Ral was sacrificed seven days after the injec-
tion date and the remainder were sacrificed individually at seven-
day intervals. One normal (control) animal was sacrificed with
each irradiated animal.

The animals used in both parts of this experiment were killed by
a blow on the head. Immediately after killing, the right femur was
removed and cracked open with a pair of forceps. A sample of bone
marrow throughout the shaft was then lifted out and placed into the
fixative. The following tissue technique was used:

Fixation Carnoy's 60 min.

Washing 95% alcohol 3-20 min. each

Clearing amyl acetate 23 hours

Imbedding 50-52 C paraffin 60 min.

Sectioning 7 niicra

Hydration xylol — H ^ O

2—6730

34 The University Science Bulletin

Staining toluidine blue 20 min.

Differentiation .... tertiary butyl alcohol 23 hours

Dehydration U ^ O — xylol

A 1% solution of toluidine blue, a thiazine dye, was used as a tissue
stain. Toluidine blue combines with basophilic substances such as
half-esters of sulfuric acid with colloidal carbohydrate of high mole-
cular weight, which may or may not be in combination with protein.
The absorption of the dyestuff cation by nucleic acids takes place by
means of the phosphoric acid group of the acids. It is believed that
treatment of the tissues with alcohol after toluidine blue staining re-
moves only the excess dye (Michaelis, 1948). Tertiary butyl alcohol
was used in destaining, for it was found to preserve the brilliance of
the dye better than ethyl alcohol.

Maximow's (1950) criteria of cell types was used for the differ-
ential counts in the experiment. All cells, both mature and imma-
ture, were classified by the term Maximow has given the adult
form in their respective series. The method for counting was
patterned after Chalkley's (1943) sampling method. This modi-
fication of his technique consisted of arranging four pointers in
the ocular of the microscope and then identifying and counting
the cell types lying under the tips of the pointers. The count
was made by random selection of a number of fields and was
terminated when the erythrocyte count had reached 100. By using
this method, data for the differential was obtained as a ratio; for
example, 60 neutrophils per 100 erythrocytes. In addition to this
technique, a method for determining changes in total cell popula-
tion was devised. This consisted in counting the number of cells
in an area of 2,500 square microns with the aid of a camera-lucida.
The average of the count made by random selection of 10 fields
was used to express the trend of change in the total cell population.

RESULTS

Weight Results

The dietary and experimental conditions for the various groups
have been mentioned in a previous section of this paper. The
weight variation data will be found in Tables II to V. These tables
give both absolute and percentage weight changes for the indi-
vidual animals used.

Table II contains the weight records for those animals placed
in Groups A and B. As shown, both groups of animals behaved
similarly as to growth changes. Both diets seemed to satisfy the

Irradiation on Bone Marrow 35

animals. The only factor which varied with these two groups was
the greater water consumption by those animals in Group A. This
can be explained by the dryer consistency of their food.

Table III contains the weight data for those animals on the syn-
thetic O.Ofo protein diet. These animals showed a marked drop in
weight by the fourth day of the experiment. As can be seen, the
weight loss was progressive until about the thirty-fifth day of
protein fast, at which time there was a leveling off. Such a trend
also appeared in the cytological changes. These animals were re-
tarded in all aspects of growth. Their bones remained small, their
external genitalia did not develop, and their fur became yellow
and shabby. Toward the latter part of the experiment, they be-
came extremely nervous and would nibble on their tails. These
animals consumed extremely small quantities of water and wasted
much of the food placed in their cages.

Table IV contains the weight data for the recovery groups (R).
These animals were maintained on the 0.0% protein diet for 35 days
and then placed on the recovery Purina fox chow diet. As shown,
these animals made a marked weight recovery. Their cytological
recovery was also very pronounced.

Table V contains weight data for animals in the radiation ex-
periment. The irradiated animals showed a progressive weight
loss throughout the experiment, while their controls showed a
slight increase in weight. Since all of the animals used were
mature, no great weight changes were anticipated in the control
group. No marked change in food or water consumption was
noted in the irradiated group, which suggests that the radium
affected the utilization of ingested food. This change was probably
due to actual tissue damage of the intestine ( Bloom, 1948 ) .

Histological Results

Tables VI to VIII contain the data on the differential counts
made of tissues taken from protein deficient, irradiated, and con-
trol animals. These data are presented graphically on Graphs I
to VII. Ordinates of the graphs show the number of cells per 100
erythrocytes, and the abscissae show time.

From Table VI it can be seen that the animals on the synthetic
protein and fox chow diet presented a very similar cytological
picture. As the animals became older, the neutrophil/erythroid
ratio increased slightly, and the activity of the bone marrow de-
creased. This decrease in activity was evidenced by the appear-
ance of a greater number of mature cells in the tissue and by the

36 The University Science Bulletin

development of more fat vacuoles. Such a decrease in activity is
noted by Maximow and Bloom (1950).

Table VII contains the data on both protein deficient and re-
covery animals. Group C, or protein deficient animals, show very
marked changes in hemocytoblast, megakaryocyte, and neutrophil
rations. The hemocytoblast level was maintained fairly well
through Group IC, but a definite change from normal occurred
within the next week. This decline in numbers increased through
Group 4C. The megakaryocyte picture not only shows a decisive
change in Group IC, but their numbers continue to increase on
through Group 4G. The lymphocyte ratio increased slightly through-
out the experiment. The neutrophil ratio also shows a marked in-
crease, though not so dramatic as the megakaryocyte change.
Another decisive change occurred in the marrow as indicated by
the absolute drop in cell numbers. This change, though pro-
nounced by two weeks on the deficient diet, was continuous
throughout the series. In these latter stages of the experiment,
approximately 50% of the previously active tissue had been replaced
by fat.

Table VII also contains the data on differential counts taken from
the bone marrow of the recovery animals. These animals are
actually a continuation of Group C, as has been previously stated.
This group showed remarkable recovery after being placed on a
diet with adequate protein. The impetus to recovery can be
recognized by the tremendous increase in the absolute number of
cells, and the increase in hemocytoblast numbers. There was
also a progressive increase in erythrocyte population. By the time
Group 2R had been sacrificed close examination was required to
distinguish the tissues from normal.

Table VIII contains the data obtained from differential counts
made on the marrow taken from the irradiated animals. Part of the
data obtained from this group is presented in Graphs V to VIII.
The number of cells per given area was observed in fat deposition
in the marrow. As in the protein deficient animals, there was a
marked increase in the megakaryocyte, lymphocyte, and neutro-
phil ratios. The hemocytoblast ratio first increased and then
declined to normal values in the animals of longer duration. Their
controls presented a cytological picture similar to the older control
animals in the protein experiment.

Irradiation on Bone Marrow 37

Geiger Count Results

Graph VIII contains the data obtained from a series of Geiger
counts made on the hve animals which had previously been in-
jected intraperitoneally with 10 micrograms of radium chloride.
According to Stranathan ( 1948 ) radium and its disintegration prod-
uct, radon, are alpha particle emitters. However, the disintegra-
tion products of radon are gamma emitters of short half life. The
range of alpha particles is so short that they will not even pene-
trate the skins of the animals; therefor, it is impossible by Geiger
count methods to make use of the alpha radiation. However, it is
possible to measure the gamma radiation from the disintegration
products of radon. In order to determine quantitatively the amount
of radium present by measurement of the gamma emission from
the disintegration products of radon their must exist a known
ratio between the amount of radon corresponding to a quantity of
radium. This is the case when radium and radon are in radio-
active equilibrium, since the ratio of the amount of radon to the
amount of radium present is constant. The radium content of the
animals can then be determined by comparing the Geiger counts
with those of a known amount of radium which is in radioactive
equilibrium if the radium injected animals are sacrificed, sealed
in containers, and laid aside for 38 days. At this time the rate of
disintegration of radon becomes equal to the rate at which it is
being formed. When this point is reached, radium and radon are
said to be radioactive equilibrium.

Hoecker and Roofe ( 1951 ) have pointed out that approximately
70^ 6f the radium injected in an animal is excreted within 15 to 20
days and that the remaining 30% is permanently retained in the
bones of the animals. This radium is constantly disintegrating to
radon, which is an inert gas. The amount of radon in the live animal
is then equivalent to the difference between the rate of its production
and elimination. The rate of elimination is dependent upon biolog-
ical excretion and radioactive disintegration. As shown in Graph
VIII the Geiger counts of the live animals indicate that very little
change in the amount of radon retained in the animals occurred after
21 days. This would definitely indicate that an equilibrium had
been attained between the rates of production and elimination of
radon in the live animals. As yet no means of calculating the
amount of radium present in a live animal has been determined.
However, if further investigation reveals a constant ratio to exist
between the amount of radon in the live animal and the amount of

38 The University Science Bulletin

radium retained in the animal's body, it will be possible to ascertain
by Geiger counts the exact amount of radium present in the live
animal.

General and Histochemical Observations of Tissues

The development of fat vacuoles in the bone marrow of the pro-
tein deficient animals has been considered previously in this paper.
In addition, it was observed that there was a decrease in mitotic ac-
tivity in the marrow. The cell types present in the tissue sections
were preponderantly mature cells, especially those in the neutro-
phil series. An occasional megakaryocyte appeared to be frag-
menting in both the protein deficient and irradiated animals. This is
indicative of their presumed function of producing blood platelets.
Tissues taken from the recovery animals showed a great deal of
mitotic activity. The marrow taken from the irradiated animals
showed little or no change in mitotic activity; however, nuclear and
cytoplasmic disintegration of some of the formed elements was ob-
served. These cells were probably the targets of alpha particles
and show a decrease in both cytoplasmic and nuclear absorption of
the dye.

The absorption of toluidine blue varied only slightly in the mar-
row taken from experimental animals and their controls. In the
protein deficient animals there was a definite drop in total tissue
absorption of the dye due to fatty infiltration. At times there ap-
peared to be a slight decrease in the cytoplasmic absorption, but this
was not consistent. The only change in cellular absorption in the
irradiated animals was noted above, that is, a decrease in ab.sorp-
tion in disintegrating cells. The majority of cells which were un-
dergoing disintegration seemed to be immature erythrocytes. This
would coincide with the relative drop in erythrocyte numbers.

DISCUSSION

For a portion of the protein experiment, Trahan (1951) made
manometric studies of the bone marrow extracted from the sacri-
ficed animals. From his studies he found an 18% drop in the Q02
of the protein deficient animals. His studies also included peri-
pheral blood studies in which he found a drop in hematocrit,
hemoglobin, erythrocytes, and lymphocytes. He also found an
increase in peripheral neutrophil count of 263%. Although many of
his data were obtained from a different group of similarly treated
animals, this data helps explain some of the data obtained in this
experiment. In both the bone marrow and peripheral studies tliere

Irradiation on Bone Marrow 39

was found a drop in erythrocyte numbers and an increase in
neutrophil numbers. The drop in tissue activity due to fatty infil-
tration is substantiated by the drop in Q02 of the tissues.

Orten and Orten ( 1937 and 1939 ) found that rats fed low protein
diets developed hypochromatic anemia with a normal red cell
count. They found this condition could be corrected by adminis-
tration of protein to the diet. Wright and Skaggs (1946), in a
similar experiment, reported a slight drop in red cell count and a
reduced hemoglobin value.

Benditt (1947) also did work with rats on protein free diets,
and found that hypoproteinemia resulted. He and his co-workers
experimented with diets varying the amino acids, and came to the
conclusion that a dietary deficiency of protein or a single essential
amino acid, with the exceptions of arginine or leucine, resulted
in a reduced number of erythrocytes and a drop in hemoglobin in
tlie peripheral blood.

Very little work has been done on the effects of diet on hemo-
poietic tissue. Brown (1954) studied the effects of protein defi-
ciency on both peripheral blood and bone marrow. His results
on peripheral studies are in close agreement with those already
stated. He did, however, find that peripheral leucopenia also
resulted and reported a drop in the peripheral platelet count. His
studies of bone marrow indicated a drop in immature cells of the
myeloid and erythroid series, and a decrease in all types of cells,
including the megakaryocytes. He also reported the myeloid/ery-
throid ratio to remain fairly constant. In this experiment there
was found a relative increase in megakaryocytes and a definite
increase in the myeloid-erythroid ratio. The methods of counting
and staining used may explain this difference in results. Brown
(1954) stained with hematoxylin and eosin, and the bone marrow
in this experiment was stained with toluidine blue. Brown selected
myeloid, erythroid, and mixed fields, and used a camera-lucida
for counting. In this experiment, the fields were selected at random
and the previously described pointer system was used for counting.

Hoecker and Roofe ( 1951 ) have found that in the bones of rats
sacrificed soon after injection, the highest concentration of radium
is in the periosteum and endosteum. In rats sacrificed several
months after injection they observed that the concentration is great-
est in the newly formed bone. They also found a similarity exist-
ing between radium deposition and optically dense areas in the
bone. They report that 96% of the total energy emitted by radium
in total disintegration is associated with alpha particle emission.

40 The University Science Bulletin

Of the remaining 4% only a fraction is absorbed by the body and
its effects can be considered neghgible.

In a series of studies done by Bloom (1948) using external and
internal sources of irradiation, it was found that cessation of mitosis,
pyknosis, and fragmentation of the cells occurred. He observed a
greater proportion of debris resulting from destruction of red blood
cells than from white blood cells. He found that the granulocyto-
poietic cells disappeared less rapidly than the erythropoietic. He
reported an initial decrease in megakaryocyte numbers with a later
return to normal and above. He concluded, "Irrespective of species
or type of radiation, erythroblasts were more sensitive than myelo-
cytes, and myelocytes more sensitive than megakaryocytes." This
was found to be the case in this experiment.

Roofe, Bingham, and Comer (1951) in a short-term experiment
using intraperitoneally injected radium chloride found that mega-
karyocyte and blood platelet numbers fell off immediately after in-
jection with megakaryocyte numbers returning to near and above
normal after 14 days. In the present study, the megakaryocyte
numbers were found to be considerably above normal by the four-
teenth day with a continuing rise to the thirty-fifth day.

Roofe and Bingham ( 1953 ) used intraperitoneally injected radium
chloride and found that in the bone marrow neutrophils and eosino-
phils increased in number with time. The erythroid series dropped
initially with a tendency to increase with time. At the same time,
peripheral polymorphonuclear leucocytes dropped, with the R. B. C.
count remaining normal. Dunlop (1942) also found an increase
in bone marrow neutrophils with a decrease in peripheral neutro-
phils. Both authors found an increase in the myeloid-erythroid
ratio in the bone marrow. This agrees with the results found in
the present study. Such a rise in bone marrow neutrophils and
increase in peripheral count suggest some factor which might cause
retention of mature granulocytes. Perhaps there is also some factor
which permits the peripheral erythrocyte members to be main-
tained in spite of relative decreases in bone marrow concentration.

According to Davidson (1950), the highly cellular organs svich
as spleen, thymus, pancreas, and bone marrow are rich in nucleic
acid, whereas organs like brain or muscle have a lower content.
In tissues with a high nuclear-cytoplasmic ratio the concentration
of DNA is high, but in tissues with a high concentration of cyto-
plasmic granules, RNA content is high. He also finds a good cor-
relation between cytoplasmic basophilia and RNA content of the
tissues. He points out that embryonic tissues have been found to

Irradiation on Bone Marrow 41

have higher nucleic acid content than adult. Since the similarity
of dye absorption in the experimental and control animals, in this
experiment, indicates little or no change in cellular basophilia, it
seems plausible to suggest that cellular nucleic acid content was
not changed by the procedures used in this experiment. Since
nucleic acids play such a vital part in cellular activities it seems
logical that their concentrations should be preserved at all costs.

SUMMARY

Two groups of experimental animals were used in this experi-
ment. The animals in the first group were subjected to protein
fast and the animals in the second group were injected intraperi-
toneally with radium chloride. Differential counts were made by
a random selection procedure on the bone marrow taken from these
animals. Toluidine blue was used as the stain to determine the
effect the protein fast and irradiation had on the nucleic acid con-
tent of the tissue.

1. Cellular basophilia and nucleic acid content is not noticeably
affected by protein fast and radiation under the conditions de-
scribed in this experiment.

2. Total bone marrow basophilia and nucleic acid content is
reduced during protein fast due chiefly to reduction in cellularity.

3. Fatty displacement of tissue occurs in bone marrow during
protein fast.

4. Mitotic activity is decreased by both protein fast and radiation.

5. Changes which take place in the bone marrow during protein
fast are overcome when animals are returned to normal protein diets.

6. The myeloid/erythroid ratio increases during protein fast and
following irradiation.

7. In the bone marrow erythropoietic elements are the most
easily affected by protein fast and internal irradiation, and gran-
ulocytopoietic elements are more susceptible than are the mega-
karyocytes.

BIBLIOGRAPHY

Benditt, E. p.

1947. Studies of amino acid utilization II. Essential amino acids as
sources of plasma protein and erythrocytes, in the hypoproteinemic
rat. Journ. Nutri., vol. 33, pp. 85-94.

Bloom, William

1948. Histopathology of irradiation from external and internal sources.
New York-Toronto-London, McGraw-Hill Book Corp., Inc.

Brown, J. Wm.

1954. The effect of protein deficiency on bone marrow and peripheral
blood of young albino rats. Univ. Kansas Sci. Bull., vol. 36
pp. 55-68.

42 The University Science Bulletin

Caspersson, T.

1947. The relation between nucleic acid and protein synthesis. Symp.
Soc. Exp. Biol., pp. 1-127.

Chalkley, H. W.

1942. Method for quantitative morphological analysis of tissues. Jour.
Nat. Cancer Inst. vol. 4, pp. 47-53.

Davidson, J. N.

1950. The biochemistry of the nucleic acids. London: Methuen and
Co., Ltd. Nevi' York: John Wiley and Sons, Inc. 1st Ed.

DUNLOP, C. E.

1942. EflFects of radiation on the blood and the hemopoietic tissues.
Arch. Path., vol. 34, pp. 592.

HoECKER, F. E. and P. G. Roofe.

1949. Structural differences in bone matrix associated with metabolized
radium. Radiology, vol. 52, pp. 856-864.

HoECKER, F. E. and P. G. Roofe.

1951. Studies of radium in human bone. Radiology, vol. 56, no. 1, pp.
89-98.

Maximov, a. a. and W. Bloom.

1953. Textbook of Histology. W. B. Saunders Co., Philadelphia & Lon-
don, 6th Ed.

Michaelis, L.

1948. The nature of the interaction of nucleic acids and nuclei with basic
dyestuffs. Cold Spring Harbor Symposia on Quantitative Biology,
vol. 12, pp. 131-146.

Orten, a. U. and J. M. Orten.

1943. Role of dietary protein in hemoglobin formation. Journ. Nutr., vol.
26, pp. 21-31.

1945. A study of hemoglobin formation following administration of cer-
tain amino acids to rats fed a diet low in protein. Jour, of Nutrition,
vol. 30, pp. 137-142.

Roofe, P. G., and H. G. Bingham, and R. Comer.

1951. EflFect of radiation from intraperitoneally injected radium chloride
upon megakaryocyte and blood platelet production in albino rats.
Kans. Acad, of Sci., vol. 54, pp. 391-394.

Roofe, P. G. and Samuel W. Lesher.

1952. Recent advances in the study and technique of anatomy. The
Smithsonian Report for 1952, pp. 327-347.

Roofe, P. G. and Hal G. Bingham.

1953. The eflFect of radiation from intraperitoneally injected radium chlor-
ide upon the hemopoietic activity of the bone marrow in albino
rats. Univ. Kansas Sci. Bull., vol. 35, pp. 945-960.

Stowell, R. E.

1947. Histochemical observations on nucleic acids. Soc. Exp. Biology,
vol. 1, pp. 190-206.

Stranathan, J. D.

1948. The Particles of Modem Physics. Blakiston Company, Philadelphia,
Toronto.

Trahan, Robert.

1951. The effect of protein-deficient diet on the oxygen consumption of
the rat bone marrow and spleen. Unpublished manuscript.
Wright, L. L. and H. R. Skaggs.

1946. Vitamin B complex studies with diets deficient in level of protein.
Proc. Soc. Exp. Biol. Med., vol. 63, pp. 327-333.

Irradiation ox Bone Marrow 43

EXPLANATION OF GRAPHS AND TABLES
GRAPHS I to VII

Ordinates show number ot cells per 100 erythrocytes.

Abscissae show time.

Group A — Purina fox chow diet animals

Group B — 18% protein diet animals

Group C — 0.0% protein diet animals

Group R — Recovery animals

Group Ra — Radium injected animals

Group N — Controls for injected animals

Line connecting groups C and R does not represent animals in the series
but is to show relationship of the two groups.

Only average values are shown on these graphs because of the small spread
in the data obtained from counts.

GRAPH VIII

The ordinate expresses the number of Geiger counts recorded from radium
injected animals as the percent of the counts recorded from a 1 microgram
standard wliich was in radioactive equilibrium.

_,.v.,^,„ -y-'

44

The University Science Bulletin
GRAPHS I to IV

GRAPH I
HEMOCYTOBLAST PROTEIN

GRAPH II
MEGAKARYOCYTE PROTEIN

HEMOCYTOBLASTS PER

MEGAKARYOCYTES PER

-

100

ERYTHROCYTES

100 ERYTHROCYTES

\,

A

14

A

B

C

R

13

1 . R

9

12

/
/
;

8

1 1

/

;

7

10

/

;
/

6
5

9
8

;

;

J

4
3
2

1

\
\
\

\
\

N

7
6
5
4
3

/"■-^.

/■

0

10 II 12 13 14 15 16
AGE IN WEEKS

10 II 12 13 14 IS 16
AGE IN WEEKS

GRAPH III

NEUTROPHIL PROTEIN

NEUTROPHILS PER

100 ERYTHROCYTES

130

/■

A

B

120
1 10

/ ■■■-.

/

/

C

B

100

/

90

eo

/
/

/
/
/

/

'•.

70

60

^^^.szJ=::=^

=:=

50
40

10 II 12 13 14 IS IS
AGC IN WEEKS

80
70
60
50
40
30
20
t 0

GRAPH IV

BLOOD CELLS PROTEIN

NO. CELLS PER 2500
SQUARE MICRONS

8

R -

\.

10 II 12 13 14 IS W
ACE IN WEEKS

Irradiation on Bone Marrow

GRAPHS V to VIII

45

GRAPH V

HEMOCYTOBLAST RADIATION

HEMOCYTOBLASTS PER
100 ERYTHROCYTES

RA-

N

I 2 3 4 5 6 7

WEEKS AFTER INJECTION

GRAPH VI

MEGAKARYOCYTE RADIATION

MEGAKARYOCTES PER
100 ERYTHROCYTES

12 3 4 5 6 7

WEEKS AFTER INJECTION

GRAPH VII

NEUTROPHIL RADIATION

NEUTROPHILS PER
100 ERYTHROCYTES

220
200
180
I 60
I 40

12 0

100

60

60

40

RA-
N -

I 2 3 4 5 6 7

WEEKS AFTER INJECTION

GRAPH VIII
GEIGER COUNTS RADIATION

POINTS REPRESENT OF STANDARD

STANDARD = I MICROGRAM RADIUM
0 - INJECTION DATE

0 I 2 3 4 S 6 7
WEEKS AFTER INJECTION

46 The University Science Bulletin

TABLE I

RAT DIETS *

General Biochemicals Amounts

Casein for Diet A, 0.0% 0.0 grams

Casein for Diet B, 18% 540 grams

Sucrose or dextrin, 93% ( Beet or Cane Sugar ) 2790 grams

Corn oil, 2% ( Mazola Oil) 60 grams

1 :20 liver extract, 1% 30 grams

Bi 16 Mg.

Bo 16 Mg.

Riboflavin 15 Mg.

Niacin 39 Mg.

Calcium Pantothenate 16 Mg.

Choline Chloride 3 grams

Haliver oil 1 drop per week per animal

L — Tocopherol Vitamin E 1 drop every 4 weeks per animal

Salts, 4% 120 grams

( by G. B. I. after Hubbel, Mendel, and Wakeman )

* Modified from diets used by Brown (1954).

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(51)

52

The University Science Bulletin

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Irradiation on Bone Marrow

53

TABLE VIII. — Cell Counts for Radiation E.xperiment Animals

Ra=: irradiated animals
N :=control animals

Animal

Hemocy-
toblast

Megakar-
yocyte

Lympho-
cyte

Eosin-
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Neutro-
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Erythro-
cyte

Time

Ra 1

Ra 2

Ra 3

Ra 4

Ra 5

Ra 6

Ra 7

N 1

N 2

N 3

N 4

N 5

N 6

N 7

2
3
5
5
4
3
2

2
3
2
1
2
2
2

4
10
13
15
16
15
12

3
3
4
2
5
3
2

22
32
26
34
22
37
26

18
17
17
16
19
22
20

6
4
4
6
4
6
3

4
4
3
4
3
3
3

132
170
202
197
196
210
186

66
72
74
70
72
68
71

100
100
100
100
100
100
100

100
100
100
100
100
100
100

7 days
14 days
21 days
28 days
35 days
42 days
49 days

7 days
14 days
21 days
28 days
35 days
42 days
49 days

THE UNIVERSITY OP KANSAS

SCIENCE BULLETIN

Vol. XXXVII, Pt. I] October 15, 1955 [No. 3

Taxonomy and Ecology of Kansas Hepaticae ^

BY

Ronald L. McGregor 2

Abstr.\ct. — This paper reports 69 taxa of hepatics for Kansas. Most are
ubiquitous species found over most of North America and elsewhere but not
previously known from Kansas. A number of significant extensions of range
are reported and more information is given for a few taxa recently reported
for North America.

Morphological variations of the taxa are discussed where they differ from
those reported in the literature or are important in problems of speciation.

The ecology of Kansas Hepaticae is considered in relation to soils, climate,
higher plant associations, invasion and succession. The hepatic flora of Kan-
sas is most closely related to that of the south, particularly the Edwards Plateau
of Texas.

INTRODUCTION

During the 90 years of botanical activity in Kansas, prior to the
present work, the Hepaticae received Httle attention. A few pubh-
cations mentioned Hverworts and a few Hsts were prepared. It is
significant that Frye and Clark ( 1937-'47 ) list one species only from
the State indicating that the general literature contained little on
this group of plants in Kansas.

The first paper to report an hepatic from Kansas was that of Car-
ruth (1873) in which he listed Marchantia pohjmorpha from Doug-
las County. Cragin (1884) noted Carruth's paper and reported
Rehoiilia hemisphaerica from Clay and Ottawa Counties. Smyth
and Smyth ( 1911 ) included 25 species of hepatics in their catalogue
of the Flora of Kansas. Lorenz ( 1915 ) reported Riccia frostii from
the State and Thompson (1948) reported Sphaerocarpus michelii
and Sphaerocarpus texanus. Gier (1949) published a list of Kan-
sars Hepaticae in which he listed 25 species based on the publica-
tion of Smyth and Smyth (1911). McGregor (1952) reported

1. Part of a thesis submitted to the faculty of the Graduate School of the University of
Kansas in partial fulfillment of the requirements for the Degree of Doctor of Philosophy.

2. Assistant Professor of Botany, University of Kansas.

(55)

56 The University Science Bulletin

Riccia rhenana and Wittlake (1953) reported Oxymitra androgyna
from the State. In most of these papers the hepatics are mentioned
only incidental to other objectives or are merely lists of species
occurring in the State, or new to the State.

For the past seven years the author has made an extensive and
intensive study of the Hepaticae of Kansas. Each of the 105 coun-
ties has been visited and all habitat types occurring in the State
have been observed. Most of these habitat types have been studied
many different times and throughout all seasons of the year. A
collection of each species was made from each location as it was
found. In addition the same species was again collected from the
same site at various times during the growing seasons. For each
collection detailed notes were made to cover collection data, abun-
dance and ecological conditions.

It is not the purpose of this work to prepare a taxonomic manual
of the Hepaticae of Kansas but rather it is an attempt to record all
species found to occur in the state, their abundance, morphological
variations and to give a detailed study of their ecology in relation
to other Hepaticae and to other vegetation. Such a study has never
been made in this region of the United States. The only works to
cover the ecology of a hepatic flora anywhere in the United States
are the important publications on Central New York State by
Schuster (1949) and the equally important work on Minnesota by
Schuster (1953). Sharp (1939) included some ecological data on
hepatics in his publication on Eastern Tennessee Bryophytes but
for the most part his work is concerned with mosses. Many other
papers have included a limited amount of hepatic ecologic data
but not a complete account for a hepatic flora in a definite area of
any size.

It was therefore considered worth-while to study the hepatic flora
of a state located in the center of the United States — one in which
the vascular flora shows a transition from the eastern forest areas
to the short-grass prairies, and at the same time, one that includes
northern, southeastern, and strong southern floristic affinities.

This work is divided into two parts. The first section is con-
cerned with the taxonomic studies, referred to above, and the sec-
ond section describes the ecology of the different taxa,

ACKNOWLEDGMENTS

The writer is indebted to Dr. R. H. Thompson for first suggesting
the problem studied and for many suggestions and criticisms. To
Dr. W. H. Horr thanks are due for invaluable assistance in collect-

Taxonomy and Ecology of Kansas Hepaticae 57

ing and unfailing aid on ecological problems. Dr. R. W. Baxter
and Dr. A. J. Mix have offered advice and assistance during the
course of the work and several graduate students of the Botany
Department, University of Kansas, have aided in collecting. Dr.
A. W. Evans, and Dr. Karl Miiller have assisted in the determina-
tion of difficult groups. To the curators of the herbaria of New
York Botanical Garden, U. S. National Museum, Harvard Univer-
sity, Missouri Botanical Garden, Chicago Natural History Museum,
American Bryological Society and Kansas State College the writer
is indebted for the loan of specimens.

TAXONOMIC STUDIES

In the following list are included all the described taxa of he-
patics now known to occur in the state of Kansas. For each are
given brief notes on the habitats, comparative frequency, distribution
by counties and other pertinent taxonomic or morphological fea-
tures.

At present the annotated list includes 69 taxa. This is believed
to include nearly all the hepatics to be found in the State since few
additional species have been found during the last two seasons.

The arrangement of taxa follows, in general, that of Schuster
(1953) in so far as order of families and genera are concerned.
The species are arranged alphabetically under each genus. Since
most collections have been made by the writer they will be
simply designated by placing an M before the collection number.
All other records are designated by usual standard abbreviations.
Unless otherwise mentioned all collections are deposited in the
Herbarium of the University of Kansas.

Anthocerotaceae

1. Anthoceros crispidiis (Mont.) Douin. Uncommon in our
area but found in prairie swales where the characteristic plant
association is dominated by Andropogon gerardi and Tripsacum
doctyloides. It has also been found once in an overgrazed pasture
near a pond. In all habitats the soil was loam or clay, moist,
and shaded.

As noted by various authors A. crispulus is very closely re-
lated to A. punctattis. Proskauer (1948) who merged the two
considered A. crispulus to be a phase or modification of A. puncta-
tus with surface outgrowths "especially in plants from relatively
dry situations." This concept may be correct but the writer
has not been able to verify it so far as Kansas material is con-

58 The Univtersity Science Bulletin

cerned. A. punctatiis has been found in several habitats much
drier than those in which material referred to A. crispultis has
been found and no sign of surface outgrowths has been noted.
In one habitat hundreds of specimens of typical A. crispulus
were found and nothing referable to A. ptinctatus could be dis-
covered. Yet, less than a mile away and on a drier prairie slope
only A. punctatus could be found. In addition to the fact that
the dorsal surface bears crowded lamellae (characteristic of A.
crispulus but not of A. punctatus) Kansas material of A. crispulus
has pseudoelaters often 3-5 cells long and measuring up to
210[JL long, while A. punctatus has pseudoelaters only 1-2 cells long
and 60-lOOiJL long. However the range of the number of cells
and length of pseudoelaters overlap in both and do not always
allow for easy separation on these points alone. Until more
information is obtained it seems best to consider the two as
separate taxa.

Distribution: Douglas County: 1 mile west of Pleasant Grove,
in bluestem prairie, July 18, 1951, M4987; July 24, 1951, M5001.
Jackson County: 1 mile west of Hoyt, bluestem prairie, August 22,
1951, M5076. Johnson County: near De Soto, prairie pasture,
October 2, 1951, R. H. Thompson. Leavenworth County: 3 miles
northeast of Tonganoxie, bluestem prairie, August 4, 1951, M5007.

2. Anthoceros punctatus L. Rather widely distributed in the
eastern half of the State where it is found commonly on moist
sandy soil in prairie dominated by Andropogon gerardi, Sorghas-
trwn nutans and Tripsacum dactyloides. It also occurs occasion-
ally on alluvial deposits of creek banks, sandy oak-hickory wood-
lands, overgrazed prairie pastures, and along paths or trails
in the sand dune areas of central Kansas.

The occurrence of the species in the state is directly related
to rainfall. A dry winter and spring such as occurred in 1952-53
prohibits the appearance of the species for the season. During
the 1953 season not a single mature plant was found while in
contrast the very wet season of 1951 revealed an abundance of
plants.

It should be noted that in Kansas the species occurs most
frequently in relatively moist, sandy prairie and less commonly
on limestone prairie. Its occurrence in woodlands is usually in
close association with such prairie areas and the occurrence on
alluvial deposits is attributed to water dispersal.

The frequency of occurrence of this species was determined
by use of meter quadrats dispersed at random in a prairie area.

Taxonomy and Ecology of Kansas Hepaticae 59

It was found than an average of 33 plants per meter quadrat
existed in a prairie of some 160 acres. Most of these occurred
around the base of Andropogon gerardi clumps. A casual study
of such an area might lead one to believe that the usual habi-
tat was along the moister drainage areas. However, the large
number of plants found along moist ditches and banks seems
to be due to a concentration of spores by drainage from the
"mother lode" which actually is dispersed over a large area
and thus might escape notice. The same condition is to be
found in forested areas.

Kansas specimens of A. punctatus seem to agree with descrip-
tions given for the species except that they are usually at the
lower range given for size of the thallus and length of the
sporangium.

Distribution: Allen County: 5 miles north of Moran, alluvial
deposit on creek bank, October 8, 1948, M2503; 1 mile south of
Humboldt, oak wooded hillside, July 7, 1949, M3334. Anderson
County: 5 miles south of Garnett, creek bank, October 8, 1948,
M2488; 2 miles south of Garnett, clay soil in oak woods, June 4,
1951, M4877. Bourbon County: 2 miles south of Uniontown, rocky,
moist hillside, October 8, 1948, M2514; 1 mile south of Uniontown,
creek bank, July 7, 1949, M3344. Cherokee County: 5 miles east
of Baxter Springs, clearing in oak woods, April 30, 1948, M1363;
5 miles east of Baxter Springs, creek bank, July 31, 1948, M1939.
Clay County: 5 miles north of Morganville, sandy bluestem prairie,
August 16, 1951, M5069. Cloud County: on soil, September 13,
1948, Rev. S. V. Fraser. Crawford County: lespedeza pasture, No-
vember 24, 1949, L. J. Gier 2952. Douglas County: 2 miles south-
east of Baldwin City, creek bank, November 2, 1947, Ml 133; 1 mile
west of Pleasant Grove, swale in bluestem prairie, July 24, 1951,
M4999, Ellsworth County: 3 miles northeast of Carneiro, low
moist prairie, May 22, 1948, M1499. Franklin County: 1 mile
southeast of Ottawa, wooded hillside, June 28, 1949, M3308. Harvey
County: 5 miles north of Burrton, cattle trail through sand dunes,
June 11, 1951, M4915. Jackson County: 1 mile west of Hoyt, blue-
stem prairie, August 22, 1951, M5075. Leavenworth County: 4
miles northeast of Tonganoxie, sandy bluestem prairie. May 7, 1948,
M1408; 2 miles west and 1 mile south of Linwood, sandy oak-
hickory woods, June 16, 1949, M3086. Montgomery County: 3
miles south of Elk City, sandy oak woods, April 4, 1942, R. H.
Thompson. Nemaha County: sandy bluestem prairie, August 9,
1948, M2018. Osage County: 6 miles south of Quenemo, moist

60 The University Science Bulletin

bank in oak woods, July 1, 1949, M3324. Ottawa County: Ottawa
County State Park, seepy area below sandstone outcrop. May 21,
1948, M1466. Reno County: 6 miles northeast of Hutchinson, cattle
path through sand dunes, June 11, 1951, M4955. Saline County: 2
miles northwest of Brookville, moist sandy prairie ravine, May 22,
1948, M1478. Washington County: 2 miles southwest of Palmer,
sandy bluestem prairie, May 27, 1951, M4827. Wilson County: 1
mile northeast of Neodesha, open sandy oak woods, April 10, 1948,
M1271. Woodson County: 2 miles northwest of Yates Center, on
shallow soil over surfacing sandstone in prairie, April 9, 1948, M1244;
Lake Fegan, sandy path in oak woods, June 19, 1949, M3191.

3. Phaeoceros laevis (L.) Proskauer. Found commonly on
shaded moist sandstone ledges, less frequently on creek banks, seepy
areas below ledges and springs. It rarely occurs elsewhere and
seldom with Anthoceros punctatus and A. crispiilus. It is an almost
constant associate with Conocephahnn conicum, Plectocolea hyalina,
and often with Reboulia hemisphaerica.

P. laevis occurs characteristically as extensive mats on the sides
of shaded, moist sandstone ledges along creek banks, ravines and
often on boulders in stream beds.

Kansas material of P. laevis seems to match closely the descrip-
tions given for the species. However, in some of the collections
from very moist habitats mature specimens often exhibit the charac-
ters of Phaeoceros carolinianus ( Michx. ) Proskauer in that the in-
volucres do not flare and marginal glandular thickenings are to be
found. These characters are by no means constant and seem to
intergrade with P. laevis in which the involucre does flare at the
summit and glandular thickenings are rare. One colony, represented
by collections M1058, M3105, M4147, M4673 and M5003, has been
studied each year from 1947 through 1953. Usually a few involucres
do not flare at the mouth and some glandular thickenings do occur
but in 1951, a year of record rainfall, most of the involucres did not
flare and glandular thickenings were rather frequent. It thus seems
best not to consider P. carolinianus as a distinct taxon. Schuster
(1953) noted these conditions and reduced carolinianus to a forma
of P. laevis. It might be best considered thus, but Kansas material
intergrades so completely that separation here seems unwarranted.

Distribution: Anderson County: 5 miles south of Garnett, creek
bank, October 8, 1948, M2489. Chautauqua County: 3 miles north-
east of Sedan, moist sandstone ledge, July 1, 1947, M1073; VA miles
northeast of Sedan, on moist sandstone rocks, July 8, 1949, M3381.

Taxonomy and Ecology of Kansas Hepaticae 61

Douglas Countij: 5 miles west of Baldwin City on moist sandstone
ledge, June 24, 1947, M1058; June 18, 1949, M3105; October 15,
1949, M4147; September 16, 1950, M4673; August 2, 1951, M5003.
Elk County: 10 miles north of Busby, sandy soil below a spring,
July 27, 1948, M1917. Franklin County: 4 miles southwest of
Ottawa, moist sandstone ledge, June 18, 1949, M3097. Greenwood
County: near Severy, September 2, 1945, R. H. Thompson; 5 miles
northeast of Fall River, moist sandstone ledge, July 17, 1947, M1075;
1 mile east of Fall River, on moist sandstone rocks, July 9, 1949,
M3413. Labette County: 4 miles north of Chetopa, creek bank,
July 2, 1948, M1620. Leavenworth County: 5 miles southwest of
Leavenworth, moist sandstone ledge, July 27, 1947, M1078. Lincoln
County: 10 miles southeast of Lincoln, moist sandstone ledge, Sep-
tember 4, 1950, M4651. Montgomery County: 3 miles southwest of
Elk City, moist sandstone rocks, and cattle trail through oak woods,
July 1, 1947, M1068; September 1, 1948, M2385; September 22,

1951, M5221. Washington County: 1 mile west of Linn, on moist
sandstone ledge in prairie ravine, August 23, 1947, M1086; August
10, 1948, M2055; 2 miles west of Linn, sandstone ledge on creek
bank, August 22, 1952, M5712, M5715. Wilson County: 3 miles
west of Neodesha, edge of pond in sandy prairie, June 30, 1947,
M1605. Woodson County: 2 miles northwest of Yates Center, moist
sandstone ledge, October 26, 1947, M1128; 8 miles south and 4
miles west of Yates Center, moist sandstone ledge, September 27,

1952, M6064. Wyandotte County: 5 miles northwest of Bonner
Springs, on moist sandstone ledge, July 27, 1947, M1080.

4. Notothylas orbicularis (Schwein.) Sulliv. Widely distrib-
uted over the eastern half of the State. It has been found abun-
dantly in a few prairies dominated by Andropogon gerardi and
Tripsacum dactyloides. These areas include sandy soils and heavy
loam soils. N. orbicidaris is found in such areas in moist prairie
swales. The species has been found twice on alluvial deposits along
stream banks, once in an overgrazed pasture and once in a thicket
of Rubus argutus in a clearing on an oak-hickory wooded hillside.

In one moist prairie swale N. orbicularis was found to average
over 150 plants per meter quadrat in an area three acres in extent.
It was not to be found on alluvial deposits in the drainage area
below this prairie or in adjacent prairies with the equivalent habitat
sites.

All collections of N. orbicularis seem to agree morphologically
with published descriptions.

62 The University Science Bulletin

Distribution: Cherokee County: 5 miles east of Baxter Springs,
alluvial deposit along Shoal Creek, August 30, 1948, M2356. Claij
County: sandy bluestem prairie, August 16, 1948, M5070. Douglas
County: 1 mile west of Pleasant Grove, bluestem prairie, July 17,
1941, M4985; July 24, 1951, M5()01. Johnson County: near De Soto,
overgrazed pasture, October 2, 1951, R. H. Thompson. Leaven-
worth County: 3 miles northeast of Tonganoxie, bluestem prairie,
August 4, 1951, M5008. Washington County: 1 mile northeast of
Linn, sandy bluestem prairie, August 21, 1951, M5074. Miami
County: Miami County State Park, river bank, October 3, 1948,
M2467. Wilson County: 1 mile northeast of Neodesha, sandy soil,
Ruhus thicket, clearing in oak woods. May 1, 1948, M1380.

Sphaerocarpaceae

5. Sphaerocarpus michelii Bellardi. Found on sandy soil near
surfacing sandstone and in bare areas between clumps of Andropo-
gon gerardi in sandy bluestem prairies. Plants are to be found in
the spring, usually maturing in April in this region.

S. 7nichelii is found most abundantly in a spring following a sum-
mer and fall of above average rainfall. In other years it is rare
or absent. In this respect the species is like S. texanus and there-
fore is not a plant characteristic of the same habitat year after year.

One site, 5 miles west of Baldwin City, Douglas County, has
been studied several years in succession. S. 7nichelii and S. texanus
are both found in this sandy, overgrazed prairie pasture. The
abundance of plants to be found is directly correlated with rainfall
and degree of removal of cover. A moist summer and fall with a
simultaneous major reduction in grass cover has twice resulted in
an abundance of Sphaerocarpus in this area. Meter quadrat studies
revealed an average of 223 plants of S. michelii and S. texanus per
meter quadrat over an area of two acres. S. texanus dominated at
the rate of 12 to 1. In normal seasons and with more cover the
abundance of Sphaerocarpus was reduced to an average of 18 plants
per meter quadrat over the same area and S. texanus was dominant
over S. michelii by a rate of over 30 to 1.

Sphaerocarpus michelii is not always easily separated from S.
texanus in our area. It has never been found in pure colonies for
S. texanus is invariably present and dominant in number of plants.
S. texanus, however, has been found many times in pure colonies.
Haynes (1910) described the chief differences separating S. texanus
from S. Sphaerocarpus (S. michelii) to he "in the more pointed
fusiform-clavate rather than obovoid involucres, in the meshes of

Taxonomy and Ecology of Kansas Hepaticae 63

the surface of spore tetrads being nearly twice as wide, and in
the high ridges forming these meshes being sinuous or crenulate-
margined or irregularly dissected, or occasionally rising into obtuse
spines at the points of intersection, but never forming sharp needle-
like spines as in S. Sphoerocarpus." Frye and Clark (1937-1947) re-
port tliat the most reliable character separating S. michelii from
S. texanus is the sharply spinose spore of the former. Frye and
Clark also give the size of the spore tetrad as 80-1 lOp. in diameter
for S. michelii and 72-170[j. in diameter for S. texanus while Haynes
(1910) gives the size as 90120[x for S. michelii and 72-17l!j. for
S. texanus.

Studies on Kansas collections of these two species indicates
clearly that the shape of the involucre is of no value in sepa-
rating these taxa. The size of the spore tetrad is likewise of little
use as the average size for both species is 108-110[;.. S. texanus
has the largest and also the smallest tetrads of the two and thus
size alone is of little significance. The only constant character which
can be used to separate the species is the size of the meshes on the
spore tetrad. In S. m'.chelii these range from 9-16.5tj. while on S.
texanus they range from 16-30[jl with the average being 21[x.

S. michelii in its best expression has the ridges separating the
spines low, their intersections with very acute or acicular spines.
S. texanus characteristically has intersections spineless or with
blunt spines. Kansas material, however, reveals a number of
specimens in which these spine characters are intermediate and
in which the ridges separating the meshes also intergrade. Haynes
(1910) illustrates considerable variation of S. texanus spores.
These variations are present in the species from our area. It
seems possible, then, that some degree of hybridization has
occurred but as yet this is only surmise.

S. michelii was first reported authentically from the United
States by Frye and Clark (1937-1947) as occurring in Austin,
Texas. Thompson (1948) reported the species from Shawnee
County, Kansas. In addition to the Kansas records listed below
the writer has collected the taxon from Arkansas, Missouri and
Oklahoma.

Distribution: Cherokee County: 5 miles east of Baxter Springs,
sandy soil, April 30, 1948, M1357. Douglas County: 5 miles west
of Baldwin City, sandy bluestem prairie, April 18, 1948, M1343.
Elk County: 10 miles north of Busby, sandy prairie, April 10,
1948, M1346. Greenwood County: 1 mile northeast Fall River,
sandy prairie, April 10, 1948, M1347. Leavenworth County: 4

64 The University Science Bulletin

miles northeast Tonganoxie, sandy clearing in oak-hickory woods,
May 7, 1948, M1407. Montgomery County: 3 miles south of
Elk City, sandy soil over surfacing sandstone, April 4, 1942,
R. H. Thompson. Shawnee County: Calhoun bluffs, north of
Topeka, sandy soil, March, 1942, April, 1942, David Clark. Wil-
son County: 2 miles northeast of Neodesha, sandy soil in oak-
hickory woods, April 10, 1948, M1345. Woodson County: 2
miles northwest of Yates Center, sandy soil in prairie pasture,
April 9, 1948, M1344.

6. Sphaerocarpits texanus Aust. Widely distributed over the
eastern half of the State. It occurs most abundantly on sandy blue-
stem prairie dominated by the big bluestem Aiidropogon gerardi.
In these prairies it occurs in bare areas between the grass or around
the base of the grass. Other habitats consist of trails or paths
through sandy, or sandy clay-loam woods, alluvial deposits of
streams, overgrazed pastures, and shallow soil over surfacing sand-
stone. It is rarely found on calcareous soils.

S. texanus is essentially a spring or early summer species and in
this region produces mature spores in April and May. During the
summer it is not to be found but in the fall it occurs from September
to November on alluvial soils of stream banks, seepy areas and other
very moist sites. It has never been found fruiting in the fall, a fact
which may be due to the lack of development of male plants. A
careful search, in the fall, has failed to reveal the presence of male
plants at that time.

Morphological variations and other taxonomic points of S. texanus
are discussed under S. michelii above.

The distribution of S. texanus in Kansas is undoubtedly greater
than indicated below since many sterile plants have been seen but
not collected. It almost certainly occurs in all counties of the eastern
third of Kansas.

Distribution: Chautauqua County: 3 miles north of Elgin, sandy
prairie pasture hilltop, April 5, 1952, M5348. Cherokee County: 5
miles east of Baxter Springs, rocky clay soil in clearing of oak-
hickory woods, April 30, 1948, M1356. Co^ey County: 2 miles
southwest of LeRoy, sandy prairie pasture, April 9, 1948, M1238.
Cowley County: 3 miles southeast of Arkansas City, sandy clay soil
in open woods, April 11, 1948, M1301. Douglas County: 12 miles
south of Lawrence, sandy prairie pasture, April 18, 1948, M1342; 5
miles northwest of Lawrence, moist soil in cattail-sedge marsh. May
14, 1948, M1420; 5 miles west of Baldwin City, sandy overgrazed

Taxonomy and Ecology of Kansas Hepaticae 65

prairie meadow, March 25, 1948, M1165; 2 miles northeast of Law-
rence, alkivial deposit on bank of Kansas River, June 12, 1952,
M5605. Elk County: 10 miles north of Busby, sandy clay soil in
prairie, April 10, 1948, M1288. Ellsworth County: 5 miles east of
Carneiro, sandy soil in prairie, April 30, 1949, M2699. Franklin
County: 1 mile northeast of Norwood, sandy prairie pasture, April
2, 1948, M1226. Greenwood County: 1 mile northeast of Fall River,
sandy soil, clearing in oak woods, April 10, 1948, M1292. Jefferson
County: 1 mile west of Medina, sandy clay pasture, April 2, 1948,
M1232. Labette County: sandy soil, oak woods, April 10, 1948,
M1268. Montgomery County: 3 miles south of Elk City, oak woods,
April 4, 1942, R. H. Thompson. Neosho County: east of Chanute,
sandy soil, February 14, 1945, R. H. Thompson; 2 miles northeast of
Chanute, sandy soil, clearing in oak woods, April 9, 1948, M1259; 3
miles east of Chanute, sandy soil. May 12, 1951, R. H. Thompson.
Rice County: 1 mile east of Geneseo, sandy prairie pasture, April 30,
1949, M2710. Riley County: near Manhattan, alluvial deposit, bank
of Kansas River, April 30, 1949, Lucille Paslay. Saline County: 2
miles northwest of Brookville, sandy prairie pasture, April 30, 1949,
M2691. Shawnee County: 7 miles southeast of Topeka, clay soil in
severely overgrazed pasture, April 23, 1949, M2686. Wilson County:
1 mile northeast of Neodesha, sandy soil over surfacing sandstone,
April 10, 1948, M1274. Woodson County: 2 miles northwest of
Yates Center, sandy prairie, April 9, 1948, M1247; March 26, 1948,
M2650. Wyandotte County: 1 mile northeast of Bonner Springs,
sandy clearing in oak woods. May 15, 1948, M1442.

LOPHOZIACEAE

7. Lophozia hicrenata (Schmid. ) Dumort. This species is rare
in Kansas having been found in two rather distant sites. In both
places it occurred on the bare, dry crest of small sandy cliffs and is
one of the most xerophytic of Kansas leafy Hepaticae.

A careful search of many suitable habitats for the species has
been made without success. It agrees morphologically with pub-
lished descriptions of the species.

Distribution: Washington County: 1 mile west of Linn, on dry
sandy soil at crest of small cliff over sandstone outcrop, August 22,
1952, M5723. Woodson County: Woodson County State Park, ex-
posed sandy soil at top of small cliff, March 26, 1949, M2643.

3—6730

66 The University Science Bulletin

jungermanniaceae

8. Jamesoniella antumnalis (D. C.) Steph. A species restricted
to rather undisturbed oak-hickory wooded hillsides where it occurs
on shaded sandstone outcrops or among mosses below decaying logs.
It also is to be found on sandy clay soils and on small cliflFs in the
forest.

Due to its habitat requirements /. autumnalis occurs most fre-
quently in northeastern Kansas where it is abundant in two sites.
It has not been found in fruit and occurs as a rather large shade
form which is light green in color except in dry seasons when it has a
decided reddish tinge. It agrees well with published descriptions of
the species otherwise.

Distribution: Douglas County: 2 miles north of Baldwin City,
sandy clay soil, hillslope in mixed woods, March 30, 1952, M5325; 5
miles west of Baldwin City, shaded sandstone outcrop, April 26,
1952, M5441; 2 miles north of Baldwin, rotten log, July 19, 1952,
M5660. Leavenworth County: 4 miles northeast of Tonganoxie,
shaded sandstone cliff, June 1, 1952, M5663. Woodson County: 2
miles northwest of Yates Center, side of moist sandstone rocks,
March 29, 1949, M2651; October 28, 1951, M5295; September 27,
1952, M6072.

9. Jungermannia lanceolata L. This species has been found but
once in Kansas and then in the northeastern quarter in an area
characterized by a flora with decided east and northeastern
affinities.

A small colony of typical appearing /. lanceolata was found on
sandy soil under an overhanging sandstone cliff. The surrounding
areas, though carefully studied, failed to reveal additional specimens.
The small colony, six inches long, has maintained itself and seems
to be enlarging.

Specimens studied agree very well with published descriptions
except that they seem regularly to produce gemmae.

Distribution: Leavenworth County: 4 miles northeast of Tonga-
noxie, sandy soil under an overhanging sandstone cliff, June 1,
1952, M5579.

10. Plectocolea crenuliformis (Aust. ) Mitt. Rare in Kansas
having been found but once. This site consisted of a shaded,
moist, overhanging sandstone ledge on which the plants occurred
as pure green mats but which were reddish-brown when the growth
extended beyond the edge of the overhanging rock and were thus
exposed.

Taxonomy and Ecology of Kansas Hepaticae 67

The one collection of this species has the leaf margins swollen,
oil-bodies 7.5ij, in diameter and numbering five to ten per cell of the
leaf. In these and other characters the specimens agree with
descriptions of the species.

Distribution: Woodso7i County: 8 miles south and 4 miles
west of Yates Center, underside of moist, shaded, overhanging
sandstone ledge, September 27, 1952, M6062.

11. Plectocolea fossombronioides (Aust.) Mitt. Rare in Kan-
sas but has been collected in five counties in the eastern third of the
State. It has been found only on the sides of moist sandstone
ledges along shaded creek banks where it forms extensive green
mats often covering several square feet of rock. It seems to
tolerate severe flooding and may prefer such conditions as it
is most abundant on rock ledges inundated by high water.

Kansas specimens are clearly bisexual, have small barely evi-
dent trigones, purpHsh rhizoids and typically crenulate perianths.
These features, characteristic of the species, serve to separate
P. fossombronioides from other species of the genus in this area.

Distribution: Chautauqua County: 3 miles northeast of Sedan,
moist sandstone ledge, March 26, 1948, M1218; April 12, 1950,'
M2400a. Douglas County: 5 miles west of Baldwin City, on
moist sandstone rocks, August 26, 1948, M2312; April 22, 1950,
M4233. Elk County: 10 miles north of Busby, sandstone rocks,
July 27, 1948, M1915. Leavenworth County: 1 mile south of
Kent, moist sandstone ledge, May 15, 1948, M1424; M1425. Wood-
son County: 2 miles northwest of Yates Center, moist sandstone
ledge, April 9, 1948, M1241; September 24, 1949, M1949.

12. Plectocolea hyalina (Lyell) Mitt. This species has the
general range and distribution of P. fossombronioides. It seems,
however, to tolerate a wider range of habitat particularly in the
abihty to grow on drier sandstone rocks. It is also much more
abundant than the above species with which it occurs in most
sites studied.

Extensive mats of P. hyalina are reddish purple in early spring
but later in the season, after unfolding of tree leaves, they be-
come hght green or nearly transparent. A characteristic of the
species is the sheathing, orbicular bracts which seem to fuse
with the perianth for a distance and from between which the
perianth projects. This seems to vary in our collections from
a rather loose to a tight ensheathing condition. The length of
projection of the perianth seems also to vary. Rhizoids are
reddish in color on drier sites and light yellowish-pink on more

68 The University Science Bulletin

moist places. Spore size varies from 13-17ij. in diameter an.l
elaters are usually Q-lOij. iu diameter.

The amount of morphological variation in the species indi-
cates that a careful detailed study of the plant is desirable.
This also has been indicated by Frye and Clark (1937-1947) and
Schuster (1953).

Distribution: Chautauqua County: 1/2 miles northeast of Sedan,
on moist sandstone ledge, April 5, 1952, M5338. Douglas County:

5 miles west of Baldwin City, on moist sandstone rocks, August
26, 1948, M2313; November 7, 1948, M2575; October 15, 1949,
M4149; October 15, 1951, M5252. Leavenworth County: 6 miles
northeast of Tonganoxie, on moist sandstone ledge, August 24,
1948, M2264. Montgomery County: 3 miles southwest of Elk City,
moist sandstone ledge and sandy soil below ledge, September 22,
1951, M5227. Woodson County: 2 miles northwest of Yates
Center, moist sandy soil at base of cliff, October 28, 1951, M5277;

6 miles south and 4 miles west of Yates Center, moist sandstone
rock, September 27, 1952, M6072.

Plagiochilaceae

13. Flagiochila aspJenioides (L.) Dumort. Rare. Found once
in the extreme southeast corner of the state in the small segment
of Ozark flora which extends just into the State. It occurred on
a steep, north facing, heavily wooded cliff. A colony some three
inches long was found at the base of a large oak. The same
colony has existed for the past several years but seems barely
to survive. A thorough search of the surrounding areas failed
to reveal more plants.

Distribution: Cherokee County: 5 miles east of Baxter Springs,
base of large oak, north facing cliff, July 4, 1948, M1746.

Harpanthaceae

14. Geocahjx graveolens {Schrsid.) 'Hees. Rare. Has been found
in only three places in eastern Kansas. In each place the plants
were growing under subcalcareous conditions usually in calcareous,
sandy areas rich in humus.

The characteristic yellow-green opaque color seems to be very
outstanding in Kansas material but the perigynia produced are few
in number and are usually small and spherical. Even mature plants
do not exhibit the prominent cylindrical perigynia found in the
species in the eastern regions of this country. Otherwise the speci-
mens agree favorably with descriptions of the species.

Taxonomy and Ecology of Kansas Hepaticae 69

Distribution: Cherokee County: 5 miles east of Baxter Springs,
sandy soil on wooded hillside, October 22, 1951, M5262. Douglas
County: 2 miles north of Baldwin, sandy clay soil in rich woods,
\lay 10, 1952, M5470. Leavenworth County: 4 miles northeast
of Tonganoxie, calcareous sandstone, wooded bluff, June 1, 1952,
M5573.

LOPHOCOLEACEAE

15. Lophocolea minor Nees. Rare. Found once in the Ozark
area of extreme southeastern Kansas. It was growing among
mosses on sandy soil below sandstone and limestone outcrops. A
careful search of the area indicated that the species was not com-
mon but scattered among clumps of mosses such as Leucohryum
glaucurn. The plants produced numerous globular masses of gem-
mae and were typical of the species.

Distribution: Cherokee County: 5 miles east of Baxter Springs,
growing in clumps of Leucohryum glaucum. May 9, 1953, M7202.

16. Lophocolea hidentata (L.) Dumort. Infrequent. Found on
moist sandstone ledges and sandy-loam soil of hillsides in wood-
lands. It never is found in abundance but occurs among mosses
on sandstone ledges or among mosses and debris on hillsides.

Kansas plants of this species are consistently sterile, light green
in color and resemble the following listed species to such an extent
that they may have been overlooked in the past.

Distribution: Douglas County: 2 miles north of Baldwin City,
on decaying leaves, wooded hillside, March 30, 1952, M5323; on
sandy soil among mosses, July 19, 1952, M5661; M5663. Leaven-
worth County: 4 miles northeast of Tonganoxie, moist sandstone
ledge, June 1, 1952, M5574. Montgomery County: 3 miles south-
west of Elk City, sandy soil among mosses, September 22, 1951,
M5228. Woodson County: 2 miles northwest of Yates Center, on
moist sandstone ledge, October 28, 1951, M5294.

17. Lophocolea heterophylla (Schrad.) Dumort. Widely scat-
tered in eastern Kansas. Where found it is usually abundant but
many suitable appearing habitats are without the species. It seems
able to tolerate a large variety of habitats from moist rotting logs
to dry sandy-clay soils of oak-wooded cliffs where it may cover the
ground over large areas. It is strange that it is not more evenly
distributed in eastern Kansas as it fruits freely and succeeds very
well in some of our most common habitats.

The species shows all the usual morphological variations described
and seems typical of the taxon as generally known.

70 The University Science Bulletin

Distribution: Atchison County: 1 mile south of Oak Mills, sandy-
clay soil among mosses, July 14, 1948, M1803. Chautauqua County:
2 miles northeast of Sedan, sandy soil among mosses and lichens,
August 10, 1950, M4395. Cherokee County: 5 miles east of Baxter
Springs, among mosses on limestone ledge, April 26, 1947, M1020;
on sandy-clay soil, July 31, 1948, M1924; August 30, 1948, M2354.
Douglas County: 2 miles north of Baldwin, base of tree, on bark,
June 20, 1948, M1588; clay soil, among mosses, June 4, 1951, M4855;
sandy-loam bank, October 15, 1951, M5251; on rotten log, March
30, 1952, M5322; 5 miles west of Baldwin City, sandy soil and among
Leucobryum glaucurn and Cladonia spp., April 26, 1952, M5439,
M5469. Johnson County: 2/2 miles south of Stanley, moist calcareous
soil, wooded cliff, August 30, 1948, M2337. Leavenworth County:
4 miles northeast of Tonganoxie, on thin soil over sandstone ledge,
June 1, 1952, M5568, M5576. Linn County: 3 miles north of Pleas-
anton, on sandy-clay soil, August 1, 1948, M1949. Montgomery
County: 3 miles southwest of Elk City, sandy soil, ledge in oak
woods, September 1, 1948, M2386; September 22, 1951, M5229.
Neosho County: 2 miles east of Chanute, sandy soil, April 17,
1949, R. H. Thompson. Shawnee County: 5 miles east of Topeka,
rotten log, April 23, 1949, M2677. Wilson County: 1 mile northeast
of Neodesha, on sandy soil among mosses and lichens, August 31,
1948, M2377. Woochon County: 1 mile northwest of Yates Center,
sandy bank, October 20, 1951, M5294.

18. Chiloscyphus pallescens (Ehrh.) Dumort. Rare. One site
known for the species in the State. At this site the plant is rather
common along a sandstone cliff which is about a mile long. Here
the plant is found on the sides of moist sandstone ledges and on
sandy soil below the ledges.

Distribution: Leavenworth County: 4 miles northeast of Tonga-
noxie, on side of sandstone ledge, October 29, 1948, M2564; on sandy
soil below sandstone ledge, June 1, 1952, M5567.

19. Chiloscyphus pallescens (Ehrh.) Dumort var. /ragi/fs (Roth)
K. Miiller. Rare. Found only with the species mentioned above
but in more moist situations. The variety differs from the species
in the larger cells of the leaves, darker color and requirement for
a more moist habitat. It seems doubtful whether var. fragilis merits
taxonomic status but needs more study.

Distribution: Leavenworth County: 4 miles northeast of Tonga-
noxie, on side of very moist sandstone ledge, June 11, 1948, M1579;
June 1, 1952, M5569.

Taxonomy and Ecology of Kansas Hepaticae 71

scapaniaceae

20. DipIophyUum apictilattim (Evs.) Steigih. Rare. Found once
in the Ozark area of the extreme southeast corner of the state. It
was growing on a rocky, clay-loam wooded hillslope. Plants were
not common and occurred only in mats of Scapania nemorosa.

Distribution: Cherokee County: 5 miles east of Baxter Springs,
rocky, clay-loam hillside, among Scapania nemorosa, April 30, 1948,
i\I1361a.

21. Scapania irrigiia (Nees) Dumort. Rare. Characteristic speci-
mens of the species were found once on a very moist depression on
the side of a sandstone cliff. The plants were not numerous and
occurred only in the depression which was kept moist by a small
spring above it.

Distribution: Leavenworth County: 4 miles northeast of Tonga-
noxie, depression in sandstone cliflF, May 7, 1948, M1390.

22. Scapania nemorosa (L) Dumort. Abundant in sandstone
areas of eastern Kansas much less common on clay-loam hillsides
and absent from all but very moist calcareous sites. Where found,
S. nemorosa may form large mats on moist sandstone ledges or be
generally distributed on soil, among mosses, or on sandy banks.
Elsewhere the species forms isolated colonies among mosses.

In our area S. nemorosa usually forms abundant one-celled, reddish
or reddish-brown gemmae. However, it may often produce few
gemmae and tliese may be light green in color or scarcely pigmented.
The expression of the denticulate leaf margin is highly variable. In
some collections the leaves are sharply spinous-dentate and in others
this is barely evident. This may be correlated with amount of mois-
ture and exposure though study fails to verify this conclusively.
The latter plants are difficult to separate from S. undulata due to
these variable conditions. S. undulata occurs also with nemorosa
though it is never as common or abundant. S. imdulata, of our area,
characteristically has 2-celled, light green gemmae, a leaf margin
with a distinct border of several rows of thick walled cells, and
ventral lobes of leaf rather indistinctly decurrent. S. nemorosa has
leaves with all cells collenchymatous, the gemmae are one celled,
and reddish-brown, and the leaf lobe is distinctly decurrent.

Distribution: Chautauqua County: 3 miles northeast of Sedan,
sandy soil, March 26, 1948, M1217; May 1, 1948, M1369; July 8,
1949, M3382; April 5, 1952, M5337. Cherokee County: 5 miles
east of Baxter Springs, rocky wooded hillside, April 30, 1948,
VI1361. Douglas County: 5 miles west of Baldwin City, on

72 The University Science Bulletin

moist sandy bank, October 18, 1947, Ml 144; November 7, 1948,
M2574; June 18, 1949, M3106; September 16, 1950, M4672; April
26, 1952, M5441. Franklin County: 4 miles southwest of Ottawa,
moist sandy bank, June 18, 1949, M3098. Greenwood Coiintij:
6 miles north of Fall River, on sandstone ledge, July 27, 1948,
M1907, M1908; 1 mile east of Fall River, sandy bank, April
13, 1949 M2671; July 8, 1949, M3412. Leavenworth County:
4 miles northeast of Tonganoxie, moist sandstone ledge, May 7,
1948, M1400. Montgomery County: 6 miles south of Elk City,
moist sandy bank, March 26, 1948, M1186; 3 miles southwest of
Elk City, sandy bank, September 1, 1948, M2389; September 22,
1951, M5226. Woodson County: 2 miles northwest of Yates
Center, moist sandstone ledge, June 19, 1949, M3197; October
28, 1951, M5285.

23. Scaponia undulata (L.) Dumort. Rare. Collections of
this species have been made several times in two similar habitats.
These consist of wooded sandstone cliffs with northern exposure
and are usually very moist. All plants were found among mosses
such as Atrichwn angustaturn, Dicranum scoparium, Entodon se-
ductrix, Pohlia nutans and Leucohryum gloucum. These mosses,
with S. undulata and often S. nemorosa, form a dense mat over
moist sandy banks.

Distribution: Douglas County: 2 miles north of Baldwin City,
sandy bank, July 19, 1952, M5659. Leavenworth County: 4 miles
northeast of Tonganoxie, sandy moss covered cliffs, August 24,
1948, M2261; M2265; June 1, 1952, M5565.

Cephaloziellaceae

24. Cephaloziella byssacea (Roth) Warnstf. Rare. Found
on the crests of moist sandy ledges which are only partially
shaded. It is usually intermingled with various mosses and species
of Cladonia. This species undoubtedly has a greater distribution
than is indicated below.

Distribution: Greenwood County: 6 miles nortli of Fall River,
crest of moist, partially shaded, sandy cliff, July 27, 1948, M1913.
Montgomery County: 6 miles south of Elk City, crest of moist
sandy chff, March 26, 1948, M1187.

25. Cephaloziella hampeana (Nees) Schiffn. Rare. Found
admixed with other hepatics and mosses on moist sandstone ledges
and seepy sandy banks particularly where limestone occurs asso-
ciated with the above habitats.

Taxonomy and Ecology of Kansas Hepaticae 73

Distribution: Chautauqua County: 2 miles northeast of Sedan,
sandy soil on seepy bank, April 5, 1952, M5343. Douglas County:
2 miles north of Baldwin City, sandy subcalcareous moist bank,
July 19, 1952, M5666. Leavenworth County: 4 miles northeast
of Tonganoxie, sandy subcalcareous bank, June 1, 1952, M5575.

26. Cephaloziella rubella (Nees) Douin. Widely distributed
in the eastern half of the State and abundant in many of the sites.
It is a distinct xerophyte and occurs on denuded crests of sand-
stone cliffs which are dry through most of the year or which
dry quickly after rainfall. It also occurs in sandy or sandy-clay,
oak-hickory forest where it is found on denuded or eroded banks,
base of trees or along the edge of road cuts and trails. It
occasionally is to be found on the sides of moist sandstone ledges.
On some of the dry crests of sandstone, ledges this is the only
hepatic to be found and often forms reddish-brown or blackish
mats which may cover several square feet.

C. rubella seems to be quite variable even in the same colony.
A little colony of the species has been collected which were
mostly autoecious. Such plants would be referred to Cephaloziella
rubella (Nees) Douin var. bifida Lindb.; however, the character
is not constant and a colony usually shows some autoecious plants
though mostly paroecious plants are to be found. This condition
is apparently not related to moisture or exposure factors.

In some of the more moist sites and particularly those in
which more humus is found one encounters plants which might
be referred to C. rubella var. sullivantii (Aust.) K. Miiller. They
have leaf cells 9-12tjL wide and walls which are a little thinner
than in C rubella. Also the involucral bracts form a ring of 7-9
lobes and are dentate on the margin while in C. rubella the ring
has 5-6 lobes which are only slightly dentate to entire. How-
ever, none of the specimens show the minute underleaves of
var. sullivanti and since the other characters seem to be quite
variable it seems best not to segregate specimens from Kansas
into varieties. If such segregation were attempted it would mean
that var. rubella, var. bifida and var. sullivantii would all occur
in the same colony.

Distribution: Cherokee County: 5 miles east of Baxter Springs,
moist sandy-clay soil, crest of cliff, October 22, 1951, M5270. Doug-
las County: 12 miles south of Lawrence, dry sandy crest of ledge,
October 13, 1951, M5243, M5250; 5 miles west of Baldwin City,
ledge above sandstone outcrop, March 12, 1952, M5321, M5324;

74 The University Science Bulletin

April 26, 1952, M5432; 4 miles north and 5 miles east of Baldwin
City, edge of road cut in sandy oak woods, May 3, 1952, M5460.
Ellsworth County: 2 miles north and 4/2 miles west of Langley,
bare sandy soil, crest of cliff, May 3, 1952, M5444, M5451. Leav-
enworth Countij: 4 miles northeast of Tonganoxie, sandy bank in
oak-hickory woods, June 1, 1952, M5564, M5565. Montgomery
County: 3 miles southwest of Elk City, on side of dry hmestone
ledge, September 22, 1951, M5224, M5225, M5226. Washington
County: 2 miles west of Linn, dry sandy soil over surfacing sand-
stone, August 22, 1952, M5717, M5721. Woodson County: 2 miles
northwest of Yates Center, side of dry ledge, October 28, 1951,
M5276, M5286; 8 miles south and 4 miles west of Yates Center,
side of sandstone rock, September 27, 1952, M6066.

Cephaloziaceae

27. Cephalozia bicuspidata (L. ) Dumort. Rare. Found on moist
sandstone ledges usually on the under side of overhanging rocks
which are kept moist by seepage. It was discovered once among
mosses on a sandy-clay hillside in oak-hickory forest.

Distribution: Douglas County: 2 miles north of Baldwin City,
sandy-clay soil, among mosses, July 19, 1952, M5662. Woodson
County: Lake Fegan, Woodson County State Park, on underside
of moist, overhanging sandstone ledge, March 26, 1949, M2644,
M2645; September 24, 1949, M4131.

28. Cephalozia catenulata (Huben.) Spr. Rare. This species
was found once on the side of a moist, sandy-clay bank in an oak-
hickory woods. The plants formed an extensive mat on the shaded
portions of these banks.

Distribution: Woodson County: Lake Fegan, Woodson County
State Park, sandy-clay bank in oak-hickory woods, March 26, 1949,
M2657.

29. Cephalozia connivens (Dicks) Lindb. Rare. Found once
on a north-facing sandstone cliff where the plants occurred on
sandy soil among mosses and lichens. A careful search showed
the plant to be rare in the area.

Distribution: Leavenworth County: 4 miles northeast of Ton-
ganoxie, north-facing sandstone bluff, on sandy soil, June 1, 1952,
M5577.

30. Cephalozia media Lindb. Rare. Collected from a moist
sandy bank and again from the side of a nearby sandstone ledge.
The plants occurred in considerable quantity on these two sites.

Taxonomy and Ecology of Kansas Hepaticae 75

Distribution: Woodson County: Lake Fegan, Woodson County
State Park, on moist sandy-clay bank, March 26, 1949, M2642; on
side of moist sandstone rocks, September 24, 1949, M4132.

Calypogeiaceae

31. CoUjpogeia meylanii Buch. Rare. Found once on the side
of a sandstone ledge on a north-facing, oak-hickory wooded blufiF.
The plants were very rare in the area.

Distribution: Leavenworth County: 4 miles northeast of Tonga-
noxie, on moist sandstone ledge, October 3, 1953, M7521.

32. Calypogeia muelleriana (Schiffn.) K. Miiller. Rare. Found
once on the side of a moist sandstone ledge in a deep ravine running
through an oak wooded area. The plants were rather generally
distributed along the ledges in the ravine.

Specimens of this species in our collections have colorless oil
bodies and seem close to C. meylanii in many respects; but after a
detailed study it was finally decided to recognize our specimens
as indicated. However, a detailed study of the species in this
country seems to be necessary.

Distribution: Woodson County: 8 miles south and 4 miles west
of Yates Center, on moist sandstone ledges, September 27, 1952,
M6063, M6065, i\I6073, M6074, M6077, M6078.

33. Calypogeia trichomanis (L. ) Corda. Rare. This species
occurs in the same general habitat as do the previous listed .species
of the genus. However, it seems to be more prevalent and perhaps
a little more dispersed. The blue-colored, oil bodies in leaf cells
separate this species readily from all closely related forms.

Distribution: Leavenworth County: 4 miles northeast of Tonga-
noxie, sandy ledge, June 1, 1952, M5571, M5572, M5580. Woodson
County: Lake Fegan, Woodson County State Park, moist sandstone
ledge, September 24, 1949, M4133.

PORELLACEAE

34. Porella pinnata L. Scattered in the southeastern quarter
of the State. The species is found in two very diflFerent types of
habitat. One is on the bark of trees or on rocks at water level of
creeks in the extreme southeast corner of the State. In such places
the plants may be partially submerged. Usually they are found
between water level and flood level. The second habitat is on
dry limestone outcrops on cliffs. This is not a common habitat
but plants have been found in two different places on these very

76 The University Science Bulletin

dry outcrops. In the latter sites P. pinnata is associated with P.
phtyplujUa and P. platijplujUoiclea.

Morphologically P. pinnata on bark at water level and on dry
limestone outcrops exhibit slight differences which one expects in
plants occupying such diverse habitats. A careful study of the
species over its entire range is, perhaps, desirable.

Distribution: Anderson County: 5 miles south of Garnett, on
dry limestone outcrop, October 8, 1948, M2485. Bourbon County:
2 miles south of Uniontown, on dry limestone outcrops, October 9,
1948, M2515. Cherokee County: 5 miles east of Baxter Springs, on
rocks, bank of Shoal Creek, July 5, 1948, M1747; bark of birch,
July 31, 1948, M1946; bark of sycamore, October 22, 1951, M5255.

35. Porella platyphylla (L. ) Lindb. Scattered in the eastern
third of the State particularly in the southern half. In some places
the species forms large mats on limestone outcrops. Abundance
depends upon moisture and shade. Dry exposed sites are without
the species and it becomes more common and abundant as mois-
ture increases.

In Kansas P. platyphylla occurs in exactly the same type of habi-
tat as does P. platyphylloidea though they are seldom found to-
gether. Schuster (1953) says that P. platyphylla is by far the
less frequent species in North America. However, in Kansas, we
find P. platyphylloidea only a little more common than P. platy-
phylla.

Morphologically the two species are quite similar and Frye and
Clark (1937-1947) reduced P. platyphylloidea to a variety of P.
platyphylla. Difficulties are met with in Kansas material in that
neither of the taxa have been found fruiting. This is apparently
true elsewhere resulting in incomplete studies being made on one
of the common species. Our determinations have been made on
mature vegetative plants. In our experience it is only the juvenile
plants or young shoots that seem to intergrade. Our findings agree
very closely with those given by Schuster (1953) on the complex.

Distribution: Allen County: 1 mile south of Humboldt, on dry
limestone ledge, July 7, 1949, M3338. Bourbon County: 1 mile
south of Uniontown, on limestone outcrop, July 7, 1949, M3347.
Cowley County: 2 miles southeast of Arkansas City, on limestone
ledge, September 2, 1950, M4646. Doiiiphan County: 2 miles
northwest of Iowa Point, Hmestone outcrop, August 2, 1948, M1972.
Greenwood County: 4 miles west of Fall Biver, limestone ledge,

Taxonomy axd Ecology of Kansas Hepaticae 77

April 13, 1949, M2673. Johnson County: 5 miles south of Stanley,
on limestone outcrop, August 30, 1948, M2330; August 3, 1950,
M4388. Miami County: Murray Lake, limestone rocks, October 9,
1948, M2530. Montgomery County: 4 miles east of Elk City, on
limestone outcrop, April 12, 1949, M2667. Wilson County: 3 miles
southwest of Neodesha, on limestone ledge, April 13, 1949, M2668.

36. Porella platyplujlloidea (Schwein.) Lindb. Distributed
over the eastern third of Kansas. Its habitat requirements are
so similar to those of P. platyphylla that no further mention of the
subject need be given. Morphological information of this species
is likewise considered with P. platyphylla.

Distribution: Anderson County: 2 miles south of Garnett, on
limestone outcrop, April 13, 1949, M2674. Bourbon County: 2
miles south of Uniontown, moist limestone ledge, July 5, 1948,
M1775. Chautauqua County: 2 miles southwest of Sedan, on lime-
stone outcrop, April 12, 1949, M2666. Cherokee County: 5 miles
west of Baxter Springs, on limestone rocks, April 25, 1942, R. H.
Thompson; on limestone outcrop, April 12, 1949, M2662. Douglas
County: 2 miles north of Clinton, on limestone ledge, September
10, 1948, M2443. Franklin County: 1 mile southeast of Ottawa,
on limestone outcrop, June 28, 1949, M3312. Leavenworth County:
5 miles northwest of Leavenworth, on limestone outcrop, September
7, 1948, M2420; 2 miles north of Leavenworth, on limestone out-
crop, October 29, 1948, M2565. Linn County: 2 miles north of
Trading Post, on Hmestone outcrop, April 11, 1949, M2659. Mont-
gomery County: 8 miles west of Sycamore, on limestone outcrop,
September 1, 1948, M2406; March 30, 1952, M5327.

Lejeuneaceae

37. Cololejeunea biddlecomiae (Aust. ) Evs. Rare. Found in
three mesic woodlands in eastern Kansas. In each case the plants
were growing on the bark of trees at the base of the trunk and in
shaded rather moist conditions. Two of these sites were on calcar-
eous cliffs and the third was in a sandstone canyon.

Distribution: Cherokee County: 5 miles east of Baxter Springs,
limestone cliff, on bark at base of hackberry tree, Oct. 22, 1951,
M5259. Leavenworth County: north edge of Military Reservation,
limestone bluff, bark of burr oak, Nov. 28, 1948, M2619. Woodson
County: 8 miles south and 4 miles west of Yates Center, sandstone
canyon, bark of Asimina triloba, September 27, 1952, M6067.

78 The University Science Bulletin

Frullaniaceae

38. Fnillania asagraijana Mont. Rare. Found but once in
the state and that in a mesic forest on the bkiffs of the Missouri
River. In this local forest, plants of F. asagraijana were found on
limestone ledges which are dry most of the year but have a period
of heavy seepage over them in early spring.

Distribution: Leavenworth County: 2 miles north of Leaven-
worth, on limestone ledge, October 29, 1948, M2566.

39. Fnillania brittoniae Evs. Common in the Ozark area of
southeastern Kansas but known from only one station outside of
that restricted habitat. This species has been found only on the
bark of trees.

Distribution: Cherokee County: 5 miles east of Baxter Springs,
bark of Quercus alba, July 31, 1948, M1930, M1932; October 22,
1951, M5257; bark of Betula nigra, M5269. Woodson County: 2
miles northwest of Yates Center, bark of Quercus macrocarpa,
October 28, 1951, M5288.

40. Frullania ehoracensis Gottsche. To be found in nearly all
mixed forests in the eastern fourth of the state. It is limited exclu-
sively to occurrence on the bark of trees especially oak. In gen-
eral a lowland or flood plain forest supports an abundance of this
species and it becomes less common as drier forest conditions are
reached. The species seems also to be more common and abundant
on forests of calcareous soils than in those on sandy soils.

In most of our material there is a decided tendency for the
plants to have caducous leaves and thus to appear leafless in late
summer or early spring. Otherwise the species exhibits the mor-
phological structures described for the taxon. In extreme south-
eastern Kansas F. eboracensis is often difficult to separate from
F. brittoniae especially when sterile or juvenile; however, they can
usually be separated on the underleaf, the upper halves of which,
in F. brittoniae, are usually dentate or dentate-crenate and are
entire or bear one tooth in F. ehoracensis. The characters of the
inflated lobules as given by Schuster (1953) to separate the two
species seem to intergrade too much to be of value in our area.

Distribution: Atchison County: 1 mile east of Potter, bark of
Quercus macrocarpa, June 2, 1949, M2816. Brown County: 5 miles
west of Hiawatha, bark of Quercus macrocarpa, June 3, 1949,
M2826. Chautauqua County: 2 miles northeast of Sedan, bark of
Quercus stellata, April 12, 1950, M4196. Cherokee County: 5 miles
east of Baxter Springs, bark of Quercus alba, July 3, 1948, M1627,

Taxonomy and Ecology of Kansas Hepaticae 79

M1628; bark of fallen oak, July 4, 1948, M1745; bark of Quercus
rubra, October 9, 1948, M2522; bark of Betula nigra, October 22,
1951, M5254. Coffey County: 4 miles west of Waverly, bark of
Quercus velutina, June 27, 1948, M1601. Cowley County: 2 miles
southeast of Arkansas City, bark of Quercus macrocarpa, September
2, 1950, M4645. Douglas County: 2 miles north of Baldwin City,
bark of Quercus alba, May 29, 1948, M1516; 4 miles northwest of
Baldwin City, bark of Quercus macrocarpa, August 26, 1948,
M2303; 5 miles west of Baldwin City, bark of Quercus velutina,
December 28, 1948, M2620; bark of Ulmus rubra, September 18,

1949, M4077; bark of Platanus occidentalis, July 19, 1952, M5658.
Labette County: 4 miles northwest of Chetopa, bark of Ulmus
americana, July 2, 1948, M1612. Leavenworth County: 1 mile
south of Kent, bark of Quercus macrocarpa, June 11, 1948, M1581;
bark of Quercus muehlenbergii, June 11, 1948, M1582. Miami
County: 3 miles east of Fontana, bark of Quercus macrocarpa,
October 10, 1948, M2531; bark of Quercus muehlenbergii, October
8, 1950, M4683; bark of Carya illinoensis, October 8, 1950, M4686;
bark of Ulmus rubra, October 27, 1950, M4690; bark of Fraxinus
lanceolata, October 27, 1950, M4691. Neosho County: 2 miles east
of Chanute, bark of oaks, April 17, 1949, R. H. Thompson. Shawnee
County: 5 miles east of Topeka, bark of Quercus macrocarpa and
Ulmus americana, April 23, 1949, M2675. Sedgwick County: 2
miles northwest of Wichita, bark of Ulmus americana, September 3,

1950, M4648. Woodson County: 2 miles northwest of Yates Center,
bark of Quercus velutina, Q. stellata, Q. marilandica and Celtis
occidentalis, October 28, 1951, M5287; bark of Quercus macro-
carpa, September 27, 1952, M6081.

41. Frullania inflata Gottsche. This species is without doubt
the most abundant hepatic in the state. It is found in all of the
eastern half where it occurs on the bark of many trees, moist lime-
stone cliffs and often on the sides of sandstone ledges. It is most
abundant in the elm floodplain forests along creeks and rivers and
only slightly less frequent in the mixed forests on river bluffs and
hillsides. It extends westward into the mid-grass prairies as far as
Quercus macrocarpa can be found along the streams of that area.

Frullania inflata is highly variable in our area. It fruits abun-
dantly and mature capsules are to be found from January to July in
seasons following average or above average rainfall but never
fruits after a very dry year. The most variable structure seems to
be the lobules. These are characteristically inflated but explanate

80 The University Science Bltlletin

or lingulate lobes are nearly always present on young parts. The
drier the habitat the more frequent the inflated lobule while the
more moist the habitat, such as seepy rock ledges, the more abun-
dant are the lingulate lobules. The size of the lobule varies from
Vr,-y2 of the size of the dorsal lobe and sometimes almost equals the
dorsal lobe in size. Such plants from Kansas have been named by
some, Frullania oakesiana Aust., a northern species, not yet found
in Kansas.

Distribution: Allen County: 1 mile south of Humboldt, on dry
limestone ledge, July 7, 1949, M3337. Anderson County: 3 miles
north of Garnett, on bark of Uhnus rubra, August 1, 1948, M1960;
2 miles south of Garnett, on Ulmus rubra, June 4, 1951, M4878.
Bourbon County: 2 miles south of Uniontown, bark of Quercus
rubra, October 8, 1948, M2519; 1 mile south of Uniontown, on
limestone outcrop, July 7, 1949, M3346; October 22, 1951, M5271.
Chase County: 1 mile south of Strong City, bark of Quercus
macrocarpa, November 27, 1948, M2602. Butler County: 5 miles
north of El Dorado, bark of Juglans nigra, August 18, 1953, M2251.
Chautauqua County: 2 miles northeast of Sedan, on bark of
Quercus stellata, April 5, 1953, M5334. Cherokee County: 5 miles
east of Baxter Springs, limestone outcrop, July 3, 1948, M1649; July
7, 1949, M3363; bark of Quercus macrocarpa, October 22, 1951,
M5267; bark of Celtis occidentalis, July 8, 1952, M5650; bark of
Quercus alba. May 9, 1953, M7200. Cowley County: 2 miles
southeast of Arkansas City, bark of Quercus macrocarpa, September
2, 1950, M4644. Crawford County: Crawford County State Park,
bark of Ulmus rubra, July 5, 1948, M1753; October 9, 1948, M2529.
Dickinson County: 6 miles northwest of Herington, bark of
Quercus macrocarpa, June 24, 1950, M4357. Doniphan County: 1
mile northwest of Iowa Point, bark of Ulmus rubra, August 2,
1948, M1965. Douglas County: 6 miles northwest of Lawrence,
bark of Ulmus americana, August 26, 1948, M2275; Lone Star Lake,
bark of Quercus macrocarpa, August 28, 1948, M2320; 2 miles
northwest of Clinton, bark of Quercus muehlenbergii, September
10, 1948, M2447; 4 miles northwest of Lawrence, bark of Quercus
muehlenbergii, August 19, 1950, M4483; 2 miles east of Lecompton,
bark of elm, July 13, 1949, R. H. Thompson; 2 miles north of
Baldwin City, bark of Quercus velutina, April 25, 1953, M7197.
Franklin County: 4 miles east of Princeton, bark of Quercus mac-
rocarpa, May 29, 1949, M2763; 2 miles south of Lane, on dry
limestone rocks, June 5, 1952, M5589. Harvey County: 1 mile
east of Annelly, bark of Quercus macrocarpa, August 19, 1948,

Taxonomy and Ecology of Kansas Hepaticae 81

M2254. Jefferson County: 1 mile north of Valley Falls, bark of
Qucrcus macrocarpa, January 22, 1953, M7000. Jewell County:
4 miles north of Burr Oak, on bark of Quercus macrocarpa, Sep-
tember 5, 1950, M4653. Johnson County: 5 miles south of Stanley,
bark of Quercus vehitina, August 30, 1948, M2335; on moist lime-
stone rocks, August 3, 1950, M4387. Labette County: 4 miles
northwest of Chetopa, bark of Betula nigra, July 2, 1948, M1616.
Leavenworth County: 1 mile south of Kent, bark of Quercus
macrocarpa, June 11, 1948, \I1580. Linn County: 3 miles north
of Pleasanton, on limestone outcrop, August 1, 1948, M1950. Lyon
County: 4 miles southwest of Emporia, bark of Quercus macro-
carpa, June 26, 1948, M1592. Marion County: 1 mile south of
Florence, bark of Ulmus americana, November 27, 1948, M2613.
Marshall County: 8 miles north of Frankfort, bark of Quercus
macrocarpa, August 9, 1948, M2032. McPherson County: 6 miles
north of Conway, bark of Ulmus rubra, July 24, 1948, M1819.
Miami County: 2 miles southeast of Fontana, bark of Quercus
macrocarpa, November 12, 1948, M2580. Montgomery County:
8 miles west of Sycamore, on limestone outcrop, September 1, 1948,
M2307; bark of Quercus macrocarpa, September 1, 1948, M2408
Morris County: 2 miles north of Council Crove, bark of Ulmus
americana, November 27, 1948, M2618. 'Nemaha County: Nemaha
County State Park, bark of Quercus macrocarpa, August 9, 1948,
M2013. Riley County: 4 miles south of Manhattan, on bark of
Quercus macrocarpa, June 24, 1953, M7480. Sedgwick County:
2 miles north of Wichita, bark of Ulmus americana, September 3,
1950, M4647. Shawnee County: 5 miles east of Topeka, bark of
trees, April 23, 1949, M2681. Washington County: 2 miles west
of Linn, bark of Quercus macrocarpa, August 22, 1953, M5714.
Wabaunsee County: 5 miles west of Eskridge, bark of Ulmus
americana, July 25, 1948, M1857. Wilson County: 1 mile north-
east of Neodesha, bark of Quercus macrocarpa, August 31, 1948,
M2379; 1 mile east of Altoona, bark of Quercus velutina, August
29, 1952, M5726. Woodson County: 2 miles northwest of Yates
Center, bark of Quercus velutina, July 5, 1948, M1792; March 26,
1949, M2640; 8 miles south and 4 miles west of Yates Center, bark
of Quercus siellata, September 27, 1952, M6082, M6070.

42. Frullania kimzei Lehm. and Lindenb. Rare. Known from
one collection from the Ozark area of the extreme southeastern
corner of the State. Here it was found on the bark of Quercus
alba. Our collection probably represents the most northwestern
record of the species.

82 The University Science Bulletin

Distribution: Cherokee County: 5 miles east of Baxter Springs,
on bark of Qiiercus alba, October 22, 1951, M5256.

43. Frtillania riparia Hampe. Rather frequent in the eastern
half of Kansas where it is found on limestone ledges and on tree
bark in forests occurring on limestone hillsides. It seems to be
entirely absent from sandstone areas except for an occasional
occurence on bark of Qiierctis niacrocarpa. Kansas specimens of
this species are often very difficult to separate from FruUania sqtiar-
rosa. In seasons of average or above average rainfall most material
seems easily referable to F. riparia. However, on very dry sites
and during dry years such as in 1952-53 most plants seem to fall
within the range of F. squarrosa. The possibility that F. squarrosa
is a xeric extreme and F. riparia a mesic extreme has been suggested
by Schuster (1953), and Kansas material seems to show that this
may be the case. In Kansas F. riparia, in mesic conditions, shows
rather small trigones and has no intermediate thickenings, the
lobules, being mostly explanate or ligulate, the leaves rather lax
and distant to underleaves. The color is usually green or reddish-
green. In xeric conditions F. riparia is reddish-brown or copperish
in color, the leaves somewhat squarrose, having large trigones,
and the underleaves large. Often the leaves are also imbricated.
The only character in these xeric plants which is absent, but present
in F. squarrosa, is the intermediate thickening between trigones.
None of our plants of the two extremes have been found with
mature capsules.

The above facts indicate, as did the findings of Schuster (1953),
that a careful detailed study of F. riparia over its range is needed.
Furthermore, careful ecological data should be taken along with
specimens. It seems possible that the finding of mature capsules
of F. riparia might help solve the problem.

Distribution: Allen County: 2 miles south of lola, bark of Quer-
cus macrocarpa, July 5, 1948, M1789. Bourbon County: 5 miles
north of Hepler, May 30, 1949, L. J. Gier, 1864; 2 miles south of
Uniontown, on limestone rocks, July 5, 1948, M1774. Chautauqua
County: 2 miles southwest of Sedan, on Hmestone rocks, April 12,
1949, M2664; bark of Quercus macrocarpa, April 12, 1949, M2665;
1 mile north of Sedan, on bark of Quercus stellata, April 5, 1952,
M5335. Cherokee County: 5 miles east of Baxter Springs, on bark
of Quercus alba, July 3, 1948, M1655; on hmestone rocks, April 12,
1949, M2663; October 22, 1951, M5260; bark of Quercus macro-
carpa, October 22, 1953, M5261. Douglas County: Lone Star Lake,
bark of Ulmus rubra, May 15, 1948, M1445; 2 miles north of Baldwin

Taxonomy and Ecology of Kansas Hepaticae 83

City, on bark of Qiiercus alba. May 29, 1948, M1517; 2 miles north-
west of Clinton, limestone ledge, September 10, 1948, M2444; 1 mile
west of Pleasant Grove, tree bark. May 8, 1949, R. H. Thompson;
2 miles north of Baldwin City, on bark of Ostrya virginiana, Decem-
ber 8, 1951, M5295. Jefferson County: 2 miles west and 5 miles
south of McLouth, bark of Qiiercus macrocarpa, October 15, 1949,
M4155. Johnson County: 2/2 miles south of Stanley, on limestone
outcrop, August 3, 1950, M4387; April 11, 1949, M2656. Linn
County: 2 miles north of Trading Post, on limestone outcrop, April
11, 1949, M2660. Miami County: Miami County State Park, on
bark of Quercus macrocarpa, October 3, 1948, M2480; 2 miles south-
east of Fontana, bark of Quercus muehlenbergii, November 12,

1948, M2581; bark of Quercus velutina, October 17, 1949, M4159,
M4160. Montgomery County: 8 miles west of Sycamore, on side
of limestone outcrop, September 1, 1948, M2409; April 5, 1952,
M5328. Osage County: 7 miles south of Overbrook, bark of Celtis
laevigata, February 22, 1953, R. H. Thompson. Shawnee County:
5 miles east of Topeka, bark of Quercus macrocarpa, April 23, 1949,
M2682. Wabaunsee County: 6 miles northwest of Pasco, bark of
Quercus macrocarpa, June 25, 1950, M4364. Wilson County: 3
miles west of Neodesha, bark of Quercus macrocarpa, April 13,

1949, M2660; 1 mile east of Altoona, bark of Carya ovata, August
29, 1952, M5727.

44, FruUania squarrosa (R., Bl. N.) Dumort. Two collections of
FruUania have been definitely assigned to this taxon. In both cases
the specimens match perfectly the descriptions given for the species
and in addition were fruiting. In Kansas the species is difficult t(j
separate from FruUania riparia and the subject is discussed under
F. riparia.

Distribution: Douglas County: 1 mile northwest of Clinton, on
limestone ledge, September 26, 1948, M2464. Leavenworth County:
2 miles north of Leavenworth, bark of Quercus macrocarpa, October
29, 1948, M2567.

FOSSOMBRONIACEAE

45. Fossombronia brasiliensis Steph. Uncommon. A species
found on moist sandy soil in bluestem prairie, over surfacing sand-
stone, sides of moist sandstone ledges and along sandy paths or
roadside cuts. It prefers exposed or recently eroded sites. Where
found it is usually abundant, though it may be present in an area
one season and absent in the next. It, like Sphaerocarpus texanus,

84 The University Science Bulletin

is most abundant in a spring following a very moist summer, fall
and winter.

F. hrasiliensis, in Kansas, seems to be a variable species particu-
larly as regards the mature spores. Frye and Clark (1937-1947)
give the size of the mature spores as 41-48[j. while we find them to
range from 29-59[;.. In one collection (M1358) the size range was
33-42[ji. with the average being 39;j. or some 2[j. below the minimum
given for the species. Another large collection (M1269) has spores
ranging from 29-40[a with the average being 35[j. or some 6[j, below
the minimum. This is in contrast with a third collection (M5339)
which has a spore size range of 46-59[jt., an average size of 52[j. which
is 4;j. above the maximum described for the species. All of our
spores have some perfectly formed meshes on the convex face.
These meshes are, however, very irregular even on spores from
the same sporangium. The number of meshes varies from 4-14 and
the size of mesh varies from 3.3-16[x. Frye and Clark [I.e.) state
that some spores are regularly reticulate with 11-12 meshes on the
convex face while usually spores have irregularly forked lamellae
without distinct meshes. None of our spores, then, agree with the
descriptions given for the species.

Another character of the spore which seems at variance is the
height of lamellae. Frye and Clark ( /. c. ) say the lamellae are low
and illustrate a spore in which lamellae are low. Our spores have
prominent lamellae which are up to 4.9[;. high. Sharp ( 1939 ) men-
tions in his key to Fossombronia species that the lamellae are high.

The inner faces of the spores are not mentioned in descriptions
of the species. In ours the inner faces are smooth, minutely granu-
late or occasionally very faintly ridged.

The pseudoperianth of F. hrasiliensis is described as turbinate,
about 1.5 mm. high, deeply cleft on one side, crispate at mouth,
deeply and irregularly sinuate lobed; the lobes rounded and entire.
This in general matches our material except that the lobes are often
more acute than rounded, have a few dentate teeth, and the sinuate
lobing appears to be quite regular. Another feature of our turbinate
pseudoperianths is the presence of a few lateral scales, usually red-
dish in color, on the sides of the structure near the base.

In many respects our F. hrasiliensis seems very close to Fossom-
bronia foveolata. Frye and Clark (1937-1947) separate F. hrasili-
ensis from F. foveolata by the former having either no meshes on
the convex surface or only 0-13 while foveolata has 15-20. Our
material seems to be intermediate between the two in these re-

Taxonomy and Ecology of Kansas Hepaticae 85

spects. Sharp (1939) separates the two by keying F. hrasiliemis
as having ridges on spores high, enclosing depressed areas 10-13^/.
in diameter and F. foveolata as having ridges on spores low, en-
closing depressed areas 7-9ij, in diameter. Again our material will
not key out following such characters.

A comparison of our material with that from other areas reveals
even more difficulties. Collections of F. foveolata by the writer,
from Arkansas, Louisiana and Texas and by Schuster from Minne-
sota and Mississippi have been studied. These show the meshes
of F. foveolata to be far less regular than described for the species.
Spores range in size from 32-61[j. in diameter and ridges are from
0-4. 2u. high. The pseudoperianth also varies and grades into that
of F. hrasiliensis. The only constant differences now observed
between the species is a tendency for the spores of F. foveolata to
have ridges lower, and less dark in color than in F. hrasiliensis and
for meshes to be a little more regular in F. foveolata. Vegetative
characters of the two species seem to be similar and equally \ari-
able.

From evidence at hand it would seem that F. hrasiliensis should
be treated as a variety or forma of F. foveolata. Until more ma-
terial is available for study, however, it seems best not to change
taxonomic rank of the plants involved at this time.

The study of Fossomhronia species in Kansas is handicapped by
the short fruiting period, which lasts barely two weeks in any given
season. Consequently sterile plants have been found frequently
which could not be identified. Most of these would probably come
within the present concept of F. hrasiliensis.

Distribution: Chautauqua County: 2 miles northeast of Sedan,
on side of moist sandstone rock ledge, April 5, 1952, M5339. Leav-
enworth County: 4 miles northeast of Tonganoxie, sandy soil in
prairie, May 15, 1948, M1438. Montgomery County: 3 miles south-
west of Elk City, sandy path in oak woods, April 10, 1948, M1269.
Wilson County: 1 mile northeast of Neodesha, sandy soil over sur-
facing sandstone, April 10, 1948, M1271. Woodson County: 2
miles northwest of Yates Center, sandy soil of roadside cut, May 12,
1951, M4812.

In addition to the above records sterile material of Fossomhronia
has been found in Allen, Cherokee, Douglas, Franklin, Greenwood.
Jefferson, Linn and Neosho counties. As mentioned above most of
these plants are probably F. hrasiliensis.

46. Fossomhronia foveolata Lindb. Rare. Habitat of the above
species. Three collections of Fossomhronia are referred to this con-

86 The University Science Bulletin

cept because they seem to match descriptions and specimens of the
species within certain Hmits. For a discussion of this species see
under F. brasiliensis.

Distribution: Chautauqua County: 2 miles north of Sedan, sandy
soil in forest clearing, April 5, 1952, M5344. Cherokee County:
5 miles east of Baxter Springs, sandy-clay soil, path in oak woods,
April 30, 1948, M1358. Woodson County: Lake Fegan, on moist
sandstone ledge, April 24, 1949, M4134a.

Pelliaceae

47. Pellia epiphylla (L.) Corda. Rare. This species was found
once in the Ozark area of extreme southeastern Kansas where it
occurred on a seepy limestone ledge in a shaded ravine. Plants
were abundant on the ledge for a distance of several feet. A search
of nearby similar sites failed to reveal more plants and repeated
visits to the first collection point, in subsequent years, have indicated
that the colony has disappeared. The site from which the collection
was made regularly dries up each summer, and is covered by leaves
and other debris. It was burned over in early spring following the
season in which the colony was discovered. This may have been
responsible for its disappearance.

Distribution: Cherokee County: 5 miles east of Baxter Springs,
moist limestone ledge, July 31, 1948, M1933.

RiCCARDIACEAE

48. Riccardia pinguis (L.) Gray. Rare. The known range of
this species in Kansas is truly remarkable. It was found once in
mouse runways in a cattail-sedge marsh in Meade County. At this
site the marsh is due to artesian water and is bounded by buffalo-
grass or short-grass prairie. The other site is in Montgomery
County where the plants are found on the sides of a small, moist,
sandy ditch below a spring in an oak forest. In the latter habitat
the plants have been observed year after year; but a search of other
similar locations in the same area and elsewhere has failed to turn
up additional stations for the species.

Distribution: Meade County: Meade County State Park, mouse
runways in cattail-sedge marsh. May 8, 1942, W. H. Horr. Mont-
gomery County: 3 miles southwest of Elk City, sandy, moist ditch
below a spring, July 1, 1947, M1270; September 22, 1951, M5222;
October 9, 1953, M7311.

Taxonomy and Ecology of Kansas Hepaticae 87

Rebouliaceae

49. Asterella tenella (L. ) Beauv. Scattered in the eastern half
of Kansas where it seems to be Hmited to sandy soils and sandstone
ledges. Where found it is usually very abundant though it is absent
from many stations where the habitat seems suitable in every way.

AstereUa tenella is often found on sandy soils on prairie hillsides
dominated by Andropogon gerardi, Boiiteloua ciirtipendtda and in
which clumps of the cactus Opiintia humifusa are conspicuous. In
such places it is extremely xerophytic. In the same area it is found
on the underside of overhanging, dripping sandstone ledges which
occur along prairie ravines or creeks. The species seem to exist
better in the more xeric stations and certainly fruits more abundantly
on such sites.

Distribution: Chautauqua County: 3 miles northeast of Sedan,
sandstone ledge, May 1, 1948, M1366; 3 miles north of Elgin, sandy
prairie hillslope, April 5, 1952, M5347. Cherokee County: 5 miles
east of Baxter Springs, sandy clay soil, clearing in oak woods, April
30, 1948, M1359. Clay County: 5 miles north of Morganville,
sandy prairie hillside, August 16, 1951, M5064. Douglas County:
12 miles south of Lawrence, sandy bank, April 18, 1948, M1337.
Ellsworth County: 5 miles east of Carneiro, sandy soil under sand-
stone ledge, April 30, 1949, M2702. Labette County: 8 miles north-
east of Cherry vale, sandy prairie bank, June 30, 1947, M1067. Leav-
enworth County: 4 miles northeast of Tonganoxie, sandstone ledge,
May 7, 1948, M1402. Montgomery County: 3 miles southwest of
Elk City, sandstone ledge, and sandy prairie hillside, July 1, 1947,
M1069. Neosho County: 4 miles northeast of Morehead, sandy
roadside bank, June 30, 1947, M1066. Rice County: 1 mile east
of Geneseo, sandy prairie bank, April 30, 1949, M2714. Saline
County: 2 miles northwest of Brookville, sandy prairie ravine,
April 30, 1949, M2694. Wilson County: 1 mile northeast of Neode-
sha, thin sandy soil over surfacing sandstone. May 1, 1948, M1381.
Woodson County: Lake Fegan, under overhanging sandstone ledge,
September 24, 1949, M4130. Wyandotte County: 1 mile northeast
of Bonner Springs, sandy bank in road cut. May 15, 1948, M1439.

50. Mannia fragrans (Balb.) Frye and Clark. Abundant on a
number of sites in the eastern half of Kansas where it is found on dry
exposed sandy prairie hillsides and on moist sandstone ledges which
are moist only in the spring. The species is very xerophytic and
occurs only on sandy soil or sandstone.

A characteristic habitat for M. fragrans consists of prairie hill-

88 The Unr'ersity Science Bulletin

sides characterized by a mixture of Andropogon gerardi, Andro-
pogon scoparius, Bouteloua curtipenduJa, Anemone caroliniana,
Opuntia humifusa and thickets of Rhus glabra. It is also commonly
found in mixed prairie of the above plants with Quercus stellata, Q.
marilandica and Q. velutina.

The plants fruit abundantly from April to June after which they
become inconspicuous, since tlie drying, shriveling, thallus margins
roll up and overlap or become strongly incurved exposing the very
black-purplish imderside. At such times the plants are easily
overlooked as they are covered by grass, dust, or otherwise partially
hidden. The application of a bucket of water causes them to unroll
quickly and after early fall rains they unroll and continue growth.

Kansas material of this species varies considerably in the position
and arrangement of antheridia. The description of the antheridia
as given by Frye and Clark ( 1937-1947 ) states that the antheridial
receptacle is sessile, distinct, oval to broadly lunate, limiting the
growth of the somewhat elongate male branch. This is true of some
of our plants in that they have an antheridial disk at the tip of the
thallus which does prohibit further growth of the thallus segment.
However, many of the male thallus branches have antheridia in a
scattered group near the tip but they also extend back from this in
the dorsal groove for Ji-Ji of the length of the thallus branch. Such
an arrangement does not prohibit increase in length of that segment
and it does continue to grow. This disagrees with the European
and American descriptions given for the species but agrees with
the findings of Schuster (1952) on material of the species from
Minnesota.

In addition our specimens show that the carpocephala are nearly
always developed at the tip of a thallus branch and on the dorsal
surface. Occasionally they develop on short, lateral ventral
branches. The former condition is characteristic of the species
as previously conceived while the latter is not. Schuster (1953)
found in Minnesota that a well-developed tendency for develop-
ment of carpocephala on ventral branches occurred. This tendency
is weakly expressed in Kansas material.

The above variations do not seem constant enough to call for
taxonomic segregation of subspecies. Much more work over the
range of the species will have to be done. It is felt that we are
dealing with a fairly common hepatic which has not received critical
attention and thus the variable characters perhaps escaped notice.

Distribution: Anderson County: 2 miles south of Garnett, sandy-

Taxonomy and Ecology of Kansas Hepaticae 89

clay soil, June 4, 1951, M4874. Chautauqua County: 3 miles north-
east of Sedan, sandy bank in oak woods, March 26, 1948, M1216;
sandy soil, prairie hillside, April 5, 1952, M5340. Coffey County:
2 miles southwest of LeRoy, sandy cliff, April 9, 1948, M1239.
Crawford County: Farlington Tree Farm, September 17, 1949,
L. J. Gier 2365. Douglas County: 3 miles southeast of Baldwin
City, sandy prairie ridge, April 2, 1948, M1225; 12 miles south of
Lawrence, sandstone ledge, October 13, 1951, M5246. Elk County:
10 miles north of Busby, sandy trail in oak woods, April 10, 1948,
M1283. Ellsworth County: 3 miles northeast of Geneseo, moist
sandstone ledge, April 13, 1948, M1309; 5 miles northwest of Lang-
ley, shaded sandy bank in prairie ravine. May 3, 1952, M5447; 6
miles southeast of Kanopolis, sandstone cliff. May 3, 1952, M5454.
Jefferson County: 1 mile west of Medina, sandy prairie slope, April
2, 1948, M1235. Greenwood County: 1 mile northeast of Fall River,
sandstone ledge in prairie ravine, April 10, 1948, M1293. Labette
County: 5 miles southwest of Dennis, sandy roadside bank, April 10,
1948, M1265. McPherson County: 2 miles northwest of Lindsborg,
sandstone ledge. May 3, 1952, M5457. Montgomery County: 3
miles southwest of Elk City, April 4, 1942, R. H. Thompson; 6
miles south of Elk City, sandy soil, March 26, 1948, M1181. Neosho
County: 2 miles northeast of Chanute, sandy slope in oak forest,
April 9, 1948, M1257. Washington County: 2 miles west of Linn,
sandstone ledge, August 22, 1952, M5713. Wilson County: 1 mile
northeast of Neodesha, sandy soil over surfacing sandstone, April
10, 1948, M1271; April 5, 1952, M5331. Woodson County: 2 miles
northwest of Yates Center, sandy prairie, April 9, 1948, M1242.

51. Plagiochasma rupestre (Forst. ) Steph. Rare. This south-
western species has been found once in Kansas extending its range
considerably north of previous records from Oklahoma. However,
a collection is in the herbarium of the University of Kansas from
Clay County, Missouri, collected by L. J. Gier 1179 on July 26, 1948,
which extends the known range to west-central Missouri. The
plants were growing on the side of a sandstone outcrop in a wooded
ravine.

Distribution: Woodson County: 8 miles south and 4 miles west
of Yates Center, sandstone outcrop, September 27, 1952, M6084.

52. Reboulia heniisphaerica (L.) G. L. and N. Common in the
eastern half of the State. It tolerates a wide range of habitat varia-
tions and is met with frequently in our area. It seems to exhibit a
preference for sandstone or sandy situations and is rather uncommon

90 The University Science Bulletin

on limestone and calcereous soils. This may be due to the shortage
of moisture so usual on limestone sites in Kansas.

R. hemisphaerica is usually to be found on the sides of sandstone
outcrops, in crevices and sometimes on moist sandy soil below
outcrops. It seldom grows upon soil unless in moist shaded condi-
tions. Even though the plant requires a moist habitat it can survive
severe drying and is therefore one of our more xeric hepatics.
During hot dry summers the thallus will curl up exposing the black-
puiple ventral surface. Normally most of the thallus will winter
kill leaving only the tips alive. These may continue growth or if
they contain antheridia and archegonia they will result in the forma-
tion of fruiting bodies.

This species is found characteristically to form rosette patterns
on rock faces and becomes gregarious only during unusually moist
seasons or when on moist soil.

Distribution: Anderson County: 5 miles south of Garnett, on
moist limestone rock ledge, October 8, 1948, M2484. Chautauqua
County: 3 miles northeast of Sedan, on side of sandstone ledge.
May 1, 1948, M1365; April 12, 1950, M4193; April 5, 1952, M5336.
Cherokee County: 5 miles east of Baxter Springs, moist limestone
ledge, April 30, 1948, M1360. Clay County: 1 mile southwest of
Longford, sandstone ledge. May 23, 1948, M1513. Cloud County:
September 13, 1948, S. V. Fraser. Cowley County: 2 miles south-
east of Arkansas City, sandy bank, September 2, 1950, M4643.
Crawford County: 1 mile east of Pittsburg, sandy bank, April 11,
1952, M5354. Dickinson County: 8 miles northeast of Solomon,
sandstone ledge, May 23, 1948, M1512. Douglas County: 12 miles
south of Lawrence, sandstone ledge, April 18, 1948, M1336; 5 miles
west of Baldwin City, sandstone ledge, October 15, 1949, M4146;
April 22, 1950, M4232; April 26, 1952, M5440. Elk County: 10 miles
north of Busby, sandstone ledge, April 10, 1948, M1284. Ellsworth
County: 3 miles northeast of Geneseo, April 13, 1948, M1308;
6 miles south of Kanopolis, sandstone ledge. May 3, 1952, M5455.
Franklin County: 4 miles southwest of Ottawa, sandstone ledge,
June 18, 1949, M3094; August 2, 1949, M3611; May 12, 1950, M4248.
Geary County: 2 miles southeast of Junction City, moist limestone
ledge, April 30, 1949, M2716. Greenwood County: 1 mile northeast
of Fall Biver, margin of sandstone outcrop, April 10, 1948, M1294.
Harper County: 1 mile west of Danville, sandy moist roadside
bank, August 18, 1948, M2233. Jefferson County: 1 mile west of
Medina, sandy bank, Nov. 16, 1947, M1136. Labette County: 5
miles southwest of Dennis, sandstone ledge, April 10, 1948, M1266.

Taxonomy and Ecology of Kansas Hepaticae 91

Lincoln County: 10 miles southeast of Lincoln, sandstone ledge,
September 4, 1950, M4650. Leavenworth County: 4 miles north-
east of Tonganoxie, side of sandstone outcrop, May 7, 1948, M1401.
McPherson County: 2 miles northwest of Lindsborg, on sandstone
outcrop, July 24, 1948, M1822; May 3, 1952, M5458. Miami County:
Miami County State Park, on limestone rocks, October 19, 1947,
M1112. Mitchell County: 1 mile northwest of Simpson, sandstone
outcrop, September 4, 1950, M4652. Montgomery County: 3 miles
southwest of Elk City, on sandstone rocks, April 4, 1942, R. H.
Thompson. Nemaha County: Nemaha County State Park, on sandy
bank, August 9, 1948, M2019. Neosho County: 2 miles northeast
of Chanute, on sandstone ledge, April 9, 1948, M1258. Osage
County: 6 miles south of Quenemo, on sandstone cliff, July 1, 1949,
M3323. Ottawa County: 2 miles southwest of Minneapolis, sandy
soil below ledge, April 14, 1948, M1316. Rice County: 2 miles
southwest of Crawford, sandy roadside bank, July 24, 1948, M1828.
Riley County: 3 miles southwest of Manhattan, limestone ledge,
April 30, 1949, M2720. Russell County: 7 miles north of Dorrance,
base of sandstone ledge, June 25, 1949, M3292. Saline County:
2 miles northwest of Brookville, on sandstone outcrop, May 22,
1948, M1480. Shawnee County: 3 miles northeast of Topeka, sand-
stone ledge, April 2, 1948, M1231. Washington County: 1 mile west
of Linn, sandstone outcrop, April 15, 1948, M1328; August 10, 1948,
M2056; August 22, 1952, M5722. Wilson County: 3 miles west of
Neodesha, limestone ledge, June 30, 1947, M1064; 1 mile northeast
of Neodesha, sandstone ledge. May 1, 1948, M1382. Woodson
County: 2 miles northwest of Yates Center, sandstone ledge, April
9, 1948, M1243; September 27, 1952, M6083.

Conocephalaceae

53. Conocephalum conicum ( L. ) Dumort. Found in the eastern
fourth of Kansas where it is largely confined to the vertical sides
of sandstone ledges along creeks or moist hillsides. It regularly
forms large, extensive mats over the surface of the rocks and will
extend on soil at the base of the rocks if the soil is moist. Less
frequently it occurs on boulders on the stream margins and hard
packed sandy alluvium. Occasionally, small patches of this species
have been found at the base of seepy limestone outcrops but such
colonies never persist for more than a season.

In Kansas Conocephalum conicum fruits very rarely. In fact
it seldom seems to produce reproductive structures and when pro-
duced they are usually winter killed. This liverwort does not

92 The University Science Bulletin

tolerate xeric conditions and is limited in Kansas to sites which
are moist even during the driest of seasons. Drought seasons
invaribly reduce the size of colonies and completely eliminates
them in limestone stations. The lack of spore production and in-
ability to tolerate drouth, in Kansas, restricts the occurrence of this
species in the State. This explains the fact that in normal moisture
seasons many apparently favorable sites do not have Conocephalum
conictim as part of their hepatic flora.

Distribvition: Chautauqua County: 3 miles northeast of Sedan,
side of sandstone ledge, July 1, 1947, M1072; July 8, 1949, 3380;
April 12, 1950, M4192. Cowley County: 4 miles southeast of
Arkansas City, seepy limestone ledge, July 23, 1947, M1076. Doug-
las County: 5 miles west of Baldwin City, side of sandstone out-
crop, June 24, 1947, M1057; April 2, 1948, M1221; April 18,
1948, M1338; April 22, 1950, M4230. Franklin County: 5 miles
northeast of Ottawa, side of sandstone ledge, October 15, 1949,
M4151. Greenwood County: 1 mile east of Fall River, on sand-
stone ledge, July 9, 1949, M3411. Leavenworth County: on side
of sandstone outcrop, July 27, 1947, M1077. Montgomery County:
5 miles southeast of Elk City, sandstone boulders, July 17, 1947,
M1074. Wilson County: 3 miles west of Neodesha, on side of
seepy limestone ledge, June 30, 1947, M1063. Woodson County:
3 miles northwest of Yates Center, sandstone cliff, June 29, 1947,
M1061; June 19, 1949, M3198. Wyandotte County: 5 miles north-
west of Bonner Springs, sandstone cliff, July 27, 1947, M1079;
May 21, 1949, M2744.

Marchantiaceae

54. Marchantia polymorpha L. Widely dispersed over the
eastern half of the State. It occurs on both sandstone and lime-
stone ledges, showing no preference for either. In our area this
species may be expected on any rocky ledge or outcrop which
is moist or from which a spring flows. The distribution, however,
is localized and the species is seldom found. The absence of
the species from most of the favorable sites is unexplainable.

Throughout late summer one can find M. polymorpha on allu-
vial deposits of our creeks and rivers. It occurs rather evenly
distributed in all such places in the eastern two thirds of the
State. Yet "mother colonies" are rarely found and are never in
abundance.

At one hill-top site, plants were found in a woods which had
been thinned, and the debris raked into small piles and burned.

Taxonomy and Ecology of Kansas Hepaticae 93

Each such burned area contained an abundance of both male
and female fruiting thalli. These occurred only on the circular
burned areas.

Our collections of this species show all the usual variations
met with in this polymorphic taxon. In a few springs at the
base of sandstone cliffs one finds the forma aquatica Nees, but it
intergrades completely with strand plants. Other plants have
been found on the sides of sandstone cliffs which could be called
forma alpestris Nees, but again intergradation seems complete
and subspecific names are apparently useless in our area.

In the list which follows showing distribution all stations,
regardless of habitat or abundance, are noted. In some of these
only a few gemma-cup-bearing dichotomies were found along
alluvial deposits of streams. The plants are not anywhere nearly
as common or abundant as the distributional data would seem to
indicate. A bryologist, unfamiliar with Kansas, might search for
days in the State and never see a single specimen of M. pohjmorpha.

Distribution: Anderson County: 2 miles south of Garnett, alluvial
deposit, June 19, 1949, M3207. Brown County: 5 miles northeast
of Hiawatha, alluvial deposit, September 7, 1948, M2434. Chase
County: 1 mile south of Strong City, bank of river, November 27,
1948, M2604. Coffey Cotmty: 2 miles north of Burlington, river
bank, November 26, 1948, M2594. Douglas County: 5 miles north-
west of Lawrence, bank of Kansas River, May 14, 1948, M1444;
University of Kansas Campus, under library steps, June 13, 1949,
M3063; 3 miles north of Baldwin City, side of sandstone rock,
October 14, 1951, M5253; May 10, 1952, M5467. Ellsworth County:
3 miles northeast of Geneseo, side of sandstone cliflF, April 13, 1948,
M1313; May 22, 1948, M1501. Franklin County: 3 miles east of
Pomona, river bank, June 8, 1948, M1576; 1 mile southeast of
Ottawa, on alluvial deposit, June 28, 1949, M3307. Geary County:
2 miles southeast of Junction City, river bank, April 30, 1949, M2715.
Leavenworth County: 2 miles northwest of Leavenworth, moist
limestone ledge, September 7, 1948, M2430; 2 miles west of Lin-
wood, sandy creek bank, June 16, 1949, M3083, M3085. Lyon
County: 1 mile north of Emporia, on alluvial deposit, November 26,
1948, M2597. Marion County: 1 mile south of Florence, river bank,
November 27, 1948, M2608. Marshall County: 'A mile east of Blue
Rapids, river bank, August 9, 1948, M2037. McPherson County:
2 miles northwest of Lindsborg, on sandstone ledge, July 24, 1948,
M1823. Miami County: Miami County State Park, alluvial deposit

94 The University Science Bulletin

along river, October 3, 1948, M2469; October 17, 1949, M4157.
Osage County: 2 miles north of Vassar, river bank, June 8, 1948,
M1572. Ottawa County: 2 miles south of Wells, on moist sandstone
outcrop, April 14, 1948, M1320; May 21, 1948, M1468. Rice County:
2 miles southwest of Crawford, on sandstone ledge, July 24, 1948,
M1830. Riley County: bank of Kansas River, October 22, 1948,
T. E. Brooks. Saline County: 2 miles northwest of Brookville, on
sandstone outcrop. May 22, 1948, M1477. Washington County: 1
mile west of Linn, moist sandstone outcrop, April 15, 1948, M1328;
August 10, 1948, M2057; August 22, 1952, M5718. Wilson County:
1 mile northeast of Neodesha, on burned over brush pile site. May 1,
1948, M1380. Woodson County: 2 miles northwest of Yates Center,
alluvial soil, June 19, 1949, M3199. Wyandotte County: 2 miles
southeast of Wolcott, below a spring on limestone outcrop, Septem-
ber 6, 1948, M2420.

Oxymitraceae

55. Oxymitra paleacea BischoflF. Found in six counties in the
southeastern quarter of Kansas where it grows only on sandy soil
over surfacing sandstone or on sandy soil in rocky bluestem prairie.
It may be found on dry sandy ledges, open places in oak woods and
along trails.

This species is an extreme xerophyte and regularly has its sides
roll up and recurve onto the dorsal surface. In such a position the
large ventral scales overlap and cover the thallus. In such a condi-
tion the plants are capable of passing through long severe dry
periods.

O. paleacea fruits in September-November during normal seasons.
A dry summer, however, delays early fall growth resulting in the
maturation of spores the following spring. Plants are always found
in gregarious colonies which may cover an area cf several square
yards.

In North America Oxymitra paleacea is known from Mexico,
central Texas, through Oklahoma and into Kansas. The Texas
plants were discovered first and named as Oxymitra androgyna by
Howe (1914). Previous to that time only one species, Oxymitra
paleacea BischoflF, was known and it had been found in Europe,
Africa and South America. Howe (l. c.) described the Texas ma-
terial as a new species and separated the North American material
on the basis of spore size (which he gave as 125-175[jl) while the
European, African and South American material was believed to
have spores 100-120pL in diameter. The European plants were uni-

Taxonomy and Ecology of Kansas Hepaticae 95

sexual and the North American were bisexual with an occasional
tendency toward the unisexual condition. In addition the walls ra-
diating from the pores were thought to be thicker in North American
material than in the other known species.

Recently Miiller (1952) reduced Oxijmitra androgyna Howe to a
synonym of Oxymitra paleacea Bischoff. In this we concur. The
writer has collected or studied specimens of Oxyrnitra from locali-
ties throughout its known range in North America. We find that the
use of spore size completely fails as a specific criterion. Wittlake
( 1953 ) has shown that spore size of Kansas Oxymitra ranges from 90
to lSO;j, thus in its lower range going below that given for the
European material, and much surpassing the maximum size of the
European spores. The writer finds North American spores to have
the range as given by Wittlake ( 1953 ) but most consistently to fall
into a range of 118-140. Spores of material which we have studied
from France and Italy range from 82-142tj. with most of them being
in a range betsveen 115-132[j.. Thus spore size in plants of the
two continents are very similar and show only that those from North
America tend to be a little larger.

The walls radiating from the pores in North American and Euro-
pean plants vary over the same range, and therefore are of no
taxonomic importance.

In North American material we have found that thalli are bisexual
and unisexual with most of it being of the former type. However,
unisexual plants are not uncommon and are found in all collections
made. The only European material we have been able to study
(France and Italy) revealed that both conditions existed, but ma-
terial was too scarce to gain any idea of the relative frequency of
each type. In American material 9 bisexual thalh were found for
every 2 that were unisexual. In some one segment was bisexual
while the sister segment was unisexual.

Wittlake ( 1953 ) describes the spore of Oxymitra paleacea ( = O.
androgyna) as having areolae too variable to consider. This we
do not find to be the case, since the spore, illustrated by Wittlake,
is characteristic and shows areolae to be quite regular in size or at
least as regular as one would expect areolae to be on spores in the
Oxymitraceae and related Ricciaceae. We find the largest areolae
to be 42[JL in greatest diameter and the smallest to be 24. How-
ever, out of a great number measured we find them to fall con-
sistendy in the range of 30-40tJ.. Wittlake (1953) also gives the
length of the projections of the spore (verrucae) as .5-2ijl while we
<ind them to be rather consistently 3.3pi. long and range from .l-Q.I^.

96 The University Science Bulletin

Oxymitra paleacea, as now understood, exhibits a striking geo-
graphic distribution. It is found in Europe, North Africa, Brazil,
Argentina, Paraguay, Mexico, and in Texas, Oklahoma and Kansas
in the United States. The Mexico record is not reported in the
literature but collections in the herbarium at the University of
Kansas by E. B. Wittlake 50241 taken on July 2, 1953, 2 miles south-
east of Tecamachalco, Puebla and McGregor 8029 from Tamuzun-
chale, San Luis Potosi, constitute valid records for Mexico.

Distribution: Chautauqua County: 3 miles east of Sedan, sandy
prairie, edge of oak woods, March 26, 1948, M1209; May 8, 1949,
W. H. Horr; 4 miles south of Sedan, sandy soil, July 8, 1949, M3370;
1/2 miles northeast of Sedan, sandy open wooded prairie, April 5,
1952, M5346. Elk County: 10 miles north of Busby, sandy soil over
surfacing sandstone, April 10, 1948, M1287. Greenwood County:
1 mile northeast of Fall Biver, edge of surfacing sandstone, April
10, 1948, M1291; 1 mile east of Fall Biver, sandy soil, April 13, 1949,
M2670; July 9, 1949, M3409; August 21, 1949, B. H. Thompson.
Montgomery County: 3 miles southeast of Elk City, sandy soil,
April 4, 1942, B. H. Thompson; 3 miles southwest of Elk City,
sandy open wooded hillside, October 25, 1947, M1119; 6 miles south
of Elk City, sandy soil, March 26, 1948, M1185; 3 miles southwest
of Elk City, sandy soil, September 22, 1951, M5230. Wilson County:
1 mile northeast of Neodesha, sandy soil, October 25, 1947, M1115;
April 10, 1948, M1272. Woodson County: 2 miles northwest of
Yates Center, sandy soil, October 26, 1947, Ml 124; April 9, 1948,
M1244; Lake Fegan, sandy soil, March 26, 1949, M2648; 2 miles
northwest of Yates Center, sandy soil, June 19, 1949, M3200; Lake
Fegan, sandy soil, edge of sandstone outcrop, September 24, 1949,
M4129; 2 miles northwest of Yates Center, shallow sandy soil over
surfacing sandstone, October 28, 1941, M5274.

Bicciaceae

56. Ricciocarpus natans (L) Corda. Bare. Found in a few
swamps, lakes and ponds in eastern Kansas where it floats on the
surface of the water. Land forms also occur at the above habitats
in late summer and fall as the water level recedes. These land
forms are also to be found occasionally on alluvial deposits along
streams in the eastern part of the State.

In Kansas R. natans (water form) fruits abundantly in early
spring and then passes into a vegetative phase during which it
reproduces actively by division and fragmentation of thalli. These
water forms persist into winter and eventually are frozen in ice.

Taxonomy and Ecology of Kansas Hepaticae 97

One station was observed through several seasons. At this site the
floating forms reguhirly are frozen in ice in early winter. This
results in death of the posterior portion of the plants. This early
freeze invariably is followed by complete thawing during which
plants undergo some growth. Through the rest of the winter
season successive freezing and thawing takes place. Never is R.
natans killed by being frozen in ice even when frozen for a period
of several weeks. The result is that by spring many small thalli
tips are present on the body of water. These grow rapidly and
produce typical fruiting plants.

In the summer and early fall many of the floating plants are left
stranded on muddy banks by receding water. These plants develop
into rosettes or irregularly gregarious colonies of the land form and
become submerged with the advent of fall rains. Such submerged
colonies exist through the winter and in the spring the tips of the
apices produce small plantlets which become detached, float to the
surface, and develop into fruiting individuals.

The finding of some strand plants along creek and river banks
indicates that the distribution of this species, at least in northeast
Kansas, is greater than tlie distribution listed below would seem
to indicate. However, we have found some strand plants on creek
banks whose drainage area is completely known and in which no
floating plants have been found. The occurrence of such plants
is not explainable unless they developed from spores carried in or
blown into the watershed area.

Distribution: Atchison County: Vi mile south of Muscotah, mud
bank of Little Grasshopper Creek, October 18, 1947, Ml 105. Doug-
las County: 1 mile north of Lawrence, alluvial deposit of Kansas
River, November 13, 1948, M2593; 3 miles northeast of Lawrence,
floating in cattail-sedge marsh, November 6, 1948, R. H. Thomp-
son; 1 mile northeast of Lawrence, floating in old ox-bow marsh,
August 27, 1949, M3893; November 13, 1949, M4163; September
23, 1950, M4676; 2 miles northeast of Lawrence, alluvial deposit
on bank of Kansas River, August 9, 1952, M56S6. Jefferson
County: 3 miles west of Williamstown, floating in cattail marsh,
November 13, 1949, M4165. Johnson County: Vi mile north of
De Soto, alluvial deposit on bank of Kansas River, July 27, 1952,
M5668. Leavenworth County: 2 miles north of Leavenworth,
alluvial deposit bank of Missouri River, October 29, 1948, M2582.
Linn County: 2 miles west of La Cygne, floating in cattail marsh,
June 4, 1949, M2855. Miami County: Murray Lake, Miami County

4—6730

98 The University Science Bulletin

State Park, alluvial deposit, October 19, 1947, M1114; Pigeon
Lake on Hugh Whitford Ranch, on alluvial deposit, April 24,
1948, M1348; 3 miles east of Fontana, bank of Marais des Cygnes
River, October 3, 1948, M2470; Pigeon Lake, mud bank, November
12, 1949, M2579; Pigeon Lake, floating. May 29, 1949, M2772;
October 17, 1949, M4158; October 8, 1950, M4681.

57. Riccia heijrichiona Hampe. Found over most of Kansas
except the northwest quarter and rare in the southwest quarter.
This species ranks with FruUania inflata and Riccia frostii as the
mostly widely distributed species of hepatic in the State. It
tolerates a wide range of habitats but is most common on
sandy soils associated with surfacing sandstone in prairie and
oak forest. In moist seasons it is scattered in limestone areas,
particularly in mixed forest and prairie sites. It is also to be
found on alluvial deposits along river banks, flood plains, road-
side ditches, and deposits from eroded fields.

R. beyrichiana is one of the most easily recognized species in
the genus; nevertheless it is quite variable in many respects.
In our specimens we find the margin characteristically has one
or two rows of stout, usually curved cilia from 30 to 300ix long.
These are always prominent at the apex but become less frequent
posteriorly. Never are they absent. Macvicar (1926) says that
specimens from England most frequently have naked margins and
Miiller (1952) concurs in this for European material. Frye and
Clark (1937-1947) report that cilia are rarely wanting, usually
few in number, in 1 or 2 series, stout and often curved. Kansas
specimens are more as described by Frye and Clark in this char-
acter but seem to show even a more pronounced tendency for
frequent, stout, curved cilia. Occasionally specimens found in
the shade or on very moist soil show a tendency for cilia suppres-
sion; but never is the margin without them.

The coloration of the thallus sides is usually greenish in Kansas
specimens except for those on drier or exposed sites where the
color may range from reddish to black-purple. Ventral scales
likewise are usually hyaline to whitish but may become purplish
on dry exposed habitats.

Thallus size varies also with the habitat. On exceptionally dry
areas, the length may be from 7-12 mm. and the width from 1-2 mm.
On more favorable sites the length is usually 5-10 mm. and the
width 2-4 mm. The thickness of the thallus, width of the dorsal
groove, and the angle of margin also vary with respect to moisture
and light conditions.

Taxonomy and Ecology of Kansas Hepaticae 99

The spore differs in some characters given by certain different
workers. Frye and Clark (1937-1947) and Howe (1923) give the
size range of greatest spore diameter as 65-140iJ. for American ma-
terial. Macvicar (1926) gives the range as 95-120tJL for English
specimens; while Miiller (1952) gives the spore size as 100-120;j.,
and Schuster ( 1953 ) 90-130[j.. Kansas material shows, on the basis
of an examination of spores from all the collections listed below,
that the range in size of mature spores is 74-138[j.. However, the
overall average of spore size is 106[j.. The spore size, as a taxonomic
character, must be used with caution, as a given colony often has
a range different from that of another. My collections M4143 and
M5446, for example, show that the range of spore size in M5446 is
75-1 12;j., and the average size is 90[j.; while M4143 has a range of
115-138[;., and an average spore size of 124[jl. Thus the spores of
M4143 average larger than the maximum limit of spore size known
for the species by Macvicar (1926) and Miiller (1952). At the
other extreme the spores of M5446 average 90ij, or the exact size of
the minimum size considered for the species by Schuster ( 1953 ) .
As a further comparison my collection M1353 shows a range of
spore size of 99-122ix and an average size of llSiJ., thus the collection
falls in line with sizes as given by all the above cited workers.

Another spore character which seems to vary is the size of areoles
on the outer spore face. Macvicar (1926) reports the size of areoles
as reaching up to 15[;., Frye and Clark (1937-1947) give a range of
10-18[;., and Miiller (1952) gives a range of IO-ISijl. Miiller (1952)
also gives the number of areolae across the face of the spore as 6-8,
and Macvicar ( 1926 ) as 7-9. We find, in our Kansas plants, that
the size of areolae varies from 7.2-18.8[j. in diameter and have 5-7
across the face of the spore. The average of areolae diameters is
15-17[j. and the average number across spore face is 5-6. Thus Kan-
sas material shows a tendency to fewer but larger areolae as com-
pared to European plants.

Another spore character, that of papillae revealed when the spore
is viewed in profile, seems also to vary. Macvicar (1926) states
that spores are highly papillose in profile while Frye and Clark
(1937-1947) refer to the papillae as being absent or obscurely
present. Most of our Kansas collections show only faint signs of
papillae, though occasionally, they are evident and up to 2.4[jl in
length. When such are observed they are found to be truncate to
obtuse.

Occasionally collections of R. beyrichiana are obtained in Kansas

100 The University Science Bulletin

(such as M1059), which are extremely atypical as far as spore char-
acters are concerned. The collection mentioned has spores in which
no complete areolae are to be found on the outer face. Instead,
rather heavy irregular ridges occur and these rarely anastomose.
The area in which this collection was made has been studied exten-
sively through several seasons and only one gregarious colony was
found with such spore characters. A tendency toward this condi-
tion has been noticed in other stations. At all such places R.
campbcUiana is common and intermingled with R. beyrichiana. It
thus seems possible that hybridization between the two might
have occurred.

It seems possible that a critical study of R. beyrichiana in North
America and Europe might show, that a part of the material (Eu-
ropean ) should be split off as a subspecies, if some of the differences
noted above prove to be constant.

Distribution: Allen County: 5 miles north of Moran, creek bank,
October 8, 1948, M2499. Anderson Coimty: VA miles northeast of
Welda, bluestem prairie, October 31, 1948, R. H. Thompson; 5 miles
south of Garnett, moist bank in woods, October 8, 1948, M2482; 2
miles south of Garnett, sandy-clay soil, June 4, 1951, M4876. Atchi-
son Coimty: Y2 mile south of Muscotah, creek bank, October 18,
1947, Ml 106. Barton County: Ji mile north of Pawnee Rock, sandy
prairie, August 30, 1950, M4629. Bourbon Coimty: 2 miles south of
Uniontown, creek bank, October 8, 1948, M2511; 6 miles south of
Fort Scott, sandy-clay prairie bank. May 9, 1953, M7201. Brown
Coimty: 5 miles northeast of Hiawatha, sandy-clay prairie bank,
September 7, 1948, M2432. Chase Coimty: 1 mile south of Strong
City, river bank, November 27, 1948, M2605. Chautauqua Coimty:
3 miles northeast of Sedan, sandy soil, March 26, 1948, M1214; VA
miles northeast of Sedan, sandy prairie, April 12, 1950, M4188;
April 5, 1952, M5342. Cherokee County: 5 miles east of Baxter
Springs, rocky-clay soil, open places in oak woods, April 30, 1948,
M1353, M1362; 3 miles southeast of Baxter Springs, sandy bluestem
prairie, October 22, 1951, M5263, M5265; 3 miles northeast of
Riverton, sandy abandoned field. May 9, 1953, M7199. Coffey
Coimty: 2 miles southwest of LeRoy, sandy prairie, April 9, 1948,
M1236; 2 miles north of Burlington, river bank, November 26, 1948,
M2596. Cowley County: 2 miles southeast of Arkansas City, sandy
prairie bank, September 2, 1950, M4642. Crawford County: Craw-
ford County State Park, moist clay bank, October 9, 1948, M2528.
Douglas County: 1 mile southeast of Lawrence, sandy soil, April 24,

Taxonomy and Ecology of Kansas Hepaticae 101

1942, R. H. Thompson; 5 miles west of Baldwin City, very moist
sandy soil, June 24, 1947, M1059; 2 miles southeast of Baldwin City,
moist sandy soil, October 11, 1947, MllOO; 5 miles west of Baldwin
City, sandy bluestem prairie, June 18, 1948, M3104; October 15,
1949, M4143; September 16, 1950, M4675; October 13, 1951, M5249;
April 26, 1952, M5434; 2 miles northeast of Lawrence, alluvial de-
posit, bank of Kansas River, June 12, 1952, M5604. Edwards
County: 2 miles east of Kinsley, sandy prairie, August 28, 1950,
M4508. Elk County: 10 miles north of Busby, sandy open oak
woods, April 10, 1948, M1286. Ellsworth County: 3 miles northeast
of Carneiro, sandy prairie, May 22, 1948, M1500; 4 miles northwest
of Langley, sandy prairie. May 3, 1952, M5446. Ford County: 4
miles southwest of Dodge City, sandy soil, August 28, 1950, M4509.
Franklin County: 1 mile northeast of Norwood, sandy soil, March
25, 1948, M1169. Geary County: 2 miles southeast of Junction City,
river bank, April 30, 1949, M2716. Greenwood County: 1 mile
northeast of Fall River, sandy soil, April 10, 1948, M1295. Hamil-
ton County: 2 miles south of Syracuse, sandy soil, August 29, 1950,
M4531. Harvey County: 3 miles north of Burrton, sandy soil, July
24, 1949, M3505; August 28, 1950, M4505; June 11, 1951, M4911;
August 25, 1951, M5083. Jefferson County: 1 mile west of Medina,
sandy soil, November 16, 1947, M1135. Johnson County: 5 miles
west of Olathe, sandy prairie, November 2, 1947, M1130. Kearny
County: Lake McKinney, edge of lake, August 29, 1950, M4590.
Labette County: 5 miles southwest of Dennis, sandy soil edge of
oak woods, April 10, 1948, M1264. Leavenworth County: 1 mile
west of Turkey Creek Station, sandy soil, December 2, 1947, Ml 142;
4 miles northeast of Tonganoxie, sandy prairie. May 7, 1948, M1405;
2 miles west of Linwood, sandy soil, June 16, 1949, M3090. Lyon
County: 1 mile north of Emporia, river bank, November 26, 1948,
M2599. Miami County: 3 miles east of Fontana, river bank, Oc-
tober 19, 1947, Ml 107. Montgomery County: 3 miles north of
Independence, sandy soil, April 24, 1942, R. H. Thompson; 3 miles
southwest of Elk City, sandy soil, July 1, 1947, M1070; March 26,
1948, M1182. Marion County: 1 mile south of Florence, river bank,
November 27, 1948, M2610. Morris County: 2 miles north of Coun-
cil Grove, river bank, November 27, 1948, M2615. Nemaha County:
Nemaha County State Park, sandy-clay soil, August 9, 1948, M1261.
Ottawa County: Ottawa County State Park, sandy soil. May 21,
1948, M1467. Reno County: 6 miles northeast of Hutchinson, sandy
soil, June 11, 1951, M4956; August 28, 1950, M4506; August 25,

102 The University Science Bulletin

1951, M5084. Rice County: 1 mile east of Geneseo, sandy prairie,
April 30, 1949, M2708. Riley County: 3 miles southwest of Man-
hattan, moist wooded hillside, April 30, 1949, M2721. Saline
County: 2 miles northwest of Brookville, sandy prairie, May 22,
1948, M1479. Sedgwick County: 2 miles northwest of Wichita,
sandy pasture, September 2, 1950, M4649. Shawnee County: 5
miles east of Topeka, rocky clay bank, April 23, 1949, M2679.
Stafford County: 1 mile west of Stafford, sandy roadside ditch,
August 28, 1950, M4507. Stanton County: 1 mile north of Saunders,
sandy soil, August 28, 1950, M4547. Washington County: 2 miles
southwest of Palmer, sandy prairie. May 27, 1951, M4830. Wihon
County: 1 mile northeast of Neodesha, sandy soil, June 11, 1952,
M5602. Woodson County: 2 miles northwest of Yates Center, sandy
prairie, June 29, 1947, M1062; May 12, 1951, M4807; 14 miles
southwest of Yates Center, sandy oak woods, September 1, 1951,
R. H. Thompson. Wyandotte County: 1 mile northeast of Bonner
Springs, sandy soil. May 15, 1948, M1440.

58. Riccia bifurca Hoffm. Rare. Found but three times in
Kansas and known only from alluvial deposits along streams. It
therefore is not a characteristic species in our hepatic flora. In
each of the three stations where the species was taken we have
since failed to locate more specimens. The occurrence of the species
on alluvial deposits indicates that plants must occur elsewhere
in our area but as yet they have not been found.

Our plants agree in all respects with descriptions of the species
and with specimens seen from other areas. Two rosettes of num-
ber M2521, however, would be referred to R. bifurca var. subinermis
Heeg on the basis of the presence of a few acutely pointed cilia
on the margin of terminal segments. This character does not
seem worthy of taxonomic recognition in this instance, and the
specimens are therefore not segregated.

Some difference of opinion is apparent in the literature as to
the correct name for this species. Frye and Clark (1937-1947)
recognize Riccia bifurca Hoffm. a.nd reduce R. arvensis Aust. to
a synonym of it. Muller (1952) also considered R. arvensis as a
synonym; but Schuster (1949, 1953) recognizes R. bifurca as a
synonym of R. arvensis. A study of North American specimens
named R. arvensis by Austin (1870) shows that they are iden-
tical with the plants in Europe known as R. bifurca. It thus is
necessary to recognize R. bifurca as the valid name.

Distribution: Cherokee County: 5 miles east of Baxter Springs,

Taxonomy and Ecology of Kansas Hepaticae 103

bank of Shoal Creek, October 9, 1948, M2521. Douglas County:
north edge of Lawrence, bank of Kansas River, May 1, 1951, R.
H. Thompson. Riley County: bank of Kansas River, October 22,
1948, Travis Brooks.

59. Riccio campbelliona Howe. Common on sandy soil in
prairie and open oak wooded areas. It is most frequent on
shallow soil over surfacing sandstone and in bluestem prairies
near such environments. Occasional plants are found on alluvial
soil along stream banks. As yet this species has not been found
on calcareous sites.

The disb-ibution of Riccia campbeUiana in the United States,
based on the literature, is somewhat confusing. The species was
described by Howe (1899) based on material from California.
Later Howe (1923) gave the range of the species as California
and Texas. Frye and Clark (1937-1947) list the species as
definitely known only from California. Jacobs ( 1951 ) reported
the species from Arkansas and Georgia, and refers to California
records. This distribution, to Jacobs, seemed to provide a re-
markable parallel to the disjunct ranges of several vascular
plants. However, both Jacobs, and Frye and Clark overlooked
a paper by McAllister, Hoglund, and Whitehouse (1932) in which
R. campbeUiana was listed from four stations in Texas. The dis-
tribution of the species, based on specimens seen by the author
or collected by him, is now known to be California, central Texas
north through Oklahoma to northeastern Kansas; then southeast-
ward to central Arkansas, to northwestern Louisiana and south-
west to central Texas. In addition to this is the record of Jacobs
( 1951 ) for Georgia. In areas of granite and sandstone outcrops
in Xexas and Oklahoma, and sandstone outcrops of Kansas and
Arkansas, the species is quite general distributed, and often is
locally abundant. The use of the species by Jacobs (l. c.) in his
discussion of disjunct ranges, therefore, seems unwarranted.

Howe (1923) gives the size range of spores of this species as
75-108[ji. but mostly 90-100[ji.. We find the maximum limit to be
118[j. and the lower limit of the range as 70[jl. The average
spore size, of Kansas material, is 96\i.. A few Kansas collections
show the outer face of the spore to be somewhat regularly areolate
with areoles up to 13. 2[;, in diameter, as compared to a size of 4-7[a
given by Howe. Otherwise Kansas material agrees with the
description of the species.

Distribution: Anderson County: VA miles northeast of Welda,

104 The University Science Bulletin

open sandy prairie, October 31, 1948, R. H. Thompson. Chautauqua
County: 3 miles northeast of Sedan, sandy soil, oak woods, March
26, 1948, M1212; 3 miles north of Elgin, sandy prairie pasture, April
5, 1952, M5350; 3 miles north of Sedan, sandy soil over surfacing
sandstone, April 5, 1952, M5353. Cherokee County: 5 miles east of
Baxter Springs, rocky clay prairie bank, April 30, 1948, M1354.
Coffey County: 2 miles southwest of LeRoy, sandy prairie, April
9, 1948, M1240. Douglas County: 12 miles south of Lawrence,
sandy prairie, April 18, 1948, M1335; October 13, 1951, M5248;
5 miles west of Baldwin City, sandy soil, April 26, 1952, M5435. Elk
County: 10 miles north of Busby, sandy prairie, April 10, 1948,
M1280. Franklin County: 1 mile northeast of Norwood, sandy oak
woods, March 25, 1948, Ml 168. Greenwood County: 1 mile north-
east of Fall River, sandy soil, April 10, 1948, M1289. Harvey
County: 5 miles north of Burrton, sandy path in sand dunes, June
11, 1951, M4917. Jefferson County: 1 mile west of Medina, sandy
prairie pasture, April 2, 1948, M1233. Labette County: 5 miles
southwest of Dennis, sandy soil at edge of oak woods, April 10,
1948, M1267. Leavenworth County: 4 miles northeast of Tonga-
noxie, sandy soil in prairie, May 7, 1948, M1404; 2 miles west of
Linwood, sandy soil in oak woods, June 16, 1949, M3087. Mont-
gomery County: 3 miles south of Elk City, sandy soil, April 4,
1942, R. H. Thompson; 6 miles south of Elk City, sandy soil in
oak woods, March 26, 1948, M1183. Neosho County: 2 miles east
of Chanute, sandy prairie, April 9, 1948, M1260. Reno County: 6
miles northeast of Hutchinson, sandy path in sand dunes, June 11,
1951, M4958. Wilson County: 1 mile northeast of Neodesha, sandy
soil edge of oak woods, April 10, 1948, M1278. Woodson County:
2 miles northwest of Yates Center, sandy prairie, April 9, ^948,
M1246; Lake Fegan, sandy soil in oak woods, March 26, 1949,
M2649; September 24, 1949, M4125; 2 miles northwest of Yates Cen-
ter, sandy prairie at edge of oak woods, May 12, 1951, M4810;
October 28, 1951, M5278.

60. Riccia canaliculata Hoffm. This species is listed here as of
questionable taxonomic standing and is based on one collection
made in Miami County, Kansas. The plants occurred on soil at
the edge of a lake.

As yet no North American specimens of the Ricciella section
of Riccia have been studied, which match the European R. canalicu-
lata with the possible exception of a collection by the author from
Texas which Miiller (personal correspondence) refers to R. cana-

Taxonomy and Ecology of Kansas Hepaticae 105

licukita. However, Crundwell (personal correspondence) says
he has seen no American material which is the same as the European
R. canaUculata. With this latter statement the writer is inchned to
agree.

The Kansas collection under consideration differs from the de-
scriptions of R. canahcuhta and with European specimens of the
species in several ways. The spores have 4-6 areolae across the
outer face while R. canaUculata has 3-4 and the areolae average 2A\).
smaller in diameter in the Kansas material. The ventral scale cells
are similar in the two, although those of our material average 4ij.
wider and some lOi^ longer. The thallus in our plants is more similar
to that of R. duplex than of R. canaUculata though it differs from
both in that the compact cells of the ventral half of the thallus has
small air chambers. R. canaUculata is not known in a water form,
while Kansas material readily grows and fruits in water culture as
well as on soil.

The present lack of critical study of the species in the iimtans
complex of the genus Riccia makes it difficult to assign the Kansas
collection to a suitable taxon. It is, therefore, provisionally assigned
to R. canaUculata to which it seems to be closely related on the
basis of spore characters.

Distribution: Miami County: 3 miles east of Fontana, Hugh
Whitford Ranch, margin of Pigeon Lake, on soil, April 24, 1948,

61. Riccia (licftjospora Howe. Found in all but the northwestern
quarter of the state. It is very common on sandy soil near sand-
stone outcrops, in sandy prairie, in certain localities in sand dune
areas of central and southwest Kansas. It is also to be found on
alluvial deposits and on rocky-clay banks. The species is found
but rarely on calcareous sites and seems to be restricted almost
entirely to sandstone or sandy areas.

This species is extremely xerophytic and survives the most severe
drought conditions. In fact, it seems to grow best on open exposed
sites which regularly dry up early in the growing season and remain
c ry until fall rains. During these periods the older parts of the
thallus die leax'ing the tips to begin growth with the advent of
moisture.

The determination of Kansas material as R. dictyospora was ac-
complished only after detailed study of the species over its ran^^e
and after careful study of R. mcalUstcri Howe. There has heln
much confusion in the literature as to the proper relationships of
these two concepts. We have reached the conclusion that R

106 The University Science Bulletin

mcaUisteri is not a valid concept and must be reduced to synonomy
under R. distyospora.

The characters which have been used to separate the taxa are
as follows:

R. mcaUisteri R. dicttjospora

1. Scales violet. 1. Scales blackish purple.

2. Margins decolorate. 2. Margin narrowly blackish

3. Hypodemial cells in distinct purple.

rows. 3. Upper surface reticulate.

4. Spores first violet but finally 4. Spores brown, rather translu-

opaque. cent.

5. Antheridial ostioles elev. 5. Anth. ostioles elev.

50-160rj.. 0-50jx.

Comparative studies show that the above characters break down
completely when used to attempt species separation. We have
studied the type specimens of the two, as well as specimens from
the entire range of the taxa and have studied them in the field over
much of their range but most importantly in the type locality of
R. nicollisteri.

It has been observed that the color of ventral scales is a
highly variable character in Riccio's and such is true with the
concepts here considered. We have observed the same colony
of these plants over several seasons and have noted that in dry
seasons they exhibit scale color of R. dictyospora and in more
moist seasons the color as given for R. mcaUisteri. The same
statement applies to color of the thallus margin. In the type
locality of R. mcaUisteri (Granite Mountain Texas) we have
observed plants growing on the sandy margin of water holes,
left by quarrying granite, which had violet scales. As one moved
away from the moist environment a progressive change in scale
color from violet to blackish purple was found. The margin of
the thallus exhibited the same transition. Even the type collec-
tion of R. mcaUisteri exhibits variation in these colorings.

The arrangement of the cells on the upper surface has been
used as a key character by Frye and Clark ( 1937-1947 ) . How
such a character came to be recognized is indeed puzzling as
type specimens and thousands of specimens seen in the field fail
to show any such distinction. We have yet to see any difference
between the two, and find them both to be reticulate, and yet to
have cells somewhat aligned in rows.

The question of spore color is also one of confusion. We find
the spores of the same colony to vary in color from spring to

Taxonomy and Ecology of Kansas Hepaticae 107

fall and from year to year. If a given colony is observed through
a season it will be noted that spores are at first yellowish after
which they become brown or violet brown and finally may be-
come opaque. Most material which has been identified as R.
(lictyospora show spores to be immature. Howe who described
both species apparently did not rely on spore characters too much
as collection identified by him as R. dictyospora have opaque
spores and others with light brown spores were identified as R.
mcaUisteri. The difficulty with spores in this complex seems to
be due to the fact that spores mature very slowly, and thus
plants which look good for specimen purposes seldom have ma-
ture spores. As indicated above the matter of spore color inter-
grades completely, and is so variable as to be useless as a taxonomic
character.

Jacobs (1951) discusses the two species and admits that they
are closely akin but states that they are quite separable. He
further states that the differences between the two are correlated
with habitat, but are associated with habitat preference rather
than habitat induction. Apparently Jacobs relies on the presence
or absence of the black margin or degree of expression of color
to separate the two so-called species. We have studied the
species in the field over the entire range of R. mcaUisteri and
over most of the range of R. dictyospora. As a result we have
failed to verify the statements of Jacobs.

Another factor which has probably led to some confusion is
the fact that Howe received specimens from McAllister (from
Texas) and grew them in a greenhouse at the New York Botanical
Garden after which he described the material as a new species
(R. mcaUisteri). This undoubtedly was somewhat responsible
for the views on coloration of the thallus.

Finally, it may be true that some of the variations noted may be,
in part, found to be grouped into geographic areas and thus perhaps
some subspecific segregation will be necessary. This can only be
decided after much more work has been done in the field, in cul-
turing, and in cytological studies.

Distribution: AUen County: 5 miles north of Moran, creek bank,
October 8, 1948, M2501. Anderson County: 4 miles south of Gar-
nett, sandy prairie, October 8, 1948, M2486. Barton County: }i
mile north of Pawnee Rock, sandy prairie, August 30, 1950, M4630.
Bourbon County: 2 miles south of Uniontown, creek bank, October
8, 1948, M2513. Brown County: 5 miles northeast of Hiawatha,

108 The University Science Bulletin

sandy roadside bank, September 7, 1948, M2431. Chase County:
1 mile south of Strong City, river bank, November 27, 1948, M2607.
Chautauqua County: 3 miles east of Sedan, sandy soil. May 1, 1948,
M1367; 4 miles south of Sedan, sandy prairie, July 8, 1949, M3372;
1/2 miles northeast of Sedan, sandy soil over surfacing sandstone,
April 12, 1950, M4202. Cherokee County: 5 miles east of Baxter
Springs, rocky clay bank, April 30, 1948, M1355. Clay County: 5
miles north of Morganville, sandy prairie, August 16, 1951, M5066.
Douglas County: 2 miles southeast of Baldvv'in City, creek bank,
October 11, 1947, M1099; 12 miles south of Lawrence, sandy prai-
rie, April 18, 1948, M1339; 5 miles v^^est of Baldwin City, sandy soil
over surfacing sandstone, June 18, 1949, M3103; September 16,
1950, M4674; August 2, 1951, M5004; October 13, 1951, M5247; 2
miles north of Lawrence, river bank, June 12, 1952, M5606. Elk
County: 10 miles north of Busby, sandy prairie, April 10, 1948,
M1279; July 27, 1948, M1916. Ellsworth County: sandy prairie, 6
miles southeast of Kanopolis, sandy prairie bank. May 3, 1952,
M5453. Franklin County: 4 miles southwest of Ottawa, sandy
prairie pasture, June 18, 1949, M3096; May 12, 1950, M4247.
Greenwood County: 6 miles north of Fall River, sandy soil in oak
woods, July 27, 1948, M1912; 1 mile east of Fall River, sandy
prairie, July 9, 1949, M3406. Harper County: 1 mile west of Dan-
ville, moist sandy roadside ditch, August 18, 1948, M2234, M2236.
Harvey County: 5 miles north of Burrton, on sandy soil at edge of
pond in sand dunes, June 11, 1951, M4916; July 24, 1949, M3506;
August 25, 1951, M5083. Jejjerson County: 1 mile east of Williams-
town, sandy prairie bank, November 6, 1948, M2569. Kiowa County:
5 miles east of Greensburg, roadside ditch, August 17, 1950, Olin
Fearing 543, 544. Leavenworth County: 4 miles northeast of
Tonganoxie, sandy soil edge of oak woods, May 15, 1948, M1438; 2
miles west of Linwood, sandy oak woods, June 16, 1949, M3089.
Lyon County: 1 mile north of Emporia, river bank, November 16,
1948, M2601. Marion County: 1 mile south of Florence, river bank,
November 27, 1948, M2612. Miami County: 3 miles east of Fon-
tana, river bank, October 19, 1947, MlllO. Montgomery County:
1 mile south of Independence, sandy soil, April 24, 1942, R. H.
Thompson; 4 miles southwest of Elk City, sandy soil in oak woods,
September 1, 1948, M2395; September 22, 1951, M5223. Morris
County: 2 miles north of Council Grove, bank of Neosho River,
November 27, 1948, M2617. Nemaha County: Nemaha County
State Park, sandy clay prairie bank, August 9, 1948, M2017. Neosho

Taxonomy and Ecology of Kansas Hepaticae 109

County: 2 miles east of Chaniite, February 14, 1945, R. H. Thomp-
son; April 9, 1948, M1262. Reno County: 6 miles west of Pretty
Prairie, sandy roadside bank, August 17, 1948, M2221; 6 miles north
of Hutchinson, sandy soil, June 11, 1951, M4957; 2 miles north of
Hutchinson, edge of pond in sand dunes, August 25, 1951, M5084.
Rice County: 1 mile east of Geneseo, sandy prairie, April 30, 1949,
M2709. Saline County: 2 miles northwest of Brookville, sandy soil
in prairie, April 30, 1949, M2690. Shawnee County: 7 miles south-
east of Topeka, sandy overgrazed prairie pasture, April 23, 1949,
M26S3. Stanton County: 1 mile north of Saunders, sandy prairie,
August 28, 1950, M4548. Sumner County: 2 miles east of Milan,
sandy-clay roadside bank, August 18, 1948, M2240. Washington
County: 1 mile west of Linn, sandy prairie pasture, August 23,

1947, M1085; 2 miles southwest of Palmer, sandy prairie. May 27,

1948, M4828; August 22, 1952, M5724. Wilson County: 1 mile
northeast of Neodesha, sandy soil October 25, 1947, M1118; 4 miles
north of New Albany, sandy prairie, July 9, 1949, M3402. Woodson
County: 2 miles northwest of Yates Center, sandy prairie, April 9,
1948, September 1, 1951, R. H. Thompson; October 28, 1951,
M52Si. Wyandotte County: 1 mile northeast of Bonner Springs,
sandy prairie bank. May 15, 1948, M1441.

62. Riccia fluitans L. Rare. Found in two places in Miami
County. In both places the plants were found in small lakes in the
Marais des Cygnes River valley. Specimens have been taken from
water and from mud banks of these stations. Our specimens agree
in every respect with the description of the species given by Miiller
(1952).

Distribution: Linn Co.: 2 miles west of La Cygne, in water,
June 4, 1959, M2S56. Miami County: 3 miles east of Fontana, in
Pigeon Lake, May 29, 1949, M2773; mud bank of Pigeon Lake,
October 8, 1950, M4679; October 27, 1950, M4696; November 30,
1951, M5293.

63. Riccia frostii Aust. A common and often abundant species
on stream banks, flood plains, and lake shores. It is to be found in
all of Kansas except the southwest quarter where it seems to be
entirely absent.

One of the remarkable features of this species is the abundance
of the plants to be found on alluvial deposits left by major floods
of the Kansas River. McGregor (1952) reported the occurrence
of this species on silt deposits after the record breaking flood of
1951. It was found that silt covered fields in the river valley con-

110 The University Science Bulletin

tained an average of 1,699,202 rosettes per acre. Hundreds of
square miles of the river valley had plants in incredible numbers
following the flood. Houses and other buildings which had been
filled with a few feet of silt soon had the surfaces of such deposits
dotted with young rosettes of R. frostii.

Our collections of this species seems to agree very well with pub-
lished descriptions and with specimens studied from other areas.

Distribution: Atchison County: M mile south of Muscotah, bank
of Little Grasshopper Creek, October 18, 1947, M1103. Barton
County: 4 miles southeast of Great Bend, bank of Arkansas River,
September 4, 1949, M3950. Chase County: 1 mile south of Strong
City, bank of Cottonwood River, November 27, 1948, M2603.
Cheyenne County: 1 mile southwest of St. Francis, bank of Re-
pubhcan River, August 28, 1947, M1088; August 13, 1948, M2144.
Clay County: south edge of Clay Center, bank of Republican
River, August 22, 1947, M1084. Coffey County: 2 miles north of
Burlington, bank of Neosho River, November 26, 1948, M2595.
Decatur County: Decatur County State Park No. 2, bank of Sappa
Creek, August 12, 1948, M2107. Douglas County: 1 mile northwest
of Lawrence, bank of Kansas River, September 27, 1947, M1092;
November 6, 1948, M2568; September 5, 1951, M5216; 2 miles
northeast of Lawrence, flooded corn field, August 9, 1952, M5685.
Ford County: 7 miles northeast of Dodge City, creek bank, Sep-
tember 5, 1949, M3976. Franklin Coimty: 1 mile north of Norwood,
creek bank, June 19, 1948, M1583. Geary County: Milford, alluvial
deposit, sandbar in Repubfican River, August 22, 1947, M1083.
Hodgeman County: 5 miles east of Grayling, creek bank, September
5, 1949, M3963. Jackson Coimty: Yi mile east of Holton, creek
bank, August 9, 1948, M2001. Jefferson County: 1 mile south of
WilHamstown, bank of Kansas River, September 28, 1947, M1096.
Johnson County: /2 mile north of DeSoto, bank of Kansas River,
September 27, 1947, M1095. Labette County: 4 miles northwest
of Chetopa, creek bank, July 2, 1948, M1614. Leavenworth County:
5 miles northwest of Tonganoxie, bed of dry lake, August 2, 1947,
M1081; /2 mile south of Lin wood, bank of Kansas River, September
27, 1947, M1093. Lincohi County: 1 mile south of Lincoln, bank of
Saline River, July 29, 1952, M5671. Logan County: 1 mile west of
Elkadeer, bank of Smoky Hill River, August 29, 1947, M1090. Lyon
County: 1 mile north of Emporia, bank of Neosho River, November
26, 1948, M2600. Marion County: 1 mile south of Florence, bank
of Cottonwood River, November 27, 1948, M2609. Marshall County:

Taxonomy axd Ecology of Kansas Hepaticae 111

J2 mile east of Blue Rapids, bank of Blue River, August 9, 1948,
M2036. Miami County: Miami County State Park, edge of lake,
October 19, 1947, MHOS; October 3, 1948, M2473. Morris County:
2 miles north of Council Grove, bank of Neosho River, November

27, 1948, M2614. Norton County: 1 mile west of Calvert, bank of
Prairie Dog Creek, August 12, 1948, M2102. Ottawa County: 1
mile south of Bennington, bank of Solomon River, July 29, 1952,
M5670. Pawnee County: 1 mile west of Lamed, creek bank, Sep-
tember 5, 1949, M3960. Pottawatomie County: St. George, bank
of Kansas River, August 22, 1947, M1082. Rawlins County: 6 miles
south of Atwood, bank of Beaver Creek, August 13, 1948, M2141.
Republic County: Scandia, bank of Republican River, August 24,
1947, M1087; August 1953, W. H. Horr. Riley County: Manhattan,
bank of Kansas River, October 22, 1948, Travis Brooks. Roo]<s
County: Stockton, bank of Solomon River, August 27, 1947, M1089.
Russell County: 1 mile south of Lucas, bank of Wolf Creek, July
29, 1952, M56S4. Saline County: Salina, bank of Smoky Hill River,
August 30, 1947, M1091. Sedgwick County: west edge of Derby,
bank of Arkansas River, November 26, 1953, M7522. Shawnee
County: 6 miles east of Topeka, bank of Kansas River, September

28, 1947, M1097; 1 mile north of Topeka, bank of Soldier Creek,
October 12, 1947, Ml 102. Sherman County: 12 miles southwest of
Goodland, bank of Smoky Hill River, September 5, 1951, M5216.
Smith County: M mile south of Claudell, bank of Solomon River,
August 11, 1948, M2094. Washington County: 1 mile west of Linn,
creek bank, August 23, 1947, M1351. Wilson County: 2 miles north-
west of Neodesha, bank of Fall River, October 26, 1947, M1123.
Woodson County: 2 miles northwest of Yates Center, creek bank,
September 25, 1948, M2460. Wyandotte County: Bonner Springs,
bank of Kansas River, September 27, 1947, M1094; September, 1951,
O. F, Fearing.

64. Riccia hirta (Aust. ) Underw. Common on sandy soils in
prairie, over surfacing sandstone, and alluvial deposits below such
areas. It is rarely found on calcareous sites and never in close
association with limestone outcrops.

This species occurs most frequently as gregarious colonies or
scattered rosettes on dry, exposed sandy locations. It is an extreme
xerophyte and regularly rolls up in early summer and remains
dormant until fall rains come when it begins growth again. Mature
sporangia are found in late fall, through the winter, and in early
spring.

112 The University Science Bulletin

R. hirta is one of the more variable species of hepatics in this
area. The variations in ciHation, length-width-thickness ratios, and
color are exceptional. Spore characters also show variations which
apparently have not been recorded. Several colonies of R. hirta
have been studied during eight successive seasons and the degree
of variation noted.

One striking variation concerns the occurrence of marginal cilia.
Usually marginal cilia are present and most abundant near the apex
and become fewer in number toward the basal end. In some speci-
mens cilia are very dense, 30-120[jl long, slender, and pointed. In
others cilia are nearly absent, 30-60[jl long, stout, and obtuse. A few
collections show numerous cilia on the dorsal surface.

The usual color of this species is glaucous-green to blue-green
fading to whitish upon drying on the dorsal surface while the sides
are concolorous to purplish. Ventral scales are usually purplish
but may be hyaline or whitish. In some specimens, from more
xeric sites, the dorsal surface of the thallus has a blue-green or
slate colored surface and blackish purple lateral sides.

Spores of Kansas R. hirta vary from 82-171[;. and have areoles
varying from Q.Q\}.-20\}.. The measurements usually expressed in the
literature are 90-135[;, and 10-13[;. for spore and areole size respec-
tively. The inner faces of spores vary from an appearance exactly
like that of the outer face to definitely less strongly areolate to
irregularly areolate.

The above mentioned variations have been found in the same
colonies. The strongly ciliate, darker pigmented, larger spore
size is correlated with dry seasons and drier periods within a
season though in a given station all variations may be encountered
in an area one meter square. The statements just made would
indicate that material in central Kansas would tend to show
variations in line with that stated for drier situations. Such is
true and, unless caution was used, might lead to unnecessary
taxonomic segregations.

This latter difficulty seems to be apparent in a paper by Jacobs
(1949) in which he described two new species of Riccia which, to
the writer, seem to be no more than ecological variations of R. hiiia
as found in sandstone areas of Kansas and the granite outcrop
areas in the Edwards Plateau of Texas.

The first of these two species described by Jacobs (I.e.) is Riccia
tenella which seems to be a gregarious form of R. hirta that occurs
on shallow sandy soil over surfacing sandstone or granite outcrops,

Taxonomy and Ecology of Kansas Hepaticae 113

in full sunlight — a habitat which readily dries out. Such sites also
become moist quickly after slight rains causing the plants to grow
for a time, become dormant awhile, and then repeat the process.
Such growth has been observed in several stations in Kansas and
invariably resultant plants match perfectly the description given of
R. tenella. These plants typically lack cilia, have a nongibbose
dorsal surface, longer terminal segments which are linear. These
are characters used by Jacobs to distinguish R. tenella from R. hirta.
Other characters, mentioned by Jacobs fall within normal range of
variation to be found in R. hirta. Through the courtesy of Dr. D. L.
Jacobs, specimens of R. tenella were obtained from the type locality
(Jacobs 1251). A study of these specimens reveals that they are
near exact duplicates of the variations of R. hirta mentioned above.
Jacobs further compares R. tenella with R. nigrella, a comparison
which is pointless to one who has seen specimens of the latter
species. A number of Kansas specimens fit the description of
R. tenella but since it is only an ecological variant of R. hirta it
seems best to consider R. tenella as a synonym of hirta.

The second species described by Jacobs (1949) is R. eldeeniae,
likewise an ecological form of R. hirta. Supposedly R. eldeeniae
differs from hirta in having smaller areolae, smaller spores with
both outer and inner face similarly sculptured. Those differences
are to be found within a single colony of R. hirta. R. eldeeniae was
described as having dorsal and marginal cilia. Such characters are
also known in several collections of K. hirta and all other characters
used to separate R. eldeeniae from R. hirta likewise fail. A collec-
tion of R. eldeeniae (Jacobs 920) has been made available for
study by Dr. Jacobs. We find the specimens to be a variation of
R. hirta as mentioned above. It thus seems that R. eldeeniae must
be reduced to a synonym of R. hirta.

From the above it is noted that R. hid a is indeed a variable
species. A detailed study of the taxon in Kansas and from stations
in Arkansas, Oklahoma, Louisiana, and Texas indicates that the
same variations occur in those areas. Only a careful cytological
study coupled with detailed field and cultural studies will show
whether taxonomic segregations are necessary. Our studies indicate
that this must be done over the complete range of the species as
now known.

Distribution: Allen County: 5 miles north of Moran, creek bank,
October 8, 1948, M2502. Anderson County: 5 miles south of Gar-
nett, creek bank, October 8, 1948, M2487; VA miles northeast of

114 The University Science Bulletin

Welda, open places in prairie, October 31, 1948, R. H. Thompson.
Brown County: 5 miles northeast of Hiawatha, sandy roadside
bank, Septemlaer 7, 1948, M2433. Chautauqua County: 3 miles
northeast of Sedan, sandy soil over sandstone outcrop, March 26,
1948, M1211; 4 miles south of Sedan, sandy prairie, July 8, 1949,
M3371; 3 miles north of Elgin, sandy prairie pasture, April 5, 1952,
M5349. Cherokee County: 5 miles east of Baxter Springs, sandy-
clay soil, edge of oak woods, April 30, 1948, M1364. Clay County:
5 miles north of Morganville, sandy prairie, August 16, 1951, M5067.
Coffey County: 2 miles southwest of Le Roy, sandy prairie, April

9, 1948, M1237. Douglas County: 1 mile southeast of Lawrence,
sandy soil. May 24, 1942, R. H. Thompson; 2 miles southeast of
Baldwin City, sandy soil, edge of pond, October 11, 1947, M1098;
5 miles west of Baldwin City, sandy prairie, November 16, 1947,
M1138; October 15, 1949, M4144; August 2, 1951, M5005; October
13, 1951, M5244. Elk County: 10 miles north of Busby, sandy
prairie, April 10, 1948, M1282. Ellsworth County: 5 miles east of
Carneiro, sandy soil, outcrop in prairie, April 30, 1949, M2698;
2 miles northwest of Langley, sandy soil over outcrop, May 3, 1952,
M5450. Franklin County: 1 mile northeast of Norwood, sandy
prairie, March 25, 1948, M1166. Greenwood County: Severy, sandy
soil, September 2, 1945, R. H. Thompson; 1 mile north of Fall River,
margin of sandstone outcrop, April 10, 1948, M1296; April 13, 1949,
M2672; July 9, 1949, M3407. Harvey County: 5 miles north of
Burrton, cattle trail in sand dunes, June 1, 1951, M4916. Jefferson
County: 1 mile east of Williamstown, sandy soil in pasture, Novem-
ber 6, 1948, M2570. Johnson Comity: 5 miles west of Olathe,
sandy bank, November 2, 1947, M1129. Kingman County: 6 miles
northwest of Kingman, sandy roadside bank, August 7, 1948, M2226.
Kiowa County: Rezeau Ranch, sandy prairie, June 12, 1951, M4922.
Labette County: 5 miles southwest of Dennis, sandy bank, April

10, 1948, M1263. Leavemvorth County: Military Reservation,
October 22, 1942, R. H. Thompson; 1 mile west of Turkey Creek
Station, sandy bank, December 2, 1947, M1143. Mia7ni County: 3
miles east of Fontana, river bank, October 19, 1947, M1109. Mont-
gomery County: 3 miles south of Elk City, sandy soil, April 4,
1942, R. H. Thompson; October 25, 1947, M1122; September 22,
1951, M5233. Neosho County: east of Chanute, sandy soil, Febru-
ary 14, 1945, R. H. Thompson. Shawnee County: 7 miles southeast
of Topeka, sandy clay prairie, April 23, 1949, M2684. Wabaunsee
County: 5 miles northwest of Paxico, at base of granite glacial drift
boulders, June 25, 1950, M4370. Washington County: 1 mile west

Taxonomy and Ecology of Kansas Hepaticae 115

of Linn, sandy prairie, August 23, 1947, M1352; 1 mile northeast of
Linn, sandy prairie bank, August 21, 1951, M5071, M5072; August
22, 1952, M5720; 1 mile west of Linn, sandy alluvial deposit, August
22, 1952, M5724. Wilson County: 1 mile northeast of Neodesha,
sandy soil, edge of outcrop, October 25, 1947, M1117; 4 miles north
of New Albany, sandy pockets in surfacing sandstone, July 9, 1949,
M2403; 1 mile east of Neodesha, sandy ledge, April 5, 1952, M5332.
Woodson County: 2 miles northwest of Yates Center, sandy oak
woods, October 26, 1947, Ml 126; Lake Fegan, shallow soil over
surfacing sandstone, September 24, 1949, M4127; 2 miles northwest
of Yates Center, sandy soil, edge of sandstone outcrop, May 12,
1951, M4809; 14 miles southwest of Yates Center, sandy soil, Sep-
tember 1, 1951, R. H. Thompson; October 28, 1951, M5282.

65. Riccia lamellosa Raddi. A species found in the eastern third
of Kansas where it occurs most commonly on sandy soil in prairies,
near sandstone outcrops and moist sandy-clay banks. It will
tolerate calcareous sites and has been found as rare rosettes on
black soil over surfacing limestone in oak-hickory woods.

In late fall and early spring this species appears as rosettes or in
gregarious colonies. During summer or dry seasons it is found as
rare dichotomies of small size but otherwise is absent. It does not
seem to withstand prolonged drought as it apparently does in Okla-
homa and Texas.

Our studies on this species in Kansas verifies the action of Miiller
(1952) in which he reduces R. austini Steph. to a synonym of R.
lamellosa Raddi. We fail to find a single character which consist-
ently could be used to separate American material from the Euro-
pean R. lamellosa. The size of the ventral scale in particular is a
rather variable feature and is determined by ecological conditions.
Distribution: Anderson County: VA miles northeast of Welda,
open places in prairie, October 31, 1948, R. H. Thompson. Chase
County: 1 mile south of Strong City, bank of Cottonwood River,
November 27, 1948, M2606. Chautauqua County: 3 miles northeast
of Sedan, sandy soil, oak woods, March 26, 1948, M1210; 3 miles
north of Elgin, sandy prairie pasture, April 5, 1952, M5351. Chero-
kee County: 3 miles southeast of Baxter Springs, rocky pasture,
October 22, 1951, M5263a. Clay County: 5 miles north of Morgan-
ville, sandy prairie, August 16, 1951, M5065. Douglas County: 1
mile southeast of Haskell, sandy soil, May 7, 1942, R. H. Thompson;
3 miles southeast of Baldwin City, sandy soil over sandstone ledge,
November 2, 1947, M1132; 5 miles west of Baldwin City, sandy soil

116 The University Science Bulletin

over surfacing sandstone, November 16, 1947, M1139; October 13,
1951, M5245. Elk County: 10 miles north of Busby, sandy prairie,
April 10, 1948, M1281. Ellsworth County: 4 miles northwest of
Langley, sandy soil over surfacing sandstone. May 3, 1952, M5449.
Greenwood County: 1 mile northeast of Fall River, sandy soil,
April 10, 1948, M1297. Jefferson County: 1 mile west of Medina,
sandy prairie bank, November 16, 1947, M1134. Johnson County:
5 miles west of Olathe, black soil over surfacing hmestone, Novem-
ber 2, 1947, M1131. Leavenworth County: 2 miles west of Lin-
wood, sandy soil, oak woods, June 16, 1949, M3088. Lyon County:
1 mile north of Emporia, bank of Neosho River, November 26, 1948,
M2598. Marion County: 1 mile south of Florence, bank of Cotton-
wood River, November 27, 1948, M2611. McFherson County: 2
miles northwest of Lindsborg, sandy prairie. May 3, 1952, M5439.
Miami County: 3 miles east of Fontana, river bank, October 19,

1947, M1113. Montgomery County: 3 miles southwest of Elk City,
sandy bank, October 25, 1947, Ml 121; sandy soil over surfacing
sandstone, September 22, 1951, M5231. Morris County: 2 miles
north of Council Grove, bank of Neosho River, November 27, 1948,
M2616. Neosho County: 2 miles east of Chanute, sandy bank, No-
vember 27, 1947, M1140; sandy soil around Santa Fe Pond, October
30, 1948, R. H. Thompson; 3 miles east of Chanute, sandy soil. May
12, 1951, R. H. Thompson. Shawnee County: 5 miles east of To-
peka, soil over surfacing limestone, April 23, 1949, M2678. Wash-
ington County: 1 mile west of Linn, sandy prairie bank, August 10,

1948, M2046; 2 miles southwest of Palmer, sandy prairie. May 27,

1951, M4827; 2 miles northeast of Linn, sandy prairie, August 22,

1952, M5719. Wilson County: 1 mile northeast of Neodesha, sandy
soil over surfacing sandstone, October 25, 1947, M1116. Woodson
County: 2 miles northwest of Yates Center, sandy soil, October 26,
1947, Ml 127; Lake Fegan, sandy prairie bank, September 24, 1949,
M4128; 2 miles northwest of Yates Center, sandy soil over surfacing
sandstone. May 12, 1951, M4808.

66. Riccia rhenana Lorbeer. This species was first reported from
North America and Kansas by McGregor (1952). In Kansas the
species has a scattered distribution but seems to grow in most abun-
dance in water below springs or in cattail-sedge marshes.

Kansas specimens exhibit all the characteristics described for the
species and match European material of the taxon. It has been
found in water and as strand plants, but never produces reproduc-
tive structures. This makes its survival in our cattail-sedge marshes

Taxonomy and Ecology of Kansas Hepaticae 117

a matter of interest as they often dry up completely and for long
periods of time. However, R. rhenana makes its appearance soon
after such marshes are refilled. It has been observed that the
apices of the plants resist rather severe dessication and this may
account for survival and spread by waterfowl.

Distribution: Douglas County: 2 miles northeast of Lawrence,
on mud, edge of cattail-sedge marsh, May 9, 1948, R. H. Thompson;
1 mile northeast of Lawrence, in water, marsh, August 27, 1949,
M3S94; 3 miles northeast of Lawrence, in cattail-sedge marsh,
November 6, 1949, R. H. Thompson; November 13, 1949, M4162;
September 23, 1950, M4677; November 30, 1951, M5294; May 8,
1952, M5461. Jefferson County: 3 miles west of Williamstown,
marsh in Kansas River Valley, November 13, 1949, M4164. Kiowa
County: 5 miles north of Belvidere, in water below a spring, June
12, 1951, M4921. Linn County: 2 miles west of La Cygne, in
cattail-sedge marsh, June 4, 1949, M2857. Meade County: Meade
County State Park, in pond fed by artesian water. May 8, 1942,
W. H. Horr; May 16, 1948, W. H. Horr 3111; August 16, 1948, M2199.
Miami County: Pigeon Lake, shallow water, March 16, 1947, M1056;
April 24, 1948, M1349; May 29, 1949, M2774.

67. Riccia sorocarpa Bisch. A rather uncommon species in
Kansas and never found in any abundance. It occurs on sandy soil
in prairie, borders of sandstone outcrops, and alluvial deposits. This
species is essentially an early spring plant, disappearing during the
summer, but found again in the late fall. It is the first species of
Riccia to disappear as the various stations dry out and thus is mesic
in its requirements.

Kansas material never seems to branch more than once and thus
appears as scattered dichotomies which fruit abundantly but do
not develop into gregarious colonies. Otherwise our material seems
typical of the species.

Distribution: Anderson County: 2 miles south of Garnett, sandy-
clay soil, open places in oak-hickory woods, June 4, 1951, M4875.
Chautauqua County: 3 miles northeast of Sedan, sandy prairie,
March 26, 1948, M1219. Cherokee County: 5 miles east of Baxter
Springs, creek bank, October 22, 1951, M5258. Clay County: 5
miles north of Morganville, sandy prairie, August 16, 1951. Craw-
ford County: 2 miles west of Hepler, field. May 1, 1948, L. J. Gier
1073. Douglas County: University of Kansas campus. May 26, 1942,
R. H. Thompson. Leavenworth County: 4 miles northeast of
Tonganoxie, sandy bank, May 7, 1948, M1403. Montgomery

118 The University Science Bulletin

County: 3 miles south of Elk City, sandy soil, April 4, 1942, R. H.
Thompson; 6 miles south of Elk City, sandy soil, oak woods, March
26, 1948, M1220. Neosho County: 3 miles east of Chanute, sandy
soil, May 12, 1951, R. H. Thompson. Shawnee County: 7 miles
southeast of Topeka, sandy soil, overgrazed prairie pasture, April 23,
1949, M2685.

68. Riccia stiUivontii Aust. Infrequent on alluvial deposits along
streams in the eastern fourth of Kansas. It is not abundant nor is
it to be found in successive years in the same stations. The species
occurs in late fall, usually after fall rains, thus the length of its
growing season is curtailed and mature sporangia rarely develop.
Kansas appears to be the western limit of the range of this species.

Distribution: Allen County: 5 miles south of Moran, bank of
stream, October 8, 1948, M2500. Anderson County: 4 miles south
of Garnett, alluvial deposit, creek bank, October 8, 1948, M2483.
Atcliison County: M mile south of Muscotah, alluvial deposit along
Litde Grasshopper Creek, October 18, 1947, M1104. Bourbon
County: 1 mile south of Uniontown, bank of Marmaton River, Oc-
tober 8, 1948, M2516. Cherokee County: 5 miles east of Baxter
Springs, alluvial deposit, bank of Shoal Creek, August 30, 1948,
M2355, M2367; October 22, 1951, M5264. Douglas County: 2
miles northeast of Lawrence, bank of Mud Creek, September 26,
1948, M2462. Leavenworth County: 2 miles north of Leavenworth,
bank of Missouri River, October 28, 1948, M2563. Miami County:
3 miles east of Fontana, bank of Marais des Cygne River, October
19, 1947, Mini; October 3, 1948, M2468; October 18, 1950, M4680.

69. Riccia trichocarpa Howe. Limited to occurrence on sandy
soil near surfacing sandstone in prairie or oak-hickory woods. The
species is an extreme xerophyte and is to be found on the driest,
exposed sandy stations.

Disti-ibution: Chautauqua County: 3 miles northeast of Sedan, on
sandy soil in oak-hickory woods, March 26, 1948, M1213. Douglas
County: 5 miles west of Baldwin City, sandy soil near sandstone out-
crop, November 16, 1947, M1137; November 7, 1948, M2575; October
15, 1949, M4145; March 30, 1952, M5326; April 26, 1952, M5433.
Elk County: 10 miles north of Busby, sandy soil in oak woods, April
10, 1948, M1285. Franklin County: 1 mile northeast of Norwood,
sandy soil, March 25, 1948, Ml 167. Greenwood County: 1 mile
northeast of Fall River, sandy soil in oak woods, April 10, 1948,
M1290. Montgomery County: 3 miles southwest of Elk City, sandv
soil near sandstone outcrop, October 25, 1948, Ml 120; 6 miles south

Taxonomy and Ecology of Kansas Hepaticae 119

of Elk City, sandy soil in oak woods, March 26, 1948, M1184; 3 miles
southwest of Elk City, sandy prairie, September 22, 1951, M52.35.
Wilson County: 1 mile northeast of Neodesha, sandy soil near
sandstone outcrop, April 10, 1948, M1273. Woodson County: 2
miles northwest of Yates Center, sandy soil in Prairie, October 26,
1947, M1125; April 9, 1948, M1245; May 12, 1951, M4811; October
28, 1951, M5279.

Excluded Species

A few species have been reported in the literature as present in
Kansas for which specimen evidence is not known. Such reports
were probably based on incorrect determinations as it seems unlikely
that some of them could occur in the State. All were listed by
Smyth and Smyth (1911) and are as follows:

1. Chiloscijphus polyanthus (L.) Corda.

2. Fossombronia angulosa Raddi.

3. Jungermannia schraderi Martins.

4. Leucolejeiinea chjpeata ( Schweinitz ) Evans.

5. Pallavicinia hjcllii Gray.

6. Pellia endiviaefolia (Dicks) Dum.

7. Preissia qttadrata (Scop.) Nees.

8. Ptilidiiim ciliare ( L. ) Hampe.

9. Riccardia latifrons Lindb.

ECOLOGICAL STUDIES

Location and Area

Kansas is located in the center of the United States and has a
generally rectangular shape. The north and south boundaries are
the parallels of 40°N. and 37°N., respectively; the eastern bound-
ary is the Missouri River and 94°38' W.; and the western boundary
is 102°1'34"W. This area— about 210 miles north-south and 410
miles east-west — contains an area of about 82,000 square miles.

Climate of Kansas

It is necessary to preface a discussion of ecology in Kansas by
considering the recorded data on temperature, moisture, winds, etc.
The present discussion of climate in Kansas is based directly on
the publication of Flora (1948) and supplemented observations of
the writer.

As a whole the State has an annual mean temperature of 55°F,
almost equal to that of Virginia, more sunshine than any state to
the east, and summer rains which average, in eastern counties,
higher than those of other states except those along the Gulf Coast.

120 The University Science Bulletin

This combination of weather elements makes the State a leading
agricultural region for grains and livestock, but tends to restrict the
development of a hepatic Hora which a consideration of weather
elements will show. The 400 mile spread of the State from east
to west results in three areas having rather distinct climates which
may be designated as eastern, middle, and western.

Rainfall: Rainfall varies from an average of nearly 36 inches in
the eastern third of the State, to about 26.5 in. in middle, and to
19 inches in the western third. This rainfall is often not equally
distributed during the season with a result that prolonged dry pe-
riods, such as occurred in 1952-53, may affect the flora more than
the long term averages would lead one to suspect.

Humidity: The average relative humidity over Kansas is less
than in any state east of the continental divide of the Rockies.
Average midday and evening humidities over the State in July
range from 30-40 per cent in the western part to 45-50 per cent in
the eastern part. This low relative humidity allows for rapid evap-
oration and drying out of most of the habitats in the State. This
has a profound effect on Hepaticae.

Temperatures: As noted previously Kansas has a mean annual
temperature of 55°F. The warmest month of the year is July with
an average mean temperature of 79°F and the coldest is January
with an average mean of 30' F. The lowest temperature ever re-
corded officially in the State was 40°F below zero and occurred
in north-central Kansas. The highest official temperature was re-
corded in the southeastern part of the State at 121 °F.

Winds: The prevailing direction of wind in all parts of the State
is from the south and southwest, April to November, inclusive. In
other months northerly winds prevail. In eastern Kansas winds
have an average hourly movement of 9 miles while they average
12 miles in southwest Kansas.

An outstanding effect of wind occurs occasionally during the
course of hot summer weather. At such times winds of rather
high velocity, low relative humidity, and with shade temperatures
of 100-110°F or higher have been known to kill foliage and destroy
growing crops. These are the "hot winds" which the writer has
observed to kill a corn crop in one day's time.

Frosts: The average last date for killing spring frosts ranges from
April 9 in extreme southeastern Kansas to May 2 in the extreme
northwestern part of the State. However, killing frosts have been

Taxonomy and Ecology of Kansas Hepaticae 121

known to occur as late as the second week of May in southeastern
Kansas and in the last week of May in extreme northern counties.

The average date of first killing frost is the first week in October
in northwestern Kansas to the last week of October in the southeast.
Killing frosts have been known in the first week of September in
the northwest to the third week of September along the southern
border.

The climate of Kansas is, then, one of great extremes and sudden
changes in temperature, precipitation, and wind. Conditions at
any one point may vary widely from year to year and very often
are extremely unsatisfactory for vegetation. It must be empha-
sized that it is the extremes of climate which most affect the occur-
rence and distribution of hepatics in the State, rather than the av-
erage which at first glance might seem to be favorable for a more
diverse vegetation including liverworts.

Vegetation Areas in Kansas ^

Studies on the vegetation of Kansas during the past 100 years
have been concerned mainly with attempts to obtain a complete
record of all species found to occur in the area. Numerous publica-
tions have been produced which give lists of species and counties
where such species have been found. As yet no one has made a
serious attempt to delimit and define vegetative areas in Kansas.
It is true that, as yet, insufficient field studies have been made
and thus it is not possible to prepare a map of vegetation areas
which would show more than broad generalizations.

A few publications have included discussions and maps which
involve vegetation areas to some extent. Some of these are Brum-
well (1941), Cockrum (1952), Fearing (1952), and Gates (1937,
1940 ) . All of these have contributed to our knowledge of vegetative
areas either directly or indirectly but none have been close to
adequately portraying the true nature of vegetation assemblages in
Kansas.

Since none of the previously published accounts of vegetation
areas in Kansas seem to be of much value, the writer here presents,
as map 1, a break-down of the major vegetation areas in Kansas.
In preparing this map it was not intended that it show all minor
zones or deal with the complexities of the numerous transition
areas. The map is based on the writer's studies, made in each
county of the State, of the distribution of bryophytes and all higher

1. Information on the Physiography of Kansas adapted from Schoewe (1949).

122

The University Science Bulletin

plant groups. Future studies will, without doubt, alter the bounda-
ries of some areas here proposed.

The small Ozark area in extreme southeastern Kansas is an ex-
tension into Kansas of the Springfield Plateau in Missouri and is the
edge of a westward dipping plain underlain by limestones of Mis-
sissippian age. The area has a relief which averages less than
60 feet, but has a number of bluffs along streams, is rocky, and has
outcrops of limestone.

This small Ozark area of some 50 square miles is botanically
one of the more interesting parts of the State. More species of

CHAUTAUQUA HILLS^
Map I. The map shows vegetation areas in Kansas.

OZARK-"

higher plants are known from this area than for any other of equal
or larger size in the State. In this area of Kansas a number of
eastern and Ozarkian taxa find the western limit of their range.
In addition a number of plants common in this Ozark area are
found only rarely elsewhere in adjacent vegetation assemblages.
Some of the taxa known in Kansas from the Ozark area only are:
Isoetes butleri, Dnjopteris hexagonoptera, Saurtirus cernuus, Arabis
laevigata, Asclepias qiiadrifolia, Ltidwigia glandulosa, Pohjinnia
iwedalia, Corniis forida, Ulmus alata, Vacciniuni stamineum var.
interius, Vaccinitim vacillans var. missoiiriensis, Hieraciwn gronovii
and Aster tiirbinelhis.

The Ozark area is characterized by oak-hickory wooded hillsides
on which the common trees are Quercus alba, Q. macrocarpa, Q.
muehlenbergii, Q. marilandica, Q. shumardii, Q. velutina, Carya
cordiformis, C. laciniosa, C. ovata, C. texana, Celtis occidentalis.
Ulmus americana, U. rubra, and Acer saccharum. The understory

Taxonomy and Ecology of Kansas Hepaticae 123

is dominated by Symphoricarpos orbiculatiis, Aesculus glabra var.
sargentii, Ostrya virginiana, Staphylea trifoUa, Cercis canadensis
and numerous other plants. Among the forested hillsides, hilltops
and bluffs are islands of tall grass prairie which are dominated by
Andropogon gerardi, Sorghastrtim nutans, Tripsacum dactyloidcs,
and Panictim spp. These prairies often have thickets or marginal
bands of Sassafras albidiim, Diospyros virginiana and Primus spp.
A few marshes, streams, and extensive rock outcrops offer a variety
of additional habitats for hepatics.

The Osage area includes parts of three physiographic divisions.
The southern third is an erosional plain whose surface slopes to the
west at an average rate of 10 feet per mile and is developed from
weak shales and sandstones. The surface is gently undulating
except for a few broad, flat-topped mesalike wooded hills which
are capped by resistant sandstone. Some of these hills rise 80 or
more feet above the surrounding country. Several wide, shallow,
flat bottomed valleys occur. The central section of the Osage con-
sists of a series of northeast-southwest escarpments which face east
and between which there are flat, to gently rolling plains. The crest
of each escarpment is capped by resistant limestone. These escarp-
ment fronts range from a few feet to more than 200 feet in height.
The northern part of the Osage is a glaciated area and for the
most part is an erosional drift-controlled surface which is gently
undulating. Near large streams the country is dissected into rough
and hilly areas with river bluffs too rough for cultivation and re-
vealing ledges of limestone and some sandstone. Interstream areas
are smooth, broad, and well rounded and become more dissected
as one approaches larger stream courses. Most valleys are wide
and open.

The Osage vegetation area consists basically of oak-hickory
wooded hillsides, creek valleys, and ravines. Uplands were origi-
nally covered with tall-grass prairie dominated by Andropogon
gerardi. It is an area much disturbed by man and original vegeta-
tion is to be found primarily in out-of-the-way places. Woods, in
general, are dominated by Quercus macrocarpa, Q. miiehlenbergii,
Q. veliitina, Carya cordiformis, C. ovata, Ulmiis americana, U. rubra
and Celtis occidentalis. In some locations significant stands of
Acer saccharum, Tilia americana, Quercus alba, Juglans nigra and
Fraxinus lanceolata are found. The understory consists principally
of Symphoricarpos orbiculatus, Ostrya virginiana, Aesculus glabra
var. sargentii, Asimina triloba, and Staphylea trifolia. Characteristic

124 The University Science Bulletin

herbs are PodophylUim peltattmi, Carex spp., Agrimonia spp., Sani-
ciila marilandica. Phlox dwaricata, Isopyrinn hiternotum, and others.

Prairies of the Osage area are dominated by Andropogon gerardu
In dry sites Andropogon scoparius and Boutdoiia cnrtipcndida be-
come evident while in moist places Tripsacum dacttjloides, Sorghas-
triim nutans, Triodia flava and Panictnn virgatiim may be locally
dominant. In all cases species of Panicwn and Poo are usual compo-
nents. A number of forbs give the prairie seasonal aspects. Some of
these are Baptisia leucophaea, Sisijrinchiwn campestre, Phlox piJosa,
Hypoxis hirsuta, Tradescantia hracteata, Senecio plattensis, Silphiwn
laciniatiim, Helianthiis spp., Ratihida pinnata, Rtidbeckia serotina,
Liatris pycnostachya, Solidago spp., Aster spp., Amorpha canescens,
Petalostemum purpureum. Psoralen esctdenta, P. tenuifora and
many others.

A number of streams dissect the Osage area and in the valleys
formed the usual Salix, Populus, Ulnius, Acer and Fraxinus stream
bank floristic assemblages are found. In the lower third of this
area significant stands of Querciis pahistris are found in lowland
woods.

The Chautauqua Hills include a belt developed on thick sand-
stones in which erosion has dissected the surface into a series of
low hills which are intersected by many small streams which have
cut deep valleys. This is an area distinguished primarily by sandy
soil and sandstone capped hills whose tops and upper slopes are
forested with Quercus marilandica, Q. stellata, Q. prinoides, and
Q. velutina. Along streams and in the northern parts of the area
many of the trees of the Osage become admixed with the above.
The prairies of the Chautauqua Hills are simliar to those of the
Osage except that they occur on sandy soils and those of the latter
are predominantly calcareous and thus drier. The sandstones of the
Chautauqua Hills extend northeastward to Leavenworth, in Leav-
enworth County. A few small areas in Douglas, Franklin, Jefferson,
Leavenworth and Wyandotte Counties should be considered as iso-
lated elements of the Chautauqua Hills. Jewett ( 1941 ) believes that
these areas should be so included in the Chautauqua Hills and the
hepatic flora corroborates Jewett's opinion.

The Flint Hills area features a range of hills about 20 miles
wide extending down the center of the area from Marshall County
in the north to Cowley County in the south. The eastern front of
these hills has a very prominent rocky escarpment several hundred
feet high. This highly dissected east-facing escarpment has ter-

Taxonomy and Ecology of Kansas Hepaticae 125

raced rocky slopes. The rocks of the area are limestone which bear
flint. The surface of the Flint Hills is gently rolling and merges on
the west with a smooth gentle slope toward the Arkansas River.
On the east the Flint Hills merge gradually with the Osage area
below the escarpment. The streams of the Flint Hills flow in steep
precipitous channels lined with rock ledges.

The Flint Hills area is best characterized as a blue-stem grass
region. On the uplands and dry places Andropogon scoparhis is the
dominant species while hillsides and moist places are dominated by
Andropogon gerardi. Bouteloua curtipendula is usually present
with both of the bluestems. Forbs are similar to those of the Osage
prairies except a larger number of legumes are present. Along the
streams are found a limited amount of forests which are tongues or
fingerlike projections of the Osage forest through the Chautauqua
Hills and from the Osage area of the north into the Flint Hills.
Trees found are usually Uhnns americana, Qiierctis macrocarpa,
Froxinus lanceolato, Acer negundo, Poptdus dehoides and Salix spp.
The Flint Hills are of little importance in hepatic distribution since
species present in the area are few in number, found only on bark
or stream banks, and consist of ubiquitous taxa.

The Dakota area consists of a maturely dissected, broad, hilly,
belt carved in Dakota sandstone and having a relief at places of
from 200-300 feet. Numerous outlying hills and mounds are found.
Stream valleys are wide but small tributaries often have abrupt
banks with outcroppings of sandstone. These often have springs or
ai-e seepy and usually shaded. This area is conspicuous due to the
frequent outcroppings of the dark Dakota sandstone. The original
vegetation was dominated by the bluestem grasses Andropogon
scoparius and Andropogon gerardi but overgrazing and drought
have allowed the more resistant short grasses Biichloe dactyloides
and Bouteloua spp. to become dominant and, in severe cases, annual
grasses, such as Hordeum pusillum, have invaded. Therefore, the
Dakota is not stabilized and is mainly one of transition between
bluestem on the east and shortgrasses on the west. Forbs also
represent a mingling of eastern and western species. No species
known in Kansas are confined to this Dakota; but since the area is
well defined, has a better soil moisture relationship than areas to
the east or west, it does form a discreet vegetation area as noted by
Gates (1937). The stream banks and valleys of the Dakota are
forested similar to those of the Flint Hills. The numerous sand-
stone exposures along with sandy soil prairies and the trees, particu-

126 The University Science Bulletin

larly Quercus macrocarpa, allows for a hepatic flora which ranks
third, behind the Chautauqua Hills and Ozark areas, in number of
plants present.

The Arkansas River is a loosely defined area consisting of the
Arkansas River valley and associated low, undulating plains along
the river and its tributaries. The soil is sandy and vegetation is dom-
inated by sand grasses both along the streams and in the extensive
sand dune formations located in the area. Doell (1938) reports
that the sand hills of Harvey, Reno and Rice counties are similar to
the sand hills westward along the Arkansas River. Doell believes
that the sand hills present a true transition between tall and short
grass prairies and that stability is never reached. In the Arkansas
River both tall and short grass associations are found. The latter
are located on drier more stable soil; but the very sandy and sand
dune parts, so characteristic of the area, though including plants of
both short and tall bluestem grass associations in addition have some
distinctive species such as Calamovilfa gigantea, Andropogon hallii
and Panicum mrgatum which are found to be locally dominant. The
forbs present are also a mixture from other areas but several are
found only in the Arkansas River vegetation assemblage. In general
the vegetation is xerophytic and grasses are the dominant plant
cover with Salix spp., Popuhis spp., and thickets of Pruntis spp.
forming the only significant woody components. Hepatics are few
in number and are found along cattle trails, through moist sandy
pastures, or around bodies of water in dune areas. All are ubiqui-
tous species.

One of the most picturesque areas of Kansas is the Red Hills. It
has red soil and outcrops of red shales, silt-stones, sandstones, includ-
ing gypsum and anhydrite, of Permian age. Barber County is espe-
cially scenic with many steep-sided buttes, pinnacles, and stream
valleys lined by steep bluffs. Vegetation of this area is xerophytic
and the most conspicuous plant, now largely removed, is the red
cedar Juniperus virginiana. Plant cover is predominantly the short
grasses Biichloe dactyloides and Boutelotia gracilis. A large number
of forbs are found, a few of which, are known only in Kansas from
the Red Hills. Few mosses and virtually no hepatics are to be
found in this area except near springs where conditions are similar
to areas in the eastern part of the state.

The High Plains area is also known as the short grass region in
which the dominant plants are the short grasses Biichloe dactyloides
and Bouteloiia gracilis. Many other distinctive grasses and forbs

Taxonomy and Ecology of Kansas Hepaticae 127

occur, some of which are found, in Kansas, only in the High Plains
area. This well-defined area is nearly devoid of bryophytes and
need not be further considered.

Habitats and Factors

Very little experimental data concerning environmental factors in
relation to bryophytes, especially Hepaticae, is available. With re-
gard to this Garjeanne ( 1932 ) says, "Though it is sometimes possible
to grow them for shorter or longer periods, in the end they always
die: this is certainly a great handicap to gaining a more intimate
knowledge of the Bryophytes." This statement while not certainly
applicable to all bryophytes is in general true and has handicapped
the study of this phase of Bryology.

In the absence of specific experimental data, careful field observa-
tions have been made on a variety of habitats and involve trans-
planting, pH, moisture, temperature, and soil studies. From these
a number of generalizations may be made. The discussions of Rich-
ards ( 1932 ) and Garjeanne ( 1932 ) encourage the use of physiologi-
cal and ecological characteristics which bryophytes have in common
with vascular plants and note that factors which affect one group will
also affect the others. In our area it is the degree to which such
factors affect various plant groups that becomes important.

Moisture conditions of both the substratum and the atmosphere
are of prime importance in the growth and survival of hepatics.
The occurrence of an abundance of hepatics in a few Kansas sites
can be directly correlated with moisture conditions of the substratum
and perhaps more importantly with that of the relative humidity of
such places.

In most of our areas the majority of species of hepatics are to be
found on moist sandstone outcrops, sandy prairie, around and below
springs or in shaded protective places which dry out slowly. It is
often possible to collect over sections of forested areas and find only
a few widely tolerant ubiquitous species. However, in the same
terrain on certain outcrops or seepy ledges a larger number of species
may be found. If the localized sites are characterized by very
seldom drying out, a good hepatic flora will be found. Any pro-
longed drouth will, however, reduce the colonies and may lead to
extermination of certain species.

A study of Kansas hepatics leads one to the conclusion that, except
in a few cases, most species are fighting for survival; and that with
the disturbance by man, some have probably disappeared from the

128 The University Science Bulletin

scene while others are barely surviving. In contrast disturbance
has probably been responsible for an increase in numbers of some
species such as members of Riccia, Sphaerocarptis, Frullania and
Aster ella.

The only species, in Kansas, which actually live in water are the
water forms of Ricciocorpiis natans, Riccia jiuitans, R. rhenana, and
possibly the plant provisionally referred to as Riccia canaliciilata.
A few exist on very wet mud banks or alluvium of streams and lakes.
Among these are land forms of the four species just mentioned and
Phaeoceros laevis, Notothijlas orbicularis, Conocephahim conicum,
Marchantia polymorpha, Plectocolea hyalina and several species of
Riccia. These latter, however, are present on such sites due to water
dispersal and are only temporary elements except for Riccia frostii
and R. stilUvantii.

A few species seem capable of withstanding severe flooding for
prolonged periods. These are usually species found on the faces
of sandstone outcrops on creek banks. Plants in this class are
Conocephalum conicum, Marchantia polymorpha, Plectocolea laevis,
Reboulia hemisphaerica and Porella pinnata which is found on the
bark of trees at the edges of streams. In addition Frullania inflata
has been observed to withstand flooding while growing on the bark
of trees found on flood plains.

The only perennial species which seem to be able to survive on
substrata which regularly dry out are: Cephaloziella rubella, C.
byssacea, Lophozia bicrenata, Mannia jragrans, Asterella tenella,
Oxymitra paleacea and several species of Riccia. Scapania nemo-
rosa is able to survive in a few places where the above occur but
actually seems to be losing out on such sites. A few annual species
appear on these locations during moist springs or falls. Among
these are: Sphaerocarpus texanus, S. michelii, Fossombronia bra-
ziliensis and Anthoceros punctatus.

Most species of hepatics in Kansas occur on sites which are so
located that moisture is present the year round and in which the
relative humidity is higher than in the surrounding area. Such
areas consist of moist outcrops in shaded ravines or along creek
banks, shaded, protected north hillsides and at the base of grass
clumps in tall grass prairie.

Moisture is, then, a critical controlling factor in the occurrence
and distribution of hepatics in Kansas.

Light seems to be another factor to be considered in determining
distribution of hepatics; but it is most diflBcult to evaluate because

Taxonomy and Ecology of Kansas Hepaticae 129

of its effect on temperature and humidity. In general light require-
ments of hepatics are not generally high. This is well shown, in
Kansas, in that most hepatics grow in shaded habitats or at least
where light is much reduced. In general the Marchantiales are
light-loving while the rest of the Hepaticae have low light require-
ments. In general, the more moist the location, the more tolerant
our species are of light. It is, therefore, difficult to evaluate the
effect of light on liverworts.

The nature of the substratum is another factor important in
hepatic occurrence and distribution. Species growing on tree bark
are limited to host plants from which the bark does not readily flake
or slough off. FruUania spp., for instance, regularly are found on
bark of Qiiercus velutina and Q. macrocarpa but the more abundant
Q. miiehlenbergii has few colonies due to the more loose, flaky bark.
If Q. miiehlenbergii is found in a moist ravine the number of colo-
nies of FruJlania on this tree increases. Thus, a moisture factor is
involved. In this case, however, the flaking of the bark seems to be
an important factor as occurrence of species on bark in a given area
is directly correlated with permanency of bark. Soft sandstones
likewise have a small hepatic flora while resistant sandstones in the
same area support a good population.

The nature of the soil is important in distribution. This is often
correlated with moisture relationships of the soil. In Kansas better
than 90 per cent of soil hepatics are to be found on distinctly sandy
sites. These soils have a more permanent moisture supply and
better aeration. Cultures have been tried on sandy soil, loam, clay
and on mixtures of these. Successful growth has been found only
on sandy soils.

A few of our hepatics seem to prefer a nitrogen-free medium.
Cephaloziella rubella and Lophozia bicrenata seem regularly to
occur in nitrogen-free sites and seldom are to be found on any other
type of habitat. Griggs ( 1933, 1935 ) gives a discussion of a nitrogen-
free medium by Cephalozia bicrenata and Cephaloziella byssacea.

The most completely investigated substrate factor is that of
hydrogen-ion concentration. A number of observations have been
made on a large number of bryophytes and the findings reported
in the literature. In Kansas most of our hepatics occur on neutral
or just under neutral pH conditions. Distribution does not seem to
be related so much to pH as to availability of calcium. Calcium
may be easily available under slightly acid conditions and a species
may react much more to calcium occurrence than to hydrogen-ion

5—6730

130 The University Science Bulletin

concentration. Some workers such as Meylan (1924) consider
species occurring in conditions with pH less than 7.0 as lacking
tolerance for lime containing rock or soil. In our area by far the
majority of our species occur on calcareous sandstones or sandy
soils in which pH is neutral, slightly acid, or shghtly alkaline. If
the work of Meylan (I.e.) was followed, most of our species would
alternately be classed as lime avoiding or lime loving as the site
might be. However, the lack of very acid stations undoubtedly
limits the hepatic flora of the State as it has been shown by Schuster
( 1949 ) that uncommon species often appear very much restricted in
occurrence by pH conditions. Thus pH probably has Httle bearing
on the distribution of hepatics now known in Kansas.

Temperature effects on our hepatic distribution is difficult to
evaluate because of its effects on humidity and moisture in general.
It has been observed that sudden drops of temperature results in
death of some members of tlie Marchantiales. Colonies of Conoce-
phalum conicum, Reboulia hemisphaerica, Riccia campbelliana and
Mannia fragrans are quickly killed back to growing tips by sudden
drops in temperature and if the drop goes from temperatures, which
have averaged above freezing to a reading of 20° F. or lower, even
the growing tips succumb. In general, though, temperature is un-
important in our area except in its effect on moisture conditions —
an effect which is often most severe.

Invasion and Succession

Though many species of hepatics seem to be attempting to survive
in Kansas habitats, several are capable of not only existing but of
invading and some successional stages can be demonstrated. The
literature on invasion and succession by Hepaticae is meager and
most of it in this country is of more or less recent publication.

Durand (1908), Skutch (1929), Emig (1930), Torrey (1932a,
1932b), Graff (1935, 1936), Sharp (1939), and Schuster (1949) all
have noted the invasion of Marchantia pohjmorpha upon areas fol-
lowing fires. This was observed once in Kansas at a station in a
forest dominated by Quercus stellata, Q. marilandica, Q. veltitina
and with an understory of Q. prinoides, Cornus drinnmondii and
Rtibus spp. During one fall much of the understory and a few
oak trees were cut. The brush and tree branches were piled in a
number of heaps over an area several acres in extent and burned.
This left several large circular burned places which the next spring
were populated by sohd growths of Marchantia polymorpha and the

Taxonomy and Ecology of Kansas Hepaticae 131

moss Funaria hygrometrica. The plants of Marchantia pohjmorpha
were small, exceptionally fertile and were to be found only on the
burned spots. This occurrence was remarkable in that M. poJij-
morpha was not previously known near the station and since has not
been located near the burned areas. The nearest permanent station
was some 25 miles away. The second year following the burning
only a few plants of Marchantia were present on these burned sites
and in following years none were discovered. In the meantime herbs
and grasses invaded the circular spots and eventually Quercus
prinoides dominated the sites.

On a few sandy-clay banks which have been recently eroded,
and in which the soil is nearly nitrogen-free an invasion by
Cephaloziella rubella occurs. In a few sites this species is soon
followed by Lophozia bicrenata. Various species of Clodonia
seem to follow these two hepatics closely and eventually lead to
occupancy by several species of mosses, later by shrubs, grasses
and often by either Mannia fragrans, or Asterella tenella. Later
the hepatics lose out in about the same order as the original
succession.

Invasion of moist sandstone ledges is initiated by Phaeoceros
laevis or Plectocolea hyalina. These may exist separately or to-
gether and very often appear to maintain their dominance of
the site. This latter happening appears most often in places
where the site is partially exposed to full sunlight. In more shaded
conditions Rebotdia hemisphaerica may soon appear and in
humid shaded places Conocephalum conicum may take over and
eventually exclude most of the earlier arrivals.

Moist limestone ledges are so rare in Kansas that little in the
way of possible invasion — much less in way of succession — can
be noted. Dry limestone ledges are plentiful and are colonized
by Fridlania inflata or Frullania riparia. Often these two are
found as mixed colonies. On the crests of such ledges Forella
platyphylla, P. platyphylloidea, and very rarely Porella pinnata
may be found in isolated stations. No other hepatics are regu-
larly found in this habitat and thus succession of hepatics does
not occur. A moss succession does obtain; but this is outside
of the scope of this paper.

Hepatics found on floodplains are often considered as invaders.
The most common hepatic on floodplains, or alluvial deposits
along streams is Riccia frostii. However, this is the regular habi-
tat of this species and should perhaps be best eliminated from a

132 The University Science Bulletin

list of invaders on such sites. Species which do more or less
regularly appear on alluvial deposits other than the above species
are: Riccia heijrichiana, R. sorocarpa, R. hirta, R. bifurca, R.
dictyospora, R. lamellosa, R. sullivantii, Notothijlas orbicularis,
Phaeoceros hevis, Plectocolea hyolina, Marchantia polymorpha
and land forms of Ricciocarpus natans, Riccia rhenana, R. fiuitans
and possibly R. canaliculota. These species do appear after the
deposits are left but they never last for more than a season, if that
long. Such habitats are useful in checking species to be found
in a water-shed but other than Riccia frostii they are of no
importance, ecologically, in this area. The statement of Schuster
( 1953 ) that Riccias are confined largely to alluvial soils, or areas
bordering alluvial soil does not apply in our area for most species
of the genus in this country. The same statement has been made
for Notothylas orbicularis by Schuster (I.e.), Sharp (1949), and
others but again it is not applicable to Kansas, since it has been
noted above the "mother lode" of this species is to be found in
the prairie. Except for Riccia frostii, the land forms of Riccio-
carpus natans, Riccia fltiitans, and R. rhenana, all the above species
invade but only temporarily so. The latter four can hardly be
considered as invaders since their usual habitat is on alluvial
sites.

The invasion of hepatics on tree bark is of regular occurrence
and involves, essentially, only the species of the genus Frullania.
These seem to persist as expanding colonies on tree bark and last
as such until crowded out by mosses. No hepatic succession
occurs in this area on bark. Other hepatics found on bark are
too rare to cite as examples of invasion.

Bryophytes are active in the revegetation af abandoned fields
in Kansas, particularly those fields with sandy soils. Mosses
are of primary importance but a few hepatics are to be noted.
The latter, however, are relatively unimportant and seem to play
a more important part in disturbed prairies and woodlands than
they do in true revegetation. Species which have been ob-
served on sites undergoing revegetation are: Riccia beyrichiaim,
R. dictyospora, R. sorocarpa, Anthoceros ptmctattis, Asterella
tenella, Fossombronia brasiliensis and Sphaerocarpus texanus.

Undisturbed prairie is inhabited by a comparative large number
of species. These are well scattered and inconspicuous. The first
stages of disturbance, however, results in many small isolated,
bare areas which are immediately invaded by several species

Taxonomy and Ecology of Kansas Hepaticae 133

of Riccia of which the more important are R. heyridnana, R.
hirta, R. dictyospora, R. campbelliana and R. sorocarpa. These
may form single scattered rosettes or more commonly form ir-
regular gregarious colonies which hold soil and help prevent
erosion and water loss. Often Sphaerocarptis texanus, Fossom-
bronia brasiliensis and Anthoceros punctatus form conspicuous
secondary invaders. Later small amounts of Asterella tenella and
Mannia fra grans may appear at about the time mosses become
evident. The mosses soon gain control, resulting in decline of
hepatics and eventual return to the original conditions of near
complete grass cover. If the conditions causing disturbance,
such as mowing, burning, or grazing remain, the area will continue
to exhibit a hepatic flora of many plants. In low moist prairie
sites the early invaders may often be Notothylas orbicularis and
Anthoceros punctatus. Such areas are soon dominated by mosses,
hepatics are excluded, and bare areas return to normal grass
cover.

Correlation With Other Vegetation

As might be expected the distribution of Hepaticae, in Kansas,
is often directly correlated with the occurrence of higher plants.
It has been found possible to drive through an area and select
stations, based on higher plant associations, which are the most
important in hepatic collecting. This fact is shown by a number
of tests in which areas were selected on the basis of higher plant
assemblages and a prospective list of hepatics for the area was
made before collecting was started. Even though such specific
areas had never been studied previously it was found that the
list of species suspected for the station was invariably 90 per
cent or more correct. After much field work had been accom-
plished it was found that the use of higher plant associations
as indicators of spots worth studying proved an efficient method
of completing distributional data in little studied counties.

As a result it has been possible to prepare lists of species which
are found in the various associations of higher plants in the State.

The oak-hickory association mentioned as occurring in the Chau-
tauqua area contains by far the largest number of hepatic species
in Kansas. This association is characteristically dominated by
Quercus stellata, Q. 7narilandica, Q. velutina, Q. prinoides and
Carya ovata. The soil is always sandy and sandstone outcrops
are frequent either as ledges or barely surfacing on slopes and
hilltops. In such forested areas the hepatic flora is dominated

134 The University Science Bulletin

by members of the Marchantiales and the genus Riccia in par-
ticular. The following lists indicated the degree of occurrence:
Species invariably found are:

Asterella tenella Riccia hirta

Frullania inflata Riccia lamellosa

Riccia beyrichiana Sphaerocarpus texanus
Riccia dictyospora

Species found, in addition to those above, if site includes dry

sandstone ledges or exposed banks are:

Cephaloziella rubella Reboulia hemisphaerica

Lophozia bicrenata

Species found, in addition to all above, if site includes moist
sandstone ledges and banks are:

Conocephalum conicum Plectocolea hyalina

Mannia fragrans Scapania nemorosa

Phaeoceros laevis

In the southern third of the Chautauqua Hills the association

would nearly always include, in addition to all above, the following:

Fossombronia brasiliensis Riccia campbelliana

Oxtjmitra paleacea Riccia trichocarpa

Species found only in this association but of rare occurrence

are:

Jungermannia lanceolata Cephalozia catemilata

Plectocolea crenuliformis Cephalozia connivens

Plectocolea fossombronioides Cephalozia media

Chiloscyphus pallescens Calypogeia meylani

Chiloscyphiis pallescens var. fragilis Calypogeia muelleriana

Scapania irrigua Calypogeia trichomanis

Cephaloziella byssacea Plagiochasma rupestre

Bluestem prairie of the Chautauqua Hills is usually sandy and is
dominated by Andropogon gerardi. Swales may have Tripsacum
doctijloides or Sorghastrum nutans as dominant vegetation. Most
of these prairies are either mowed once a year for hay or are grazed.
If grazing is well controlled the two uses result in prairie of about
tlie same characteristics. In such prairies, hepatic species usually
expected occur.

Species invariably present on flats or hilltops are:

Anthoceros punctatus Riccia hirta

Asterella tenella Riccia sorocarpa

Riccia beyrichiana Sphaerocarpus texanus
Riccia dictyospora

Taxonomy and Ecology of Kansas Hepaticae 135

Species found in swales in addition to all above are:
Notothylas orbicularis Riccia lamellosa

In southern third of area the following are expected:
Oxijmitra paleaca Riccia trichocarpa

A number of other species have been found on these prairies but
are rare and not of usual occurrence.

In the extreme southeastern corner of Kansas is found a bit of
the Ozark Plateau, here referred to as the Ozark area. As men-
tioned before this area consists of rocky, oak-hickory forests with
islands of rocky bluestem prairie. It is an area that ranks only
second to the Chautauqua Hills in abundance of hepatics but it
breaks down into fewer groups when correlation of hepatics with
other vegetation is made. The groups are:

Species found in upland oak-hickory forests on bark, soil, or dry
rock ledges:

Porella platyphylloidea Frullania riparia

Friillania brittoniae Riccia beyrichiana

Frullania eboracensis Riccia hirta

Frullania inflata

Species found on moist, north-facing, rocky, wooded hillsides
occurring on bark, soil, and moist rock ledges:

Plagiochila asplenioides Frullania brittoniae

Geocalyx graveolens Frullania eboracensis

Lophocolea minor Frullania inflata

Lophocolea heterophylla Frullania kunzei

Diplophyllum apictilatum Frullania riparia

Scapania nemorosa Pellia epiphylla

Porella platyphylloidea Asterella tenella
Cololejeunea biddlecomiae

Species found in the prairie islands are:

Anthoceros punctatus Riccia dictyospora

Sphaerocarpus michelii Riccia hirta

Sphaerocarpus texanus Riccia lamellosa

Riccia beyrichiana Riccia sorocarpa

The Osage area in general has comparatively few hepatics both
in number of species and quantity of any species. The only excep-
tion is the genus Frullania which is to be found in any oak-hickory
area. The dominant trees of this association are consistently
Quercus macrocarpa, Quercus muehlenbergii, Qtiercus veliitina,
Carija cordiformis and Canja ovata. The oaks are by far the most
abundant. On the bark of these trees Frullania inflata can always

136 The University Science Bulletin

be found and occasionally F. eboracensis and F. riparia will be
observed. Moist, mossy, north-facing cliffs or banks in this associa-
tion often have several species of hepatics, among which Lopho-
colea heterophyUa and Scapania nemorosa are the most common.
Often they are the only species present. If the moss Leucohnjum
glaucum is present on these banks a careful search of the area may
reveal certain other forms. The following species, in addition to
the two named above, may be found:

Jamesoniella autumnalis Scapania undulata

Geocalyx graveolens Cephalozia bicuspidata

LopJiocolea hidentata

In a few places in the northern part of the Osage area there are
places where the habitat is intermediate with that of the Chautauqua
Hills. Such places have a larger hepatic flora but in general the char-
acteristics of such sites are much more similar to types of the Chau-
tauqua Hills and are thus considered with the latter area.

The prairie of the Osage area, if sandy, characteristically has a
few species of which Riccia beyrichiana, R. dictyospora, R. hirta,
R. lamellosa and R. sorocarpa are the only ones usually found. The
rate of occurrence varies greatly.

The Flint Hills has such a small hepatic flora that no associations
can be made. The only consistent correlation is the occurrence of
Frullania inflata in all places where Quercus macrocarpa is found.
This oak occurs along a few streams and bluffs.

The Dakota is primarily a prairie association on sandy soil, with
numerous sandstone ledges, boulders, or flat surfacing sandstone
exposures. Most of the hepatics in the area are found near the edge
of the exposed sandstones. However, prairies dominated by
Andropogon gererdi and Andropogon scoparius have an abundance
of liverworts in some places. Species usually found are Riccia bey-
richiana, R. hirta, R. dictyospora, Asterella tenella, Mannia fragrans,
Sphaerocarpiis texanus and more rarely Anthoceros ptmctatiis and
Notothylas orbicularis. On the bark of trees along streams or ra-
vines will be found the ever present Frullania inflata, if Quercus
macrocarpa is present.

All other areas of Kansas have so few hepatics that no correlation
is possible. Those found in such areas usually are associated with
springs or alluvial deposits on streams. The latter type stations are
very unstable and thus allow for little in the way of comparisons.

Taxonomy and Ecology of Kansas Hepaticae 137

Phytogeography

The hepatic flora of Kansas is a flora composed mainly of ubiqui-
tous, widely distributed species which occur to the east, north, west
and south of the state. This is, as might be expected, due to the
location of the region in nearly the exact center of the United States.
The environment is such that only those species with a wide degree
of habitat toleration find it possible to survive.

The hepatic flora of the State is decidedly dominated by members
of the Marchantiales whose species, though not more numerous, are
much more abundant. They form the characteristic component of
the State's hepatic vegetation and in many sites an observer finds
only this group represented.

A few species seem to indicate that a relationship to the southern,
southeastern and southwestern hepatic floras is evident.

These species are:

Frullania kunzei Riccia campbelliana

Oxymitra paleacea Riccia dicUjospora

Plagiochasma rupestre Riccia trichorcarpa

Plectocolea fossombronioides Sphaerocarpus michelii

Frullania kunzei is listed by Frye and Clark (1937-1947) from
North Carolina, Arkansas, Louisiana, Mississippi, Alabama, Florida,
Georgia, South Carolina, West Indies and South America. In
Kansas it is found in the Ozark Plateau area in extreme southeastern
Kansas. It thus is not far removed from Arkansas.

Oxymitra paleacea has been reported in Texas and Oklahoma
and the writer has collected it in those two states and in six counties
in Kansas. This species, then, is clearly southern in its affinities rang-
ing into Mexico and South America. It also occurs in North Africa
and Europe.

Plagiochasma rupestre is reported by Frye and Clark (1937-1947)
as occurring in North America in New Mexico, Arizona and Mexico.
However, there are specimens in the herbarium at the University
of Kansas from Texas, Oklahoma and extreme western central Mis-
souri. Our Kansas collection, therefore, completes the gap from
Missouri to Mexico.

Plectocolea fossombronioides has a range according to Frye and
Clark (1937-1947) of Connecticut, Ohio, Illinois, North Carolina,
West Virginia and New Jersey. Thus Kansas seems far out of range
for this species; however, Wittlake (personal communication) re-
ports that he has specimens (verified by Evans) from Arkansas.

138 The University Science Bulletin

Kansas material ( also verified by Evans ) is, then, not far out of the
known range of the species but nevertheless is an important exten-
sion of range.

Riccia camphelliana was reported by Frye and Clark (1937-1947)
in California; by Jacobs ( 1951 ) in Georgia and Arkansas; by Mc-
Allister, Hoglund and Whitehouse (1932) in Texas; and it has been
collected by the writer in Oklahoma and Kansas.

Riccia trichocarpa has been reported by Frye and Clark (1937-
1947) from Cahfornia, Oregon, Texas and Oklahoma in the United
States. It occurs in several counties in Kansas and Schuster ( 1953 )
reports a single station in Minnesota. The species ranges into
lower California and is definitely southern in its affinities.

Sphaerocarpiis michelii is a species which has probably been over-
looked or misinterpreted. Frye and Clark (1937-1947) report the
species from Texas and Thompson ( 1948 ) reported it from Kansas.
The writer has collected it in Arkansas, Oklahoma and several places
in Kansas. Indications to date, therefore, are that the species is
southern in its range.

Other species show some relationship to southern and south-
western floras. The two stations of Porella pinnata on dry limestone
ledges, mentioned previously, is an example of the habitat of this
species in the South. Sphaerocarpiis texanus seems to be mainly
southern in its range at least in abundance.

About the only indication of a relationship to northern hepatic
floras is the occurrence of Scapania irrigua from a glaciated region
in northeastern Kansas.

A few species might reveal relationship with other areas, particu-
larly the south, if taxonomic and distributional details were known.
In general much more field work needs to be done before many
phytogeographic interpretations in the Hepaticae can be made.

BIBLIOGRAPHY

Austin, Coe F.

1869. Characters of some New Hepaticae (mostly North American),
together with notes on a few imperfectly described Species. Proc,
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1929. Hepatics and Sites: A Short Study in the Ecology of Hepatics.
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1941. An Ecological Survey of the Fort Leavenworth Mihtary Reservation.
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Campbell, D. H.

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Cabruth, J. H.

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CocKRUM, E. Lendell.

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Cragin, F. W.

1884. Miscellaneous Notes. Bull. Washburn Lab. of Nat. Hist., vol. 1,
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DoELL, Jacob H.

1938. The Flora of the Sand Hills of Central Kansas. Univ. Kansas Sci.
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Dltrand, E. J.

1908. The development of the sexual organs and sporangia of Marchantia
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Emig, W. H.

1930. A Society of Bryophytes. The Bryologist, vol. 33, no. 1, 1930,
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Fearing, Olin S.

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Frye, T. C. and L. Clark

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Graff, Paul W.

1935. Plant invasion following fires. Torreya, vol. 35, 1935, pp. 137-141.

1936. Invasion of Marchantia polymorpha following forest fires. Bull.
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Griggs, R. F.

1933. The Colonization of the Katmai ash, a new and inorganic "soil".
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Haynes, Caroline C.

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Jewett, J. M.

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THE UNIVERSITY OF KANSAS

SCIENCE BULLETIN

Vol. XXXVII, Pt. I] October 15, 1955 [No. 4

A Pollen Profile Study of the Muscotah Marsh ^

W. H. HORR

The University of Kansas

Abstract: This paper treats of a pollen profile of a raised artesian marsh
in Atchison County Kansas, samples being taken at 2-foot intervals.

The marsh can be divided into five distinct strata. Abies pollen was
found at all levels from 36 feet to 12 feet, and is the only tree to appear
in an unbroken succession of levels. Herbaceous plants replaced the woody
plants in the marsh and immediate environs between the 22 and 20-feet
levels. The vegetative successions were not clear cut until the herbaceous
plants become dominant.

The successions were Abies to Abies-Picea-Larix, to all conifers, to
Cyperaceae-Chenopodiaceae-Gramineae, to Cyperaceae-Gramineae, indicating
a succession of weather conditions from wet-cool, to moist-warm to wet-cold
to warm-dry.

The raised artesian marsh from which the material for this
study was obtained is located at the east edge of the flood plain
of The Delaware River, VA miles south of Muscotah in the NE/l
sec. 16, T. 6 S., R. 17 E., Atchison County, Kansas (Horr &
McGregor 1948)-. Artesian water shows up in several places
both north and south of this locality. At present this marsh covers
about seven acres but there are definite indications that at earlier
dates it covered a much greater area. It seems probable that
the source of this water is a buried glacial till, as such a till
has been tapped 6 miles east and /2 mile south of this locaHty
at a level approximately 100 feet above that of this marsh. An
artesian well at the edge of this marsh, which has been flowing
for over 30 years, had enough pressure to raise the water 20
feet above the level of the ground when it was drilled and
cased. It is evident that the pressure has been reduced con-
siderably in the last few years, as is indicated by a reduction

1. This work was supported by a Kansas University Research Grant. Roland Ander-
son served as research assistant.

2. Horr, W. H. & R. L. McGregor, 1948. A Raised Marsh Near Muscotah. Kansas:
Trans. Kan. Acad. Sci., Vol. 51. No. 2.

(143)

144 The University Science Biilletin

in the flow of water from both the well and the surface of the
marsh, and also by the fact that from the fall of 1952 to the
fall of 1953 the material in the marsh settled and became so
compact that it was very difficult to take samples from the
lower levels.

The material in this marsh has been laid down on a flood
plain of stratified clay, sand and coarse gravel. The water supply
has apparently been continuous since the start of this marsh
but there are indications that the flow of water has varied
considerably during the time of the development of this marsh.
The surface vegetation is made up of typical marsh plants with
the sedges being strongly dominant. (Horr & McGregor, op.
cit.) During the drought of 1952-53 grazing cattle kept the sedges
cropped off and the grasses and dicotyledonous plants increased
greatly. R. L. McGregor, The University of Kansas, isolated
pollen from this marsh in 1950.

METHODS

Samples were taken with a Davis punch-type sampler at inter-
vals of 2 feet. Certain strata, composed of sand, clay and vegeta-
tion, were too hard to be handled with this sampler. A /4-inch rib
was attached to one side of the cylinder with which to expand the
hole as the sampler was forced downward. This was effective until
the fall of 1953 when the settling of the marsh, due to a reduction
of water pressure, made it necessary to resort to levers to get
through the harder strata.

The samples were examined for pollen, seeds, leaves, mollusk
shells and other material which might help to give a more complete
record of the fauna and flora of this area back through the ages as
it was being recorded by nature. Numerous water pockets made
it difficult to get a good series of samples at consecutive levels.
The nature of the material obtained made it necessary to use a
rather severe sodium hydroxide-acetic anhydride treatment to free
the pollen for study. Neither alcohol nor detergents were effective
on this material.

RESULTS

The profile in this marsh can be divided into five rather distinct
strata, based on the physical make-up of the deposit:

1. From 1 to 10 feet the plant material, which is only partially decom-
posed, is very coarse.

2. From 10 to 22 feet the vegetation is much more decomposed and re-
duced to rather small fragments.

Study of the Muscotah Marsh 145

3. From 22 to 26 feet is a very dense stratum composed mostly of clay
and a considerable amount of sand with about 8% of the total dry
weight being organic. (Analysis made by The Kansas Geological Sur-
vey Service)

4. From 26 to 32 feet is plant material of extremely small sizes, mixed
with some clay and sand.

5. From 32 to 36 feet is a mixture of clay and plant matter much like
that found in 3 but with a much greater percentage of sand.

Marl and mollusk shells are common in the first 26 feet with their
greatest concentrations where there is clay present.

In the first stratum various genera of Cyperaceae furnished from
86% to 90% of the total pollen identified at the various levels. The
Gramineae furnished about 7% and about 5% were unidentified ex-
cept to determine that they were mostly Compositae. Chenopo-
diaceae contributed 2% at 6 feet and Poptilits 4% at 8 feet.

In the second stratum Cyperaceae provided most of the polleri
down to 18 feet, then dropped to 30% at 20 feet but did not appear
at any lower level. Gramineae provided 9% of the pollen at 12 feet
and dropped to 3% at 18 feet but was not found at any lower level.
Abies furnished most of the tree pollen, starting with 3% at 12 feet
and increasing to 70% at 22 feet. All of the other woody plants pro-
vided small amounts of pollen at various levels with all of them,
except Qiiercus, showing at 22 feet.

In the third stratum Abies again provided most of the total pollen.
All of the other woody plants except Picea and Quercus provided
small quantities of pollen. Chenopodiaceae provided 11% at 26
feet and there was about the usual number of unknowns.

In the fourth stratum at 28 feet Abies pollen dropped sharply to
12% and increased to 21% at 30 feet. Populus showed the most
pollen with 36% at 28 feet and 31% at 30 feet. All of the other woody
plants contributed small amounts of pollen at 30 feet, and all except
Piniis, Tsuga and Picea, appeared at 28 feet. Chenopodiaceae
contributed 10% of the pollen at 28 feet with unknowns averaging
about 10%.

In the fifth stratum the profile was again dominated by Abies
pollen, with Picea and Larix the only genera to appear at all levels,
and Quercus the only woody plant which did not appear. Coryhis
and Tsuga appeared only at 36 feet while Pimis provided 5% of the
pollen at 34 feet and 2% at 36 feet. The only herbaceous pollen to
appear in this stratum was Chenopodiaceae which contributed 5%
at 32 feet, and the unknowns which averaged about 10%.

146 The University Science Bulletin

DISCUSSIONS

It seems obvious that these strata are the result of factors other
than the vegetation, and that the chmate was the most important
factor. Strata 1 and nearly all of 2 are dominated by the remains
of herbaceous plants but there is a rather sharp dividing line at
about 12 feet. Above this level there has been enough water present
to prevent the decay of the vegetation but below this level the
vegetation has been reduced by decay to a brownish black mass
of very small plant fragments. This indicates a more favorable
oxygen supply such as would be the case if the vegetation was not
submerged in water. The sharpness of the line separating these
two strata rules out the possibility of its being due to the difference
between the ages of these strata.

The silt which makes up 92% of the third stratum could have been
brought in by flood waters from the Delaware River or by surface
erosion of the higher ground to the south and east of this marsh.
The absence of any appreciable amount of silt above this stratum
might be explained by the upward growth of the marsh being rapid
enough to keep it above the source of the silt. However, in view
of the fact that the next lower stratum has only a small amount
of silt, this does not seem like a very valid explanation.

As stated above, the fourth stratum contains very little silt and
the plant material has been reduced to very small particles. At
certain places conifer wood and leaves were encountered in large
quantities. This wood is much better preserved than the other
plant material found in this stratum.

The fifth stratum is much like the third, except that the decay of
the plant material is more nearly complete and there is considerable
wood present. It becomes quite evident that the absence of large
quantities of silt in the fourth stratum is the only thing which
separates the lower 14 feet into 3 strata.

The interpretation of the pollen profile in terms of climatic condi-
tions is somewhat confused by the fact that we are dealing with relic
vegetations. Vegetative changes in the surrounding area may have
preceded the changes in the marsh by a great many years. This is
well illustrated by the mollusk shells recovered at various depths.
Dr. A. A. Leonard of The University of Kansas examined the shells
and stated that some were pleistocene forms. He also stated that
"snails now living there have not been found in the midwest in any
other place south of the northern part of Iowa."

The pollen appearing most consistently in this study is Abies

Study of the Muscotah Marsh 147

which is now found at all levels up to 12 feet. This pollen indicates
a rather definite series of cHmatic changes, especially when corre-
lated with other pollens studied. With Abies furnishing 53% of the
pollen at the beginning of the marsh it is apparent that the climate
was cool and moist. The presence of the other conifers and the
relative absence of deciduous trees at this level all indicate at least
a moist climate and one that was much cooler than at present.
The increase in Abies pollen to 67% at 34 feet followed by the de-
crease to 60% at 32 feet might not be too significant were it not
for the drop to 21% at 30 feet. More significant than this is the
abrupt increase in all of the otlier tree pollens at this level. Es-
pecially significant is the sudden increase in Corylus, Populus and
Qiierctis pollen at the 30 and 28-foot levels. These are trees of
warmer and drier climates. The further drop of Abies pollen to 12%
at 28 feet, the disappearance of Finus, Picea and Tsiiga coupled
with the increase of Corylus and Populus pollens indicate that there
was a considerable increase in temperature, and probably a decrease
in moisture. However, the decrease in Quercus pollen does not
follow the expected pattern. The sudden increase in Abies pollen
to 63% at 26 feet coupled with the reappearance of Pinus, Tsuga
and Larix and the sudden drop in Corylus and Populus indicate
that the climate had changed back to a more cool and moist period.
This trend continued at the 24-foot level where the Abies pollen
reached 85% and again reversed itself at the 22-foot level where
all of the woody plants, except Quercus, were present. The most
striking change came at 20 feet where the Abies pollen dropped
to 20% and all other woody plants, except Larix, disappeared and
the pollen picture became dominated by Cyperaceae and Cheno-
podiaceae, each of which furnished 30% of the total pollen. This
definitely marks the transition from a forest to a grassland formation
brought about by a continuing decrease in moisture and an increase
in temperature. The continuance of Abies up to the 12-foot level
probably can be explained by the continued presence of cold water
coming up from below and maintaining these trees as a relic vegeta-
tion long after Abies had disappeared from the surrounding land
areas. The influx of large amounts of Chenopodiaceae pollen,
along with that of Cyperaceae at 20 feet, indicates competition
between the herbaceous plants for dominance of this area as the
trees disappeared and herbaceous plants took over. The Cyperaceae
won the struggle because this is a much more favorable habitat
for them. There are no species of Chenopodiaceae growing on
the raised portion of this marsh at the present time.

148 The University Science Bulletin

After the grasses became established at 18 feet there seems to
have been httle change in their prevalence. Today there are more
grasses and Compositae than were present in 1951, due to the
destruction of the tall sedges and the decrease in surface water.

The relative absence of Quercus in the upper 20 feet poses an
interesting question. One would assume that Quercus pollen would
have been carried in from the surrounding area and, unless there
was a very drastic change in the climate, there should have been
a considerable number of these trees present. The distribution
of Finus pollen is quite different from that which is usually found;
in fact, it is drastically different from that which Roland Anderson
(University of Kansas) found in a similar situation 2/2 miles to the
south of this locahty. There, Tinus furnished 100% of the pollen
at the lowest level and continued upward in varying amounts for
15 feet where it contributed 39%. No determinations have yet been
made above that level. It is altogether possible that the erratic
appearance of Finus pollen in small quantities can be attributed to
the fact that there might not have been any pine trees growing
at this marsh and that the Finus pollen which is present was wind
carried.

Because of the many papers written on pollen analysis of bogs
and marshes, the author felt that a review of the literature would
be superfluous in this paper. However, the author felt that this
work was worth doing, because no other marsh in this region has
yet yielded fossil pollen by which the post-Pleistocene flora and
climate could be studied. Further attempts will be made to recover
pollen from an artesian marsh in southwestern Kansas where all
preliminary tests have been negative.

SUMMARY

1. This a study of pollen profile of a raised artesian marsh in
Atchison County, Kansas, made from samples taken at 2-foot inter-
vals.

2. The material in this marsh can be divided into five distinct
strata.

3. Ahies pollen is found at all levels from 36 feet to 12 feet and
is the only tree to appear in an unbroken succession of levels.

4. Herbaceous plants replaced the woody plants in the marsh
and the immediate environs between the 22 and 20-foot levels.

5. The vegetative successions were not clear cut until the herba-
ceous plants became dominant at the 20-foot level.

Study of the Muscotah Marsh

149

6. The successions were: Abies, to Ahies-Picea-Larix, to all
conifers, to Cyperaceae-Chenopodiaceae-Gramineae, to Cyperaceae-
Gramineae.

7. The indicated weather conditions are: wet-cool, to moist-
warm, to wet-cold, to warm-dry.

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Silt: Only Pinus pollen has been found below 36 feet.

Table 1. — Pollen profile of the Muscotah Marsh taken at 2-foot intervals.
These percentages are based on 100 pollen grains from each level.

THE UNIVERSITY OF KANSAS

SCIENCE BULLETIN

Vol. XXXVII, Pt. I] October 15, 1955 [No. 5

The Anatomy and Modes of Preservation of the Genus
Cardiocarpus spinatiis Graham

By Elmer A. Roth

Abstract: Four modes of preservation of the fossil seed genus, Cardiocarpus
spinatus Graham, are described as follows; petrifactions, nucellar coal balls,
cuticular remains, and impressions. Details of morphology and anatomy as
determined from the coal ball seeds and nucellar coal balls are illustrated and
discussed. New infomiation is presented about the sclerotesta, endotesta, and
nucellus tissue. An elaboration is made of the structure of the sarcotesta and
gametophyte. A new term "nucellar coal balls" is suggested for certain speci-
mens preserved as free nucellar areas of the seeds. A review of theories relative
to the lack of embryos in Paleozoic seeds is given. A brief discussion is made
with respect to the preservation of nucellar coal balls in coal.

INTRODUCTION

Brongniart (1828) described the genus Cardiocarpon as "fruits
compressed, lenticular, cordiform or reniform, terminated by a
somewhat acute point." Since Brongniart's investigations, many
others have contributed to the study, but his work has become the
foundation for the study of fossil seeds as we know them today.

The purpose of this paper will, in essence, be twofold. First, to
present a description of some new material which Dr. S. H. Mamay
of the U. S. Geological Survey has called "miniature coal balls." I
would like to enlarge on Dr. Mamay 's proposed descriptive nomen-
clature and submit the term "nucellar coal balls" for these loose
"casts" or "nutlet" appearing specimens ( figure 4 ) , because the pre-
served organic material appears as a miniature coal ball, and the
preservation is of the nucellar region of the seed.

Secondly, to add to our current state of knowledge of the seed
Cardiocarpus spinatus Graham based on additional well-preserved
material found in coal balls from the coal fields of West Mineral,
Kansas, and the Clay Products Company mine one mile southeast of
What Cheer, Iowa.

(151)

152 The University Science Bulletin

HISTORY

Andrews and Felix ( 1952 ) , in their excellent work on the gameto-
phyte of Cardiocarpus spinatus Graham, present a brief history of
the genus, however, some value may be gained by future workers
if the history of the genus is enlarged upon in this paper.

Among diose who investigated the genus Cardiocarpon (now re-
ferred to as Cardiocarpus) prior to 1900 are included Brongniart
(1828, 1874, 1881) who estabhshed the genus and described a
number of species, and Dawson (1865, 1896) who, while working
with C. cornutum, arrived at the conclusion that the "fruits" de-
scribed by Brongniart were in reality seeds, because of

"2-cotyledons, a strong membranous or woody tegemen, surrounded by a
fleshy outer coat with a notch at the apex which represented the foramen or
micropyle of the ovule."

Hooker and Binney (1855), Lesquereux (1880), White (1899),
and Williamson (1877) were concerned with the history, distribu-
tion, and descriptions of various species of Cardiocarpon, as well as
other carboniferous seeds and seed-bearing plants. Lindley and
Hutton (1833, 1837) investigated the paleozoic seeds of Great
Britain, and described several species of Cardiocarpon found in that
region. Newberry ( 1853, 1873 ) worked a great deal with compres-
sions and impressions and has presented a general discussion and
description of the genus. He also refers to and illustrates an impres-
sion of a cordaites stem bearing several, what he calls, "cardiocarpa."
These "cardiocarpa" appear to be impressions of the nucellar area of
a Cardiocarpus-Xy-^e seed. Solms-Laubach (1891) presents a dis-
cussion of the affinities of paleozoic seeds and theories relative to
their lack of embryos.

Many of the specimens found and described prior to 1900 were
none too well preserved. In most instances specimens were found
as compressions and/or, impressions with little or no cellular tissue
preserved, making it quite diflRcult to present adequate descriptions.
The lack of perfect specimens, and the relatively inefficient tech-
niques did not, however, prevent these early investigators from
producing work which has become the sound foundation for the
investigations being carried on today.

Since 1900, many investigators have delved into the field of fossil
seeds, some adding little to the knowledge previously presented,
others making a definite contribution to our meager accumulation
of information. Upon looking over the works of recent investi-
gators we find that Andrews and Felix (1952) have produced an

Study of Cardiocarpus spinatus 153

excellent description of a well-preserved gametophyte in the species
Cardiocarpus spinatus Graham; Arber (1906, 1914) has worked
on a revision of seed impressions in British Coal Measures; Arnold
(1938, 1947) presents an excellent discussion on paleozoic seeds;
Chamberlain (1935), and Coulter and Chamberlain (1917a, 1917b)
present a phylogeny and comparison of the Cordaitales with the
cycads, Ginkgo, and other gymnosperms and also offer (1917b) a
theory regarding the lack of embryos in paleozoic seeds. Graham
(1935) describes the species Cardiocarpus spinatus; Oliver (1902,
1903) worked with the relationships of Cardiocarpus to seeds of
the cycads and Ginkgo; Reed (1946) presents an excellent discus-
sion as well as some illustrations of some seeds found at the coal
mines near What Cheer, Iowa, and considers them to be com-
parable to Cardiocarpus spinatus Graham, and finally Scott ( 1909a,
1909b) and Seward (1917) list and describe various species in the
genus and present some morphological descriptions.

Here again, as in early day investigations, the specimens of seeds,
in general, were too poorly preserved to present adequate descrip-
tions of the entire seed. The gametophyte, and the sclerotesta are
the two zones of the seed that have received the most attention,
and of these only the gametophyte, Andrews and Felix (1952),
has been adequately discussed. Most investigators, Andrews and
Felix (1952), Graham (1935), Reed (1946), admit that the lack
of proper preservation has made complete descriptions of the seeds
quite difficult. Even though poorly preserved seeds are the rule
rather than the exception, the use of the parlodion peel method,
the gelatin method, and improved microscopic instruments have
resulted in more complete descriptions of all types of fossils than
has heretofore been done.

TYPES OF PRESERVATION

The seeds, generally, are preserved in coal balls along with other
organic material (fig. 8). It is through this type of preservation
that cell and tissue structure is best preserved.

Nucellar, or miniature, coal balls occur directly in the coal. Many
times these may be found lying loose in weathered areas of the coal
fields, or where the "washings" from the coal are dumped. As shown
in figures 4, 5, they are heart-shaped, platyspermic, resembling the
description presented by Brongniart ( 1828 ) . In diameter they vary
from 8mm x 8mm to 15mm x 12mm with an average of 10.5mm x
8.5mm (figs. 3, 6, 7).

154 The University Science Bulletin

Another form of preservation may be found as impressions in
shale. It is from this form of preservation that most of the early
day investigations were done.

One other form to be considered is that of cuticle remains. This
form usually does not show the true appearance of the seed, but has
been found in close association with the cuticle of Cordaianthiis,
being quite abundant in the shale layer at Lone Star Lake, Kansas.

DEGREE OF PRESERVATION

The degree of preservation of organic matter in the nucellar coal
balls varies a great deal. Of the 114 specimens collected at West
Mineral, Kansas, 30 showed well-preserved gametophytic tissue
(fig. 13). Of these, 14 specimens show the presence of structures
resembling archegonia at the micropylar end. Forty-four were
found to have been infiltrated with a varying amount of pyrite.
Some were so heavily impregnated that the study of the tissue was
an impossibility. In others, the penetration of pyrite was very slight,
and, though sometimes making microscopic study somewhat diffi-
cult, did not completely obscure the tissue from view. Of the 44
nucellar coal balls found impregnated with pyrite, 3 were found
to have the sclerotesta preserved as the outermost tissue. The re-
mainder of the 114 specimens collected were found to be true casts
with the organic material being completely replaced. Here, as in
many specimens found embedded in coal balls, remnants of the
megaspore membrane and of the nucellus may be distinguished as
dark brown lines around the megaspore area.

The outermost part of these nucellar coal balls is, generally, the
megaspore membrane, however, some specimens may have as the
outer covering a portion of the nucellus tissue, endotesta, or sclero-
testa. Evidence on hand indicates that the "nucellar coal balls"
consist of the nucellus area of the type found in Cardiocarpiis. Any
one of the 114 specimens will fit into any vacated area of a nucellus
in any of the coal balls in our collection ( fig. 6 ) . Since Cardiocarpiis
spinatus Graham is the sole species so far discovered in Kansas
coal balls, there is little doubt that they represent the nucellus area
of this species.

When preservation is in the form of impressions one seldom, if
ever, finds organic tissue preserved. As a rule it seems apparent
that the compression of the outer sarcotesta has given rise to the
erroneous concept of a winged seed.

Study of Cardiocarpus spinatus 155

DESCRIPTION

In considering, specifically, Cardiocarpus spinatus Graham, I find
that those investigating the species since Graham's original descrip-
tion, Andrews and Felix (1952) and Reed (1946), have added much
to our meager knowledge. However, it is felt, and admitted by
authors concerned, that most of the descriptions were given from
relatively poorly preserved material. I have been fortunate in
having available specimens with almost perfect preservation of the
various tissues. It is because of this excellent, well preserved ma-
terial that I am presenting an additional, more complete, description
of the species than has heretofore been published.

Since the seeds are bilaterally symmetrical almost all sections ex-
posed in coal ball cuttings are usually oblique. This makes it diffi-
cult to establish definitely the true orientation of the seed, and,
when the gametophytic tissue is found, it is difficult to determine
the numbers, true position, and structure of archegonia (when
present), and finally to establish definitely true cellular structure of
the tissues exposed. I have been fortunate in that I have been
able to obtain almost perfectly preserved specimens removed intact
from coal balls, and therefore able to determine the exact plane of
the sections made.

Graham ( 1935 ) notes that the sarcotesta of his specimen is almost
completely destroyed, so he does not present a description of this
tissue. Reed (1946) presents a general description, and illustrates,
by camera lucida drawings, some of the tissues involved. Andrews
and Felix ( 1952 ) present a somewhat clearer description, but note
that in most specimens only the sclerotesta is preserved with the
sarcotesta being seldom preserved intact.

In light of the statements concerning the lack, in most cases, of
the preservation of the sarcotesta, it is of interest to note that in a
number of seeds on file in the paleobotanical collection at the Uni-
versity of Kansas the sarcotesta and the epidermal layer are well
preserved (figs. 2, 9, and 10). In general, these seeds show well-
preserved tissues at all layers in the seed.

Epidermis: The epidermis is one cell thick. The cells are thin-
walled, rectangular, averaging 70[x x 49[;t in diameter. No stomata
have been observed although occasionally there are found, at irregu-
lar intervals, structures in the epidermis that are suggestive of
sunken stoma. Surrounding the cavities in these areas are small,
parenchymatous, isodiametric cells, distinct from the normal cells
found in the outer sarcotesta. Little can be said about the true

156 The University Science Bulletin

nature of these areas, but future specimens should be carefully
scrutinized for further infonnation and enlightenment as to their
anatomical nature, (figs. 9, 10, and 11).

Outer sarcotesta: This tissue is generally 15 to 20 cells thick.
There may be a greater or lesser amount in some specimens, de-
pendent upon the development of the spines of the sclerotesta.
Andrews and Felix ( 1952 ) state the cells to be about 165[a in diam-
eter. The cells of our specimens average 168[x which is a favorable
comparison. The cells are more or less isodiametric, parenchyma-
tous, and closely packed (fig. 11). The tissue, as preserved in the
coal balls, has a characteristic amber color with occasional cells
being filled with a resinouslike substance having a somewhat
darker brown color than the cell walls ( fig. 8 ) .

Inner sarcotesta: The cells of this tissue differ from those of the
outer sarcotesta only in that they are about one-third the size of
the outer sarcotesta. The cell walls are amber colored, and the
lumen area of each cell is generally empty. The tissue, for the most
part, varies from 40 to 50 cells thick, again dependent upon the de-
velopment of the sclerotesta (fig. 11). Andrews and Felix (1952)
state these cells to average 55[;, in diameter. The cells of our speci-
mens average 42[jl x 42pi, in diameter which is a favorable comparison.

Sclerotesta: Graham (1935) describes the sclerotesta as being
made up of two layers; an outer layer of thick-walled more or less
isodiametric cells. He states this layer to be of variable thickness
with spines at the outer surface. The second layer he designates as
the inner layer, and this layer is made up of moderately thick-
walled cells, elongated in the direction of the axis of the seed. Reed
( 1946 ) notes that the sclerotesta is exceedingly thick-walled, roughly
isodiametric cells, and has a formation of spines. She does not
mention the sclerotesta as being made up of two layers. Andrews
and Felix ( 1952 ) note that the sclerotesta is the conspicuous portion
of the integument, and that it is characterized by its spiny out-
growths.

A number of specimens in our collection are well enough pre-
served to show some hitherto undescribed features of this tissue.
Due to the fact that most seeds, when exposed by cutting a coal
ball, are found to be cut at right angles to the broad plane (text,
fig. 2), the true significance and structure of this tissue has been
difficult to ascertain. It was indeed fortunate, and an extreme piece
of luck that Dr. Baxter, while working with a coal ball from the
coal fields of West Mineral, Kansas, exposed an intact seed in such

Study of Cardiocarpus spinatus

157

a manner that it was possible to orient properly the seed with ref-
erence to its broad plane, and to begin making peels, at the very
outset, on the broad plane.

A total of 44 peels were made, making it possible to determine
the true location of the spiny projections of the sclerotesta. This
was further corroborated by making 3-dimensional peels of other
seeds in the collection.

It is quite evident in virtually every specimen that, as Graham
indicated, the sclerotesta is made up of two distinct tissues — an
outer and an inner. It is also quite evident that the inner tissue is,
in turn, made up of two distinct zones.

ISC

Cardiocarpus spinatus Graham

Fig. 1 — Diagrammatic longitudinal view of the broad plane.

Fig. 2 — Diagrammatic longitudinal view of the narrow plane. E — Epider-
mis, OS — outer sarcotesta, IS — inner sarcotesta, OSC — outer sclerotesta, ISC —
inner sclerotesta, EN — endotesta, N — nucellus, M — megaspore, V — vascular
bundle, SP — spine of outer sclerotesta (note chamber).

Outer sclerotesta: It was found by the author that when observ-
ing this region of the testa, one must use caution in interpreting the
general structure and geometry of the cells. Both Graham (1935),
and Reed (1946) indicate that the cells in this tissue are roughly
isodiametric, however, I have found the cells to be vertically elon-
gated in the direction of the vertical axis of the seed, being seven
to eight times longer than broad. The cells are extremely closely
packed, sinuous, and almost hyphaelike in appearance. The cell
walls are very thick, and the lumen area small. In a vertical cut
at right angles to the broad plane, the cells in the spines tend to

158 The University Science Bulletin

become horizontally elongated, while a transverse cut of the seed
shows the cells as isodiametric. The fact that a high percentage
of seeds exposed in coal balls show a transverse, or oblique-trans-
verse plane probably explains the usual concept of "isodiametric"
cells.

The development of the spines in Cardiocarpiis spinatus Graham
is variable with the maturity of the seed. These projections may
be slight in a relatively immature seed, projecting only into the
inner sarcotesta. However, in the more mature seeds, the spines
have been observed to extend to, and to abut on, the epidermal
layer ( text fig. 2, and fig. 1 ) . It is beHeved that these variations in
the development of the spines represent, as indicated previously,
the maturity of a given seed, and should not be construed as having
taxonomic value.

Also of import is the location of the spines on the sclerotesta. As
indicated earlier, dependent upon the cut of the seed, one sees dif-
ferent aspects in the projection of the spines. In a cut at right angles
to the broad plane, the spines are seen to project into the sarcotesta
as far out as the epidermal layer, ( text fig. 2, and fig. 1 ) . However,
in peels taken from the broad plane it may be noted that as a
true median section is approached the projection of the spines
lessen until they appear to be little more than a serration of the
sclerotesta, (text fig. 1, and fig. 2). This constant variation in the
development of the spines has been further verified by the use of
3-dimensional peels.

Another characteristic noted is that the outer sclerotesta is
chambered as well as spined. This is evidenced by the presence
of "islands" of parenchymatous tissue of the sarcotesta surrounded
by the extensions of the sclerotesta (figs. 9 and 11).

The coloration of the outer sclerotesta is distinctive in being a
very dark brown, the intensity due, undoubtedly, to the extremely
thick-walled cells.

Inner sclerotesta: While the sclerotesta is made up of two dis-
tinct tissues, the inner sclerotesta has itself been found to be further
divided into two distinct zones. These will be referred to as the
outer zone of the inner sclerotesta, which is the zone abutting the
outer sclerotesta, and the inner zone of the inner sclerotesta which
is the inner layer of cells abutting the endotesta.

The outer zone of the inner sclerotesta: The tissue in this zone
averages 20 to 25 cells in thickness. The cells are sinuous, vertically
elongated, presenting a nearly isodiametric appearance in trans-
verse, or oblique-transverse, sections, and appearing as sinuous

Study of Cardiocarpus spinatus 159

tubes in longitudinal section. They are relatively thick-walled
near their juncture with the outer sclerotesta, but tend to become
less thick-walled as they approach the inner zone. The cells have
a distinct amber color and appear to be filled with a tan, resinouslike
substance.

The inner zone of the inner sclerotesta: This zone is immediately
adjacent to the endotesta. The zone appears to be six to eight cells
thick, but may have been slightly thicker in the living state. This
is evidenced by broken cell walls near the region of the endotesta.
The tissue is distinguished from the rest of the inner sclerotesta by
the thicker cell walls, and the fact that the lumen of the cells is
consistently empty.

Endotesta: Graham (1935) makes no reference to any layer
between the inner sclerotesta and the nucellus area in his descrip-
tions of the seed. Reed (1946) mentions an inner fleshy layer lying
between the sclerotesta and the nucellar area. However, her indi-
cation of this inner fleshy layer in her figure 2 is apparently only
the inner sclerotesta. Andrews and Felix (1952) note the presence
of an endotesta but state that the cellular tissue is poorly preserved,
and make no attempt to describe it.

This tissue in our specimens is, in general, not too well preserved.
However, in coal balls M-34, and 782-B, we have found some of the
endotesta preserved so as to present some idea as to the depth of the
tissue as well as cell shape and structure. The tissue appears to be
15 to 20 cells in thickness. This undoubtedly was variable, de-
pendent upon the maturity of the seed. In transverse, or oblique-
transverse, sections the cells are shown as isodiametric in appear-
ance. In a longitudinal view the cells appear thin-walled, paren-
chymatous, rectangular to rhomboidal, varying from 56[;. x 28tJi.
to 154[j. X 84[jt. with an average of 112ix x 42[j.. Delimiting the tissue
on its inner surface is a single layer of thicker-walled cells which
served, probably, as an epithelial lining (fig. 14).

The cells are a rich amber color, and the lumen area is filled with
a light brown resinouslike substance. This is a strikingly different
tissue than that found in any other region of the testa making it
quite easy to identify when preserved.

Nucellus: Graham (1935) notes that this tissue is poorly pre-
served, and does not attempt a description of it. Reed (1946) pre-
sents a limited discussion of this area, and Andrews and Felix ( 1952 )
mention the presence of this tissue but offer no discussion.

Here, as in the endotesta, the amount of tissue preserved is lim-
ited, but occasional seeds have provided material good enough to

160 The University Science Bulletin

give us an idea of its anatomical structure. In observing a mature
nucellus, the author has found the cells to be moderately thick
walled, closely packed, and with a coloration not unlike that of
the endotesta. The cells vary in diameter from 28[;. x 28ijl to
56[j,x28[jL with an average of 38pi, x 28[jl (figs. 14 and 15). As illus-
strated in fig. 15, the tissue is remarkable in apparently showing
the presence of nuclei.

Megaspore membrane: Graham (1935) and Andrews and Felix
(1952) mention the presence of this membrane, but offer little
discussion as to its composition. Reed (1946) states that it is
thick and conspicuous, and that

. in many seeds it appears firm and round and apparently outlines
faithfully the shape and extent of the female gametophyte or endo-
sperm

Little can be added to the information heretofore presented. Its
identification in our specimens appears as a brownish colored mem-
brane lying internally to the nucellus and abutting the external
layer of cells of the gametophyte.

Gametophyte: Graham (1935) notes that there is no gameto-
phytic tissue preserved in any of his specimens. Reed ( 1946 ) states
that little of this tissue is present in her specimens, and therefore
offers no discussion of it. With reference to her specimens, she
indicates that

there are occasional instances of tissue which may be endosperm,
but most of the tissue within the membrane is obviously intruded
material . , ."

Andrews and Felix ( 1952 ) were directly concerned with the game-
tophyte of C. spinattts Graham, and their excellent description has
left little to be added. Due to the fact we have on file a number
of nucellar coal balls with well-preserved gametophytic tissue (fig.
13), a little elaboration, but mostly corroboration, will be given
here with reference to the work done by Andrews and Felix.

In general, our specimens agree with the dimensions Andrews and
Felix give for their seed, ( fig. 12 ) . With reference to cellular detail
of the gametophyte, we would like to add that the cells are delicate,
parenchymatous, rectangular to isodiametric, varying from TOpi. x 56[jl
to 126[ji. X 84[JL with an average of 84ijl x 84[x. The archegonia present
in our specimens agree in size with that of Andrews and Felix.
The archegonia in their specimen attained a maximum diameter
of 0.5mm, and ours varies between 0.5 and 0.6mm. In no specimen
could be found more than two archegonia, and these were at the

Study of Cardiocarpus spinatus 161

apex of the megaspore, equidistant on either side of the pollen
chamber.

Vascular sysleni: The vascular system seems to be adequately
covered by Seward (1917), Coulter and Chamberlain (1917a),
Graham (1935), Scott (19()9a), Oliver (1902), and others, and
needs little further discussion here other than to present a short
summary of the works of one of the investigators. For this purpose,
the work of Oliver (1902) will be considered, because he was more
concerned with the genus as a whole and not with a particular
species. He states that

". . . as the supply bundle enters at the chalaza and traverses the sareo-
testa, it gives off a pair of bundles whieh run along the inner limit of that layer
to the micropyle. The plane in which these two bundles run is the plane of
flattening, generally designated the principal plane of the seed. The main
bundle continues to the base of the nucellus, where it expands into the tracheal
plate. From the margins of this plate a number of nucellar strands pass off
peripherally in the wall of the nucellus and extends a variable distance in the
direction of the pollen chamber . . ."

In this same publication, Oliver illustrates various genera of the
paleozoic seeds with their vascular systems.

The best indication of the vascular system in any of my specimens
may be seen in figure 2, which is a peel of seed M-6, and indicates
the entrance of the main vascular bundle at the chalaza, and clearly
shows the traversing strand in the sarcotesta.

SYSTEMATIC PROBLEMS

Due to the confusion that exists in the classification of paleozoic
seeds, there is a definite need for a reconsideration of the various
genera and species. It is important, where possible, to combine
seeds having essentially the same internal characteristics as con-
forms to size, shape, and integumentation in order to eliminate
superfluous generic names. Seward (1917) in considering this
state of confusion, notes that

the bionomics of paleozoic plants is a subject worthy of more
serious attention than it has so far received . . ."

He indicates that the search for morphological characters would,
in addition to other things,

. . . not only extend our knowledge of the morphology of ancient types,
hut would stimulate comparative study, and, incidentally, relieve the dullness
of pure description . . ."

He specifically notes, with reference to systematic problems, that,
• . . one of the difficulties inseparable from the study of fossil plants,

6—6730

162 The University Science Bulletin

namely the identification of impressions and casts with specimens exhibiting
anatomical characters, is particularly well illustrated by seeds. The descrip-
tion of a genus based on mere external fonns may sometimes be extended
without great risk of error to include species found on anatomical characters,
but on the other hand, there are many instances in which — despite a general
resemblance in form and size between petrifactions and impressions — lack of
evidence of generic identity reciuires the employment of distinctive
names . . ."

There are many examples one could present to show superfluous
names, and few investigators interested in this work are not familiar
with most of them. In the consideration of Cordaites seeds, one
is faced with the problem of certain descriptions indicating that
some of them possessed wings. Arnold (1947) indicates that in
general those flattened seeds with a narrow winglike border belong
to the Cordaiteae. He further states that the seeds of the Cardio-
carpales are flattened, heart-shaped seeds with the nucellar region
being surrounded by an integument of which the outer layer is
"laterally expanded into an encircling wing." He specifically states
that

It should be noted that in the Cordaiteae both the seeds and the
pollen grains are winged, a fact that stresses the important role of wind in
the life cycle of the plant

The evidence for a winged Cardiocarpus is based on those
specimens preserved as compressions. It is primarily through
this type of preservation upon which the concept that those seeds
of Cardiocarpus are flattened and heartshaped, is based. Numerous
intact specimens found in coal balls from West Mineral, Kansas,
and the coal mines near What Cheer, Iowa, compels me to state
that both concepts are erroneous, and I feel that there is no valid
evidence of a wing being present on seeds of this group, nor are
any members of the group flattened and heartshaped in external
appearance.

The outline of a wing, as indicated by those specimens preserved
in compressions, is quite obviously due to the compression of the
sarcotesta. The evidence for the seeds being flattened and heart-
shaped has arisen from investigators seeing the exposed nucellar
area, which is flattened and heartshaped. However, seeds, taken
intact from the coal, and from coal balls, indicate that none pos-
sessed wings, and that they are more spherical than flattened in
external appearance, and not heartshaped. Any specimen found to
be definitely winged should be placed in the genus Samaropsis
providing other characters conform to those attributed to that genus.

Study of Cardiocarpus spinatus 163

Identifying characters: One of the greatest needs in the study of
fossil seeds is a need for a classification plan that would follow
natural relationships rather than purely artificial characters. Just
what should be considered in developing such a key is difficult to
ascertain. In a classification plan for modern seeds Martin (1946)
states that external characters are many and often varied among
species of living seeds. Accordingly, Martin chose the embryo as
the principal criterion in his classification, stating;

". . . the significant types of internal arrangement incline to be relatively
few and comparatively stable.

In considering a classification plan for paleozoic seeds, the com-
plete lack of embryos renders impossible their use as a criterion for
classification. Therefore, other more prominent and identifiable
characters must be considered. The shape of the seed, platyspermic
or radiospermic, has been used with some success in this way,
Scott ( 1909a ) and Seward ( 1917 ) . I also find that the testa seems
to be well adapted for use as an identifiable character. Here the
numbers of layers of tissues, their thickness, cellular size and shape,
distinctive formation, i.e., sclerotesta spiny or not spiny, certainly
have a distinct importance. The arrangement of the vascular sys-
tem, when identifiable, and the external appearance of the seed,
where possible, are other characters w^hich may be used.

DISCUSSION

Theories on the lack of embryos in paleozoic seeds.

Solms-Laubach ( 1891 ) gives as a probable explanation of the lack

of embryos in paleozoic seeds that,

in these forms the development of the embryo proceeds slowly, as
is the case at the present day with Ceratozamia, Ginkgo, and Gnetum.

Accordingly, petrifaction may have usually preceded embryo
formation.

Scott ( 1909a ) notes the absence of embryos as being a,

. . . common character to all known paleozoic seeds, though in many
the archegonia are preserved. . ."

He continues wdth,

' • • . yet the seeds, as a rule, are evidently mature and not mere ovules.
Neither, considering the frequent presence of pollen grains in the pollen cham-
ber, can we assume that all the seeds found happen to be barren. It is possible
that, though pollinated, they were not yet fertilized, the latter process taking
place an appreciable time after the seed was shed; or again, it may be that a
resting stage followed immediately on fertilization before any marked develop-

164 The University Science Bulletin

ment of the embryo had taken place. In any case it appears that in these
ancient seeds the period of rest came much earHer with reference to the
growth of the embryo than in most of our recent seeds. We find a rehc of the
old conditions in li\ing cycads, in which the embryo is often scarcely to be
recognized wlien (he seed is ripe and ready for sowing.

The most complete hypothesis with reference to the lack of an
embryo in paleozoic seeds, is presented by Coulter and Chamber-
lain ( 1917b ) . It is known that embryos can be preserved in petri-
factions, as witnessed by their abundant occurrence in the seeds
of the Mesozoic Bennettitales. Coulter and Chamberlain do not
agree with Oliver's ( 1902 ) suggestion that perhaps the long intra-
seminal period of the embryo seed plants had not been developed
in the paleozoic. On the contrary, they believe

the abundant endosperm invested by an efficient testa seems to be
against this view . . . while ... it should be remembered that
the proembryo of the more primitive gymnosperms is quite extensive, and that
even were it preserved it would hardly be recognized as an embryo. . ."

In the same discussion they postulate,

that paleozoic seeds matured ( developing testa ) before fertilization,
became detached after pollination, and subsequently developed a proembryo
which did not invade the endosperm before germination, or invaded it in a
very feeble way as compared with the extensive invasion in modern gymno-
sperms. . .," and that, "• • • checking the growth of the enibyro if it
is related to the organization of a testa, may occur at any stage; and if the testa
is formed before fertilization (as may have been true in paleozoic seeds), the
growth of the embryo may well have been checked at some stage of the pro-
embryo. That even a well-organized testa does not check the growth of the
embryo, however, is evidenced by the fact that even among existing gymno-
sperms, as cycads, the true embryo (as distinct from an extensive proembryo)
may escape from the seed as soon as it is developed and grow as continuously
as do the embryos of pteridophytes. In such a case, however, the chance
of finding preserved a recognizable embryo invested by the testa would be
as exceptional as of finding any very brief stage in the life history. It would
seem, therefore, that in paleozoic seeds there was either no resting period
between intraseminal and extraseminal development of the very young plant,
or it occurred in the proembryonic stages. . ."

Scott (1909b) has suggested as probable
". . . that the nursing of the embryo had not yet come to be one of the
functions of the seed, and that the whole embryonic development was rele-
gated to the geniiination stage. . ."

Coulter and Chamberlain also note that

since belief in the absence of embryos from paleozoic seeds rests as
yet upon negative evidence, it should not be forgotten that the sections of
suitably preserved seeds are far too few in number to overcome the uncer-

Study of Cardiocarpus spinatus 165

tainty of negative evidence or the possibility of abortive seeds. In fact, com-
paratively few seeds have been found attached to the shoots that bore them,
and seeds that have fallen off at the iniinature stages suggested by all pub-
lished figures of sections would be abortive seeds, so far as experience with
the living plants is concerned. Only good seeds remain on the plant, and
there is no record of a section of an attached paleozoic seed. Moreover, if
the seeds germinated at once, as do those of the cycads, no embryos would
be found except in attached seeds. The seeds of Bennettitales that show con-
spicuous embryos are all attached seeds, and it is altogether probable that such
seeds of Cycadofilicales and Cordaitales will also show embryos . . ."

There have been Httle or no improvements of late on the fore-
going postnhitions for we today have yet to find an embryo or
even an attached paleozoic seed showing preserved internal struc-
ture. With the small bits of information available with reference
to these paleozoic seeds, Coulter and Chamberlain's theory appears
to be well founded, but it must remain only a theory until such
time as the paleobotanical record either proves or disproves it. It
does allow, however, a good working hypothesis to carry on re-
search in this field for the time being.

Comparison of Cardiocarpus with Ginkgo and the Cycads

To attempt to place Cardiocarpus in its proper taxonomic niche
of the family tree of plants is no easy task. Even with the knowl-
edge at hand, no definite placement can be made until such time
when embryonic development can be determined and definite at-
tachment to the parent plant can be established. However, a com-
parison with some modern seeds can be made to show similarities
in development and probable phylogenetic relationships.

To begin with, the nucellar attachment may be compared. We
find in the modern day cycads and Ginkgo that only the upper por-
tion of the nucellus is free, whereas in Cardiocarpus the only at-
tachment of the nucellus is found to be at the chalaza. There is
a distinct similarity in the development of the integument in all
three groups of seeds in that there is an inner fleshy layer (endo-
testa), a middle stony layer (sclerotesta), and an outer fleshy
layer ( sarcotesta ) . In comparing the vascular system of the three
groups it may be seen that the vascular system of Cardiocarpus is
somewhat more complex than that of the living cycads and Ginkgo.
It is interesting to note that the entrance of two vascular strands
into the ovule and testa, is similar in the three groups. These are
well illustrated and compared by Oliver ( 1902 ) , and by slight modi-
fications of the vascular strand of Cardiocarpus the trend towards

166 The University Science Bulletin

the cU'velopmcnt of the system in tlie cycads and Ginkgo may be
seen.

Further comparisons between the cycads, Ginkgo, and Cardio-
carptis inchide the prominent nucelhis with a relatively large
pollen chamber surmounted by a beak or "tent pole", all being
quite similar in the three groups, though the pollen chamber of
Cardiocarpus is the larger. With reference to the nucellar area
we find that in all three groups the nucellus is platyspermic and
"heart-shaped" following very closely the description Brongniart
(182(S) gave for the genus Cordiocarpon.

Additional comparisons we can make, are, for the moment,
hypothetical, because of the lack of sufficient knowledge of paleo-
zoic seeds. However, a free nuclear period following fertilization
is a character probably shared by the three groups. The presence
of swimming sperms known for the cycads and Ginkgo probably
was a character present in the seeds of the Cordaitales. With
reference to integumental development, Seward (1917) states,

". . . in both extinct and modern seeds the balance of evidence would
seem to be in favour of attributing a single rather than a double origin to
the integument .

Coulter and Chamberlain (1917a) in comparing the majority of

known paleozoic seeds with those of angiosperms, state,

". . . it is evident that the seeds (of the angiosperms) have advanced in
the reception of pollen by the micropyle, and in the nutrition of the embryo;
but that the general structure of paleozoic seeds is much more complex, with
their elaborate vascular system, pollen chamber, and often highly differ-
entiated testa

These comparisons are not made to indicate a direct lineage
of development of modern gymnospermous seeds from the paleozoic
seeds, but are here recorded to show similarities and possible
heritage. This line of thought has been held for a number of
years by prominent paleobotanists and no recent investigator has
postulated a better comparison. Until such time as future dis-
coveries provide the answers to the innumerable questions pre-
vailing in the paleobotanical records, concepts as to development
and phylogenetic relationships of present day plants with those
of primeval forms must remain purely theoretical.

Theory of the preservation of Nucellar Coal Balls in Coal.

The preservation of the nucellar area of Cardiocarpus seeds in
coal, as shown in figures 4, and 5, in addition to the finding of
numerous similar specimens lying in the weathered areas of

Study of Cardiocarpus spinatus 167

coal fields (fig. 3), raises the question as to why the nucellar
region was preserved and not the testa. It seems logical that the
hardiest cells or tissues would be preserved while the more deli-
cate cells or tissues would disintegrate.

It is known that since tissues rapidly decay in air, only those
plants living in or near water have much chance of preservation
as fossils. The exceptional facilities available for plant preservation
during the Upper Carboniferous period is a probable explanation
for the numerous fossil plants found in the Coal Measures. Arnold
(1947) indicates that,

the chief factors governing the extent of tissue preservation in fossil
plants are ( 1 ) the kinds of tissues composing the plants, and ( 2 ) the con-
ditions to which they were subjected preceding and during fossiliza-
tion

He further states that,

. the disintegration of plant tissues proceeds according to a rather
definite selective order of decomposition. First to disappear are the proto-
plasmic contents of the cells . . . Those tissues consisting of thin-walled
cells usually disappear next. These are followed in order by wood and bast
fiber cells, suberized cork cells, and cutinized spores and epidermal frag-
ments . .

The foregoing being true, why were nucellar coal balls pre-
served in coal minus their integument? Again we go to Arnold
(1947) to find a partial answer. He indicates that the escape
from serious damage of cells by crushing may be caused by a
previous infiltration of mineral matter into the cell cavities, and
where cell cavities have become completely filled, the structure
may suffer little or none at all. The explanation as to why thin-
walled cells are not crushed when the wood cells are, is that the
thin walls are more permeable to mineral matter in solution.
Arnold states that,

the efiFect of mineral infiltration is to add strength to the tissues,
whereas in the less permeable thicker-walled cells the cavities remain empty,
rendering them more susceptible to collapse .

Here, then may lie the nucleus of a final answer to the problem.
Undoubtedly mineral infiltration into the nucellar region of the
seeds (through the micropyle opening) was quite rapid to pre-
serve the tissue as we find it in the nucellar coal balls. Some infil-
tration may have occurred simultaneously in the sarcotesta, but to
a lesser degree, or the sarcotesta resisted infiltration because of the
presence of the cuticle layer externally to the epidermal layer. The
cells of the sclerotesta, on the other hand, being sclerotic in nature,

168 The University Science Bulletin

resisted infiltration of any mineral matter. During the coal forming
process, the noninfiltrated integuments of the seed were compressed
and carbonized while the infiltrated nucellar areas were preserved
as miniature coal balls.

SUMMARY

The anatomy and modes of preservation of the genus Cardio-
carpiis have been studied, with special attention to the anatomy of
Cardiocarpiis spinattis Graham.

Special attention has been given to the sclerotesta of the seed.
This portion of the integument is made up of two distinct tissues:
the outer sclerotesta is chambered and spined, the spines often
extending to the epidermis in sections taken through the narrow
plane with little or no extension of the spines in sections parallel
to the broad plane, while the inner sclerotesta is made up of two
distinct zones.

Additional information has been given on the tissues of the
sarcotesta; an account has been given of the endotesta and the
nucellar tissue that these tissues are quite distinct from other tissues
of the seed.

A new mode of preservation has been described and illustrated.
The specimens concerned in this new mode of preservation have
been designated as "nucellar coal balls."

A review of the theories relative to the lack of embryos in pale-
ozoic seeds is given, and a theory has been postulated with respect
to the preservation of nucellar coal balls in coal.

ACKNOWLEDGEMENTS

The author wishes to express his thanks and appreciation to
Dr. R. W. Baxter, Department of Botany, University of Kansas, for
his advice and criticism during the course of this study, and for
the use of a large number of specimens collected by him; to Dr.
H. N. Andrews, Henry Shaw School of Botany, Washington Univer-
sity, St. Louis, Missouri, for his interest and advice on the project,
and for the loan of his specimens; to Dr. S. H. Mamay, of the
U. S. Geological Survey, for his generosity in permitting the use
of certain of his specimens and allowing me to illustrate them in
this work.

Study oi" Cardiocarpus spinatus 169

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Lesquereux, L.

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170 The University Science Bulletin

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1873. Descriptions of fossil plants from the Coal Measures of Ohio.

Geological Survey of Ohio, vol. 1, part 2, section 3, Paleontology,

1837, pp. 359-385, pi. XLI, figure 4.

Oliver, F. W.

1902. On some points of apparent resemblance in certain fossil and
recent gymnospemious seeds. New Phytol., vol. 1, pp. 145-154.

1903. The ovules of the older gymnosperms. Ann. Bot., vol. 17, pp.
625-629.

Reed, F. D.

1946. On Cardiocarpon and some associated plant fragments from Iowa
Coal Fields. Bot. Gaz., vol. 108, pp. 51-64.

Scott, D. H.

1909. Studies in Fossil Botany. London.

1909. The Paleontological record. II. Plants. Darwin and Modern
Science, Cambridge University Press., pp. 200-222.

Seward, A. C.

1917. Fossil Plants, vol. Ill, pp. 1-656, Cambridge.

Solms-Laubach, H. G. Z.

1891. Fossil Botany, p. 119, Oxford.

White, D.

1899. Fossil Flora of tlie lower Coal Measures of Missouri. U. S. Ge-
ological Survey Monograms, Washington, 1899, pp. 265-267, pit.
LXXII, figure 3.
Williamson, W. C.

1877. On the organization of the fossil plants of the Coal Measures.
Philosophical Transactions Royal Society of London, vol. 167, part
1, 1877, pp. 213-270, pis. 14-16, figures 116-113.

Study of Cardiocakpus spinatus

171

Cardiocarpus spinatus Graham

Fig. 1. Transverse-oblique seetion through tlie narrow plane showing the
spines of the sclerotesta extending to, and abutting on, the epidermis. X 5?».

Fig. 2. Median-longitudinal section through the broad plane. Mieropyle
opening is visible at top center. At bottom center are shown vascular strands
entering the sclerotesta, and branching into the inner sarcotesta. X olL

Fig. 3. Nucellar coal balls as found at West Mineral, Kansas. All consist
of just the nucellar region except for the specimen second from right in bot-
tom row which shows some attached sclerotesta. X 2/j.

172

The University Science Bulletin

Cardiocarptis spinahis Graham

Fig. 4. Nucellar coal ball embedded in coal. X SJL Courtesy — Dr. S. H.
Mamay, U. S. Geological Survey.

Fig. 5. Nucellar coal ball embedded in coal collected from West Mineral,
Kansas. X 3J2.

Fig. 6. Nucellar coal ball from West Mineral, Kansas, showing its exact
fit into a vacated nucellar area of Cardiocarpiis spinatus Graham in a coal ball
from the What Cheer Mine, Iowa. X 2/2.

Fig. 7. Enlargement of nucellar coal balls from West Mineral, Kansas.
Specimen in upper left hand corner has retained the sclerotesta as its outermost
covering. X 5/2.

Study of Cardiocarpus spinatus

173

Cardiocarpus spinatus Graham

Fig. 8. Oblique-transverse section photographed from etched surface of
coal ball. Tissues from outside towards center are: Outer sarcotesta (black
cell contents), Inner sarcotesta (cells smaller and empty), spiny sclerotesta,
fibrous inner sclerotesta, endotesta, and nucellus. X 7.

Fig. 9. Oblique-trans\'erse view showing well preserxed tissues of the
testa. Note chamber in the outer sclerotesta in the upper right hand comer
of illustration. X 6)2.

Fig. 10. Photomicrograph of epidennal cells. X 20.

Fig. 11. Enlargement of a portion of fig. 9 showing, from right to left:
The large cells of the outer sarcotesta, the smaller cells of the inner sarcotesta,
chambered outer sclerotesta, and inner sclerotesta. X 10.

174

The University Science Bulletin

'^^-*i#'-^:

•^^%

Cardiocarpus spinatus Graham

Fig. 12. Oblique-transverse section through narrow diameter of a seed con-
taining a gametophyte. X 6/'2.

Fig. 13. Peels of gametophytic tissue from nucellar coal balls. An arche-
gonium may be seen immediately left of the micropylar region in upper peel.
Pentagonal pattern of bottom peel appears to be due to mineralization. X 6.

Fig. 14. Oblique-transverse section through narrow diameter showing extent
of endotesta and its epithelial Hning. From left to right the tissues are: Outer
aclerotesta (large black band), inner sclerotesta (lighter wavy portion immedi-
ately internal to the outer sclerotesta), Endotesta (wide band of cells light
in color). Epithelial lining (dark band, one-cell thick, on inner surface of endo-
testa), Nucellus tissue, Megaspore membrane. X 20.

Fig. 15. Photomicrograph of nucellus tissue. Note the appearance of nu-
clei in some of the cells of the tissue. X 40.

THE UNIVEESITY OP KANSAS

SCIENCE BULLETIN

Vol. XXXVII, Pt. I] October 15, 1955 [No. 6

The Distribution and Subspeciation of Erebia epipsodea
Butler (Lepidoptera: Satyridae)*

BY

Paul R. Ehrlich
INTRODUCTION

This is the second in a series of papers on the subspeciation
and distribution of the species of Erebia found in North America.
The purpose of these papers, and those to follow, is to summarize
on the specific level our knowledge of the genus in the Nearctic
region. The author's intention is to provide for future workers
on Erebia an organized modern analysis of the individual species
on which they can base further studies.

Erebia epipsodea Butler is appropriately known as the "Common
Alpine". It is our most widely spread species, and in collections
it far outnumbers all other species with the possible exception of
E. discoidalis Kirb\'. The fact that it is an extremely variable
species had given rise to the myth that it was devoid of sub-
specific variation, an idea so widely accepted that such a distinct
race as E. e. remingtoni Ehrlich remained undescribed years after
its characteristics had been noted and it had been figured in
color in our most popular book on butterflies (Holland, 1931).
Recently two additional subspecies have been described,' demon-
strating that epipsodea shows as much geographic variation in
North America as any Erebia.

Historical Background

Since the description of epipsodea by Butler in 186S, the litera-
ture on the species has been characterized by short notes on
distribution and biology, both in periodicals and books. A brief
glance at the synonymies will suffice to explain to tlie reader why
the author has not attempted a detailed discussion of material
published.

*Contribution No. 883 from Department of Entomology, University of Kansas, Lawrence.
1. Ehrlich, 1954, Jomrial of the Kansas Entomological Society, vol. 27, p. 80.

(175)

176 The University Science Bulletin

Materials and Methods

The author has personally examined over 1600 specimens of
epipsodca and has received data from an unknown additional
number. The work is based largely on the author's collection of
some 500 specimens, probably the most comprehensive in existence,
supplemented by extensive material from the American Museum
of Natural History, the Canadian National Collection, and the
collection of F. M. Brown. The large collection of C. F. dos Passos
was consulted, but was not worked on in detail as was the afore-
mentioned material. The material in several smaller collections
was also examined.

In order to be able to present a unified discussion of the dis-
tribution and subspeciation of epipsodea the whole picture has
been discussed in detail at the end of the paper, and only briefly
under the individual subspecies. Each subspecies is first treated
separately in order to prevent nomenclatorial confusion.

All measurements are length of one primary wing, base of
subcosta to apex in millimeters, taken to the nearest millimeter.

Acknowledgments

The author wishes to thank the following people who kindly
supplied him with specimens, data, and/or advice: J. B. Baker,
C. Bird, K. Bowman, P. F. Bruggemann, F. M. Brown, O. Buchholz,
H. K. Clench, H. de Lesse, C. F. dos Passos, J. D. Eff, T. N. Free-
man, A. G. Gabriel, N. W. Gillham, W. A. Hammer, J. C. Hop-
finger, R. Leuschner, L. M. Martin, E. G. Munroe, Mr. and Mrs.
F. W. Preston, C. L. Remington, P. S. Remington, D. B. Stallings
and J. B. Ziegler.

The author is most grateful to Dr. Alexander B. Klots, Research
Associate of the American Museum of Natural History, Mr. F. Martin
Brown of Colorado Springs, and Dr. R. R. Sokal of the Department
of Entomology, University of Kansas, all of whom have given
very extensive assistance on many facets of the work. Finally
the author wishes to express his deep appreciation to Dr. F. H.
Rindge, Curator of Lepidoptera of the American Museum, who
gave up much of his valuable time to read this manuscript and
to give advice on a great many taxonomic problems.

SYSTEMATIC DESCRIPTIONS

Erehia epipsodea is easily identified by both the facies and the
genitalia. Its habitus is actually unlike that of any other Nearctic
Erehia, but as a result of superficial examination an occasional

Distribution and Subspeciation of Erebia 177

si3eciineii of Erebia disci Thunberg has found its way into series of
epipsodea. Any specimen of epipsodca should be determinable
without resort to genitalic dissection through the use of the plates
and descriptions of the present paper or of any other work which
deals with the Nearctic Erebia. However if it should prove neces-
sary to employ the genitalia to determine the specimen, the follow-
ing distinctions should serve: in the male, epipsodea is the only
Nearctic species with a toothed juxta and aedeagus; in the female,
it is the only Nearctic species with a greatly reduced clapet (the
scooplike projection of the genital plate which is characteristic of
a great many Erebia ) .

A total of 71 genitalic preparations of epipsodea have been made
by tlie author, including both sexes of all subspecies. No significant
intersubspecific differences have been found.

The life history of epipsodea was described by Edwards in 1897,
in his "Butterflies of North America" (3rd series, Erebia 3). Habits
and habitat will be discussed under the individual subspecies.

Erebia epipsodea epipsodea Butler

Figs. lA, IB, 2A, 2B, 3A, 38

Erebia epipsodea Butler, 1868, Catalogue of diurnal Lepidoptera . . .
family Satyridae in the . . . British Museum, p. 80, pi. 2, fig. 9;
Edwards, 1872, Synopsis of North American butterflies, p. 26; Edwards,
1877, Trans. Anier. Ent. Soc., vol. 6, p. 34; Edwards, 1884, ibid., vol. 11,
p. 289 {pa?-tim); Edwards, 1897, Butterflies of North America, 3rd series,
Erebia 3, figs. 1-5, a-g (partim), op. cit., supplementary notes; Mead,
1875, in Wheeler, report upon geographical explorations and survevs
. . . vol. 5, p. 775; Scudder, 1875, Bull. Buffalo Soc. Nat. Sci., vol.
2, p. 243; Strecker, 1878, Butterflies and moths of North America, p. 151;
Strecker, 1879, in Ruffner, Annual report upon explorations and surveys in
the department of the Missouri, appendix, S. S. of the annual report of the
Chief Engineers for 1877, p. 1856; Elwes, 1889, Trans. Ent. Soc. London,
p. 326 (partim); Elwes, 1898, ibid., p. 190; Cockerell, 1890, Ent. News,
vol. 1, p. 58; }vlaynard, 1891, A manual of North American butterflies, p.
114 (partim); Skinner, 1898, A synonymic catalogue of North American
Rhopalocera, p. 36; Skinner, 1905, ibid., supplement no. 1, p. 14; Chap-
man, 1898, Trans. Ent. Soc. London, p. 230, pi. 14, fig. 43; Holland,
1898, The butterfly book, p. 210, pi. 25, fig. 28 (partim); Holland, 1915,
The butterfly guide, p. 133, pi. 66, fig. 2 (partim); Holland, 1931, The
butterfly book, revised ed., p. 204, pi. 25, fig. 28, pi. 61, figs. 25, 26 ( par-
tim); Dvar, 1902, Proc. Ent. Soc. Washington, vol. 5, p. 129; Dyar, "1902"
(1903), Bull. U. S. Natl. Mus., vol. 52, p. 29 (partim); Wright, 1906,
The butterflies of the West Coast, p. 190 ( partim ) ; Elrod, 1906, The but-
terflies of Montana, p. 114, figs. 87, 88; Oberthiir, 1909, Etudes de Lep-
idopterologie comparee, vol. 3, p. 295; Weymer, 1911, in Seitz, Macro-
lepidoptera of the world, vol. 5, p. 238, pi. 51e (partim); Blackmore, 1927,
Checklist of the Macrolepidoptera of British Columbia, p. 8 (partim);
Klots, 1930, Bull. Brooklyn Ent. Soc, vol. 25, p. 156; Klots, 1951, A field
guide to the butterflies, p. 76 (partim); Gaede, 1931, in Strand, Lepidop-
terorum Catalogus, p. 554 ( partim ) ; Warren, 1936, Monograph of the
genus Erebia, p. 239, pi. 38, fig. 358, pi. 83, figs. 1164, 1170, 1171 (par-
tim); Cross, 1937, Proc. Colorado Mus., vol. 16, p. 11; Davenport and

178

The University Science Bulletin

])t'tliier, 1937, Ent. Americana (new series), vol. 17, p. 169; dos Passos,
1939, Bull. Cheyenne Mountain Mus., vol. 1, pt. 2, p. S-13; dos Passos,
1951, Jour. New York Ent. Soc., vol. 56, p. 161; Brown, 1942, Colorado
College Publication, general series no. 233, p. 17; Brown, "1951"
(1952), Lepidopterists' News, vol. 5, p. 119; Eff, 1947, Lepidopterists'
News, vol. 1, p. 91; EfF, "1948" (1949), ibid, vol. 2, supplement, p. iv;
EflF, "1949" (1950), ibid., vol. 3, p. 92; EfF, "1951" (1952), ibid., vol. 5,
p. 92; C. L. Remington, 1948, Lepidopterists' News, vol. 2, p. 37; Rem-
ington, 1952, Psyche, vol. 59, p. 66; de Lesse, 1949, Rev. Francaise d'En-
tomologie, vol. 16, p. 181; Bowman, 1951, Canadian Jour. Zool., vol. 29,
p. 130 (partim); Jones, 1951, Ent. Soc. British Columbia, occasional paper
no. 1, p. 9jpartim); Ehrlich, 1952, Ent. News, vol. 63, p. 225; P. S.
Remington, "1952" (1953), Lepidopterists' News, vol. 6, p. 103.
Maniola epipsodea, Kirlay, 187l, A synonymic catalogue of diurnal Lepidop-
tera, vol. 1, p. 65.

Photographs by C. F. dos Passos.

Fig. 1. Erebia epipsodea Butler, type, Rocky Mountains.
A. Upper side. B. Under side.

Erebia Rhodia Edwards, 1871, Trans. Amer. Ent. Soc, vol. 3, p. 273; Ed-
wards, 1872, List of species collected . . . in Hayden, Preliminary
report of U. S. Geological Survey of Montana and adjacent territory, fifth
annual report of progress, p. 467; Brown, 1934, Jour. New York Ent. Soc,
vol. 42, p. 161; Ehrlich, 1952, Ent. News, vol. 63, p. 230.

Erebia epipsodea var. rhodia, Saunders, 1885, Trans. Royal Soc. Canada, sect.
4, p. 98.

Erebia epipsodea? var. brucei Elwes, 1888, Trans. Ent. Soc. London, p. 326;
Elwes, 1898, ibid., p. 190; Edwards, 1897, Butterflies of North America,
3rd series, Erebia 3, figs. 6, 7; Weymer, 1911, in Seitz, Nhicrolepidoptera
of the world, vol. 5, p. 238; Warren, 1936, Monograph of the genus
Erebia, p. 241, pi. 83, fig. 1171; Cross, 1937, Proc Colorado Mus., vol. 16,
p. 11; Davenport and Dethier, 1938, Ent. Americana (new series), vol.
17, p. 169; Ehrlich, 1952, Ent. News, vol. 63, p. 230.

Erebia brucei, Holland, 1931, The butterfly book, revised ed., p. 204, pi. 61,
fig. 20.

Male: length of one primary wing 19-24 mm., average (152
specimens) 21.7 mm.

Above, ground color light to deep chocolate brown. Primaries
with deep orange-brown submarginal patches. Patches in cells Rg,
Ml, Mg, M3, Cui, narrowest in cell Cui, widening craniad with
widest in cell M^, then narrowing slightly to cell R,. Largest patch
3-6 mm. wide. Patches in cells M^, Mo, M3, and Cui, usually

Distribution and Subspeciation of Erebia 179

enclosing black ocelli. Snbapical ocelli (those in cells M, and M,)
always pupilled with white, sometimes fused. Ocellus in cell M.
smallest, rarely pupilled, sometimes absent. Ocellus in cell Cu^
smaller than subapical ocelli, sometimes pupilled, rarely absent.
Veins separating patches ground color, patch in cell Cu^ sometimes
separated from other patches by a wide strip of ground color along
vein Cu^. Outline of patches varying in sharpness, sometimes very
indefinite. Secondaries with submarginal patches in cells Mj, M.^,
M3, and Cuj. Patches concolorous with those on primaries. Patch
in cell Ml often reduced, very indefinite, or absent. Largest patch
in cell M2, with those in cells Mg and Cuj progressively smaller.
Black ocelli of varying size, often pupilled with white, are found
in patches of cells Mo, M3, and Cuj; patch in cell M^ rarely with
an ocellus, never with a pupilled ocellus. Patches often reduced
to rings around the ocelli.

Below, ground color a duller and lighter brown than above. Pri-
maries with patches suffused with ground color, especially in cells
R5, M3, and Cuj. Subapical ocelli reproduced as above. Ocellus
in cell M3 absent, that in cell Cuj absent or represented by a black
point. Apex and sometimes outer margin flecked with gray scales.
Secondaries traversed by a dark brown discal band, crenate distad,
irregular basad. Prominence of the band varying with the amount
of gray scaling in the basal and limbal areas. Patches reduced
to light rings around the ocelli. Ocelli in cells M.,, M3, and Cu^
usually present but extremely variable in size, ocellus in cell M^
less frequently present. Pupilling very varied. In some specimens
there is a faint dark marginal band. Fringes concolorous with
ground color.

Head, thorax and abdomen brown-black. Antennae above brown,
flecked with white scales; below white with yellow brown club.

Female: length of one primary wing 20-23 mm., average (34
specimens) 21.6 mm.

Above, ground color and patches somewhat lighter than in the
male. Ocelli larger and more heavily pupilled. Rarely with a
black point in cell Rg and/or Cu,.

Below, ground color slightly lighter than above. Ocelli repro-
duced below as in male. Basal and limbal areas of secondaries and
veins of discal area scaled with gray. Discal band not much more
prominent than in male because of its lighter color.

Otherwise similar to male.

180 The University Science Bulletin

Erebia epipsodea epipsodea is found from the high mountains of
northern New Mexico north in the Rockies to western Alberta and
eastern British Cohmibia (see map). It occurs in the Hudsonian,
Canadian, and upper Transition zones. It is found in small numbers
in the arctic-alpine zone, but Brown states (in lift.) "I have a strong
suspicion that where they (epipsodea) are abundant at treeline you
will always find that the treeline is false ... In many places in
the state (Colorado) the treeline has been artificially depressed by
snowslides or avalanches." In Colorado E. e. epipsodea occurs in
open grasslands and less frequently in sage flats. It is found over
the western part of the state above 7,000 feet up to 11,800 or 12,000
feet, chiefly between 8,500 and 10,000 feet. In general in Colorado
epipsodea flies in June around 8,500-9,000 feet, July around 11,000,
and extends well into August at 12,000 feet. The interesting aber-
ration of epipsodea which has no ocelli on the upper surface
(brucei Elwes) is found mainly at high elevations in Colorado. Mr.
and Mrs. F. W. Preston found it quite frequent among typical
epipsodea just below the summit of Loveland Pass at an elevation
of about 11,950 feet, in early August of 1952. It was impossible to
separate the brucei from the epipsodea in flight, and they appeared
to be randomly distributed in the epipsodea population. Some 66
epipsodea and 11 brucei were taken in three days, but these num-
bers indicate a larger proportion of brucei than was actually present
as some seconds of epipsodea were released. The Prestons kindly
sent the author two of the brucei which they captured and a series
of the epipsodea from the same locality. Among the latter the
author was pleased to find two males which are definitely transi-
tional to brucei.

In southern Wyoming, Klots (1930) found epipsodea in the lower
and upper Canadian zone (8,500-9,600 feet) in late June and early
July in small glades in the forest, and more abundantly in the
open meadows above the forest belt. The author has before him
fresh specimens taken in Albany county in southern Wyoming at
7,500 feet in early July, and in Sheridan and Johnson counties in
northern Wyoming at 7,800 feet, also in early July. Almost all of
the material which has been seen from Yellowstone National Park
has been taken in July. An interesting series in the author's collec-
tion from Kiowa, Montana, just east of Glacier National Park,
dated July 10, 1952, consists of 15 worn and battered males and a

Distribution and Subspeciation of Erebia 181

single fresh female, indicating a much earlier emergence. Banff,
Alberta specimens seem to be taken mainly in mid-July.

Butler (1868) gives the type locality of Erebia epipsodea as
"Rocky Mountains." This unfortunately is not sufficiently restricted
for modern systematic work, and so every effort has been made to
discover more exactly the area where the type was captured. Due
to the great individual variations of epipsodea the type itself cannot
be accurately placed in one of the subspecies on the basis of its
facies. Mr. A. G. Gabriel, Senior Scientific Officer of the Depart-
ment of Entomology of the British Museum (Natural History)
kindly looked up the registration of the type and found that it was
one of 50 butterflies presented to the Museum by the Earl of Derby
in 1845. The present Earl of Derby (in lift.) informed the author
that he had no information on where or how his ancestor had ob-
tained the specimens. Mr. Gabriel, however, was able to discover
four of the original 50 butterflies in the British Museum collection.
They were a pair of Colias christina Edwards, a female Colia.s skin-
neri Barnes, and a female Speyeria eurynome Edwards. On consul-
tation with Dr. Alexander B. Klots, one of our foremost authorities
on the Nearctic Rhopalocera the following was decided: assuming
that the above specimens were taken in the same general area as
the type of epipsodea then the type locality of epipsodea would be
somewhere in the area from Red Desert gap in the Rockies in south-
ern Wyoming north to Jasper National Park in the Canadian
Rockies. Because of the great separation of christina and skinneri
in Montana there is a little better evidence for the eastern side of
the divide in the Canadian Rockies north to Jasper. This hypoth-
esis is supported by the fact that it seems improbable that specimens
would be reaching England in 1845 from the far northern Rockies
or the Cascades. As will be discussed later, material from that
part of the Canadian Rockies is somewhat transitional, but appears
to be closest to the subspecies from Colorado, therefore in this
paper Erebia epipsodea epipsodea is the name applied to the sub-
species which ranges from Colorado north, and typical epipsodea
will be considered as coming from Colorado.

Erebia rhodia Edwards was described from Colorado and is
therefore a synonym of Erebia epipsodea epipsodea Butler. Erebia
epipsodea? var. brucei Elwes described from Summit county, 12,000
feet, Colorado is an aberration and is also a synonym of E. e.
epipsodea.

182 The University Science Bulletin

Erebia epipsodea reniingtoni Ehrlich

FiRs. 2C, 2D, 3C, 3D

Ereh'ui cpipsndca, Holland, 1898, The butterfly book, p. 210 (partim); Hol-
land, 1900, Ent. News, vol. 11, p. 387; Holland, 1915, The butterfly
guide, p. 133 (partim); Holland, The butterfly book, revised ed., p. 204,
pi. 61, figs. 24, 27 (partim); Dyar, "1902" (1903), Bull. U. S. Natl.
Mus., vol. 52, p. 29 (partim); Weynier, 1911, in Seitz, Macrolepi-
doptera of the world, vol. 5, p. 238 (partim); Gibson, 1920, Report
of the Canadian Arctic expedition 1913-18, vol. 3, pi. I, p. 18i; Gaede,
1931, in Strand, Lepidopterorum Catalogus, p. 554 (partim); Warren,

1936, Monograph of the genus Erebia, p. 241 (partim); McDunnough,

1937, Canadian Ent., vol. 69, p. 18; Jones, 1951, Ent. Soc. British
Columbia, occasional paper no. 1, p. 9 (partim); Klots, 1951, A field
guide to the butterflies, p. 76 (partim).

Erebia epipsodea remingtoni Ehrlich, 1952, Ent. News, vol. 63, p. 225; P.
S. Remington, "1952" (1953), Lepidopterists' News, vol. 6, p. 105.

Male: length of one primary wing 21-22 mm., average (14
specimens) 21.6 mm.

Female: length of one primary wing 20-22 mm., average (7
specimens) 21.1 mm.

E. e. remingtoni differs from E. e. epipsodea in the reduction
in size and number of the ocelli on both surfaces of both sexes.
Approximately half of the males of this subspecies have no ocelli
on the underside of the secondaries, and the discal band is often
very dark and prominent. The number of white pupilled ocelli
are also greatly reduced in the males, with the subapical "ocelli"
sometimes being merely black disks. The patches in the males
are very variable, and are occasionally reduced to traces on the
upperside of the secondaries.

Erebia epipsodea remingtotu has thus far been seen from the
extreme northwestern corner of British Columbia; southeastern
Yukon Territory; along the Yukon River Valley from Dawson,
Yukon Territory, to Fort Yukon, Alaska, on the Arctic Circle; and
central Alaska from the Yukon border to Fairbanks.

According to Bruggemann (in litt.), remingtoni in the Dawson
area flies "in meadows, both dry and wet, open, grassy hillsides,
along roads and in grassy openings in the woods, both deciduous
and coniferous, up to elevations of about 4000 feet". The dates
of the type series run from June 11 to August 22.

This is the most distinct of the subspecies of epipsodea. It was
described by the present author ( 1952 ) from 121 specimens, from
a number of British Columbian, Yukon, and Alaskan localities.
The type locality is Dawson, Yukon Territory. Since the descrip-
tion, several more specimens and records have come to light (see
map), including 10 males and a female from Eagle, Alaska, in

Distribution and Subspeciation of Erebia 183

the collection of the Los Angeles County Museum. Mr. Lloyd
M. Ahirtin has examined these specimens, and writes that they
are typical of the subspecies. Two records given by Edwards
( 1897 ) are believed to be referable to Erebia disa ssp. These
are St. Michaels and Nushagak, Alaska.

Erebia epipsodea hopfingeri Ehrlich

Figs. 2E, 2F, 3E, 3F

Erebia epipsodea, Elwes, 1889, Trans. Ent. Soc. London, p. 326 (partim);
Maynard, 1891, A manual of North American butterflies, p. 114 (partim);
Edwards, 1897, Butterflies of North America, 3rd. series, Erebia 3 (partim);
Wright, 1906, The butterflies of the West Coast, p. 190, pi. 24, fig. 262
(partim); Blackmore, 1927, Checkhst of the Macrolepidoptera of British
Columbia, p. 8 (partim); Warren, 1936, Monograph of the genus Erebia,
p. 241, pi. 38, fig. 357, pi. 83, figs. 1163, 1165, 1169 (partim); Leighton,
1946, Univ. of Washington Publ. Biol., vol. 9, p. 55; Hopfinger, "1948"
(1949), Lepidopterists' News, vol. 2, supplement, p. iii; Hopfinger, "1949"
( 1950), ibid., vol. 3, p. 88; Hopfinger "1950" ( 1951 ), ibid., vol. 4, p. 93.

Erebia epipsodea hopfingeri Ehrlich, 1954, Jour. Kansas Ent. Soc, vol. 27, p. 80.

Male: length of one primary wing 22-25 mm., average (30 speci-
mens) 23.5 mm.

Female: length of one primary wing 23-25 mm., average (16
specimens ) 24.3 mm.

E. epipsodea hopfingeri is characterized by its large size, propor-
tionately small patches, and very dark chocolate brown ground color.
It is separated from E. e. epipsodea by these characters, and from
E. e. remingtoni by its complete set of ocelli and to a lesser degree
by the above characters. It is consistently larger than remingtoni
but sufficient fresh material of remingtoni is not available to make
a comparison of ground colors feasible.

Type Material: Holotype male and allotype female, Black Canyon
lUOO feet, South of Methow, Okanogan County, Washington, May
24, 1952 (J. C. Hopfinger). There are 53 male and 13 female para-
types as follows: 20 males and seven females, Black Canyon, 1000
feet, South of Methow, Okanogan County, Washington, May, June
(various dates) 1944, 1946, 1952 (J. C. Hopfinger); six males (geni-
talic slides nos. 13 and 83, P. R. Ehrlich), Tiffany Lake 7000 feet.
Northwest of Conconully, Okanogan County, Washington, June,
July (various dates) 1947, 1949 (J. C. Hopfinger); one male, Gold
Creek, Washington, June 27, 1943 (J. D. Eff); two males. Gold
Creek, Washington, June 18, 1944; one male and one female, Spo-
kane Falls, Washington Territory, May 30, 1882 (Stretch); one male,
Brewster, Okanogan County, Washington, June 10, 1944 (W. A.
Hammer); 17 males (genitalic slides nos. 91, 92, 93, and 94, P. R.
EhrHch) and four females (genitalic shde no. 87, P. R. Ehrlich),

184 The University Science Bulletin

Brewster, Washington, May, June (various dates) 1938, 1947, 1949,
1952 (J. C. Hopfinger); three males and one female, Wallowa Lake,
Wallowa National Forest, Oregon, July 23, 1940 (J. H. Baker);
one male, Wallowa Lake, Oregon, June 30, 1938; one male, same
locality, July 26, 1949. The holotype and allotype are in the Ameri-
can Museum of Natural History, ex collection P. R. Ehrlich. Of
the above paratypes five males and four females are in the collec-
tion of J. C. Hopfinger and one male is in the collection of J. D. Eff.
The remaining paratypes are in the collection of the author, some
of which will be distributed.

E. epipsoclea Jiopfingeri has been seen from eastern Washington,
northeastern Oregon, south-central British Columbia and northwest-
ern Idaho. Because of the uncertainty of the extent of the sub-
species in Idaho and British Columbia (to be discussed later) only
Oregon and Washington specimens have been included in the type
series.

The author was pleased to name this subspecies after his friend,
Mr. John C. Hopfinger of Brewster, Washington, who collected
most of the type series.

Erehia epipsoclea freemani Ehrlich

Figs. 2G, 2H, 3G, 3H

Ercbia sinc-ocellata Skinner, 1889, Canadian Ent., vol. 21, p. 239; Aaron, 1890,
Ent. News, vol. 1, p. 12; Edwards, 1897, Butterflies of North America, 3rd
series, explanation of pi. Erehia 3; Ehrlich, 1952, Ent. News, vol. 13, p. 230.

Erehia epipsoclea, Lvman, 1896, Canadian Ent., vol. 28, p. 274; Carv, 1906,
Proc. U. S. Natl. Mus., vol. 31, p. 448; Turner, 1906, Ent. Monthly Maga-
zine, vol. 42, p. 119; Turner, 1906, The Entomologist, vol. 39, p. 165;
Brodie, 1929, Trans. Royal Canadian Institute, vol. 17, part 1, p. 83; Gaede,
1931 in Strand, Lepidopterorum Catalogus, p. 554 {partim); Warren, 1936,
Monograph of the genus Erehia, p. 241 (partim); Brooks, 1942, Canadian
Ent., vol. 74, p. 33; Bowman, 1951, Canadian Jour. Zool., vol. 29, p. 130
(partim); Klots, 1951, A field guide to the butterflies, p. 76 (partim).

Erehia episodea (sic). Turner, 1906, The Entomologist, vol. 39, p. 142.

Erehia epipsodea freemani Ehrlich, 19.54, Jour. Kansas Ent. Soc, vol. 27, p. 80.

Male: length of one primary wing 19-23 mm., average (54
specimens) 21.3 mm.

Female: length of one primary wing 20-22 mm., average (14
specimens) 21.1 mm.

This subspecies differs from the other three subspecies of epip-
sodea in the lighter orange color and sharper outline of the patches.
The sharper outline is especially evident on the underside of the
females. E. e. freemani is the brightest in habitus of the four sub-
species of epipsodea. The females often have a very heavy gray
overscahng on the underside of the secondaries.

Distribution and Subspeciation of Erebia 185

Type Material: Holotype male, Lloydininster, Alberta, June 8,
1946 ( P. F. Bruggemann ) ; allotype female, same locality, June 13,
1948 (P. F. Bruggemann). There are 60 male and 17 female para-
types as follows: 15 males and seven females (genitalic slides nos.
28 and 49, P. R. Ehrlich), Lloydminster, Alberta, June 1-13, 1944,
1946, 1947, 1948 (P. F. Bruggemann); 10 males (genitalic slides
nos. 96, 97, and 98, P. R. Ehrlich) and one female, Harlan, Sas-
katchewan, June 8-14, 1939, 1946, 1947 (P. F. Bruggeman); one
male and one female, Rivercourse, Saskatchewan, June 10, 25, 1940;
six males and one female, Attons Lake, Cut Knife, Saskatchewan,
June 8-9, 1940 (A. B. Brooks); one female, Cravan, Saskatchewan,
June 27, 1920 ( C. H. Young ) ; one male and one female, Starblanket,
Saskatchewan, May 27, June 1; seven males and four females, Mini-
ota, Manitoba, June 3-10, 1919, 1921 (H. Gibbon); three males,
same locality and collector, no dates; seven males (genitalic slide
no. 99, P. R. Ehrlich), Beulah, Manitoba, June 10-July 16; seven
males, McCreary, Manitoba, June 10-July 10, 1928, 1931; three
males and one female, Manitoba, Canada. The holotype and allo-
type are in the American Museum of Natural History, ex collection
P. R. Ehrlich. Of the above paratypes seven males and one female
are in the collection of the American Museum of Natural History,
13 males and three females are in the Canadian National Collection,
four males are in the collection of the Academy of Natural Sci-
ences of Philadelphia, three males are in the collection of Otto
Buchholz, one male is in the collection of F. M. Brown, ex collection
A. M. N. H., ex collection P. R. Ehrlich, one male is in the collection
of J. D. Eff, ex collection A. M. N. H., ex collection P. R. Ehrlich,
and one male is in the collection of J. C. Hopfinger, ex collection
P. R. Ehrlich. The remaining paratypes are in the collection of the
author, some of which will be distributed.

E. epipsodea freemani is the subspecies of the Canadian Prairie-
Parkland. According to Bruggemann (in lift.) it is the commonest
Satyrid in the Lloydminster district, flying in June along the
roads, in pastures (both dry and wet), on grassy hill sides and
along the margins of poplar "bluffs". Farther north at Harlan,
Saskatchewan, where the main feature of the vegetation is an
outlier of the northern coniferous forest, freemani flies at the same
time and in the same habitats as at Lloydminster but is slightly
less numerous. It occurs also on the grassy muskegs, but never
inside the stands of conifers, where it is replaced by E. disa
mancina Hewitson, which is rarely, if ever, encountered in the

186 The University Science Bulletin

open. The subspecies also flies at Cold Lake (80 miles due north
of Lloydminster) at the edge of, and around Primrose Lake
(100 miles due north of Lloydminster) within the northern conif-
erous forest, in habitats similar to those around Harlan. Brug-
gemann (in litt.) states "As a rule only the males are seen fly-
ing, rather leisurely, but somewhat erratically. As is the case
with several other northern Satyrids, e. g. Oeneis alberta alberta
Elwes and Edwards, O. uhleri variina Edwards, and Erebia dis-
coidalis Kirby, the females are sluggish fliers and, when fully
gravid, often unable to take off".

The subspecies has thus far been seen from southwestern
Manitoba, the southern two thirds of Saskatchewan and the
southern plains of Alberta. Because of the uncertainty of the
extent of the subspecies in Alberta only Saskatchewan and Mani-
toba specimens have been included in the type series.

The subspecies was named in honor of Dr. T. N. Freeman,
co-ordinator of the Northern Insect Survey, as a slight token
of appreciation of his many favors.

Erebia sine-ocellata Skinner was described as a "var. of epip-
sodea" from a specimen taken at Fort Qu'Appelle, Saskatchewan
and a specimen without data. From the discussion of the varia-
tion of epipsodea which is included in the short original descrip-
tion it is obvious that sine-ocellata was described as an individual
variant ", and all subsequent workers have considered it an aber-
ration, usually as a synonym of brucei Elwes. The type specimen
is, indeed, a transition towards brucei. Because of the ineptness of
the name and because the type of the subspecies would be an
aberrational specimen, sine-ocellata was not considered available
and freemani was proposed for the new subspecies.

GEOGRAPHICAL VARIATION

Because of the tremendous individual variation exhibited by
epipsodea it was necessary to assemble large numbers of speci-
mens before any significant geographical variation could be dis-
cerned. Anyone familiar with the species knows that no two
series are just alike, and material from one locality may be quite
different from material from another locality a few miles away.
Thus the problem has been to disregard the minor extrinsic varia-
tions and to discover the broader trends of (theoretically) genetic

2. See Bull. Zoo. Nomencl., vol. 10, pp. 438-44.5, 1953.

DlSTRinUTIOX AND SUBSPECIATION OF ErEBIA 187

variation. The criteria employed to differentiate between the two

types of variation are mainly those of Mayr, who states ^

"There is frequently one good indicator which permits us to differentiate
phenotypical from geographic-racial characters. The geographic race has,
in general, a fairly wide range, including a number of different habitats.
The phenotypical Miriant, on the other hand, usually has a rather small
range at any one locality, but crops up again and again wherever the
same or a similar combination of environmental factors occurs".

Each of the four races delimited in this paper fulfills the above
requisites.

One of the first characters studied was size as indicated by the
length of one primary wing. Table 1 shows sample sizes, mean
lengths and standard errors of this character for series of male
cpipsodca from 14 localities. In several cases small series from
several localities were combined into a single series for this analysis.
These were: Miniota, Manitoba, which contains specimens from
Miniota, Beulah, and McCreary, Manitoba; Brewster, Washington,
which contains specimens from Brewster, Black Canyon, and Tif-
fany Lake, Washington; Jesmond, British Columbia, which includes
specimens from Jesmond, Pavilion Lakes, Clinton, and Lilloet,
British Columbia; Osoyoos, British Columbia, which includes speci-
mens from Osoyoos, Summerland, Penticton, Hedly, Keremeos, and
Grand Forks, British Columbia; and Kamloops, British Columbia,
which includes specimens from Kamloops, Vernon, and Salmon Arm,
British Columbia. On entering the locality means on a map, it is
evident that the size of the individuals tends to increase as one
goes from east to west.

On subjecting these means to an analysis of variance a highly
significant F value ( F = 26.23 indicating P < < .01 ) was found.
This would indicate that there is true differentiation between the
localities with respect to wing length. It was estimated that 52.28%
of the total variance of size in the study is attributable to dif-
ferences between localities.

The individual means in the analysis were then analysed by
methods proposed by Tukey.^ These methods deal with establish-
ing significance between means and groups of means in an analysis
of variance, taking into account the fact that the tails of any
normally distributed array of means would appear significantly
different if examined by customary t-tests. A rigorous significance
level of P < .01 was adopted,

3. 1942. Systematics and the origin of species, p. 63.

4. Biometrics 5: 99-114, 1949.

188 The University Science Bulletin

The following groups were found: group 1, subgroup 1 — Hall
Valley, Miniota, Harlan; group 1, subgroup 2 — Didsbury, Tolland,
Banff, Rock Creek, Glacier; group 2 — Wallace, Cranbrook, Jesniond,
Osoyoos, Brewster, Kamloops. The means are arrayed above in
order of increasing size from Hall Valley to Kamloops. The long-
est gap in the array is between groups 1 and 2; the other gap,
between subgroups 1 and 2 of group 1 is shorter, although still
significant. Analyses of variance of the segregates gave the follow-
ing results: group 1, subgroup 1 — F = 4.35, P > .05; group 1, sub-
group 2— F = 1.45, P > .05; group 2— F — 1.94, P > .05. It is thus
seen that there is no significant differentiation between means within
these groups.

Biologically the author would draw the following conclusions
from these data. First that there are two areas of steep gradient
in the general east-west (small-large) size cline; the first around
eastern Alberta and the second in the eastern half of the southern
arm of the area marked "zone of intergradation" on the map ac-
companying this paper. Second that the type series of hopfingeri
was drawn from the area of maximum size of the species, and
that large size is a valid character for this subspecies. It should
be noted, however, that certain other populations whose mean size
was not significantly different from hopfingeri (Brewster) have
been placed in the zone of intergradation on the basis of other
characters. Finally it is the author's opinion that the Hall Valley
specimens represent a specialized high altitude form. In summary,
although the pattern of size variation in the populations studied
supports the author's subspecific arrangement in a general way,
there is no exact concordance.

Before continuing with the discussion of geographic variation in
epipsodea it would be best to explain the map which accompanies
this paper. The map shows a black dot for each record of epipsodea,
without regard for subspecies; the shaded areas show the theoretical
range of each subspecies taking into consideration life zones. The
area marked "zone of intergradation" is an area from which we
have but few specimens with insuflficient data. Most of the speci-
mens which the author has seen from this area are intermediates
between two of the subspecies. The boundaries on the map are
purely arbitrary, as it is doubtful if there are any natural inter-
subspecific boundaries in epipsodea.

The color, definition, and size of the patches are important geo-
graphically varying characters in epipsodea. In specimens from the
Canadian prairies (freemani) a large percentage of the indi-

Distribution and Subspeciation of Erebia

189

EPIPSODEA
REMINGTONI

FREEMANI
HOPFINGERI

'.'^ ZONE OF INTERGRAOATION

"'-'-°'.L

Map, courtesy of the McKinley Publishing Company.
Distribution of Erebia epipsodea Butler.

190 The University Science Bulletin

viduals have light colored patches. By dividing the material
arbitrarily into specimens with light patches and specimens with
dark patches the following results were obtained. Of 65 specimens
from the prairies 70.8% had light patches, while of 445 specimens
from the other populations of epipsodea only 2.5% had light patches
( difference significant at P < < .001 ) . In the prairie specimens
the patches also tended to be more clearly defined, with less suffusion
of ground color around the edges. Of 14 prairie females 12 (85.7%)
had clearly outlined patches on the underside of the secondaries,
while of 47 females from other populations only 4 ( 8.5% ) had clearly
outlined patches ( difference significant at P < < .001 ) , the re-
mainder having the patches suffused with ground color and poorly
defined.

The i^roportionately small size of the patches is a diagnostic
character of E. e. hopfingeri. This will serve in most cases to sepa-
rate hopfingeri from its closest relative epipsodea. However, ma-
terial from Banff, Alberta in the author's collection is very similar
to hopfingeri in this respect, although very close to epipsodea in
all others. Because the type locality of epipsodea may fall in the
Banff area it has been necessary to decide whether this intermediate
material was closer to the Colorado or Washington races. Because
in habitus the Banff material appears more like that from Colorado,
and because it seems preferable to have the nominate subspecies in
Colorado, from which State we have so much material, the author
has included the Banff material in the Colorado subspecies under
the name epipsodea.

The reduction or loss of ocelli is characteristic of £. e. remingtoni.
The only indication that this character may fall into a cline is that
a series of specimens from the Jesmond-Clinton, British Columbia
area shows a maculation very similar to that of remingtoni while
retaining the large, dark appearance of hopfingeri. Whether or not
there is a series of epipsodea populations running northward through
western British Columbia and more or less connecting this southern
population with remingtoni at Atlin is not known. Much of this
area has never been collected, and the author strongly suspects that
such a link may exist. The map has been drawn to show this theoret-
ical link as existing.

In using ground-color as a character the author has been limited
to employing reasonably fresh specimens. Fortunately, where it
was most important, recent specimens have been available. This
is in the comparison of epipsodea to hopfingeri. In this case hop-
fingeri definitely is a darker insect.

Distribution and Subspeciation of Erebia 191

Neither hopfingeri or freemani could be considered as a "strong"
subspecies, and hopfingeri is probably the weaker of the two. How-
ever they both cover considerable geographic area and, in spite of
the great individual variability of the species, are quite readily
recognizable, especially in series. The problem of whether hop-
fingeri was merely a response to the lush environment of the Cas-
cades and neighboring country has been one of no small magnitude.
However the author feels that the combination of three characters,
size, color, and patch size, plus its extensive range strongly indicate
that liopfingeri has a genetic basis.

A DEVELOPMENTAL HYPOTHESIS

Erebia epipsodea appears to be one of our oldest Erehia resi-
dents. Its nearest relative in the Palearctic is probably E. medusa
Schilfermiiller and Dennis, but the two species are extremely dis-
tinct. Erehia vidleri Elwes has perhaps been in North America as
long as, or longer than epipsodea since it is not considered as having
close relatives in Asia. Erebia magdalena Strecker, E. fasciata Butler,
and E. ijoungi Holland if not conspecific with Asiatic forms are at
least very closely related to them [E. erinnyn Warren, E. semo
Grum-Grshimailo ^ and E. dabanensis Erschoff respectively). The
remaining Nearctic Erehia, rossii Curtis, disa Thunberg, discoidalis
Kirby, theano Tauscher and callias Edwards are represented by
subspecies in the Palearctic region.

It is assumed that the present distribution and subspecific pattern
of epipsodea has come about since the last (Wisconsin) glaciation.
On the basis of the present distribution of epipsodea races I would
like to suggest the following pattern of subspeciation. At the
height of the Wisconsin, epipsodea existed in a single race much
like the Colorado subspecies today, somewhat to the south of the
southern end of its present range. With the retreat of the ice,
epipsodea split into three aggregates, one adapted to cold conditions
advancing north along the main chain of the Rockies, and two
adapted to more temperate conditions advancing northward
through what is now the Great Basin and the Great Plains. With
the warming and drying of these regions, epipsodea slowly died
oflF in the southern ends of its range, partially or completely isolating
the three populations in the northern end of the range. Because
of the reduction or cessation of the gene flow between these

5. The status and distribution of the rare and often misidentiiied Erebia avinoffi Holland
is still much in doubt.

192 The University Science Bulletin

populations they were able to evolve into three separate subspecies
in response to their different environments. While hopfin^eri moved
into the Cascades and freemani spread through the Canadian plains,
epipsodea continued to move north through the mountains, eventu-
ally evolving into remingtoni at the extreme northern end of its
range. Finally hopfingeri and freemani moved in and made contact
with the main epipsodea-remingtoni chain, resulting in the dilution
of the subspecies and the formation of intermediate populations.
The extent of gene flow between the four subspecies at the present
time is not known, but since the species is not a strong flier it is
probably rather sluggish.

TABLE 1. — Length of Primary Wing in Males

Series N

Miniota, Manitoba 25

Harlan, Saskatchewan 29

Didsbury, Alberta 18

Banflf, Alberta 30

Glacier National Park, Montana 17

Rock Creek, Colorado 16

Tolland, Colorado 30

Hall Valley, Colorado 30

Wallace, Idaho 17

Brewster, Washington 30

Cranbrook, British Columbia 30

Osoyoos, British Columbia 30

Kamloops, British Columbia 8

Jesmond, British Columbia 15

Mean ± S. E.

( in mm. )

21.16 ±

.125

21.34 ±

.159

21.72 ±

.195

22.03 ■+■

.155

22.29 -t-

.166

22.19 -H

.188

21.87 ±

.104

20.73 ±

.166

22.76 ±

.250

23.47 ±

.157

22.97 -^-

.189

23.33 ±

.200

23.62 ±

.263

23.00 ±

.258

DiSTHIBUTlON AND SUBSPECIATION OF ErEBIA

193

Fig. 2. Upper sides of specimens of Ercbia epipsodea. A, B. E. epipsodea
epipsodea. A. Male, Tolland, Gilpin county, Colorado. B. Female, same
locality. C, D. E. epipsodea remingtoni paratypes. C. Male, Dawson, Yukon
territory. D. Female, same locality. E, F. E. epipsodea hopfingeri. E. Holo-
type male. F. Allotype female. G, H. £. epipsodea freemani. G. Holotype
male, H. Allotype female.

7—6730

194

The University Science Bulletin

B

D

S

Fig. 3. Under sides of specimens shown in figure 2.

THE UNIVERSITY OF KANSAS

SCIENCE BULLETIN

Vol. XXXVII, Pt. I] October 15, 1955 [No. 7

Some Taxonomic and Morphological Studies of Two

Genera of North American Dictyopharidae,

( Fulgoroidea, Homoptera )

By Kathleen C. Doering ^

Abstract: This paper deals with the taxonomy of two genera, Desert a Ball
and Acinaca Ball and Hartzell of the subfamily Orgerinae (Dictyopharidae,
Homoptera). A key to the North American genera of the subfamily
Orgerinae is given. In the genus Deserta four older species have been re-
described and the genitalia of both sexes dissected and described in those species
where material was available. Two species, fuscata and pinturensis are herein
described as new. Two keys for identification of the six species of Deserta are
given, one dealing with characters common to both sexes and one key for
identification of male specimens only, based on characteristics of the male
proctiger and genitalia. A synoptic table of the characteristics of the species
of Deserta is also included. The genus Acinaca still remains, as far as our
present knowledge goes, a monotypic genus. Acinaca liirida Ball and Hartzell
has been redescribed, the genitalia of both sexes described and illustrated and
new distribution records from New Mexico and Arizona are added to the one
state record, namely California.

The characters of most value for classification in these two genera of Or-
gerinae are the shape and length of the head process, the increased size and
development as well as shape of the flaplike extension of the tenth segment,
the anal flap.

The morphological aspects of the paper include mainly a study of the
genitalia and terminal abdominal segments. It is expected that other features
of morphology studied but not specifically described in this paper will con-
tribute to a larger and more comprehensive morphological paper dealing with
the comparative morphology of the entire family.

INTRODUCTION

Since a very excellent and complete bibliography for each species
of the insects studied in this paper has been given by Z. P. Metcalf
in the General Catalogue of the Hemiptera, Fascicle IV, Fulgo-
roidea, Part 8, Dictyopharidae, 1946 it was deemed unnecessary to

1. Contribution number 885 from the Department of Entomology, University of Kansas

(195)

196 The University Science Bulletin

repeat these references. Therefore only snch references that have
appeared since this date have been inckided under each species
heading. At the end of the paper are cited the references which
have been most useful in this study.

Because of the bizarre appearance of some of the Orgerinae a
drawing of the entire insect has been made for at least one species
in each genus. As regards characteristics of taxonomic significance
each species has been extensively illustrated. All drawings have
been made to the same scale for a given character so that difference
in size of drawings is significant.

In making the keys and descriptions certain specific measure-
ments, not easily described in words, have been used. To indicate
these the plan was adopted of placing double letters with a line
drawn between showing the extent of the distance, on the key
diagrams on Plate I. In the keys the measurement was referred to
by stating the letter combination, such as a-a, b-b, etc. Single
letters on the diagrams represent particular points mentioned in a
key or description.

Key to North American Genera of Orgerinae
(Modified from Ball and Hartzell)

1. Callosity behind eye; vertex elongate or angulate 2

No callosity behind eye; head rounded or if elongate, scimitar-
shaped if seen from the side 5

2. ( 1 ) Vertex elongate, produced in front of eye more than one half

its length 3

Vertex angulate, produced in front of eye not more than one

half its length Orgerius Stal

3.(2) Cephalic process truncate at the extremity; apical plate of
process large, pentagonal, its length almost as wide as

eye 4

Cephalic process as seen from the side, beaklike, the apex
obliquely rounding from above, the lower angle slightly
produced; apical plate reduced, its length much less

than width of eye Deserta Ball & Hartzell

4.(3) Cephalic process gradually tapering as seen from both top
and side; apex as seen from side slightly rounded,

Orgamara Ball
Cephalic process parallel-margined; apex as seen from side
squarely truncate and slightly enlarged,

Yucanda Ball & Hartzell
5.(1) Vertex angulate, produced in front of the eyes at least one

third its length 6

Vertex broad and short, scarcely longer than its basal width;
the lateral carina of the front closely margins the eye
clear to its juncture with the lateral margin of the vertex . . 7

Taxonomic and Morphological Studies 197

6. ( 5 ) Head twice the length of the eye; as seen from the side widen-
ing toward the truncate apex; eye and pronotiim widely

separated Acinaca Ball 6< 1 lartzell

Head distincdy less than twice the length of the eyes; as seen
from the side, distinctly narrowing and roundingly trun-
cate; eyes contiguous with pronotinn Ariclia Ball & Hartzell
7.(5) Fore and middle femora and tibiae broadly foliaceous, cari-

nated Loxophora Van Ruzee

Fore and middle femora and tibiae normal 8

8.(7) Lateral carinae of the frontal tablet uniting some distance

below the apex of the head Ticida Uhler

Lateral carinae of the frontal tablet converging but not

uniting before joining the vertex . . Timonidia Ball & Hartzell

Genus Deserta Ball and Hartzell

Deserta Ball, E. D. and Hartzell, A., Ann. Ent. Soc. Amer., vol. 15, pp. 141-142,
1922. Genotype: Orgamara bipunctata Ball, by original designation.

ORIGINAL DESCRIPTION

"This genus resembles Yucanda and Orgamara since the head is
produced into a definite snoutlike process. It differs by having the
apex of this process beaked and curved downward instead of being
truncate and rounded. Vertex with sharply carinate parallel sides
until just before apex where the margins become elevated phlanges
which converge to a triangular point. Genal area extended in front
of eye as a lateral compartment conspicuously separated from the
frons by a prominent carina, behind eye forming an oval callosity.
Frons with conspicuous lateral intermediate carinae separating the
central elevated tablet bearing a median carina down the entire
length, from the two lateral depressed compartments in which occur
numerous round pores. Pronotum collar like, completely encircling
mesonotum anteriorly and with a distinct central tablet set off by
prominent lateral carinae; a median carina equally prominent.
Prominent median and lateral carinae on mesonotum. Tegulae
lacking. The elytra brachypterous, irregularly reticulate. Hind
wings absent. Legs simple; hind tibiae with seven strong spines."

GENERAL MORPHOLOGY

The genus is a difficult genus to classify due to the fact that
specific differences although numerous, are of smaller magnitude
than in many taxonomic groups. Hence there are no obvious char-
acters on which to base keys. Ball (1922) states that most of the
species which he described have been beaten from species of
Artemisia of the western states. The exception, known to date.

198 The Univkhstty Science Bulletin

is Deserta raptoria BaW which was collected from ral)bit brush
(Chrysothaiiinus). This ecological information probably means
that each species of Deserta has a different plant as a host. If these
host plants are mostly of the same genus then the close similarity
of the insect species is not too unexpected.

Each species of Deserta can only be identified by a combination
of numerous small differences which are best followed by studying
the following synoptic chart in combination with the accompanying
illustrations.

For quick identification the length of the head process seems to
be the best character for classification. Male specimens also are a
little more readily identified because in addition to head process
they show slight differences externally in the tenth or anal segment.
A separate key for males has been made based on differences in
this latter structure.

Genitalia for both sexes are described below. But due to the
complexity of the male phallic structures, which seem to have a
double-walled theca which hides most of the aedeagus, the small
size of these genitalia and the difficulty of arranging dissected ma-
terial of this type always in the same or a standard position, compari-
sons are not too reliable.

Chief variations in the male phallus are found in the curious,
large, dorsal membranous lobes ( protrusible glandlike structures ) of
the theca and also in the apices of the aedeagal hooks. The female
genitalia are complex in that they show varying degrees of scleroti-
zation interspersed with membranous areas. The size of the teeth
of the dorsal sclerotized lobe of the first valvula seems to be the
most useful character.

An attempt was made to use the number of lateral spines on the
hind tibia as a specific character. For species identification it
proved unsatisfactory as several species varied between six, seven,
or eight on different specimens. In one case there were nine for one
leg and six for the other leg of the specimen. Numerous cases
occurred of this variability between the two legs of the same speci-
men. In generic identification, however, it may be useful to know
that the spines vary from six to eight on each leg with the most
frequently occurring number being seven.

Female Terminal Abdominal Segments and Genitalia. The eighth
tergum is like the preceding terga. The eighth sternum is mostly
membranous with a broad, extremely short, somewhat sclerotized
median plate, the lateral edge of which forms a heavily sclerotized

Taxonomic and Mohi'iiolocucal Studies 199

slender curved arm attached to the anteromesal angle of each large
first valvifer. These large bilohed valvifers occupy most of the
sternal region but do not meet at midline; each is partially divided
at base by a membranous wedge into two lobelike plates.

The ninth tergum is distinct and forms posteriorly a narrow ring
which runs ventrad beneath the tenth segment.

The tenth segment forms a broad flaplike plate which extends
over the dorsal margins of the third valvulae. The eleventh segment
is tubular, fitting into a hollow area of the tenth segment and bear-
ing an apical hollow closed process called the anal stylus.

The first valvula is broadly attached to the first valvifer and is
bilobed, the ventral lobe being mostly membranous, the dorsal lobe
strongly sclerotized and with the outer margins bearing six teethlike
projections, the two apical ones large and the others diminishing in
size toward the base. A lateral armlike extension or ramus of the
first valvula extends from the anterolateral angle of the first valvula
and articulates to the ninth tergum; ventrad it extends on to the
dorsal lobe of the valvula for approximately half its length as a
sclerotized grooved rod.

The second valvulae are two triangular lobes, united at base and
deeply telescoped cephalad; the dorsal combined surfaces form a
very broad, sclerotized, somewhat triangular plate which is ex-
tended forward at mid-line as a slender arm and whose lateral
apices are articulated to the vertical barlike second valvifer; the
combined ventral bases of the valves form a partially sclerotized
rectangular broad plate lying under the triangular sclerotized dor-
sal plate. Dorsad the second valvifer bears the third valvulae and
also gives off a small sclerotized extension which articulates to the
ninth tergum just behind the articulation of the ramus of the eighth
segment. Ventrad the second valvifer leads directly into the ramus
of the second valvula ( Bonhag and Wick, 1953 ) which appears as
a sclerotized rod along the exposed lateral margin of the valvula
for two thirds its length. This ramus of the 2nd valvula appar-
ently moves on the concave surface of the ramus of the first \alvula.

The third valvula is quadrangular in flattened position, posteriorly
is deeply notched, thus separating a dorsal membranous lobe King
completely protected by the anal plate, from a ventral sclerotized
lobe which partially conceals the first valvulae. The apical margin
of the valve bears a cluster of prominent setae.

Male Terminal Abdominal Segments and Genitalia. The ter-
minal segments of abdomen are strongly telescoped, especially

200 The University Science Bulletin

dorsally so that as viewed from the side only the posterior margin
of the ninth segment shows as a somewhat trianguhir plate. When
dissected out the ninth segment consists of a single sclerotized ring.
Attached to the ninth segment posteroventrad are the large spoon-
shaped freely movable gonostyli, each with a prominent recurved
hook on the dorsal margin and another lateral hook near base.
The tenth and eleventh segment form the proctiger ( See Bonhag
and Wick, 1953 ) . The tenth segment is prominent, at base formed
into a large tube, thus causing the eighth and ninth terga to be
telescoped under the seventh, the ventral apical margin of the
tube extended ventrad and caudad over the phallus as a flaplike
plate (anal flap). The eleventh segment is a reduced ringlike
segment attached by a conjunctiva to the caudal end of the upper
margin of the tube. The eleventh segment bears a single posterior
projection, the anal stylus.

Male Genitalia. Each gonostylus is attached by a conjunctiva
to a slender, transverse, sclerotized bar, from the center of which
arises a perpendicular heavily sclerotized rod (probably the fused
gonocoxopodites ) ; the dorsal terminus of the rod attaches firmly
to the base of the aedeagus at the point where the ejaculatory duct
extends into the aedeagus.

A complex phallic structure is protected dorsally and laterally
by the enlarged proctiger and ventrad by the paired gonostyli.
The aedeagus is greatly reduced and mostly hidden by a double-
sleeved theca, with the outer sleeve shorter than the inner one so
that the latter is exposed where it joins the aedeagus. The theca,
probably formed as an extended fold of the conjunctiva of the
ninth segment, is supported at its base by a sclerotized brace
which is attached to the ninth segment. The apical two thirds of
the dorsal outer sleeve wall of the theca consists of two pairs of
membranous lobes, the outer pair being large and conspicuous, the
inner pair smaller. In normal position these lobes collapse into
accordion-pleatlike folds. When placed in certain fluids, such as
caustic potash, followed by acetic acid they become first extended
and then fold together again. The ventral surface of the outer
thecal sleeve forms a partially sclerotized basal plate; in some
species two well sclerotized bars form a track under each aedeagal
hook and two median membranous, pointed lobes extend caudad.
At base the aedeagus appears as a strongly, sclerotized tube and
apically bears a pair of greatly, elongated sclerotized hooks, whose
curved apices extend beyond the caudal margin of the theca.

Taxonomic and Morphological Studies 201

Key to the Species of the Genus Deserta

( For both sexes )

1. From above head process in front of eye (e-c, plate II, dia-

gram 1 ) approximately three times length of eye ( d-d ) ;
from sideview process extended to a narrowed beaklike
apex due to upper margin bending suddenly at apex of
laterofrontal carina ( x, plate I, diagram 1 ) and acutely
meeting the slightly angled lower margin (y); part of
frons in front of eye (e-e, plant II, diagram 9) longer

than part behind ( f-f ) 2

From above head process not over 2/2 times length at eye;
from side view distance from origin of bend to apex
shorter, upper margin (except bipunctata) gradually
rounding to apex; part of frons in front of eye equal to
or shorter than part behind 3

2. Lateral margins of vertex flaring but not greatly elevated

at apex; apical plate of head process from cephalic view
pentagonal but distinctly wider than long; uniform gray

color, slightly irrorate raptoria (Ball) p. 209

Lateral margins of vertex elevated and suddenly constricted
at apex so lateral compartments of frons are distinct;
apical plate of head process pentagonal but length and
width equal; darker, fuscous dots more numerous,

fuscata n. sp. p. 205

3. ( 1 ) From side view lower apical angle of head process produced

into a rounded knob, the upx^er margin of process flared

and abruptly bent ventrad .. : bipunctata ( Ball ) p. 204

From side view lower margin of head process straight or
almost so, apex not forming a beak or knob, upper margin
not flared, gradually rounded to meet lower margin ... 4
4.(3) Part of frons in front of eye (e-e) shorter than part behind
(f-f); apical plate of head process from cephalic view
pentagonal, its vertical length greater than its width,

obscura (Ball) p. 207
Part of frons in front of eye equal to part behind; apical

plate broader than long 5

5.(4) Laterofrontal carina and genal compartment in front of
eyes conspicuous, vertex flared at apex obscuring frontal
compartments; central tablet of frons occupying most

of apical portion pinturensis n. sp. p. 208

Laterofrontal carina and genal compartment not conspicuous,
lateral margins of vertex constricted anterior to eyes,
narrowing to a pointed apex; central tablet of frons not
distinctly widened at apex obesa ( Ball ) p. 206

202 The University Science Bulletin

Key to the Species of the Genus Deserta
(Key to Males, based on external genitalia)

1. Anal flap with upper surface bulbous at base, greatly pro-
duced caudad; measured from dorsal view its length
at a-a ( plate III, diagram 20 ) more than twice the length
of anal tube at b-b; lateral margin distinctly notched just
before apex and extended ventrad as a distinct phalange, 2
Anal flap shorter, its length from a-a less than twice the
length of anal tube at b-b; lateral margin at most only
shallowly notched and shghtly expanded ventrad 3

2.(1) Anal flap more bulbous at base; apical margin rounded,

ninth sternum without dark patch pinturensis n. sp. p. 208

Anal flap slightly bulbous; apical margin truncate, dark
fuscous patch occupying center of 9th sternum,

bipunctata (Ball) p. 204
3.(1) Anal flap shorter, its length less than IM times length of

anal tube; apical margin of flap concave 4

Anal flap length over 1/2 times the length of anal tube; apical

margin rounded 5

4.(3) Anal flap with upper surface flat, neither bulbous nor con-
cave; apical margins shallowly and broadly concave;
lateral margins parallel-sided, and not expanded,

ohesa (Ball) p. 206
Anal flap with upper surface concave through middle; apical
margin bilobed, lateral margins parallel, then converging

before apex obscura ( Ball ) p. 207

5.(3) Anal flap witli upper surface flat, apex blunt, ventrad margin
somewhat more deeply notched and expanded ventrad as

in bipunctata fiiscata n. sp. p. 205

Anal flap with upper surface slightly concave at base, apex
rounded, ventral margin not notched at side or expanded,

raptoria (Ball) p. 209

The symbols used in table I require special explanation as follows:

Explanation of Table I

Line 2. Species marked " — " have margins of vertex constricting slightly
but without abruptly bending downward at apex, species marked "0" grad-
ually constricting from eyes to apex; those marked "A" have the margins
bulging outward between eyes, constricted through middle, then flaring
broadly at tip.

Line 3. Species marked "0" are only sflghtly elevated.

Line 4. Species marked " — " have the anterior extremity of vertex pointed,
exposing frons on either side.

Line 5. Species marked " — " have the vertex less than one-half as wide as
long at its mid-point.

Line 6. The symbol "0" indicates tliat the suture and compartment is
only moderately conspicuous; symbol " — " that the sutures are not elevated and
prominent and the genal compartment does not even show.

Taxonomic and Morphologic:al Studies

203

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204 The University Science Bulletin

Line 7. Symbol " — " indicates longer than wide; symbol "0?" indicates
mere trifle wider; symbol "0" indicates length and width equal.

Line 9. Species marked " — " have the lower margin curved and apex of
head produced beaklike; species marked "0" have the lower margin slightly
angulate but not produced; species marked "A" have lower margin straight
and apex distinctly produced as a knoblike process.

Line 10. Symbol " — " indicates more slender bod\', its length twice its
width; s\mbol "0" indicates body ovate, with length approximately one and
three-fourths times its width.

Line 1L Species marked " — " have the part of frons in front of eye dis-
tinctly longer than part behind; species marked "0" have the part in front of
eye shorter than the part behind.

Line 12. Symbol "0" indicates lateral carinae converging to a pointed
apex with lateral compartments conspicuous, at tip each one equal to half the
width of median compartment.

Line 13. Symbol " — " indicates that the lateral carinae are parallel and
central tablet before eyes is not so wide as in other species.

Line 14. Symbol " — " indicates length of flap is eiiual to or shghtly shorter
than the tube.

Line 15. Symbol "0" indicates flap is concave at base.

Line 16. Symbol " — " indicates apical margins deeply indented; symbol
"0" indicates margin broadly concave.

Deserta bipiinctata (Ball)

Orgamara bipiinctata. Ball, E. D. Proc. Biol. Soc. Washington, vol. 22: p.
199, 1909; Ball, E. D. and Hartzell, A. Ann. Ent. Soc. Amer., vol. 15, pp.
142-143, 1922.

Size: Length of female 6.5 mm, width, 2.6 mm.; length of male
6.25 mm., width 2.3 mm.

Color: This species is dull fuscous, irrorate with brown and has
darker patches occurring on the wings. A large dark brown spot
occupies the center of the ninth sternal plate.

In the original description Ball ( 1909 ) describes the species as
having "a pair of round white spots just inside the apices of the
abbreviate elytra and a pair of light stripes extending from these
to the apex of the abdomen." In a revised description ( 1922 ) these
markings were not mentioned. Perhaps less emphasis was placed
on these light marks since the same pattern frequently occurs in
most of the other species although in varying degrees of intensity.
In bipiinctata the markings are particularly conspicuous.

Structural Details: This species has a moderately long head
process and the body distinctly irrorate. The proctiger of male is
large with the anal flap (a-a) two and one-half times length of
anal tube (b-b); the anal flap is bulbous at base, each lateral
margin is greatly expanded caudad, and the apical margin is trun-
cate, and deeply notched on each side. Other structural compari-

Taxc^xomic axd Morphological Studies 205

sons can be found in the synopsis chart, Table I, p. 203, and the il-
kistrations on Plates I, II and III.

Types: Originally described from seven specimens, collected
from St. George, Utah, by E. D. Ball, July 24, 1908. One male
specimen from the syntype series is hereby designated lectotype
and one female from the same series as lectoallotype. All types
in the United States National Museimi. Three homotypes, one
male and two females, collected from Leeds, Utah, August 9,
1936, by R. H. Beamer in the Snow Entomological Museum.

Distribution: Leeds and St. George, Utah. Also recorded by
Ball and Hartzell from Ravena, California, probably on Artemisia
trident at a.

Deserta fuscata n. sp.

Size: Length of female 5.85 mm, width 2.2 mm.; length of male
6.3 mm., width 2.2 mm.

Color: General color much darker than other species, the female
being darker than the male. Background yellowish-tan with green-
ish tinge in some areas. The entire body heavily irrorate with dark.
The cells of the tegmina dark fuscous; in the female the mesal half
of each tegmen darker than the lateral half. The female particularly
resembles bipunctata by possessing a pair of conspicuous, elevated
longitudinal cream-colored stripes, bordered mesad in each ab-
dominal segment by a blackish-brown spot.

Structural Detaih: Resembling D. raptoria in size and form,
somewhat longer and more slender with lateral margin of body at
middle less rounding. Length and shape of head process similar to
raptoria, narrowing to a raptorial beak, which is a trifle longer than
in fuscata. Apical plate of head from cephalic view pentagonal, with
length and width equal.

The anal flap of proctiger of male shorter than in raptoria or
bipunctata with its length about one and one-half times the length
of the tube; its ventral margii;i is flared and its caudal margin dis-
tinctly notched on each side.

For other differences consult the chart on p. 203 and the illus-
trations on Plates I, II, III and IV.

Genitalia: Male: The laterodorsal pair of membranous lobes of
the theca are much larger than the mesal pair and both pairs
are deeply pleated and capable of much expansion. The paired
membranous apical lobes of the ventral thecal wall are prominent.
The apices of the aedeagal hooks are bluntly pointed, sharply re-

206 The University Science Bulletin

curved and show a light colored bar and slight constriction at the
base of the curved portion of the hook.

Female: The i^roportion of meinbranization and sclerotization of
the valvulae is shown in the drawing on plate III. The dorsal lobe
of the third valvula is more sclerotized than in raptoria, having sev-
eral folds overhanging more membranous areas. On the dorsal lobe
of the first valvula the second tooth from apex is smaller, the third
tooth more slender and the sixth tooth larger than in raptoria.

Types: Described from holotype male, allotype female, eleven
paratype males, and fourteen paratype females, Cajon Pass, Cali-
fornia, August 6, 1936, collected by R. H. Beamer: additional para-
types identified as follows; one female paratype, Mojave, California,
July 7, 1933, (R. H. Beamer); one female paratype, Monrovia, Cali-
fornia, July 27, 1935, (R. H. Beamer); one female paratype, Los
Angeles County, California, July, ( Coquillett ) . These types are in
the Snow Entomological Museum, University of Kansas.

Distribution: California.

Deserta obesa ( Ball )

Orgamaro obesa. Ball, E. D. Proc. Biol. Soc. Washington, Vol. 22, p. 199,
1909; Ball, E. D. and Hartzell, A. Ann. Ent. Soc. Anier., Vol. 15, pp. 142-
143. 1922.

Size: Length of female 6 mm.; length of male 4.5 mm., width
1.95 mm.

Color: In the orginal description described by Ball as follows:
"Pale straw, slightly washed with orange and sprinkled with smoky
brown, giving the whole insect a dirty orange caste. Cephalic
process pale, sparsely dotted with fuscous, about three pairs of
heavier spots along the outer carinae just below the apex. Clypeus
reddish orange. Pronotum and scutellum pale, sparsely dotted.
Elytra with nervures pale straw, the cells smoky."

Structural Details: This species has a shorter head process than
other species except obscura. From obscura it differs by the process
being proportionately narrower. The proctiger of male is least
developed in the species; length of anal flap is only one and one-
fourth times the length of the anal tube, its dorsal surface is not
bulbous or concave, sides are parallel and apex broadly concave.

Other structural details can be found in the synopsis chart, Table
I, p. 203 and the illustrations on Plates I, II, and III.

Types: Originally described from three specimens from Parowan
and American Fork, Utah, collected by E. D. Ball. One male

Taxonomic and Morphological Studies 207

specimen from the syntype series, Parowan, Utah, July 21, 1908
is hereby designated lectotype. All types in the United States
National Museum.

Distribution and Host Plant Records: Parowan and American
Fork, Utah. Collected from sage brush in July and August (Ball
and Hartzell, 1922).

Desert a ohsciira (Ball)

Orffamara ohsciira. Ball, E. D. Proc. Biol. Soc. Washington, Vol. 22, p. 199,
1909; Ball, E. D. and Hartzell, A. Ann. Ent. Soc. Amer. Vol. 15, pp. 142-
143, 1922.

Size: Length of female 5.1 mm., width 2.5 mm.; length of male
4.8 mm, width 2.1 mm.

Color: General color yellowish gray. Described by Ball as being
"white, finely sprinkled with dark points, giving the whole insect an
ashy gray appearance. Front greenish, the clypeus rusty orange."

Structural Details: Head process short, part in front of eyes
being only about t\\'ice length of the eye, stouter than in ohesa.
Proctiger of male less developed as in ohesa, the anal flap only one
and one-third times the length of the anal tube, its dorsal surface
slightly concave instead of bulbous, the ventral margin scarcely
expanded, the apical margin angulately concave and not notched
at sides.

Other structural details can be found in the synopsis chart. Table
I, p. 203 and the illustrations on Plates I, II, III and IV.

Genitalia: Male: The lateral membranous lobes of the dorsal
thecal wall larger than in raptoria and proportionateK" larger than
middle lobes. The sclerotized bars of the ventral thecal w^all are
roundingly pointed and the paired membranous ventral apical lobes
are longer than in raptoria. The aedeagal hooks taper to a fine point
and have sharp finely pointed serrate subapical margins.

Types: Described from eight specimens, collected by E. D. Ball,
from Nampa, Idaho. These types are in the United States National
Museum.

Two homotypes, collected at Somerville, Utah. July 5, 1931, by
R. H. Beamer. These specimens are in the Snow Entomological
Museum.

Distribution and Host Plants Records: Collected from sagebrush.
Recorded by Ball and Hartzell from the following localities: Nampa,
Idaho; Wells, Nevada; Marysvale and Chadburn's, Utah; Doyle
and Chilcoot, California. Somerville, Utah, is now added to this list.

208 The University Science Bulletin

These authors make the statement that this species seems to be
confined to the state of Nevada and a narrow fringe of adjoining
states and only on the higher levels where sagebrush grows. They
reported that nymphs and adults were collected at lower altitudes
dining July with adults all out by July 20th but that in the mountains
nymphs were still present up to early August and adults up to the
last of the month.

Deserto pinturensis n. sp.

Original Description

Size: Length of female 7 mm., width 3.1 mm., length of male 7
mm., width 2.7 mm.

Color: General color straw yellow, sparsely irrorate with light
brown, especially on vertex between eyes. Mesal half of each teg-
men clouded with fuscous. No sharply contrasting spots or stripes
present on abdomen. The conspicuous brown patch in center of 9th
sternal plate found in bipunctata, obesa, and obscura is lacking in
this species.

Structural Details: Resembling D. bipunctata but head process
from above shorter and stouter, the part in front of eye being ap-
proximately two and one-fifth times the length of the eye; from side
view lower margin of head process straight and apical angle not
produced as in bipunctata; apical i^late of process broader than long.

Proctiger of male long as in bipunctata, the anal flap about two
and one-half times the length of the anal tube, strongly bulbous
at base, ventral margin distinctly expanded and caudal margin
roundingly pointed through middle and deeply notched on each side.

GefiitaJia: Male: The outer pair of lobes of the dorsal thecal
wall are developed as long, tapering sacks which are capable of
much expansion due to the deep accordion-pleatlike folds of their
membranous walls and the helical twisting of their apices. The
ventral thecal wall shows two pairs of membranous apical lobes
instead of the usual one pair. The apices of the thecal hooks are
drawn out into long, fine slightly recurved spines.

Female: The dorsal lobe of the third valvula is less membranous
than in such species as raptoria and the ventral lobe is slightly more
hirsute than in other species. The pattern of the teeth of the dorsal
lobe of the first valvula shows the third tooth smaller than usual,
thus making the third, fourth and fifth teeth approximately equal
in size, and the sixth tooth only slightly smaller than these three.

Taxonomic and Morphological Studies 209

Types: Described from holotype male, allotype female, seven
male paratypes and ei<2;ht female paratypes, all identified by the
following data; Pintura, Utah, Angust 11, 1929, R. H. Beamer
collector: additional paratypes identified as follows; one male
paratype, Order\'ille, Utah, August 27, 1950, H. C. Severin, col-
lector; one female paratype, Lone Pine, California, July 27, 1947,
R. H. Beamer, collector. These types in Snow Entomological Mu-
seum, University of Kansas.

Deserta raptoria Ball

Dcserta raptoria. Ball, E. D. Bull. Brooklyn Ent. Soc. vol. 32, p. 176, 1937.

Size: Length of female 6 mm., width 2.3 mm.; length of male
6.15 mm., width the same.

Color: A uniform, lighter gray species, sparsely irrorated, mark-
ings Jiot so dark as in fuscata. Reticulations and apical portions of
elytra light.

Structural Details: This species is distinguished from other spe-
cies except fuscata by the longer, strongly hooked head process.
From fuscata it is distinguished by its more ovate body shape, lighter
color and apical head plate being wider than long.

The proctiger of male is moderately developed, the anal flap (at
a-a ) approximately 1% X the length of the anal tube ( b-b ) , not bulb-
ous at base, ventral margin only slightly flared, apical margin
broadly rounded, indistinctly notched on each side.

For other structural features see chart I, page 203 and illustra-
tions on plates I, II, III and IV.

Gentalia: Male: The two pairs of membranous lobes from dorsal
wall of theca subequal, with the lateral lobes not greatly larger than
middle lobes and also much smaller than in ohscura. Sclerotized
bars of the ventral thecal wall truncate instead of pointed, the paired
membranous apical lobes prominent but shorter than in ohscura.
The aedeagal hooks roundingly truncate at apex, the apical fourth
of each hook strongly bent back against the basal portion, the sub-
apical serrations of outer margin blunt and inconspicuous.

Female: Variations in the degree of membranization and sclero-
tization of the three pairs of valves is best studied by an examination
of drawings 1, 4, 6, 10, plate III. The chief differences between
species seems to center around the teeth of the dorsal lobe of the
first valvula. In this species the second tooth from apex is larger,
the third tooth broader and the sixth tooth is more minute than in
other species.

210 The University Science Bulletin

Types: Described from holotype female, allotype male and four-
teen paratypes, collected by E. D. Ball, Tehachapi Pass, California,
Jmie 30, 1931. Ball failed to indicate, however, which female was
the type. Therefore one female from the type series is herein desig-
nated as lectotype. One male, same data, is herein designated as
lectoallotype. These types are in the United States National
Museum. A large series of homotypes from Mint Canyon, Cali-
fornia is in the Snow Entomological Museum, University of Kansas.

Distribution and Host Plant Records: California. Collected by
Ball from a heavy stand of Chrysothaninus (Rabbit Bush).

Genus Acinaca Ball and Hartzell

Acinaca Ball, E. D. and Hartzell, A., Ann. Ent. Soc. of Amer., vol. 15, pp. 141-
142, 1922.

Genotype: Acinaca lurida Ball and Hartzell, by original desig-
nation.

generic description

Closely resembling the palaearctic genus Tigrahauda Oshanin but
apparently differing according to Ball and Hartzell by having the
laterofrontal carina of the face curving abruptly upwards and ending
in the dorsum, and vertex not furrowed, whereas in Tigrahauda
the carina runs parallel to ventral margin of the cephalic process and
vertex is furrowed.

Among nearctic genera Acinaca resembles Yucanda Ball and
Hartzell in shape of head process as viewed from the side but is
easily distinguished by the process being strongly compressed and
much shorter than in Yucanda. From all other genera readily
identifies by the scimitar shaped head process.

Body oval, head conspicuous, posterior region behind eyes con-
siderably lenghtened and narrowed into a pseudo-neck, thus widely
separating eye and pronotum. No callosity behind eye. Vertex
narrow, median carina on basal half only, lateral margins carinate,
erect, at base constricted, dilated between eyes, then beginning at
eye angle converging to a pointed apex; a deep transverse bend
across vertex just anterior to eye, which tilts the head process up-
ward. Head process as seen from the side flat, length and width
subequal, apex truncate, whole process strongly inclined upward.
Central tablet of frons narrow, almost linear, tricarinate, carinae
sharply delineated; lateral compartments of frons bearing large,
round pores. Laterofrontal carina joining the vertex at a point
one third the distance from apex of vertex to eye line. Apical plate

Taxonomic and Morphological Studies 211

of liead from cephalic view four-angled, widest near ventral surface
where it is one-fourth as wide as its length.

Pronotum greatly naiTowed anteriorly with middle third angu-
larly produced forward, lateral thirds reduced to an extremely short
collar, numerous round pores occurring on each side of a distinct
median carian; no central tablet or lateral carinae present. Meso-
notum small, an arcuated carina present following posterior line of
pronotum, a short lateral carina on each side branching from this
arc and a faint median carina indicated. Elytra brachyterous, finely
and irregularly reticulated with veins scarcely raised. Pores on
abdominal segments large and extending less than half way to
median line. Femora and tibiae of the first and second pairs of
legs somewhat compressed and edges slightly dilated; hind legs
with three or four lateral spines on each tibia.

Acinaca lurida Ball and Hartzell

(Plates I and III)

Acinaca lurida. Ball, E. D. and Hartzell, A. Ann. Ent. Soc. Amer., vol. 15,
p. 146, 1922.

Size: Length of female 3.5 to 3.8 mm., width 1.9 to 2.2 mm.;
length of male 3.3 mm., width 1.9 mm.

Color: A strikingly bicolored species with all of head except
clypeus and labium, as well as the central third of pronotum and
the anterior third of mesonotum light greenish yellow, rest of body
including tegmina uniformly reddish-brown or bright testaceous.
Anterior and middle femora red-testaceous, striped with pale
yellow lines; hind legs with tips of tibial spines dark. Central
tablet of frons with two narrow dark brown stripes separated by
the pale, elevated median carina. All facial carinae, except latero-
frontals etched in dark brown, including the extreme apical mar-
gins of the head process.

Structural Details: A small, globose species; with prominent
head and reduced thorax. Cephalic process extending in front of
eye a distance equal to length of head. Dorsal plate of head process
triangular, at base abruptly inclined dorsad at an acute angle to
basal part of vertex. From side view head process appears as a
broad scimitar-shaped blade with blunt truncate apex. Frons
narrowed and central tablet between eyes strongly elevated, all
carinae sharply accented. Clypeus with distinct median and
lateral carinae.

Posterior margin of ninth segment of the male projected ventro-
posteriorly as an angulate armlike plate. Proctiger of male moder-

212 The University Scienc;e Bulletin

ately developed, the ventral extension of the tenth segment (anal
Hap) not much longer than dorsal length of the segment; upper
surface of anal flap concave or hollowed almost to apex, its lateral
margins not flared as in Deserta and its apical margin slightly
notched through middle.

Genitalia: Male: Ventral wall of theca bilobed at apex, dorsal
wall ending in two pairs of membranous lobes, which are capable of
a certain amount of pleating and extension. Exposed portion of
thecal hooks are bulbous at base, then suddenly are constricted
just before apex into a slender recurved finger, and on each side
bear a row of five minute short spines.

Female: The ventral lobe of first valvula is triangular, semi-
membranous and fused with first valvifer; the dorsal lobe has its
basal half which is bordered mesad by the rodlike ramus, folded
under the sclerotized toothed portion, the three apical teeth broad,
flat, approximately equal in size, the fourth tooth from apex much
smaller and the basal tooth reduced to a minute spine.

The second valvulae are united at base as in other genera; the
ramus of each valve is projected along the outer edge of the valve
as a broad, heavily sclerotized bar while the basal half of the free
mesal border appears as a membranous triangular area; the rest
of the valve is moderately sclerotized.

The third valvula is quadrangular in flattened position, posteriorly
is deeply notched, thus separating a dorsal fingerlike lobe from
the broader rectangular ventrolateral lobe; the posterior third of
the latter lobe is covered by prominent setae.

Types: Described from three pairs from Mojave, California,
July 30, 1912, E. D. Ball, collector. These types in the National
Museum, Washington, D. C. Two homotypes, male and female,
compared by Ball with the types are in the Snow Entomological
Museum. These specimens bear the following data; Prescott,
Ariz. Aug. 7, 1932, R. H. Beamer collector. Also in the latter
collection are two topotype females, bearing the data, Mojave,
California, July 7, 1933, R. H. Beamer collector.

Distribution: Specimens have been collected from Arizona, Cali-
fornia and New Mexico.

For this study specimens were on hand from Arizona as follows;
Prescott, 5 females and' 6 males, same data as homotypes above,
from Prescott on July 29, 1933, 5 females and 7 males, two females
from Miami, Aug. 6, 1944, one female from Indian Hot Springs,

Taxonomic and Morphological Studies 213

Aug. 6, 1941, one female, Kingman, July 29, 1936, all specimens
collected by R. H. Beamer.

From California specimens were on hand as follows: one male,
Aug. 1, 1935 and one female x\ug. 6, 1936, Cajon Pass, two males and
three females July 18, 1940, Campo, R. H. Beamer collector; in
addition Ball and Hartzell state that they studied specimens from
Ravena, California. In New Mexico a large series of 15 males
and 26 females was collected on July 22, 1936 from Silver City by
R. H. Beamer.

LITERATURE CITED

Ball, E. D.

1937. Some new Fulgoridae from the western United States, Bull. Brook-
lyn Ent. Soc., vol. 32, pp. 171-183, 1937.
1922. Review of the Desert Leaflioppers of the Orgerini. (Rhynchota,
Fulgoridae). Ann. Ent. Soc. Amer., vol. 1.5, pp. 137-152, 1922.

BoNHAG, Phillip, and Wick, James R.

1953. The functional anatomy of the male and female reproductive sys-
tems of the milkweed bug, Oncapeltus fasciatus ( Dallas ) , ( Heterop-
tera, Lygaeidae). Jour. Morph., vol. 93, pp. 177-284, 1953.

Fexnah, R. G.

1945. The external male genitalia of Fulgoroidea. Proc. Ent. Soc. Washing-
ton, vol. 47, pp. 217-226, 1945.

Kramer, Sol

1950. The Morphology and Phxlogeny of Auchenorhynchous Homoptera.
llhnois Biol. Monographs, vol. 20, (no. 4), pp. 1-109, 1950.

Marks Edwin P.

1951. Comparative Studies of the Male Genitalia of the Hemiptera
( Homoptera-Heteroptera ) . Jour. Kansas Ent. Soc. \()1. 24, pp. 134-
141, 1951.

Metcalf, Z. p.

1946. General Catalogue of the Hemiptera, fasc. IV Fulgoroidea; i^art 8
Dictyopharidae, pp. 118-119 and 219, 1946.

214 The University Science Bulletin

PLATE I

1. Lateral view of head process of Deserta raptoria Ball (paratype male).

2. Same view of Deserta fuscata, n. sp. (holotype male).

3. Same view of Deserta obesa (Ball), (male lectotype).

4. Same view of Deserta pinturensis, n. sp. (holotype male).

5. Same view of Deserta obscura (Ball), (homotype male).

6. Same view of Deserta hipunctata (Ball), (male lectotype).

7. Cephalic view of head process of Deserta obscura ( Ball ) .

8. Same of Deserta bipunctata (Ball).

9. Same of Deserta fuscata n. sp.

10. Same of Deserta obesa ( Ball ) .

11. Same of Deserta raptoria Ball.

12. Same of Deserta pinturensis n. sp.

13. Dorsal view of Deserta bipunctata ( Ball ) .

14. Dorsal view of Acinaca lurida Ball and Hartzell.

14a. Ventral view of head of Acinaca lurida Ball and Hartzell.

14b. Lateral view of head of Acinaca lurida Ball and Hartzell.

14c. Dorsal view of head of Acinaca lurida Ball & Hartzell.

14d. Dorsal view of second valvulae of ovipositor of Acinaca lurida Ball &

Hartzell.
14e. Flattened view of first valvula of ovipositor of Acinaca lurida Ball &

Hartzell.
14f. Flattened view of third valvula of ovipositor of Acinaca lurida Ball &

Hartzell.
14g. Dorsal flattened view of male proctiger of Acinaca lurida Ball & Hartzell.
14h. Foreleg of Acinaca lurida Ball & Hartzell.

14i. Cephalic view of head process of Acinaca lurida Ball & Hartzell.
14j. Ventral view of male genitaha of Acinaca lurida Ball & Hartzell.
14k. Lateral view of male genitalia of Acinaca lurida Ball & Hartzell.
141. Lateral view of apical segments and male genitalia of Acinaca lurida Ball

& Hartzell.

Taxonomic and Morphological Studies

215

PLATE I

13 OESERTA
BIPUNCTATA

216 The University Science Bulletin

PLATE II

1. Dorsal view of head of Deserta pinturensis, n. sp. (holotype male).

2. Same view of Deserta ohesa ( Ball ) , ( male lectotype ) .

3. Same view of Deserta hipunctata (Ball), (male lectotype).

4. Same view of Deserta raptoria Ball, (paratype male).

5. Same view of Deserta ohscura (Ball), (homotype male).

6. Same view of Deserta fuscata n. sp. (holotype male).

7. Ventral view of frons of Deserta ohscura (Ball).

8. Same view of Deserta ohesa ( Ball ) .

9. Same view of Deserta raptoria Ball.

10. Same view of Deserta hipunctata (Ball).

11. Same view of Deserta fuscata n. sp.

12. Same view of Deserta pinturensis n. sp.

Taxonomic and Morphological Studies

21'

PLATE II

aOESERTA RAPTORIA ,0 DESERTA BIPUNCTATA

IIOESERTA FUSCATA

12 DESERTA PINTURENSIS

218 The University Science Bulletin

PLATE III

1. Lateral view of apical segments and female genitalia of Deserta raptoria
Ball.

2. Ventral view of terminal abdominal segments and ovipositor of Deserta
raptoria Ball.

3. Lateral view of terminal abdominal segments and ovipositor of Acinaca
lurida B. and H.

4. Flattened view of third valvula of ovipositor of Deserta raptoria Ball.

5. Dorsal view of second valvulae of ovipositor of Deserta pinturensis n, sp.

6. Flattened view of first valvula of ovipositor of Deserta raptoria Ball.

7. Flattened view of third valvula of ovipositor of Deserta fuscata n. sp.

8. Flattened view of first valvula of Deserta pinturensis n. sp.

9. Dorsal view of second valvulae of Deserta fuscata n. sp.

10. Same view of Deserta raptoria Ball.

11. Flattened view of tliird valvula of ovipositor of Deserta pinturensis.

12. Dorsal flattened view of male proctiger of Deserta fuscata n. sp.

13. Lateral view of apical abdominal segments of the male of Deserta fuscata
n. sp.

14. Flattened view of first valvula of ovipositor of Deserta fuscata n. sp.

15. Lateral view of apical abdominal segments of the male of Deserta pin-
turensis n. sp.

16. Dorsal view of male proctiger of Deserta pinturensis n. sp.

17. Lateral view of apical abdominal segments of the male of Deserta ob-
scura (Ball).

18. Same view of Deserta raptoria Ball.

19. Same view of Deserta bipunctata (Ball).

20. Dorsal view of male proctiger of Deserta obscura (Ball).

21. Same view of Deserta raptoria Ball.

22. Lateral view of apical abdominal segments of the male of Deserta obesa
(Ball).

23. Dorsal view of male proctiger of Deserta obesa (Ball).

24. Same view of Deserta bipunctata (Ball).

Taxonomic and Morphological Studies 219

PLATE III

SEGMENT XI
AHHL STYLUS

STERNUM VII

RAMUS OF FIRST
VALVULA

SECOND VALVULA
THIRD VALVULA
-FIRST VALVULA

FIRST VALVIFER

THIRD VALVULA
SEGMENT X

2 DESERTA HAPTORIA

A DESERTA HAPTORIA

FIRST VALVIFER
P DESERTA RAPTORIA
SEGMENT VIII SEGMENT IX SEGMENT XI

-ANAL STYLUS

220 The University Science Bulletin

PLATE IV

1. Dorsal view of ninth segment and male genitalia which have been pulled
forward and laid out flat of Deserta pinturensis n. sp.

2. Lateral view of apical segments and male genitalia of Deserta pinturensis
n. sp.

3. Lateral view of male genitalia, showing membranous dorsal lobes of the
theca distended, of Deserta pinturensis n. sp.

4. Foreleg of Deserta piiiturensis n. sp.

5. Lateral view of male genitalia with dorsal lobes of theca collapsed of
Deserta obscura (Ball).

6. Lateral view of apical segments and male genitalia of Deserta fuscata n. sp.

7. Lateral view of male genitalia of Deserta raptoria Ball.

8. Dorsal view of tenninal abdominal segments of female of Deserta raptoria
Ball.

9. Same view of Deserta pinturensis n. sp.

10. Dorsal view of phallus of Deserta fuscata n. sp.

IL Ventral view of phallus of Deserta obscura (Ball).

12. Same view of Deserta raptoria Ball.

13. Dorsal view of tenninal abdominal segments of female of Deserta fuscata
n. sp.

14. Dorsal view of phallus of Deserta raptoria Ball.

15. Ventral view of phallus of Deserta fuscata n. sp.

16. Same view of Deserta pinturensis n. sp.

I

Taxonomic and Morphological Studies

221

PLATE IV

THE UNIVERSITY OF KANSAS

SCIENCE BULLETIN

Vol. XXXVIL Pt. I] October 15, 1955 [No. 8

Biological Studies and Taxonomic Notes on the Genus

Stwngijlogaster Dahlbom.='' (Hymenoptera:

Tenthredinidae )

Robert E. Beer
University of Kansas, Lawrence

INTRODUCTION

Observations on the biology of insects constitute an important
source of information which is useful not only in establishing
phylogenetic relationships but also in formulating methods of con-
trol for species that are economically important. Contributions
to the knowledge of the bionomics of sawflies are of sporadic
occurrence in the literature; but when such studies are aggre-
gated and perused by a worker interested in the group, it is
possible for him to draw conclusions which might otherwise be
impossible without many years of accumulation of his own data.
This paper constitutes a contribution to the knowledge of the
bionomics of two species of North American sawflies with particular
emphasis upon the developmental stages.

Although considerable work has been done on the classification
of adult sawflies, the association of adults with their develop-
mental stages as well as the association of sexes still remains as
a field of investigation worthy of considerable attention. This
situation is particularly applicable to the western species of saw-
flies, as the published information has been concerned largely
with species found in the eastern United States. Yuasa (1922)
estimated that more than eighty percent of the Nearctic species
of sawflies are yet to be described in their developmental stages,
and the situation has changed little since the appearance of his
paper.

♦Contribution No. 880 from the Department of Entomology, University of Kansas.

(223)

224 The University Science Bulletin

It is a generally accepted hypothesis that the sawflies represent
the most generalized group of the order Hymenoptera. Because
of this status the group has been an important center of interest
to hymenopterists, and it may be found that observations on
the bionomics of species within the group will contribute to the
better understanding of the phylogeny of the Hymenoptera.

As a group the sawflies are most abundant in areas character-
ized by a temperate climate, but they are well represented by
species showing a preference for a subtropical or tropical en-
vironment. For example, in the genus Strongijlogaster from which
two species are treated in this paper, approximately one sixth of
the described species are indigenous to subtropical and tropical
areas, the remainder being recorded only from temperate regions.
With the advent of the use of synthetic organic insecticides in
recent years, field testing has focused the attention of many eco-
nomic entomologists upon the shortcomings of limited knowledge
of the developmental stages of many common insects. It is often
imperative that specific information regarding the life cycle,
ecological preferences and physiological attributes of a pest be
understood in order to direct effectively control measures. Thus
many insect groups originally investigated through morphological
characters of the adults alone are being restudied to supplement
this information with biological data of all stages of development.
In many instances a population of insects, which was believed
originally to be comprised of a single species, subsequently has
been found to have more than one constituent when physiological
characteristics were investigated, a situation which often may be
verified by detailed study of the differences in immature forms.
There are many species of sawflies which are impossible to dis-
tinguish at the present time by morphological characters of the
adults alone, but conspicuous differences may be found in morpho-
logical characters of immature forms, ecological preferences and
physiological limitations.

Due to their rather common occurrence, immature stages of
sawflies have interested naturalists for many years. Many refer-
ences were made to various forms before the time of Linnaeus.
It is probable that more studies would have been made were
it not for the difficulty of rearing and the emphasis which was
given by early workers to studies of adult insects only. Among the
more noteworthy contributions to the literature dealing with im-
mature forms of sawflies are the works of Dahlbom, Dyar, Cam-

Notes ox the Genus Stroxgylogaster 225

eron, Konow, Middleton, MacGillivray, Lepeletier, Westwood and
Yuasa.

Methods and materials: The problems facing an individual in-
terested in the investigation of the biology of sawflies are exceed-
ingly numerous. The most difficult problem encountered is due
to a one year life cycle which is characteristic of the majority of
species. Such a situation prevails in numerous other insect groups,
to be sure, and may often consume more time than one year, but
one can only envy those investigators who are able to keep con-
tinuous cultures going in all stages of development throughout the
year.

Much of the rearing discussed in the following pages was con-
ducted in the laboratory utilizing shell vials, and glass jars of
various sizes as rearing chambers. Glass jars rather than screen
cages were used in order to maintain humid conditions. Food
materials were placed either in a cotton-plugged vial vase within
the rearing chamber or else the base of the stem or leaf was
wrapped in wet cotton before being placed in the chamber in
order to preserve the material in a fresh condition for as long as
possible. Rearing chambers which were used out of doors con-
sisted of one inch and two inch glass tubes as pupation containers
and cloth covered lamp chimneys as rearing chambers. Some
one inch stender dishes and three inch petri dishes were also used
as rearing chambers.

It was found expedient to use shell vials as rearing chambers
for the first few instars of the Strongijlogaster larvae. Later instars
were transferred to larger containers with food material in a vase
as described above. It was found advisable to use cotton tightly
wrapped with cloth as plugs for the vials because last instar larvae
escaped through simple cotton plugs. Attempts were made to rear
Strongijlogaster larvae to maturity in the laboratory by allowing
the larvae to enter old fern twigs on the surface of a layer of soil
in the rearing chamber jar. This method was unsuccessful pri-
marily due to the lack of control of molds which evidently de-
stroyed the larvae during their long periods of quiescence in the
pupation cells.

Outdoor rearing chambers which consist of glass tubes one inch
in diameter inserted into the ground to a depth of two and one-
half inches served as rearing containers too, with one-half inch of
tubing left projecting above the ground surface, and this capped
with brass screen.

8—6730

226 The University Science Bulletin

All specimens upon which descriptions are based are in the
Snow Entomological Museum at the University of Kansas. For
preservation of larvae some specimens were placed directly in
KAAD solution and after twenty-four hours transferred for per-
manent storage into seventy-five percent ethyl alcohol. Other
specimens were killed in alcohol after which they were immersed
in boiling water for one quarter to one half a minute and following
this treatment were returned to seventy percent ethyl alcohol for
permanent storage. Many individual specimens were dissected
and mounted in balsam on glass slides. Clearing and, in some
cases, staining of slide mounted material was found advisable.
Adult specimens were mounted on pins or preserved in alcohol,
the latter specimens being used for slide mounts.

The terminology utilized in descriptions is that followed by
Yuasa (1922) and Ross (1937), and the system of classification
proposed by Ross (1937) has been adopted for use in this paper.

TAXONOMY

The systems of arrangement of the higher categories of sawflies
are almost as numerous as the authors working with the group.
Perhaps the first important comprehensive treatment of the group
was made by Konow ( 1905 ) , the essentials of his arrangement
being that adopted by Ross ( 1937 ) and subsequently by the writer
for the purposes of this paper. In 1898, Ashmead developed a
classification based upon wing venation and relative body propor-
tions. MacGillivray, in 1908, after an exhaustive study of wing
structure proposed a classification based upon this character alone.
Rohwer (1911) followed Konow's groupings fairly closely. Enslin
(1911), although emphatic in his denunciation of Konow's system,
did not refute the most important concepts. A summary of the
subject has been fairly well covered by Yuasa ( 1922 ) and Ross
(1937). It might be appropriate, however, to review the status
of the major categories so that the position of the species included
in this paper may be clarified.

The order Hymenoptera is clearly separable into two distinct
suborders, the Symphyta and the Apocrita, based largely upon the
manner in which the abdomen joins the thorax. These two groups
are identical in definition to the Chalastogastra and Clistogastra,
names which are used by some authors. Ross (1937) recognized
two definite series within the suborder Symphyta which he calls
the Strophandria and the Orthandria, the distinction being the ro-
tation of the male genitalia 180° on the median axis before eclosion

Notes on the Genus Strongylogaster 227

and supporting characters of wing \enation in the Strophandria.
He places the single superfamily Tenthredinoidea in the Stro-
phandria series, with the Siricoidea, Megalodentoidea and Ceph-
oidea in the Orthandria series.

The superfamily Tenthredinoidea is segregated into twelve fam-
ilies on characters of the antennae, spurs of front tibiae and thoracic
structure. The largest of these families is the Tenthredinidae which
is separated into ten subfamilies. It is in the division of this fam-
ily into units that the greatest divergence of opinion occurs among
workers with the group. Among the more recent studies is the
exhaustive work of H. H. Ross ( 1937 ) . His study led him to con-
clude that the hypothetical ancestor for the family had a prepectus,
bidentate mandibles, simple antennae, proximal anal cell of ante-
rior wangs contracted in the middle and separated from the distal
anal cell by a crossvein and the media of the front wings joining
the radius at the fork of the radial sector.

Careful consideration of these characters shows that the sub-
family Selandriinae approaches the primitive condition more closely
than any other group. Ross recognized six genera in the nearctic
assemblage of Selandriinae, including the genus Strongylogaster.
Adults are separated from other members of the subfamily on the
nature of the prepectus. Two species belonging to this genus are
treated at length in this paper, and these represent a very primitive
condition. The larvae of species of Strongylogaster are medium to
large in size, finely but distinctly annulate, green in color with
various characteristic brownish markings sometimes occurring on
the head capsule and pleura. Thoracic legs are well developed
and normal in form. Uropods are setiferous and occur on abdominal
segments two to eight inclusive, and postpedes are present on the
tenth abdominal segment. Specific characters used in larval identifi-
cation include pigmentation of the head capsule, number and
arrangement of setae on the clypeus, labrum, palpi, mandibles,
pleura and uropods, antennal coloration and structure and com-
parative size of thoracic leg segments.

There follows a report of observations made on two species of
Strongylogaster.

Strongylogaster tibialis Cresson

Strongylogaster tibialis Cresson, 1880, Trans. American Ent. Soc, vol. 8, pp.
19-20.

Female: Shiny black. Body covered with dense pile of short
hairs, less conspicuous on dorsum of abdomen. Length 7.5 mm.
Alar expanse 18 mm. Wings hyaline; nervures brown to black.

228 The University Science Bulletin

Head. Shiny black with dense covering of short hairs. Antennae
fihforni; second segment as long as broad; remaining segments de-
creasing in lengdi from third to ninth inclusive. Clypeus honey-
colored with proximal margin and narrow distal margin black,
angularly emarginate, clothed with long, pale hairs. Labrum
rounded truncate, honey-colored, densely pilose. Palpi amber,
faintly infuscate. Mandibles black at base, toothed margin reddish-
brown, pilose, bidentate. Ocelli reddish, ocellar area a triangular
elevation, caudal margin of anterior ocellus with Y-shaped furrow
which joins transverse post ocellar furrow connecting caudal mar-
gins of posterior ocelli. Cervical sclerites shiny black. Thorax:
Tergum of prothorax constricted forming narrow black collar on
meson, dorsolateral angular expansions honey-colored, posteroven-
tral angles shiny black. Tegulae and cenchri honey yellow; remain-
der of thorax shiny black, densely pilose. Prepectus a raised shoulder,
separated from episternmn by a deep furrow. Abdomen: Shiny
black except narrow caudal margins of tergites two through nine
inclusive, pleurites and median and caudal portions of sternites
which are honey-colored. Pile short on dorsum, longer on pleura
and venter. Sheath shiny black, angulate, projecting slightly be-
yond cerci. Legs: Black, except tips of femora, basal third of hind
tibiae, front and middle tibiae and tarsi which are amber; tarsi with
slight infuscation. Tarsal claws bifid, inner tooth short. Wings:
Hyaline with principal nervures and stigma of front wings dark
brown or black, the remaining nervures amber. Anal crossvein
perpendicular. Hind wings with two middle cells.

Male: Length 6.5 mm. Alar expanse 13.5 mm. General descrip-
tion similar in most characters to female; entire venter of abdomen
and all leg segments except hind tibiae and tarsi honey colored.

There is very little variation among individuals of this species,
the pigmentation of the legs being the only character noted which
shows any appreciable variation.

The above descriptions were made from specimens now deposited
in the Snow Entomological Museum, University of Kansas. The
holotype female, from Nevada, is in the collection of the Academy
of Natural Sciences of Philadelphia.

A study of the biology of this species was initiated by the present
author in the spring of 1947 and observations made during this
study are presented in the discussion which follows.

Notes on thk Genus Strongylogaster 229

DISTRIBUTION

Specimens exaininecl In the w liter liave been taken in tlie follow-
ing localities: Washington: Forks (Jnly), Clallam Co. Oregon:
Oney (June). California: Mill Valley (Feb., Mar.) and Lagunitas
(Mar.) Marin Co.; San Francisco Co. (Apr.), (no specific locality
indicated); Pescadero (Apr.). San Mateo Co.; Point Richmond
(Apr.), Contra Costa Co.; Berkeley (Mar., Apr., May) and Oakland
Hills (Apr., May, June), Alameda Co.; Carmel (Feb., Mar., Apr.)
and Sequel Creek (May), Santa Cruz Co.; Lake Tahoe (June) and
Glen Alpine Creek (July), Eldorado Co.; Yosemite (May, July),
Merced Co.; and Giant Forest (July), Tulare Co.

It is quite probable that this species is distributed along the
entire Pacific Coast from British Columbia to San Diego, as no
natural barriers to its migration occur in this area. The host plant,
Pteridium aquilinum, has a continuous distribution throughout this
region, and it seems logical to surmise that the occurrence of the
species would be limited only by the arid, host free areas east of
the coastal ranges of southern California, Sierra Nevada mountains
of central and northern California and Cascades of Oregon and
Washington. Cameron identified a specimen in his collection taken
at Volcan de Chiriqui, Panama, as this species. Since his specimen
was not compared with the type, the identification may have been
incorrect.

This species has been taken at sea level and elevations in the
neighborhood of 5000 feet, occurring at many intermediate levels.
Specimens taken at high elevations as well as those collected in
Oregon and Washington, were obtained later in the spring than
specimens collected from the central coastal region of California.
This, of course, is the expected situation, indicating a later emer-
gence of adults in areas of higher latitude or altitude.

Behavior of Adults

This species, in common with most sawflies, is a very poor flier.
Both sexes, when disturbed from a resting position on a leaf, fre-
quently collide with vegetation while trying to escape. Their flight
is erratic and because of this ineptness, alighting becomes a hit-or-
miss proposition. The outstretched legs snag the vegetation as the
insect flies over it and a crude landing is made which at times
somersaults the individual, but more often the insect appears to
throw itself upon the leaf or branch. Immediately after alighting
the sawflies are very easy to catch by hand as they are not suffi-
ciently oriented to take flight effectively.

230 The University Science Bulletin

The males spend a great deal of time in the vicinity of the host
phnits, making frequent, short, circular flights around a favored
fern frond and returning to alight on the tip where they remain
for a period, later repeating the performance. There appears to
be a definite selection of a specific fern frond for a resting station
from which short flights are made. While resting on the tip of a
frond they are very alert to the flight of other insects, and it is
quite possible that the females are thus spotted. The antennae
are almost continuously in motion with a vibrating or quivering
appearance. The front legs are extended, elevating the anterior
portion of the body. Males are easily distinguished from females
in the field by their smaller size and the narrow, paler abdomen.

Females spend much of their time on the host plant searching
for oviposition sites. They walk down the main stalk of the frond
after alighting on the tip, frequently proceeding as far down as
two inches above the ground level. The frond is then reascended
with some searching along lateral branches. Individuals have been
observed for as long as thirty minutes during which time they
crawled up and down a single frond searching along its main stem
and lateral branches for a place sui*^able for oviposition. Flight is
usually launched from the tip of the frond, although at times the
sawfly will take wing from the upper surface of a branchlet.

When a satisfactory spot is found on the stem, the female pre-
pares for ovipositing by facing downward. The abdomen is then
humped so that the saw penetrates the epidermal tissues of the
stem at or near a right angle. This position is maintained, with
slight movements of the abdomen for about one minute. The ab-
domen is then flattened out against the stem with the hind legs
often extended stiffly in a position at right angles to the body and
not grasping the stem, and with the abdomen thus appressed to
the stem there is a visible quivering for a ten second interval dur-
ing actual egg deposition. Immediately after laying the egg the
insect renews its search for another oviposition site. During much
of the searching the females carry the abdomen in an arched posi-
tion typical of the attitude assumed for the initial thrust described
above. Frequently an incision is started but abandoned before
egg laying is completed.

There appears to be a definite preference shown for depositing
eggs in grooves of the most succulent portions of the stems and
branches, as more than eighty percent of the eggs seen were so
located.

Notes on the Genus Strong ylogasteh 231

The maximum number of eggs laid by one captive female during
a twenty-four hour period was ten, the maximum number laid by
a single female being twenty. This latter number was deposited
by an individual in the following sequence: eight during the sec-
ond twenty-four hours of captivity, nine during the following
twenty-four hours, and three during the ensuing twenty-four hours.
It might be stated here that during the first twenty-four hour period
of captivity of this particular female, the rearing chamber was
furnished with a vase containing two species of ferns, nei*^her of
which was the known host species. No oviposition was attempted
on these ferns.

Most captive females deposited eggs if supplied with suitable
material. On many occasions males and females were caged
together but copulation was never observed. Captive insects
did not at any time exhibit any antagonism toward one another.
This fact was also observed in the field where two females would
frequently encounter each other while walking along the fern stems
searching for oviposition sites. On such occasions they merely
bypassed without abnormal behavior or one or both would re-
verse its direction of progress.

Although feeding by the adults was never observed in the
field, they often were observed to feed on sap exuding from
fronds of P. aquiliniim which were kept in vases in the rearing
chambers. They also fed upon honey water applied in droplets
to the sides of the chamber. Males have been kept alive for
five davs and females for seven davs. Cause of death could
not be determined in any instance. Both sexes when captured
in a net or taken in hand habitually feign death by drawing
the legs to positions closely appressed to the body and curling
the abdomen inward so that the tip almost touches the thorax.
This pretense of death is often maintained for a minute or more
following their original disturbance.

Life History

This species, in connnon with most sawflies, has a one year
life cycle. Adults emerge in April and early May in the San
Francisco Bay region of California at which time the eggs are
laid. Larvae have usually disappeared by early June and remain
in the pupation cells in rotten wood until the following spring.

Egg stage: Eggs are deposited singly and in a random dis-
tribution on the tender, succulent stalks and bases of the branches
of the host plant. They are inserted beneath the thin, hairy,

232 The University Science Bulletin

epidermal layer of the plant which serves as a protective cover-
ing for the egg. The insertion is not made into the tissues under-
lying the epidermis, and the egg displaces the epidermis causing
it to stretch outward over the egg making it clearly visible as
a raised object on the smooth stem surface. The entire egg is
not covered by the parchmentlike epidermis, one end being ex-
posed where the incision slit is made. Occasionally, due to faulty
insertion by the female, eggs are entirely exposed with crumpled
epidermal tissue of the fern clustered at the uppermost end of
the egg. Oviposition is conducted ordinarily so that the egg
is in a position parallel to the stem or branchlet. When deposition
occurs at the thickened area of juncture of a lateral branch with
the main stem of the frond, slight deviations from the parallel
may be noted. Eggs are inserted only on the thickened portions
of the frond, rarely being placed on stems less than one eighth
of an inch in diameter. More than one hundred eggs have been
seen in the field by the writer, and none were placed on the
frond lower than two inches above the ground level.

When first laid, eggs are glistening pale yellow, oval in shape,
averaging 1.1 mm. long and 0.5 mm. in greatest diameter. As
the embryo grows the egg case stretches, increasing in all dimen-
sions. Twenty-four hours after deposition the color is darker
yellow with an oval-shaped opacity near the center and slightly
larger at the caudal end. After forty-eight hours small brown
specks representing the eye spots become visible at the anterior
end of the egg. Color at this time is a yellow-orange. Seventy-
two hours after deposition the egg is about 1.5 mm. long and
0.6 mm. in greatest diameter. Eye spots are clearly visible as
black dots. Mouth parts appear brownish and the alimentary
tract appears as an opaque area more definite in outline toward
the posterior end. Movement of the embryo within the egg case
may be noticeable at this time, particularly evidenced by a ro-
tating motion of the anterior part of the body with the eye spots
rolling from side to side. There is also a frequent withdrawing
of the anterior end of the embryo from its normally appressed
position to the egg case. Hatching occurs ninety-six hours after
deposition, the larva emerging from a dorsal, longitudinal split
which extends from the anterior end half the length of the egg.

The above incubation period is typical of specimens reared in
the laboratory where the temperature remains fairly constant near
72° F. In the field the incubation period is of course variable,

Notes ox the Genus Strongylogaster 233

depending upon prexailing temperatures. Eight days from egg
deposition to hatehing is the longest interval observed by the
writer in the field.

Larva: First instar: Upon emergence from the egg the larva
is very pale yellowish-white, almost transparent with the head
capsule appearing large for the body. The body tapers caudad.
Length 1.8 mm.; width of head capsule 0.35 mm. Ocellarae and
ocularia black; mouth parts brownish.

Second instar: Length 4.0 mm.; width of head capsule 0.5 mm.
Body almost transparent; blood giving green color to dorsum.
Head capsule opaque white with small, fuscous, V-shaped area
on dorsum of vertex and another fuscous masklike patch on front
extending over distance between ocularia. Tibiae and tarsi of
thoracic legs faintly fuscous.

Third instar: Length 9.0 mm.; width of head capsule 1.0 mm.
Body green, paler ventrad; median, dorsal, longitudinal band darker
green, the color being imparted by contents of the dorsal vessel.
Head capsule opaque white, the infuscate mask of second instar
not present, middorsal, dark area on vertex present. Tibiae and
tarsi of thoracic legs fuscous.

Fourth instar: Length 11.5 mm.; width of head capsule 1.5
mm. Color same as previous instar with body slighly darker
green. Head capsule opaque white or faintly amber with narrow,
fuscous, middorsal line on vertex.

Fifth instar: Length 17.0 mm.; width of head capsule 1.9 mm.
Color same as previous instar with head capsule opaque white
with greenish, amber or faintly fuscous tinge and without median-
dorsal, fuscous line on vertex. In the late active phase of this
instar, just prior to excavating the pupal chamber, the body be-
comes reddish in color, particularly in the abdominal region.

Molting of all instars is accomplished after a dorsal, longitudinal
splitting of the head capsule and thorax. The head splits along the
epicranial stem and the larva pulls out of the exuvia by lifting the
head upwards. Exuviae are commonly seen on the host plants.
The caudal portion of the exuvia is cemented in place, prior to the
anterior splitting, by extruding the viscous terminal portions of
the alimentary tract. The larva, after pulling its head free from
the old integument, merely crawls out of its skin, the entire process
being completed in a very short time. A detailed description of the
fifth instar larva follows.

Head subglobose, opaque white with a faint greenish tinge. An-

234 The University Science Bulletin

tennae prominent, five-segmented, broadly conical, with segments
tvt^o through five inclusive black; first segment and narrow distal
margin of second, third and fourth white; all segments with irregu-
larly spaced sensory pits. Antacoriae large, with diameter equal
to diameter of ocularia. Ocellarae shiny black, diameter approxi-
mately one half the diameter of ocularia, situated slightly antero-
dorsad of center of ocularia and bulging somewhat. Clypeus trun-
cate with six strong setae. Labrimi rounded truncate with shallow,
median indentation distally, and with eight prominent setae. Maxilla
with a five jointed palpus, the distal margin of fourth segment armed
with six or seven stout, truncate bristles; one stout bristle on lateral
margin of first, second and third segments, and two smaller bristles
on fifth segment; apical segment without setae; galea flattened,
curved inward to a rather abrupt point, the anterior margin with
numerous small pits; lacinia flattened, margin with eight or nine
broad spines. Labial palpi with three segments. Labium rounded
truncate, bearing broad spines. Mandibles brown, quadridentate;
edges of teeth serrate.

Prothorax with conspicuous, dark brown, winged spiracles, the
spiracles absent on mesothorax and metathorax. Thoracic legs
strong and prominent and normal in form, five-segmented; coxa
nearly as broad as long; femur with apical lobe; tibia cylindrical;
tarsal claw without teeth.

Abdomen elongate, tapering caudad. Uropods on segments two
through eight inclusive and ten. Spiracles winged, located on seg-
ments one through eight inclusive; main trunk of trachea connecting
spiracles visible as a white, threadlike line situated midlaterally.
Abdominal segments seven-annulate with sparse and inconspicuous
setae on annulations two, four and seven. Spiracular opening on
second annulet. Small cylindrical glandubae sparsely distributed,
being present only on setiferous annulets and most abundant on
lateral folds immediately above uropods. All uropods armed with
approximately twenty-two pale and inconspicuous setae.

The larvae, although normally colored green, fade to white in
alcohol. Glandubae were not observed in living specimens as high
magnification is required in order that they be seen. It is quite
possible that they are retractile and that they are extruded when
the larvae are killed in boiling water. They are present on all
segments of the body with exception of the head capsule, and are
more numerous on the abdomen than on the thorax.

Behavior of larva: Immediately after emerging from the egg the

Notes on the Gexus Sthoncylocaster 235

vouiig larva migrates upward on the fern frond until the small,
curled tip of tender new growth is reached. Protection is songht
within these cinls of branchlets, the larva positioning itself usnally
upon the hairy under side of a leaflet within the coil. Here feeding
begins, the larva remaining in or near the curl of tender tip growth
until the fourth instar. When the fourth instar is reached," larvae
may be found on exposed branchlets, reposing on either the upper
or lower surfaces of the leaflets. It is in this and the succeeding
instar that the greatest damage is inflicted upon the plants, as the
larger larvae are voracious feeders.

In all instars the larvae, when disturbed vigorously wave the
abdomen back and forth, throwing it from side to side and up over
the thorax, while maintaining a grasp on the plant with the claws
of the thoracic legs. It is possible that the glandubae function on
such occasions by emitting substances repellant to natural enemies,
but the writer has not been able to detect any peculiar odor or see
any abnormal secretion at such times. Larvae in late instars fre-
quently drop to the ground when disturbed, but in the first three
instars it is extremely difficult to force them to lose their grasp on
the host plant.

After the ultimate stage is entered the larva feeds voraciously for
a few days and then becomes quite ambulatory, the first move being
to descend from the fern frond. On the ground a search begins
for a piece of dead wood suitable for penetration and construction
of a pupal chamber. If not furnished with such material, larvae in
captivity will wander around in the rearing chamber until death
occurs. When a suitable piece of dead wood is found the larva
immediately begins excavating a tubular cavity. The cavity is
entered and plugged with frass, sealing the larva inside a tight-
fitting chamber. This entire process is often completed in as little
as three hours. For the remainder of its life as a larva it remains
fairly quiet, at times moving slightly. Thoracic legs are pulled
inward and are held close against the body. The body becomes
shortened in length and somewhat shriveled in appearance. Larvae
removed from the pupal chamber immediately after the frass plug
is constructed have in every instance failed to construct another
chamber. Often larvae make several beginnings at excavating a
chamber before finally succeeding, the abandoned excavations being
at almost all stages of completion. They have never been seen
to abandon a chamber once the frass plug is completed. The cell
proper is just large enough to accommodate the larva, being ap-

2,36 The University Science Bulletin

proximately 13 mm. long and 3 mm. in diameter. The fras.s ping
varies in length from 5 to 10 mm. Old, dead fern stems from the
previous season's growth which usually abound in areas where the
larvae develop are frequently entered. Such stems are often pene-
trated two or three inches before the plug is constructed, and on
some occasions when this material was used in rearing chambers
more than one larvae entered the same end of the stem forming
pupation cells in line, sealed off from adjacent cells by frass plugs.
Larvae remain green in color and shortened in length as well
as somewhat shriveled in appearance in the chamber until early
spring at which time metamorphosis to the pupal stage occurs.
Because of the prolonged prepupal period, the larvae are very
susceptible to attack by molds which frequently account for their
death. Much difficulty has been encountered by the writer in
rearing this species because of this fact. The most advisable
method of handling larvae when they reach maturity appears to
be the use of aseptic techniques, confining the larvae in sterile vials
plugged with sterile cotton and with a piece of clean cork for con-
struction of the pupal chamber.

Pupa: The pupal stadium is quite brief and its appearance is
that of the typical exarate configuration.

Stimmary of life cycles: The eggs which are inserted on succulent
stems of the host plant, Fteridiiim aqiiilinum, hatch in from four
to eight days. The first instar larva which feeds on the tender
apex of the frond lasts from two to four days. Larvae remain in
the second instar for two to three days, continuing feeding under
the protection of curled leaves at tips of the stems of the host plant.
Duration of the third instar is one to three days. The fourth instar
endures a four to seven day period. The fifth instar, which is the
ultimate stage, is divisible into three distinct phases, the first being
a phase of voracious feeding, the second a phase of searching for
a pupation site and excavating the pupation cell and the third a
phase of quiescence. The feeding phase lasts four or five days
followed by a brief period of boring into dead wood for construc-
tion of the pupal chamber where the larva becomes quiescent for
a period of about ten months. The pupal stage is of short dura-
tion, lasting from ten to fourteen days, after which tlie sawfly
emerges as the winged adult. Thus the insect normally is in the
egg for one week, an active larva for two to five weeks, a quiescent
larva for about ten months, a pupa for two weeks and an adult for
two or three weeks.

Notes on the Genus Strongylogaster 237

Host Plants

Several species of ferns were placed in rearing chambers con-
taining adult insects to test the selection of a host for oviposition
and feeding. Although the sawflies thus confined were observed
to feed upon plant juices exuding from breaks in the epidermis of
many of the ferns, no attempts were made by the females to ovi-
posit on any plant but the recognized host, P. aquiUnum. Larvae
were also confined in rearing chambers provided with various
species of ferns. In all cases, they starved to death without even
sampling the material, unless the known host was included.

Although a limited number of plants was utilized in this survey
for possible hosts, it seems appropriate to conclude that this sawfly
is confined to a single host.

Natural Enemies

More than one hundred larvae, taken from the field when nearly
mature, failed to reveal any parasitism by other insects. It has not
been in the writer's experience actually to witness the attack upon
larvae or adults in the field by insect or arachnid predators or birds,
although some such natural enemies no doubt exist. Old fern
fronds have been taken which contained dead larvae in their pu-
pation chambers, all indications pointing to the fact that their
demise was caused by fungus attack. Diseases of neither adults
nor larvae have thus far been seen.

Economic Importance

Larvae of this species may at times become very destructive to
ornamental plantings of the host plant. Bracken fern is used rather
extensively in the San Francisco Bay region in landscape planning
of rock gardens and hillside terrace gardens. These are often sub-
ject to the attack of this sawfly. The writer has seen large beds
of such plantings completely denuded of foliage with only the
skeletons of branches remaining after the ravages of the sawfly
larvae. A simple inspection of the damaged plants often fails to
reveal the culprits because the protective coloring of the larvae as
well as the habit of the older larvae of dropping to the ground
when disturbed which makes them difficult to see. Their presence
may be rather easily verified, however, by the occurrence of whitish
exuviae on the damaged branchlets. The injury to the fern plants
is often blamed on slugs and snails, and much bait for these pests
has been distributed by home owners in attempts to rid the ferns
of the defoliator.

238 The University Science Bulletin

Control

S. tibialis is easily controlled with certain sprays or dusts. Rote-
none used at two pounds of 4.5 percent material per one hundred
gallons of water will kill larvae effectively. Basic lead arsenate
sprays at two poimds per one hundred gallons of water are also
effective. DDT dusts and sprays are not effective even with spray
applications as high as one pound of actual material per one hundred
gallons of water.

Strongylogaster distans Norton

Strongylogaster distans Norton, 1868, Trans. American Ent. Soc, vol. 2, p. 220.

Female: Length 9.0 mm. Alar expanse 18.5 mm. Head, thorax
and abdomen except tergites of third through ninth inclusive which
are dark amber, black. Entire body clothed with dense pile. Wings
hyaline, minutely but densely setiferous; nervures black to brown;
stigma brown. Head: Black, densely pilose, minutely and uni-
formly punctate. Antennae stout, filiform; segments decreasing
in length from third to ninth; terminal segment bluntly rounded;
last seven segments densely clothed with short, stiff, black bristles;
hairs on first two segments long; second segment as broad as long;
first, one and one-half times as long as broad. Ocelli reddish;
channels at their sides not reaching back of head. Eyes in anterior
aspect slightly converging below, separated by a distance greater
than their height. Clypeus black, emarginate, densely covered with
long hairs. Labrum rounded truncate, amber-colored, with long
hairs, nearly concealed by clypeus. Mandibles black, densely pilose,
bidentate. Palpi pale, slightly fuscous, hairy. Cervical sclerites
shiny black; ventromesal portion V-shaped, not meeting on meson.
Thorax: Black, shining, densely clothed with sericeous pile; pre-
pectus separated from episternum by a furrow. Tegulae, auxilliary
sclerites and pronotum pale; anepimeron and katepimeron black;
cenchri pale, distinct; post tergite shiny black. Abdomen: Broad,
robust, the last three segments tapering to tip, dark honey-colored;
tergites of first two segments black; first tergite divided forming
narrow inverted V on meson; tergites three to nine marked with
black patches medially in triangular pattern pointing caudad but
with posterior margins of tergites always pale; venter honey-colored
with lateral areas of sternites black forming two dark longitudinal
bands on the venter. Cerci broad at base, projecting beyond sheath.
Legs: Honey-colored except coxae, trochanters of middle and hind
legs, proximal portions of middle and hind femora and distal por-

Notes on the Genus Strongylogaster 239

tion of hind tibiae, which are bhick; middle and hind tarsi dusky
brown. Chiws with stout inner tooth one-half as long as outer
tooth. Wings: Hyaline, minutely setiferous on both surfaces; setae
dark brown, more numerous toward tips. Nervures pale amber to
dark brown, stigma dark brown. Lanceolate cell without crossvein,
markedly constricted near base, veins M and Im-cu parallel; M
joining R close to fork of Rs; Rs -j- m distinctly curved; hind wings
with two middle cells.

Mole. Length 9.5 mm. Head, thorax and first two abdominal
tergites shiny black, remainder of abdomen long and slender,
reddish-amber; legs honey-colored with basal margins of coxae
black and hind tibiae distally and hind tarsi slightly infuscate.
Labrum honey-colored, retracted, almost concealed by clypeus.
Other characters as in female.

There is less variation in the male than in the female of this
species. The greatest deviation from the condition considered as
normal in the male is variance in over-all dimensions. A few males
have been examined which have small, black patches on the dorsum
of abdominal segments three, four and five. Females show a great
degree of variation in the amount of black on the abdomen, many
having the abdominal tergites almost entirely black while others
are almost entirely honey-colored in this area.

The above description was made from four female and four
male specimens in the Snow Entomological Museum, University of
Kansas. The holotype, female, from California, collected by A.
Agassiz, is in the collection of the Cambridge Museum of Compara-
tive Zoology.

A biological study of this species was undertaken concomitantly
with the study of the preceding species.

Distribution

Specimens examined by the author have been taken by various
collectors in the following localities: British Columbia: Nanaimo
Biological Station (June). Washington: Forks (July), ClaUain
Co. and Port Angeles (May). Oregon: Homestead Inn, Mt. Hood
(July), Wallawa Mts. (July), Baker Co. and Hood River (July).
California: Carrville (May), Trinity Co.; Shingletown (May),
Shasta Co.; Yorkville (May), Mendocino Co.; Big Bend Mtn. (May),
Butte Co.; Meadow Valley (June), Plumas Co.; El Dorado Co.
(June), (specific location not indicated); Lake Tahoe (June), An-
gora Lake (June) and Fallen Leaf Lake (July), Placer Co.; Muir

240 The University Science Bulletin

Woods (Apr.), Mt. Tamalpais (Apr.), Lagunitas (Apr.), Ross
(Apr.), Dillon Beach and Taylor State Park (Apr.) and Bear Valley
(May), Marin Co.; Bubbs Creek Canyon (July) Fresno Co.; Yo-
semite Valley (May), Merced Co.; Giant Forest (July) and Sequoia
Nat. Park (June), Tulare Co.; Leona Heights (Apr.) and Berkeley
(Mar., Apr., May, June), Alameda Co.; Ben Lomond (June), Santa
Cruz Co.; Carmel (Mar.), Monterey Co.; Bear Valley (June) and
San Bernardino Mts. (July), San Bernardino Co.; and Keen Camp
(June), Riverside Co. These distribution records appear to indicate
that this species ranges throughout the Pacific Coast region, prob-
ably following the same distributional pattern as S. tibialis. This
species has been taken at altitudes ranging from sea level to 9,700
feet, the latter being the Bubbs Creek record for a male specimen
taken in July.

Behavior of Adults

S. distans is very similar to S. tibialis in many aspects of adult
behavior, and it seems redundant to repeat many of the observations
made except where some variance or contrast occurs. Collection
records, as in the preceding species, indicate that protandry is the
rule, a rather common situation in many sawfly species.

Females of this species do not wander about on the stems and
branches as do the females of S. tibialis, but confine their search for
oviposition sites to the tight curls of leaves at the very tip of the
fern frond. In preparing to oviposit, the abdomen is characteris-
tically humped and the egg depositing mechanism is applied at
right angles to the surface of the plant tissues. Egg insertion takes
approximately one minute, after which time the female flies to an-
other frond to repeat the performance. Before alighting upon a
frond of suitable maturity for ovipositing, females very often circle
it several times, the flight circle diminishing in radius each time or
they zigzag in short swoops of descending flight. Egg deposition has
been observed only on the succulent growth of the very tip of the
frond, although first instar larvae have been taken from similar
succulent tips of lateral branches. In captivity a female will ovi-
posit in any tightly coiled portion of the frond. In the field no more
than a single larva has been taken from a single coil of plant
growth, but confined females have deposited as many as five eggs
in one frond tip.

Food of adults of this species appears to be the same as for S.
tibialis, individuals having been observed to take sap from the host
plant and honey water artificially supplied in the rearing chamber.

Notes on the Genus Strongylogaster 241

Males and females have been kept alive in confinement for a maxi-
mum of six days and probably could have been kept longer if a more
elaborate rearing chamber had been prepared.

The maximum number of eggs deposited by a confined female
over a twelve hour period was nine with the total number of eggs
laid by a single female during her lifetime of confinement ( five days
in this instance) being twelve.

Life History

This species requires one year to complete its life cycle. Like
the species discussed previously in this paper, the adults emerge
in early spring in the San Francisco Bay region of California, ap-
pearing concurrently with S. tibialis.

Egg stage: As previously noted, eggs are deposited in the coiled
apices of the main frond or lateral branches of the host plant,
Pteridium aquilinum. Insertion is made between the closely spaced
branchlets comprising the whorl, the saw blade puncturing the
epidermis of the main stem, deposition being made in this spot. The
egg may be parallel or diagonal to the main stem and often is only
slightly covered with the epidermis of the plant. The portion of
the frond where eggs are laid is closely knit with epidermal hairs
making the egg accessible only to the very smallest of arthropods.
It is thus quite effectively protected from natural enemies.

It may be of interest to cite here from notes taken on oviposition
activities of one female sawfly and subsequent inspection of these
activities.

A female of this species was observed ovipositing on the curled
tip of a P. aquilinum on Berkeley campus of the University of Cali-
fornia. The sawfly, after completing the egg laying process was
captured and taken into the laboratory for positive verification of
its identity. The tip of the fern upon which oviposition was
observed was also taken into the laboratory for examinatin, careful
note being taken of the exact spot where the sheath was observed
to have been inserted. In the laboratory a detailed inspection of
the bracken was made, care being taken not to tear or injure the
plant tissues in any way. Although a careful search was made the
egg was not located. On April 25th the bracken was dissected and
the egg was located in one of the uppermost, curled, young leaves,
inserted partially beneath the epidermis of the main axis in the
center of the whorl which itself was only 2 mm. in diameter and
situated 4 or 5 mm, from the tip of the frond. Because of the
numerous epidermal hairs on leaves and stems the egg was well

242 The University Science Bulletin

concealed and protected. At the time of discovery it was 1.5 mm.
long and 1.0 mm. in greatest diameter, a pale yellowish-green in
color. This egg hatched on April 27th.

Every egg sought for was almost as difficult to find as the one
mentioned above.

Shortly after deposition the egg is 1.35 mm. long and 0.8 mm. in
greatest diameter, tapering equally to both ends. Color is glisten-
ing opaque white with a faint greenish tinge. The egg case in-
creases in size by stretching as the embryo develops. The dark
eye spots and mouth parts are the first definite larval characters
to become visible in the embryo. Movement of the embryo within
the transparent egg case is similar to that noted in S. tibialis. At
72° F. constant temperature eggs hatched in five days.

Larva. — First instar: Length 2.5 mm.; width of head capsule
0.5 mm. Early in this instar the larva is completely colorless except
for black ocellarae and ocularia and brownish mouth parts. Later
in the instar the head capsule becomes slightly infuscate and the
green blood imparts a greenish tinge to the dorsum of the thorax
and abdomen.

Second instar: Length 6.5 mm.; width of head capsule 0.8 mm.
Color is the same as in the late phase of the previous instar.

Third instar: Length 7.5 mm.; width of head capsule 1.1 mm.
Color of thorax and abdomen a slightly darker green than previous
instar; head capsule opaque white with a greenish tinge and slightly
infuscate on vertex.

Fourth instar: Length 14 mm.; width of head capsule 1.7 mm.
Color of thorax and abdomen is pale green with the dorsal vessel
imparting the effect of a dark green median dorsal stripe delimited
laterally by conspicuous white stripes. Main tracheal trunk con-
necting spiracles appears as a threadlike, white, lateral line. Head
capsule is opaque white with a greenish tinge and with two, small,
fuscous patches on dorsum of vertex.

Fifth instar: Length 19 mm.; width of head capsule 2.0 mm.
This is the ultimate larval instar and is identical in appearance to
the previous instar. Immediately prior to entering dead wood for
pupation the larva becomes a purplish-brown color.

Larvae cast their skins in the same manner as described for S.
tibialis. The following is a description in detail of an early fifth
instar larva of S. distans:

Head subglobose, opaque white with greenish tinge, infuscate
pattern of irregular shaped, small blotches on dorsum of vertex;

Notes on the Genus Strongylogaster 243

front and genae sparsely setiferous. Ocellarae and ocularia black.
Antennae conical broad at base, five-segmented, all segments
opaque white, segments two, three and four with slitlike sensory
pits, one and five with small, round pits. Antacoriae large, round,
nearly ecjual in diameter to ocularia. Clypeus truncate, with four
stout spines. Labrum rounded truncate with shallow involution at
apex, armed with ten stout spines. Maxilla with five-jointed palpus,
distal margin of fourth segment with five stout bristles, three stout
spines on first and one on anterolateral margin of third segment,
four smaller setae on second segment; galea flattened, curved in-
ward, anterior margin with numerous pits; lacinia flattened, anterior
margin with ten to thirteen elongate and broad spines. Labial
palpi with three segments. Labium rounded truncate. Mandibles
quadridentate, brown.

Thorax with prominent, winged, dark brown spiracles located
midlaterally on prothoracic segment. Spiracles absent on remainder
of tliorax. Thoracic legs normal in form, five-segmented, setiferous,
tarsal claws not toothed.

Abdomen robust, tapered slightly caudad. Uropods occur on
segments two through eight inclusive and ten. Spiracles dark
brown, winged, conspicuous, narrowly ovate; prothoracic spiracle
largest, abdominal spiracles all of same size, situated slightly ven-
trad to a midlateral position on second annulet. Abdominal seg-
ments seven-annulate, setae in sparse, single-row arrangement on
second, fourth and seventh annulations.

Glandubae distributed over most of body, on abdomen occurring
dorsally and dorsolaterally only on setiferous annulations; most
numerous ventrolaterally on thorax and abdomen; none on head
capsule.

Behavior of Larva

Upon emergence from the egg the larva remains in the vicinity
of the apical whorls of leaves of the host clinging to the densely
haired lower surfaces of the leaflets upon which they feed. The
body is usually stretched out flat against the under surface of the
leaflet. Larvae remain on apical leaves of fronds until the fourth
instar is reached, at which time the desire for a much greater vol-
ume of less succulent food material appears to stimulate a descent
of the frond to lateral branches with more mature leaflets. Larvae
of the two species, S. tibialis and S. distons, frequently may be
found side by side on fern branchlets, late instar larvae being
differentiated most readily by the more conspicuous, dorsolateral,

244 The University Science Bulletin

white, longitudinal stripes on the thorax and abdomen of S. distans.
This character is not always reliable in differentiating the two
species, but an examination of the antennae with a hand lens will
disclose either the pale type of S. distans or the black segments of
S. tibialis. Also the presence of two, distinct, fuscous patches on
the dorsum of the vertex characterizes S. distans.

The ultimate larva of S. distans may feed for five or six days
before search is initiated for suitable dead wood material for pen-
etration and construction of the pupation cell. Shortly after feed-
ing is discontinued the color of the larva metamorphoses from
green to purplish-brown. A tunnel is excavated in dead wood by
chewing away the material with the mandibles, pushing the frass
to the exterior with the palpi, thoracic legs and uropods. Often
the tunnel entrance can be detected by the accumulation of frass
beside it or below it on the ground. Extent of penetration of wood
and construction of the pupal chamber within is very similar to
that observed in S. tibialis although the diameter of the tunnel is
slightly larger, in order to accommodate the more robust larva.
Typical cells are almost 16 mm. long and 3.5 mm. in diameter.
As in S. tibialis, the chamber proper, sealed off at the entrance
and with a frass plug, is not lined in any manner with silken threads
or other larval exudate, the quiescent larvae being in naked con-
tact with the wood. However, the sides of the chamber are
smoother than the forward part of the tunnel beyond the plug.
Larvae examined several months after entering wood were green-
ish-brown in color, shrunken in length and somewhat shriveled in
appearance. Very little movement was elicited when disturbed.
If suitable dead wood material is not placed in the rearing chamber
for availability to ultimate larva, death inevitably ensues after a
prolonged ambulatory search has been made for this requirement.
Death is probably due to starvation, for once the larva ceases to
feed it cannot be induced to resume feeding and the continued
wandering about no doubt starves the tissues which normally
would require little nutrition with the larva quiescent in the pupal
chamber. Larvae are very similar to S. tibialis in behavior pat-
terns, having the same characteristic habits of dropping to the
ground or whipping the abdomen back and forth when disturbed.

Pupa: Due to the destruction of cultures by molds, a detailed
study of this stage of development has not been made. However,
this stage has been reared by T. O. Thatcher in the laboratory.
Doctor Thatcher was kindly permitted the inclusion in this paper

Notes on the Genus Strongylogaster 245

of his notes taken on the species: "Three hirvae were collected 31
Max- 1946, 6 mi. N. Willets, California, by F. D. Thatcher as they
were boring into unseasoned redwood lumber. One survived long
enough to bore completely into the wood. It appeared as a
shrunken larva until 8 Nhirch 1947; it was a pupa when examined
11 March 1947. The larvae were green, with a rather felty ap-
pearance similar to Pieris rapae. The pupa was bright blue-green
in head, thorax and legs and brown neutral in abdomen. On 17
March 1947 the thorax, head and legs had turned very dark, almost
black and the abdomen had lost most of its brownish tinge. The
pupa was active when disturbed. The adult emerged on 21 March
1947. It had black head and thorax and red-brown abdomen; the
insect's left wings did not fully extend." This specimen was subse-
quently identified by the present author as a male Strongylogaster
distans Norton.

Summary of life cycles At 72° F. constant temperature eggs
hatched in five days. With this temperature maintained the dura-
tion of instars is as follows: First instar, two to four days; second
instar, one to two days; third instar, two to four days; fourth
instar, four to seven days; fifth instar, four to six days. Time
interval for the fifth instar is for active larva only, the quiescent
phase of this instar being in the neighborhood of ten and one
half months. The true pupal stage endures approximately ten
days. Thus, the species is in the egg stage normally for about one
week, an active larva for two to four weeks, a quiescent larva
for about ten months and a pupa for two weeks. Longevity of
the adult in the field has not been determined, although its life span
is probably two or three weeks judging from collection records.

Host Plants

It seems unnecessary here to repeat data on host plant selection,
for the same information given in the discussion of S. tibialis
applies. The only known host for S. distans is common bracken,
Pteridium qinliniim. Neither oviposition nor larval feeding has
been observed on other species of ferns.

Natural Enemies

The greatest natural enemy of this species, as in S. tibialis,
appears to be fungus attack upon the aestivating and hibernating
larval stage. Insect parasites and arthropod predators have not
to date been observed to interfere with this sawfly in any of

246 The University Scienc:e Bulletin

its cicvelopinental nor mature stages. No diseases other than
fungus attacks noted above have been seen.

Economic Importance

This savvHy occins commonly in association with S. tibialis and
contributes to the damage wrought upon ornamental plantings
of the host plant, and with this consideration it may be desig-
nated as a definite pest. This sawfly as well as the preceding
species, may be considered as being beneficial in the respect
that it destroys the foliage of a plant of common occurrence in
range and pasture lands which is known to be toxic to livestock.
However, the writer has not seen defoliation of stands of bracken
to a degree where the activity of larval feeding appreciably re-
duced the plant stand in a range or pasture area.

The greatest and most conspicuous plant damage appears to
occur in small, isolated stands of bracken, such as are encountered
in ornamental garden plantings. These are subject to prolific
egg deposition.

Control

The same methods as recommended for the previous species
also apply to the control of S. distans. For this information the
reader is referred to the earlier section on control.

ACKNOWLEDGMENT

The author is indebted to Dr. Ray F. Smith of the University
of California for assistance rendered in comparing specimens of
both species included in this paper with the types.

SELECTED BIBLIOGRAPHY
Enslin, E.

1911. Zur systematik der Chalastogastra. Deutsch. Ent. Zeitschr.
1911, pp. 434-439.

ROHWER, S. A.

1911. A classification of the suborder Chalastogastra of the Hymenop-
tera. Proc. Ent. Soc. Washington, vol. 13, pp. 215-224.
Ross, Herbert H.

1937. A generic classification of Nearctic sawllies ( Hymenoptera, Sym-
phyta). Illinois Biological Monographs, vol. 15, no. 2, pp. 1-173.

YuASA, Hachiro

1922. A classification of the larvae of the Tenthredinoidea. Illinois
Biological Monographs, vol. 7, no. 4, pp. 1-172.

Notes ox the Genus Strongylogaster

247

PLATE I

Fig. 1 — Stron^ii]op,aster tibialis Cresson. Fifth instar larva on branch
of host plant, Ptcridium aquilinum.

Fig. 2 — Strongylogaster distans Norton. Fourth instar larva on branch
of host plant, P. aquilinum.

PLATE II

5 ' 9

Larval Characters

Fig. 1 — Strongylogaster disfans. Lateral aspect of anterior portion of
body of fifth instar lars'a.

Fig. 2 — S. tibialis. Third abdominal segment.

Fig. 3 — S. distans. Third abdominal segment.

Fig. 4 — S. distans. Maxilla and labial palpus.

Fig. 5 — S. tibialis. Maxilla.

Fig. 8 — S. distans. Antenna and antacoria.

Fig. 9 — S. tibialis. Antenna and antacoria.

Adult Characters

Fig. 6 — S. distans. Saw.

Fig. 7 — S. tibialis. Saw.

Notes on the Genus Strongylogaster 249

PLATE III

Portion of frond of Pteridium aquilinum showing damage caused by larval
feeding of Strongylogaster tibialis and S. distans.

THE UNIVERSITY OE KANSAS

SCIENCE BULLETIN

Vol. XXXVII, Pt. I] October 15. 1955 [No. 9

Trombiciila giirneyi Ewing and Two New Related
Chigger Mites ( Acarina, Trombiculidae ) ' -

By Richard B. Loomis

Abstract. — The larvii of T romhicula ^unietji Ewing is redescribed. Trom-
hicula hamcrtoni Radford is~donsidered a synonym. This chigger mite is an
inhabitant of eastern woodlands. Trombiciila gunicyi campestris subsp. nov.
is described from larvae taken on grassland vertebrates of the western parts
of Nebraska, Kansas and Oklahoma. Trombiciila kansasensis sp. now, closely
related to Trombiciila giirneyi, is known at present only from Kansas. The
author considers these two species as a separate group, the gurneyi group, of
the genus Trombiciila, sensii Into.

INTRODUCTION

Taxonomic and ecological studies of the larval chigger mites of
the genus Trombiciila from the central United States have disclosed
two closely related species, which cannot be placed in any of the
described subgenera, as defined by Wharton and Fuller (1952:42).
One of the species, Trombiciila gurneyi Ewing, has been found in
the southeastern and central states. Trombiciila liamertoni Rad-
ford, from an unknown locality, but probably from one of the
southeastern states, is considered synonymous with T. gurneyi. A
new subspecies, Trombiciila gurneyi campestris, is described from
the western parts of Nebraska, Kansas, and Oklahoma. The second
species, Trombiciila kansasensis sp. nov., is known from the state
of Kansas. Trombiciila kansasensis has been found in the same
localities as Trombiciila gurneyi and in western Kansas, both spe-
cies were recovered from the same hosts. Trombiciila kansasensis
and T. gurneyi have both been found to occur on reptiles and mam-
mals, while the latter species was also recovered from birds. The

1. Th° studies upon which this paper is based were conducted in part und°r a contract
N6 ori 220 Task Order II, between the Universit>' of Kansas and the Office of Naval Re-
search.

2. Contribution No. 877 of the Department of Entomology, Uni\ersity of Kansas.

(251)

252 The University Science Bulletin

known period of larval activity of T. gtirneyi is spring to late fall
and that of T. kansasensis is summer to late fall.

Since these two species cannot be placed in any of the named
subgenera, I propose that they be considered together as the
"gurneyi" group of the genus Trouibicula. The characters shared
in common include the bifurcate palpal claw, having axial prong
internal; scutum roughly rectangular; sensilla flagelliform with
distal branches; galeal seta nude; two pairs of sternal setae; leg
III with coxa having only one branched seta and without long,
nude, whiplike setae; body setae total 44; and palpal femur with
seta branched.

ACKNOWLEDGMENTS

I wish to thank Dr. E. W. Baker, Insect Identification Section,
United States Department of Agriculture, for the loan of a cotype of
Tromhicula gurneyi and Dr. G. Owen Evans, Custodian of Acarina,
British Museum (Natural History) and Dr. Charles D. Radford
for sending descriptions of critical characters of T. hamcrtoni. For
aid in the assembly of larvae reported below, grateful acknowledg-
ments are extended to the following persons formerly or now at
the University of Kansas: Mr. D. A. Crossley, Jr., Mr. Harold A.
Dundee, Dr. Henry S. Fitch, Mr. Robert B. Finley, Jr., Mr. J. Knox
Jones, Jr., Mr. Ervin H. Kardos, Mr. Louis J. Lipovsky, Mr. Olin
L. Webb and other members of the departments of Entomology and
Zoology. The writer also wishes to thank Mr. Crossley, Dr. Fitch
and Dr. Charles D. Michener of the University of Kansas for care-
fully reading and criticizing the manuscript.

ACCOUNTS OF SPECIES AND SUBSPECIES

The terminology used throughout the paper is that of Wharton,
et ill (1951). All of the measurements are in microns. The de-
scription of each new form is based upon the holotype, with dif-
ferences among the paratypes indicated in parentheses. Unless
otherwise indicated, the specimens listed are in the Snow Ento-
mological Museum, University of Kansas, with the slide numbers
preceded by the initials KU. The number in parentheses in the
listing of specimens denotes the number of larvae examined.

Trombictila gurneyi gurneyi Ewing

Figures 1-5; Table I

Tromhiciihi gurneyi Ewing, Proc. Biol. Soc. Washington, vol. 50, p. 169, Oct.
28, 1937, type from Priest Bridge, Patiixent River, Maryland, host Etimeces
fasciatus, April 24, 1937; Fuller, Zool. Verhandehngen, no. 18, p. 99, 1952;
Wharton and Fuller, Mem. Ent. Soc. Washington, no. 4, pp. 65-66, Dec. 10,
1952.

Some Ciiiccer Mitfs 253

Eutrombicula gurnetji, Ewing, Jour. Washington Acad. Sci., vol. 28, p. 294,
1938; Radford, Parasitolosy, vol. 34, p. 67, 1942.

Trombicula luiuicrtoni Radford, Parasitology, vol. 34, p. 62, fig. 28, 1942,
( tvpe from unknown localitv, host probably Elaphe guttata, from the London
Zoo, March 8, 1939): Radford, Proc. Zool. Soc. London, vol. 116, p. 589,
figs. 15-16, 1946; Radford, IVoc. Zool. Soc. London, vol. 117, p. 275, 1947;
Fuller, Zool. Verhandelingen, no. 18, p. 99, 1952; Whartcm and Fuller,
Mem. Ent. Soc. Washington, no. 4, p. 66, 1952. New synonymy.

Acariscus giimet/i, Ewing, Proc. Ent. Soc. Washington, vol. 45, pp. 57-59,
1943 (in part).

Trombicula (Eutrombicula) gurneiji, Thor and Willmann, Das Tierreich, Lief.
71, p. 291, 1947.

Diagnosis: Larva with body setae total 44, dorsal setae 22, be-
ginning 2-6-6, ventral setae 22, with two pairs of sternal setae; eyes
2/2 on faintly visible ocular plate; scutum roughly rectangular,
wider than long, with sint^ous margin, PL seta nearly twice length
of AL seta, AM seta short, average 27tj. (25-30ij. ); sensilla Hagelli-
form with 8 to 10 distal branches; galeal seta nude; palpal femur
with branched seta, tibia with three nude setae; palpal claw bi-
furcate, axial prong internal; leg I with tarsala * short ( lS[i. ) ; leg III
coxa with 1 seta, without whiplike setae. Similar to Trombicula
gurnciji campestris subsp. nov. and Trombicula kansasensis sp. nov.,
with differences listed under their diagnoses.

Description of hrva: Based upon a cotype, USNM 1262, with
differences among other specimens indicated. Condition of cotype
examined, poor, lacking sensillae and anteromedian seta. Specimen
originally prepared in KOH, thus making many characters indistinct
or obscure.

Body: Cotype 187 by 150 (partially engorged), larvae from
Kansas (unengorged) 170 by 148 and (engorged) 394 by 297;
color yellow to orange (engorged) to whitish (unengorged) in
life. Eyes 2/2, red in life; posterior lens indistinct, ocular plate
faintly outlined.

Dorsal setal formula 2-6-6-4-2-2, total 22; humeral seta measures
46, anterior dorsal seta 40, posterior dorsal seta 35. \^entral setal
formula 2-2-6-4-4-4, total 22, anterior seta measures 35, anterior
ventral seta 28, posterior ventral seta 27. Total body setae 44.

Scutum: Shape roughly rectangular, wider than long, lateral
margin emarginate, posterior margin poorly defined in cotype, but
sHghtly sinuous, punctae small and moderately numerous, postero-
lateral seta nearly twice length of anterolateral seta; bases of sen-
sillae nearly equidistant between anterior and posterior margins,
and nearly in line with mid-points between lateral setae. Sensillae

* See Audy (1952:152) for the use of tarsala and microtarsala in place of spur and
microspur on the tarsi.

254 The Univehsity Science Bulletin

(absent in cotype) flagelliforni, with 8 to 10 branches on distal
two thirds in specimens from Kansas and other central states. For
scn^al measnrements, see Table I.

Gnathosoma: CheHceral blade long and slender, slightly cnrved
with one prominent dorsal tricuspid cap and a prominent ventral
projection; galeal seta nude. Capitular sternum with one pair of
branched setae. Palpal femur with one branched seta; genu with
one nude seta; tibia with three nude setae; tarsus with one stout
basal tarsala (7;ji.), one slender nude seta (subterminala), and live
branched setae; palpal claw bifurcate, prongs nearly equal in
length, inner prong axial, ventral and curved slightly inward.

Legs: Leg I coxa, trochanter and basifemur each with one
branched seta; telofemur with five branched setae; genu with
four branched setae, three genualae and a microgenuala; tibia
with eight branched setae, two tibialae and a microtibiala; tarsus
with approximately twenty branched setae, tarsala ( 13;j. ) , micro-
tarsala, subterminala, parasubterminala and pretarsala. Leg II
coxa and trochanter each with one branched seta; basifemur with
two branched setae; telofemur with four branched setae; genu
with three branched setae and a genuala; tibia with six branched
setae and two tibialae; tarsus with approximately sixteen branched
setae, tarsala ( 13[jl ) , microtarsala and a pretarsala. Leg III coxa
and trochanter each with one branched seta; basifemur with two
branched setae; telofemur with three branched setae; genu with
three branched setae and a genuala; tibia with six branched setae
and a tibiala; tarsus with approximately fourteen branched setae
(no long, nude, whiplike setae).

Taxonomic remarks: Although the writer was not able to examine
the holotype of Tromhicido hamertoni, a description of it was given
to me by Dr. Evans of the British Museum (Natural History).
This description compared favorably in every detail with cotype
of r. giirneyi and with specimens referred to T. g. gurneiji from
the eastern half of the United States. Radford (1942:56) stated
that this type of Trombicida hamertoni was found in a "tube con-
taining the lungs or air sac of a Corn snake (Elaphe guttata)".
The tube was sent to Dr. Radford by Col. A. E. Hamerton, Pa-
thologist of the London Zoo. The locality from which this snake was
secured is unknown. If the identification of the snake was correct,
and the larva actually came from the snake in question, it is prob-
able that this Elaphe guttata was originalh- from one of the south-
eastern states, since the range of Elaphe guttata guttata (Linnaeus)

Some Chiccer Mites 255

is confined to the Gulf and Atlantic coastal states, extending no
farther westward than Louisiana and Kentucky.

GeofirapJiic distribution: Known from Maryland (type locality),
central Florida (Lake County), southern Louisiana (St. Tammany,
and St. Charles Parishes), eastern Texas (Titus-Red River County
line and Travis County), southwestern Arkansas (Little River
County), eastern and central Oklahoma (Haskell and McClain coun-
ties north to Creek County), eastern Kansas (Bourbon County
north to Doniphan County ) and southeastern Nebraska ( Nemaha
County). Intergrades with Tromhicida ^iirneyi campestris subsp.
nov. are known from south-central Kansas (Barber County).

Ecology: Larvae of thig subspecies have been taken in eastern
Kansas on thin black rectangular plastic plates (chigger samplers)
which were placed upon rotten stumps, logs and associated decaying
wood surrounding them. They were found to be abundant in this
habitat in deciduous forests in May to early October. The hosts
of tlie larvae were found to be those species which commonly in-
habit woodlands and regularly come into contact with this larval
habitat. The negative evidence supplied by examination of many
different grassland dwelling hosts and by chigger sampling of other
habitats in the known season of T. g. gurneyi, supports the idea
that decaying wood, especially logs, stumps and standing dead trees,
is the habitat of the free-living stages, and that the hosts are infested
with larvae when they move over these surfaces or rest on them.
The nymphs and adults which have been taken from the center of
large decaying logs in northeastern Texas, in March, support this
theory.

In eastern Kansas, the fi\'e-lined skink, Eumeces jasciatus Lin-
naeus, seems to be the principal host. This lizard is not only
abundant in the known habitat of the larvae, but it has been found
to harbor numerous larvae throughout the season of larval activity.
As many as 100 larvae have been found on a single adult skink.
The larvae attach to these lizards between the toes, and on the
axilla, groin, head and neck under protective scales. The daily and
seasonal activity of both the lizards and the larvae seem to be ap-
proximately the same.

The western part of the range of Eumeces jasciatus is superim-
posed on the map (fig. 1) of Tromhicula gurneyi, and shows the
similarity of ranges between this skink and the eastern subspecies,
T. g. gurneyi. This western part of the range of E. jasciatus also
closely coincides with the extent of larger stands of deciduous wood-

256 The University Science Bulletin

lands. West of this range the woods which are present are usually
limited to a few scattered trees in the stream valleys. This habitat
does not seem suitable for Eumeces fasciatus. The intergrades be-
tween T. g. gurneyi and T. g. campestris however were taken in
this type of habitat. It is probable that T. g. gurncf/i will be found
throughout most of the range inhabited by the five-lined skink.

The usual sites of attachment on mammals seemed to be on the
body, base of the tail and the upper part of the legs. They were
especially common in the areas surrounding the anus and genitalia.
The larvae were attached under the lateral and ventral scales of
snakes.

In northeastern Kansas, the larvae appear in late April and early
May, closely following the appearance of warm spring weather.
They are numerous in the late spring and summer, decreasing in
number on hosts and chigger samplers in late September and early
October, disappearing completely in November. The amount of
moisture present in the fall seems to determine the abundance of
active unengorged larvae and the time of their disappearance. The
lower temperature in October and November also seems to con-
tribute to the decline and the complete disappearance of the larvae.
A single unengorged larva was obtained from nest material of the
wood rat, Neotomo floridona, on February 17, 1952, in central Okla-
homa (McClain County). This larva probably hatched from an
egg in the nest material as it was heated in a Berlese funnel to
recover arthropods. No larvae of Trombiciila gurneyi were re-
covered from five wood rats examined from the same locality in
March and early April, the first larva being taken on April 14, 1952.

Larvae of this subspecies were common on lizards in eastern
Oklahoma in early April (April 8, 1950).

Specimens examined: Total 355 larvae, as follows. Maryland.
Prince Georges Co.: Priest Bridge, Patuxent River, Eumeces fasci-
atus, April 24, 1937, USNM 1262 (cotype). Florida. Lake Co.:
Plymouth, on decayed oak tree, Aug. 21, 1947 (1). Louisiana. St.
Tammany Parish: 8 mi. SE Slidell, April 30, 1954, Eumeces inexpec-
tatus (8) and Sceloporus undulatus (2). St. Charles Parish: 17 mi.
SW New Orleans, Lampropeltis getuhis, May 1, 1954 (2). Texas.
Titus Co. — Red River Co. border: 3 mi. NW Talco, Sulphur River
bottoms, adults from decaying logs, March 26, 1948 (21). Travis
Co.: 5 mi. W Austin, Uta ornata, April 23, 1954 (1). Arkansas.
Little River Co.: 5 mi. SE Ashdown, May 3, 1954, Eumeces fasciatus
(4), Eumeces laticeps (5). Oklahoma. Cherokee Co.: 2 mi. W

Some Chigger Mites 257

Cookson, Sceloporus ttndtilatiis. May 6, 1950 (4). Creek Co.: 7 mi.
SW Tulsa, Parthenes Park, Eitmeces fasciatus, April 8, 1950 (8).
Haskell Co.: 4 mi. S, 3 mi. E Quinton, Eumeces fasciatus, April 9,
1950 (1). McClain Co.: 8 mi. SW Norman, Neotoma foridana,
April 14, 1952 ( 1 ) and nest, Febr. 17, 1952 ( 1 ) . Rogers Co.: 6 mi. E
Catoosa, Eumeces fasciatus, April 8, 1950 (12). Wagoner Co.: 2 mi.
S Okay, Eumeces fasciatus, April 8, 1950 (3).

Kansas. Bourbon Co.: 1 mi. W Ft. Scott, Peromyscus leucopus,
Sept. 4, 1947, (1); 2 mi. E Hiattville, Elaphe ohsoleta. May 10, 1953
(1). Doniphan Co.: 2 mi. N White Cloud, Peromyscus leucopus,
Aug. 21, 1948 (2). Douglas Co.: 1 mi. W Clinton, Eumeces fasci-
atus, May 10, 1952 (2);~1^2 mi. E Eudora, Eumeces fasciatus, April
20, 1950 (5), April 30, 1950 (8) and May 11, 1952 (2); Lawrence,
Elaphe ohsoleta, July 22, 1951 (6) and Scalopiis aquaticus, Nov. 10,
1948 (1); 3/2 mi. E, 4 mi. S Lawrence, May 15, 1948, Eumeces fasci-
atus, (1) and Elaphe ohsoleta, (1); 5 mi. N, 1 mi. E Lawrence,
Univ. Kansas Nat. Hist. Reserv., Chigger samplers, all in 1952, May
25 (5), May 31 (1), June 30 (11), July 16 (2), Aug. 20 (2), Sept.
3 (4), Sept. 17 (10), Sept. 23 (1), Oct. 2(4), -Eumeces fasciatus.
May 9, 1950 (40), May 11, 1950 (8), May 13, 1952 (4), May 16,
1948 (2), May 21, 1952 (5), June 4, 1952 (1), June 20, 1952 (1),
June 28, 1952 (2), June 30, 1952 (2), July 15, 1951 (12), Aug. 16,
1952 (5), -Crotalus horridus, Sept. 2, 1953 (7), -Elaphe ohsoleta.
May 16, 1948 (6), Sept. 10, 1952 (1), -Blarina hrevicauda, July 2,
1952 (1); 20 mi. SW Lawrence, Sciurus niger, Sept. 20, 1953 (1);
Lone Star Lake, Eumeces fasciatus, April 24, 1948, (4); ^2 mi. N
Pleasant Grove, Eumeces fasciatus. May 20, 1949 (55); VA mi. S,
VA mi. E Pleasant Grove, Eumeces fasciatus, April 30, 1950 (7).
Jefferson Co.: 8 mi. N, 1 mi. E Lawrence, Elaphe ohsoleta, June 27,
1952 (2); 2/2 mi. E, 4 mi. N Williamstown, Peromyscus leucopus,
April 29, 1952 (2). Johnson Co.: 1 mi. E Sunflower, Eumeces fasci-
atus, April 30, 1950 (8). Miami Co.: 3 mi. E, 1 mi. S Fontana,
Pigeon Lake area, Elaphe ohsoleta, Oct. 12, 1948 (1), -Eumeces
fasciatus. May 13, 1950, (6), May 26, 1951 (5) and May 31, 1953
(1), -Eumeces laticeps. May 13, 1950 (7), May 26, 1951 (2),
-Sciurus carolincnsis. May 31, 1953 (3), -Chigger Sampler, May 30.
1952 (4), Sept. 9, 1953 (4). Nebraska. Nemaha Co.: 3 mi. S, 1^2
mi. E Peru, Peromyscus leucopus, Oct. 9, 1953 (1).

9—6730

258 The University Sc:ience Bulletin

Tromhicula gurneyi campestris subsp. nov.

Figure 1; Table I

Types: Larvae: Holotype, KU. 3924, and 22 paratypes, KU.
3802-04, 3807-11 and 3925-39, Snow Entomological Museum, Uni-
versity of Kansas, from 13 miles south, 6 miles east of McDonald,
Rawlins County, Kansas, taken from three thirteen-lined ground
squirrels, Citelhis tridecemlineatiis (Mitchell), field no. RL 490807-9,
shot on August 7, 1949, by R. B. Loomis and Robert E. Elbel.

Diagnosis: Larva similar to Trombicula gurneyi gurneyi Ewing,
but differs principally in the longer anteromedian ( AM ) scutal seta,
average 37[;- (35-40[j.), several scutal measurements average larger
with sensilla longer having branches nearer distal end. See table
I for comparison of scutal measurements.

Description of larva: Similar to T. g. gurneyi Ewing, except for
the following characters including measurements of the type series.

Body: Holotype (partially engorged) 300 by 250, (engorged 380
by 331), color, yellow to whitish in life. Eyes 2/2, red in life,
ocular plate faint. Humeral seta measures 43, anterior dorsal seta
36, posterior dorsal seta 35, ventral setae 22, anterior and posterior
sternal setae measure 33, anterior ventral seta 29, posterior ventral
seta 31.

Scutum: Sensilla flagelliform, long with several basal barbs and
10-13 long distal branches. Scutal measurements of holotype:
AW- 66, PW- 76, SB- 31, ASB- 22, PSB- 16, AP- 18, AM- 36, AL-
25, PL- 44, S- 52. See table I for the measurements of additional
specimens.

Gnathosoma: Palpal genual seta occasionally with a single
branch.

Legs: Tarsalae I and II short, 12-13[a.

Taxonomic remarks: The morphological differences between the
two subspecies are slight; however these characters seem to be
constant throughout each of the known geographic ranges. The
specimens considered as intergrades are geographically and mor-
phologically intermediate.

The name campesiris refers to the plains and field habitat of this
subspecies, in contrast to the woodland habitat of the typical sub-
species.

Geographic distribution: Known from southwestern and south-
central Nebraska (Hitchcock and Webster counties), western and
central Kansas (Cheyenne, Rawlins, Wallace, Jewell, Seward and

Some Chigger Mites 259

l^arber counties), eastern Colorado (Yuma County), and western
Oklahoma (Woods, Harper and Harmon counties). Intergrades
with T. g. gurneiji are known from south-central Kansas in Barber
County.

Ecology: Tromhicula gurneyi campestris seems to have ecolog-
ical requirements different from those of typical gurneyi, being
foimd on small mammals, birds and reptiles that inhabit the grass-
lands of the high plains. The burrows and underground nests of
small grassland mammals seem to provide the habitat of the free-
living stages. The nests and surrounding materials maintaining
favorable temperature and humidity, may actually approach the
microhabitat of the easteVn subspecies, which lives in decaying
wood.

Larvae of this subspecies have been taken on 8 species of mam-
mals, 1 species of bird, and 7 species of reptiles. The single bird
record, from the burrowing owl, Speotyto ciinicuJaria, as well as
the other records all support the theory that the free-living larvae in-
habit nests and burrows.

The larvae of this subspecies have been taken on hosts as early
as April 12 in southwestern Oklahoma and were common from
July to October in western and central Kansas.

The usual site of larval attachment upon mammals was observed
to be around the anus and genitalia. Larvae were found under
the anterolateral and ventral scales of snakes and on the legs and
wings of birds.

The larvae considered intergrades between the two subspecies
were obtained in a valley supporting a stand of timber, mostly
elm and cottonwood. Numerous dead trees were present which
afforded the habitat of decaying wood characteristic of the eastern
subspecies. This locality is approximately 5 miles east of the near-
est station of typical campestris.

Specimens examined: Total, 231 larvae, as follows. Colorado.
Yuma Co.: 20 mi. W St. Francis, Kansas, Masticophis flogellum,
Aug. 6, 1949, KU 8911. Nebraska. Hitchcock Co.: 13 mi. S, 2
mi. W Tren^-on, CiteUus tridecemlineatus, Aug. 8, 1949, KU 3909-10
(total 3). Webster Co.: 3 mi. E Guide Rock, Heterodon nasicus,
Aug. 5, 1951, KU 3908. Kansas. Barber Co.: 2 mi. E Aetna, Di-
podomys ordi, Sept. 14, 1953 (6); 4 mi. S Aetna, Neotoma micro-
pus. July 25, 1952, KU 7078, Sept. 15, 1953 (2); -Peromyscus
leucopus, July 26, 1952, KU 3881 and Sylvilagus fJoridanus, Sept.
14, 1953 (1); 4 mi. S, 2 mi. E Aetna, Dipodomys ordi, Aug. 22,

260 The University Science Bulletin

1949, KU 3877-80; 5 mi. S, 3 mi. E Aetna, Masticopltis flageUum,
Sept. 14, 1948, KU 324, 3832-75 and 4; lOJ^ mi. W Hardtner, Me-
lanerpes erythrocephahis, KU 3883-87 and Sijhilagiis fioridanus,
KU 3882, July 26, 1952, considered intergrades between Tromhicida
g. gurneyi and T. g. campestris; 17 mi. W Medicine Lodge, Ari-
zona elegans, Sept. 14, 1948, KU 3876. Cheyenne Co.: 3-5 mi. N
St. Francis, Onychomys leucogaster, July 23, 1948, KU 287-88,
3757; 4 mi. E, 4 mi. N St. Francis, Onychomys leucogaster, KU
289, 3772-86, 3912-17, Total 30, and Peromyscus manicuJatus, KU
3758-71, July 24, 1948; 6 mi. S, 2 mi. E Benkelman, Nebr., Perogna-
thus hispidus, Aug. 7, 1949, KU 3787-89, 3918-23. Jewell Co.: 1 mi.
E, M mi. N Lovewell, Lampropeltis triangidum, July 3, 1951, KU 3831
(3); 4 mi. W Lovewell, Eunieces ohsoletus, July 10, 1951, KU
3830. Rawlins Co.: 3/2 m. S Atwood, Perognathiis hispidus, Aug.
8, 1949, KU 3820; 6 mi. S Atwood, Perognafhus hispidus, July 30,
1948, KU 1592, 3800-01, 4M mi. E Atwood, Crotalus viridis, Aug.
7, 1949, KU 3815-19 (total 8); 9 mi. W, VA mi. S Atwood, Pero-
myscus maniculafus, Aug. 10, 1949, KU 3825; near Beardsley,
Citellus tridecendineattis, July 26 and 28, 1949, KU 3790-99; 9
mi. S Beardsley, Crotalus viridis, KU 3823 (2) and Peromyscus
maniculatus, KU 3824, Aug. 9, 1949; 13 mi. S, 6 mi. E McDonald,
Citellus tridecemlineattis, KU 3802-11, 3821-22, 3924-41, -Cynomys
ludovicianus, KU 7116, and -Speotyto cunicularia, KU 3812-14
(total 5), Aug. 7-8, 1949. ?>eward Co.: 12 mi. NE Liberal, Perog-
nathus hispidus, Sept. 10, 1948, KU 3826-29. Wallace Co.: 3 mi.
W Sharon Springs, Peromyscus manictdatus, July 4, 1949, KU 3750-
56 (total 9).

Oklahoma. Harmon Co.: 8 mi. N, 3 mi. W Vinson, Eumeces
ohsoletus, April 12, 1950, KU 3903. Harper Co.: 10 mi. N Buffalo,
Citellus trideccmlineatus, April 12, 1950, KU 3904-07. Woods Co.:
6 mi. S, 2 mi. W Aetna, Kansas, Heterodon platyrhinos, Oct. 7,
1951, KU 3888-89, -Masticophis flageUum, Aug. 22, 1949, KU 3S91-
3902; I'A mi. S, 5 mi. E Aetna, Dipodomys ordi, July 26, 1952, KU
3890.

Trombicula kansasensis sp. nov.

Figure 6; Table I

Tijpes: Larvae: Holotype, KU slide number 3701 and 16 para-
types, KU 3702-3710 and 3713-3719, taken from two bull snakes,
Pituophis catenifer (Blainville), trapped on October 17, 1950,
by Dr. Henry S. Fitch, and 12 paratypes, KU 3720-30, taken
from a deer mouse, Peromyscus maniculaius (Wagner), field

Some CiirccjER Mites 261

number RL511()20-3, obtained on October 21, 1951, by R. H.
Loomis, all from the University of Kansas Natural History Reser-
vation, 5 miles north and 1 miles east of Lawrence, Douglas County,
Kansas.

Diagnosis: Larva similar to Troinhiciihi gunieiji Ewing, but
differs in having longer anteromedian AM scutal seta, average
47[JL (45-50[j, ); longer tarsalae I and II (16;;.); longer sensilla, average
69\). (64-74;j. ); larger scutum (see table I); and eyes 1/1, without
visible ocular plate.

Description of larva: Similar to Tronihicula gurneiji in most
characters, but differs as-iollows.

Body: Holotype (engorged) 478 by 366, color yellow to whitish
in life; eyes 1/1, red in life, ocular plate absent, diameter of lens
6[A. Humeral seta measures 53, anterior dorsal seta 43, posterior
dorsal seta 53, anterior sternal seta 43, posterior sternal seta 41,
anterior ventral seta 34, posterior ventral seta 40.

Scutimi: Punctate; large (see table I); sensilla long with
several basal barbs and 9-10 long distal branches. Scutal measure-
ments of holotype: AW- 72, PW- 86, SB- 37, ASB- 27, PSB- 18,
AP- 19, AM- 49, AL- 30, PL- 50, S- 68. See table I for scutal
measurements of additional specimens

Gnathosoma: Palpal claw bifurcate, with deep cleft between
longer inner axial prong and smaller outer prong.

Legs: Tarsalae I and II, 16;j-.

Geographic distribution: Known only from western (Wallace
County), south-central (Barber County) and northeastern (Doug-
las County) Kansas.

Ecology: Trombicula kansasensis and T. g. gurneyi have been
taken in the same general area in Douglas County, but they
were found on different hosts collected in different habitats. Trom-
bicula kansasensis and T. g. campestris were both taken from the
same individual hosts in Barber and Wallace counties, which in-
dicates a close similarity of habitat for these latter kinds of larvae.

Larvae were taken from hosts in July to mid-October. The
species seems to be common in open rocky situations, and the
type series was obtained from hosts inhabiting an abandoned lime-
stone quarry at the University of Kansas Natural History Reser-
vation. The hosts from Barber County were taken in sandstone,
gypsum canyons.

The common sites of attachment of the larvae \\'ere under the
anterolateral and ventral scales of snakes, and on the bodies of

262 The University Science Bulletin

the mammalian hosts. The hirvae from the paratype host, Pero-
myscus manic iiJatus, were found along the penis, near the anus
and surrounding the base of the tail.

Specimens examined: Total, 69 larvae, as follows. Kansas.
Barber Co.: 4 mi. S Aetna, Neotoma micropiis, Aug. 22, 1949,
KU 3743, Oct. 6-7, 1951, KU 3735-36, Sept. 15-16, 1953 (5), -Pero-
mijscus leucopiis, Sept. 15, 1953 (3) and Peromtjscus maniculatus,
Oct. 7, 1951, KU 3737. Douglas Co.: 5 mi. N, 1 mi. E Lawrence,
Univ. Kansas Nat. Hist. Reserv., Coluber constrictor, Oct. 5, 1949,
KU 3732, -Crotalus horridus, Sept. 5, 1949, KU 3731, -Pituophis
catenifer, Oct. 17, 1950, KU 3701-19 and (7), -Thamnophis sirtalis,
Aug. 25, 1949, KU 3733, -Peromijscus maniculatus, Oct. 20, 1951,
KU 3720-30 and (1). Wallace Co.: 3 mi. W Sharon Springs,
Peromyscus maniculatus, July 4, 1949, KU 3738-52 and (1).

Distribution of Paratypes

Paratypes of the new species and subspecies will be sent to
the United States National Museum; the Rocky Mountain Labora-
tory; the British Museum (Natural History); the South Aus-
tralian Museum, Adelaide; Dr. G. W. Wharton, University of
Maryland; Dr. Charles D. Radford, Manchester, England; the
Museum National d'Histoire Naturelle, Paris, France; the Army
Medical Service Graduate School, Washington, D. C.; and Dr.
J. R. Audy, Institute for Medical Research, Kuala Lumpur, Malaya.

Literature Cited

Audy, J. R.

1952. Trombiculid Mites Infesting Bats in Malaya with Descriptions of
Three New Species. Bull. Raffles Mus., Singapore, no. 24, pp.
132-159.

Taylor, E. H.

1935. A Taxonomic Study of the Cosmopolitan Scincoid Lizards of the
Genus Eumeces. Univ. Kansas Sci. Bull., vol. 23, pt. 1, pp. 1-643.

Wharton, G. W., D. W. Jenkins, J. M. Brennan, H. S. Fuller, G. M. Kohls
and C. B. Phillip

1951. The terminology and classification of trombiculid mites (Acarina:
Trombiculidae). Jour. Parasit., vol. 37, pp. 13-31.

Wharton, G. W. and H. S. Fuller

1952. A Manual of the Chiggers. Mem. Ent. Soc. Washington, no. 4,
pp. 1-185.

Some Chigger Mites

263

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264 The University Science Bulletin

FIGURE 1

Map showing the known locaUties of Trombicula gurneyi Ewing, in the
central United States, and the known range of the five-Uned skink, Eumeces
fasciatus ( Linnaeus ) , after Taylor ( 1 935 : 205 ) .

Some Chigger Mites

265

FIGURE 1

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97

266 The University Science Bulletin

FIGURES 2-6

Tromhicula giirneyi gurneyi Ewing

Fig. 2. Dorsal aspect of body.

Fig. 3. Ventral aspect of body, showing the nude setae on the legs.

Fig. 4. Gnathosoma.

Fig. 5. Scutum and eyes.

Tromhicula kansasetisis sp. nov.

Fig. 6. Scutum and eye.

Some Chigcer Mites

267

FIGURES 2-6

THE UNIVERSITY OP KANSAS

SCIENCE BULLETIN

Vol. XXXVIL Pt. I] October 15, 1955 [No. 10

Comparison of the Effects of Administration of Propyl-
and lodothiouracil to Pregnant Guinea Pigs ^

Roy R. Peterson 2
Department of Anatomy, University of Kansas, Lawrence

Abstract. From the differences in the response of the thyroid glands of
fetal guinea pigs to propyl- and iodothionracil administered in approximately
equimolar doses to the mother during the last 30 days of pregnancy, it is con-
cluded that, like propylithiouracil, iodothionracil passes the placental barrier.
The goitrogenic activity of iodothionracil is about one thirteenth that of propyl-
thiouracil.

Both drugs cause an increase in litters with one or more dead offspring,
and an increase in fetal mortality. Propylthiouracil was more deleterious than
iodothionracil. Fetal mortality was reduced by administering thyroxine simul-
taneously with propylthiouracil or by giving the propylthiouracil throughout
pregnancy.

It has been reported that, although 5-iodo-2-thiouracil lowers
the rate of oxygen consumption in the rat approximately the same
amount as thiouracil ( Barrett and Gassner, 1951 ) , the iodo- form
does not cause significant goiter formation in this species (Barrett,
Gassner and Dittmer, 1951; Bradley, 1954). Extreme sensitivity
of fetal guinea pigs to goitrogenic agents is indicated by the very
large goiters found in offspring of propylthiouracil-treated mothers
( Peterson, Webster, Rayner and Young, 1952; Peterson and Young,
1952). It was felt that the goitrogenic activity of iodothionracil
could be better investigated in this more sensitive test animal,
provided the drug passed the placenta. Information relating to
possible effects of propyl- and iodothionracil on the outcome of
pregnancy would also be gained from such experiments.

1. This investigation was supported in part by research grant RG-3017 from the Na-
tional Institutes of Health, Public Health Service, and in part by grant 44 from the Uni-
versity of Kansas Research Fund to William C. Young.

2. Research Fellow of the National Institutes of Health, Public Health Service. Present
address: Department of anatomy, Washington University School of Medicine, St. Louis.
Missouri.

(269)

270 The University Science Bulletin

METHODS

Pregnant guinea pigs were treated as follows beginning on the
38th day of gestation: Seven were given O.lf propylthiouracil'^
and 10 were given 0.15% iodothiouracil ^ in the drinking water.
Ten untreated pregnant guinea pigs served as controls. From
measurements of the daily water consumption the average drug
intake was found to be 35 mg or 206 micromols of propylthiouracil
per day, and 70 mg or 276 micromols of iodothiouracil per day.

On the day of delivery the thyroid and pituitary glands of the
offspring were removed, dissected free of excess fat, weighed
on a torsion balance, and fixed in 0.9% saline solution saturated
with mercuric chloride to which formalin was added just before
use in the ratio of 1:10. The thyroids of the mothers were also
removed and fixed at this time. The thyroid glands were em-
bedded in paraffin, sectioned at 10^;., stained with hematoxylin and
eosin, and examined microscopically.

This investigation deals in part with possible effects of treat-
ment with antithyroid drugs on the outcome of pregnancy. In
earlier experiments data pertaining to this facet of the problem
were accumulated. These data, previously unpublished, are pre-
sented here for comparison with the data resulting from the
present experiments. The treatments employed are described in
detail in previous publications (Peterson, Webster, Rayner and
Young, 1952; Peterson and Young, 1952). In the experiments
described in the first of these papers the animals received pro-
pylthiouracil before mating and throughout pregnancy while in
the experiments set forth in the second paper the animals received
propylthiouracil alone or in combination with thyroxine during the
last 30 days of gestation.

All guinea pigs were from our own stock kept under the follow-
ing conditions: The temperature of the laboratory was main-
tained between 70° and 75° F. Food and water were available
at all times. The diet consisted of a mixture of oats and rabbit
pellets supplemented with alfalfa hay and fresh green vegetables
once daily.

3. The propylthiouracil used in this investigation was supplied by Lcderle Laboratories,
Pearl River, New York.

4. The iodothiouracil used was supplied by CIBA Pharmaceutical Products Inc., Sum-
mit, New Jersey.

Effects of Antithyroid Drugs on Guinea Pigs 271

RESULTS

The average birth weight was significantly lower than the
control value in both treated gronps ( Tab. 1 ) . The relative thyroid
weight of offspring from propylthiouracil-treated mothers was, on
the average, 91 times that of the control offspring. In the off-
spring from iodothiouracil-treated mothers the relative thyroid
weight was approximately 7 times that of the controls. The differ-
ences between the thyroid weights of offspring from control and
treated animals were of the same general magnitude as the above
fignres when calculated on' the basis of absolute weights. The
relative pituitary gland weights of the offspring in both treated
groups were significantly heavier than the control value. How-
ever, these differences were not apparent if one considered the
absolute weights.

The thyroid of the untreated female guinea pig at the termination
of pregnancy has follicles of variable size lined with a very flat
epithelium (Fig. 1). After 30 days of treatment with propyl- or
iodothiouracil during pregnancy there was detectable only little
stimulation of the maternal thyroid (Figs. 3 and 5). Most of the
thyroids from treated mothers could not be distinguished from
those of untreated mothers. However, the thyroids of three fe-
males in each of the treated groups showed some stimulation. The
degree of stimulation was slightly greater in the three iodothiouracil-
treated females than in the three propylthiouracil-treated females.

The thyroids of offspring from propylthiouracil-treated mothers
showed the extreme stimulation typical of such glands (Fig. 4).
The follicles, practically devoid of colloid, were lined with a high
columnar epithelium which showed spur formations. The thyroids
of offspring from iodothiouracil-treated mothers were made up of
large follicles containing colloid (Fig. 6). The epithelium was of
variable height, ranging from low cuboidal to columnar types.
There was histological evidence of some stimulation in most of
the glands from this group, but in none of the glands was the
stimulation so great as that caused by propylthiouracil.

Data bearing on the effect of antithyroid drugs on the outcome
of pregnancy are summarized in Table 2. There was a percentage
increase in the number of litters with one or more dead offspring,
and a percentage decrease in the young born alive following pro-
pylthiouracil treatment during the last 30 days of pregnancy.
This effect was greater with propylthiouracil treatment than with
iodothiouracil treatment. Thyroxine administered simultaneously

272 The University Science Bulletin

with propylthiouracil eliminated fetal mortality in the four females
so treated. When propylthiouracil was given throughout pregnancy
the percent of litters with dead offspring was essentially the same
as that of the control group, and the percent of young born alive
was only slightly less than the control value. Litter size was not
affected by any of the treatments.

DISCUSSION

The use of the fetal guinea pig as an assay animal for the goitro-
genic activity of antithyroid drugs is probably a more sensitive
test than any yet devised, because the magnitude of the response is
so great that goitrogens undetectable in the rat give a measurable
response in the fetal guinea pig. In addition the treatment of small
numbers of pregnant guinea pigs yields an adequate sample of target
organs for statistical analysis. The time necessary for a full response
is relatively short, i. e., 30 days. It is obvious that in order to be
assayed in this manner, the drug must be able to pass the placental
barrier of the guinea pig. It has been demonstrated that propyl-
thiouracil passes the placenta (Peterson and Young, 1952). From
the evidence obtained in this study it is judged that iodothiouracil
also passes the placenta of the guinea pig and that its goitrogenic
action on the fetal thyroid is approximately one-thirteenth that of
propylthiouracil.

The effects of these two antithyroid agents on the outcome of
pregnancy are not easy to explain. It is apparent that in the dosages
employed, both propyl- and iodothiouracil increase fetal mortality.
In the case of propylthiouracil at least, this effect can be overcome
by simultaneous administration of thyroxine in appropriate amounts.
Comparable experiments have not yet been carried out using
iodothiouracil. We attribute the fetal mortality observed under
short term treatment to a pharmacological reaction in the fetus
which can be counteracted by exogenous thyroxine. If this is a
pharmacological reaction, why is it not manifested when the anti-
thyroid agent is given throughout pregnancy? No explanation for
this enigma is possible with the present data.

Effects of Antithyroid Drugs on Guinea Pics

273

SUMMARY AND CONCLUSIONS

From the differences in the response of the thyroid glands of
fetal guinea pigs to propyl- and iodothiouracil administered in ap-
proximately equimolar doses to the mother during the last 30 days
of pregnancy, it is concluded that, like propylthiouracil, iodothiou-
racil passes the placental barrier. The goitrogenic activity of
iodothiouracil is about one thirteenth that of propylthiouracil.

Both drugs cause an increase in litters with one or more dead
offspring, and an increase in fetal mortality. Propylthiouracil was
more deleterious than iodothiouracil. Fetal mortality was reduced
by administering thyroxine simultaneously with propylthiouracil or
by giving the propylthiouracil throughout pregnancy.

REFERENCES

Barrett, H. W., and F. X. Gassner

1951. Endocrinology, vol. 48, pp. 197-204.

Barrett, H. W., F. X. Gassner, and K. Dittmer

1951. Endocrinology, vol. 48, pp. 189-196.

Bradley, W. O.

1954. Science, vol. 119, p. 912.
Peterson, R. R., R. C. Webster, B. Rayner, and W. C. Young

1952. Endocrinology, vol. 51, pp. 504-518.

Peterson, R. R., and W. C. Young

1952. Endocrinology, vol. 50, pp. 218-225.

TABLE 1. — Effect of Treatment of the Mother on the Weight of the Thyroid
and Pituitary Glands of the Newborn Guinea Pig

Treatment

Number

of
mothers

Number

of
young

Birth weight

Thyroid

weight

(mg./lOOgm)

Pituitary

weight

(mg/lOO gm)

Untreated

10

30

96.3±2.8i
(30)

31.4rt-1.3l

(30)

6.3±0.24i
(29)

Propylthiouracil .

7

22

71.9±4.0*

(17)

2853.5±572.6*
(13)

7.5dr0.44**
.(8)

Iodothiouracil ....

10

29

63.3±3.0*
(29)

227.5±35.3*

(25)

7.9±0.35*
(15)

' The ^ values are the standard errors of the means. The figures in parentheses are
the number of animals on which the averages are based.

* The difference from the control value is significant at the 1% level.
** The difference from the control value is significant at the 2% level.

274

The University Science Bulletin

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Effects of Amituvkoiu Drugs on Glinla Pics

PLATE 1

275

-'- -T '^^

Fig. 1. Thyroid of an untreated control female. X 200.

Fig. 2. Thyroid of an offspring from an untreated control female. X 200.

Fig. 3. Thyroid of a propylthiouracil-treated female. X 200.

Fig. 4. Thyroid of an offspring from a propylthiouracil-treated female.

X 200.
Fig. 5. Thyroid of an iodothiouracil-treated female. X 200.
Fig. 6. Thryoid of an offspring from an iodothiouracil-treated female.

X 200.

THE UNIVERSITY OP KANSAS

SCIENCE BULLETIN

Vol. XXXVII, Pt. I] October 15, 1955 [No. 11

A Taxonomic Revision of the Family Gelastocoridae

( Hemiptera )

BY
Edward L. Todd *

Abstract. — This paper constitutes a revision of the family Gelastocoridae
(Hemiptera). The nature of the paper is primarily taxonomic, but infor-
mation on the biology of some of the species of the family is included.

Prior to this work, the subfamily Gelastocorinae contained two genera and
the subfamily Nerthrinae contained six genera. In this paper there is considered
to be but a single genus in each of the two subfamilies. Prior to this work,
seventy-one species were recognized in this family. Of this number, fifteen
are placed in synonomy, one species reduced to subspecific status, one species
renamed, and three varieties or subspecies are raised to specific status. Sixteen
new species are described in this paper and all other species a\ailable to the
worker are redescribed.

The new species are: Gelastocoris viridis (Mexico, Guatemala), Nerthra
huenoi (Brazil), N. iciUiamsi (Brazil), N. corudis (Guatemala), IV. munni
(Mexico), N. hungerfordi (Central America), N. hracchialis (Mexico), A'.
gurneyi (Bougainville Island), N. omani (Guadacanal Island), N. niidata
(Australia), N. macrostijla (Solomon Islands), N. rohusta (New Guinea), N.
tasmaniensis (Tasmania), .V. walkeri (AustraHa), N. simiosa (AustraHa).

Genera placed in the synonomy of Nerthra Say are: Mommyx Laporte,
Phintias Stal, Matinus Stal, Peltopterus Guerin-Meneville, and Glossoaspis
Blatchley. In the Gelastocorinae, Montandonitis Melin is placed in the
synonomy of Gelastocoris Kirkaldy.

Nertlira lenehrosa is the new name used for Mononijx ohscurus Melin
which is a homonym of Mononyx ohscurus Stal.

Gelastocoris andinus peruensis Melin, Mononyx amplicollis var. ecuador-
ensis Melin and Mononyx fuscipes var. rudis Melin are raised to specific rank.

Gelastocoris variegatus (Guerin-Meneville) is treated in this paper as a
subspecies of G. oculatus (Fabricius).

The new synonomy at the specific level is as follows: Gelastocoris cucul-
latus Martin, a synonym of G. hungerfordi Melin; G. duplicatus Martin, a
synonym of G. major Montandon; G. calif orniensis Melin, a synonym of G.

* Currently with Entomological research Branch, Agricultural Research Service, U. S.
Department of Agriculture, Washington, D. C.

(277)

278 The University Science Bulletin

oculatus ociilatus (Fabricius); G. bergi DeCarlo, G. vianai DeCarlo, G.
bolivianus DeCarlo and G. pamguayensis DeCarlo, synonyms of G. nebulosus
(Giu'rin-Mrneville); G. luarfiiiczi DeCarlo, a synonym ol G. ftiscus Martin,
Montcindonius willineri DeCarlo, M. mansosotoi DeCarlo and A/, bridawlli
DeCarlo, synonyms of G. angulatus (Melin); Mononyx niger Melin, a
synonym of N. ranina ( Herrich-Sehjiff er ) ; A/, stali Melin, a synonym of N.
grandicoUis (Gennar); M. projcctus Distant, a synonym of N. indica (Atkin-
son); Ghssoaspis brunneti Blatchley, a synonym of N . rugosa (Desjardins).
Including this paper, there are seventy-four species and one subspecies
recognized in this family.

TABLE OF CONTENTS

PAGE

Introduction 280

Acknowledgments 281

Biology and Ecology 282

Phylogeny 285

Taxonomy of the Gelastocoridae 287

Previous studies of the Gelastocoridae 287

Taxonomic value of certain characters 289

Techniques 292

Family Characteristics 295

Key to Subfamilies and Genera 295

Gelastocorinae Champion 296

Genus Gelastocoris Kirkaldy 296

Key to the species of Gelastocoris 297

Gelastocoris oculatus ocidatus ( Fabricius ) 298

Gelastocoris oculatus variegatus ( Guerin-Meneville ) 311

Gelastocoris rotundatus Champion 314

Gelastocoris hungerfordi Melin 316

Gelastocoris vicinus Champion 319

Gelastocoris bufo ( Herrich-Schiiffer) 322

Gelastocoris amazonensis Melin 325

Gelastocoris major Montandon 327

Gelastocoris peruensis Melin 329

Gelastocoris nebulosus (Guerin-Meneville) 331

Gelastocoris fuscus Martin 336

Gelastocoris viridis n. sp 338

Gelastocoris angulatus ( Melin) 339

Gelastocoris apureensis Melin 342

Nerthrinae Kirkaldy 343

Genus Nerthra Say 343

Key to the species of Nerthra 347

Nerthra stygica Say 354

Nerthra mexicana ( Melin) 356

Nerthra martini Todd 358

Nerthra usingeri Todd 360

Nerthra parvida ( Signoret ) 362

Nerthra quinquedentata ( Melin) 363

Nerthra buenoi n. sp. 365

The Family Gelastocoridae (Hemiptera) 279

PAGE

Ncrthra mptnria ( Fabricius ) 366

Nertlira nmina ( Herri ch-Scliiiffer) 368

NertJtra nepaeforniis ( Fabricius) 371

Nerthra terrestris (Kevan) 373

Nerthra horealis ( Melin ) 375

Nertlira tcncbrosa n. n 376

Nertiira unicornis ( Melin) 378

Nerthra peruviana ( Montandon ) 379

Nerthra montancJoni (Melin ) 380

Nerthra amcricana ( Montandon) 381

Nerthra wiUiamsi n. sp 383

Nerthra amplicollis (Stal) 384

Nerthra ecuadorensis ( Melin ) 387

Nerthra planifrons ( Melin) 388

Nerthra ater ( Melin ) 389

Nerthra lata ( Montandon ) 389

Nerthra rudis ( Melin ) 390

Nerthra conidis n. sp 392

Nerthra fuscipes ( Guerin-Meneville ) 393

Nerthra manni n. sp 396

Nertlira hungerfordi n. sp 398

Nerthra hracchialis n. sp 400

Nerthra grandicoJlis (Germar) 401

Nerthra indica (Atkinson) 405

Nerthra turgiduJa ( Distant) 406

Nerthra lohata ( Montandon ) 407

Nerthra serrata ( Montandon) 409

Nerthra asiatica ( Horvath) 410

Nerthra spissa ( Distant) 412

Nerthra rugosa ( Desjardins ) 412

Nerthra macrothorax ( Montrouzier) 414

Nerthra nervosa ( Montandon) 416

Nertlira hirsuta n. sp 417

Nertlira laticoUis (Guerin-Meneville) 418

Nerthra gurneyi n. sp 419

Nerthra mixta ( Montandon ) 420

Nerthra oniani n. sp. 422

Nerthra femoralis ( Montandon) 423

Nerthra nudata n. sp 425

Nerthra annulipes ( Horvath ) 426

Nertlira anipliata ( Montandon ) 426

Nertlira macrostijla n. sp 428

Nerthra rohusta n. sp 429

Nerthra liiteovaria ( Distant ) 430

Nerthra tuhercidata ( Montandon) 431

Nerthra alaticuUis ( Stal ) 432

Nertlira adspersa ( Stal ) 434

Nerthra stall ( Montandon ) 435

Nerthra elongata ( Montandon) 436

280 The University Science Bulletin

PAGE

Nerthra tasmaniensts n. sp 437

Nerthra suherosa ( Erichson ) 438

Nerthra walkeri n. sp 439

Nerthra sinuosa n. sp 440

Nerthra grandis ( Montandon ) 440

VI. Literature Cited 442

VII. Index 444

VIII. Illustrations 447

INTRODUCTION

The family Gelastocoridae, as considered in this work, is com-
posed of two subfamihes, each of which has within its hmits a single
genus. The total number of species is 74, of which 13 belong to
the genus Gclastocoris and 61 to the genus Nerthra. The identi-
fication of the species in this family has always been most difficult.
This has been due, in part, to the similarity of many species and the
extreme variability of color and of certain morphological characters
within a single species. On the other hand, the difficulty has,
in part, also been due to the inadequacy of many of the original
descriptions, especially of the earlier workers, and the lack of com-
prehensive keys. It has been the aim of the writer to assemble the
literature pertaining to the Gelastocoridae; to present the known
facts concerning the biology and the ecology of the species of this
family; to establish techniques which would facilitate the study
of the Gelastocoridae; to evaluate the characters used in the taxo-
nomic study of these insects; to study geographical distribution as
an aid to our knowledge of their phylogeny; to determine the
phylogenetic relationships between the Gelastocoridae and other
cryptocerous Hemiptera; to discover the natural relationships among
the species within the family; and finally to present keys which
would permit the determination of genera and species of this
difficult family. One of the greatest obstacles to the accomplish-
ment of these aims has been the inaccessibility of type specimens.
The types, for the most part, are scattered in the various European
museums. Limited material available for study, particularly from
certain areas of the Eastern Hemisphere, has been another hin-
drance. Even so, the writer has been able to study what is without
question the largest collection of Gelastocoridae ever assembled.
In addition to the very large collection in the Francis Huntington
Snow Entomological Collection at the University of Kansas, the
writer has received, through the efforts of Doctor H. B. Hungerford,

The Family Gelastocoridae (Hemiptera) 281

specimens from nearly all the major entomological collections in
this country and from some of the collections in Europe.

In this study, it has been necessary to place in synonomy a number
of genera, which in my opinion were not based upon sound
generic characters. It is unfortunate that it was not currently
possible to retain more of the existing generic names, at least as
subgenera, but, as it will be explained at another point (see pages
296, 343), many of the characters previously utilized have certainly
developed independently in rather distantly related species and
therefore do not delimit natural groups. This is particularly true
in the subfamily Nerthrinae. Such characters are very useful in
the formation of keys, but obviously of considerably less im-
portance in the development of a natural system of classification. It
is the hope of the writer that this work will simplify the task of
specific determination and especially, that it will stimulate other
workers to undertake further studies concerning the taxonomy,
biology and ecology of the Gelastocoridae.

ACKNOWLEDGMENTS

The writer wishes to take this opportimity to express his appre-
ciation for the assistance received from various sources during the
progress of this study.

To Doctor H. B. Hungerford of the University of Kansas, who
suggested this study to me, I am especially grateful. Not only has
he generously given his time to the discussion of various problems
that occurred from time to time, but also, through his influence
specimens were received from most of the major collections in this
country and from several European museums. Further, his notes
on the specimens in many of the European museums have greatly
facilitated many aspects of this investigation.

To Doctor C. D. Michener of the University of Kansas, I am
deeply indebted for the encouragement he has given to me and for
the consideration he has given to the many questions of procedure
and taxonomy.

To the others at the University of Kansas, and especially to Doc-
tors Kathleen Doering and R. H. Beamer, I am sincerely appreciative
for their willingness to give assistance at all times.

Thanks are due to Doctor O. Lundblad of the Museum of Stock-
holm, Stockholm, Sweden, through whose kindness the writer was
able to study those types of Douglas Melin which are located in
that museum; to Doctor R. L. Usinger of the University of Cali-
fornia, who not only sent material from his personal collection.

282 The University Science Bulletin

but also made a number of comparisons between drawings sent by
this writer and types located in the British Museum in London,
England; and to Doctor Eugene Seguy of the Museum of Paris,
for his comparison of a specimen with a type located in that In-
stitution and for sending a paratype of one of the species I had
not been able to study.

I would also like to thank the curators of the various museums
and collections who have provided specimens for this study. As
it has been necessary to abbreviate the names of these institutions
in the distributional data of the descriptions in this work, a list of
these institutions and the abbreviations used is given at this point.

United States National Museum at Washington, D. C, (USNM);
American Museum of Natural History at New York City, N. Y.,
(AMNH); Museum of Comparative Zoology at Harvard University,
Cambridge, Mass., (MCZH); California Academy of Science at
San Francisco, California, (CAS); J. C. Lutz Collection, Philadel-
phia, Pennsylvania, (J. C. Lutz Coll.); Carnegie Museum at Pitts-
burgh, Pennsylvania, (Car. Mus. ); Museum of Prague, Prague,
Czechoslovakia, (Mus. Prague); University of Michigan Collection
at Ann Arbor, Michigan, (Mich. Coll.); Purdue University Collec-
tion at Lafayette, Indiana, ( Purdue Coll. ) ; British Museum ( Natural
History), London, England and Rijksmuseum van Natuurtijke His-
toric, Lieden, Netherlands. All specimens not indicated by one of
the above abbreviations or unless otherwise stated are located in
the Francis Huntington Snow Entomological Collection at the Uni-
versity of Kansas.

BIOLOGY AND ECOLOGY

Little is known concerning the biology of the Gelastocoridae.
The life history of only one species, Gelastocoris ociilattts (Fabri-
cius), is known to any extent. Doctor H. B. Hungerford's study of
this species (Univ. Kansas Sci. Bull., 1922, vol. 14, pp. 145-171.) has
shown that the oval eggs are deposited in sand with the cephalic
end uppermost. The egg, which is white when dry and amber or
ferruginous when moistened, blends perfectly with the sand. One
female probably deposits as many as 200 eggs during a season.
Doctor Hungerford recovered 192 nymphs and eggs from one fe-
male. The number of eggs laid per day may vary from 1 to 13.
The incubation period is from 12 to 15 days. There are five
nymphal instars. The total developmental period is from 60 to
100 days. In the laboratory, mating occurred almost daily from
May to November, but it is not known whether this is true when

The Family Gelastocoridae (Hemiptera) 283

the insects are in their natural habitat. Millspaugh (Field and
Laboratory, 1939, vol. 7, No. 2, pp. 82-83. ) suggests that G. ciicid-
lafiis Nhutin {=G. hungerfordi Melin) has a definite period during
which mating is more commonly observed than at other times dur-
ing the season. The exact number of generations of G. ocidatiis
(Fabricius) is unknown, but probably varies from one in the
extreme northern part of its range to several in the southern part
of its range. It is not recorded in the literature how G. ocidatiis
( Fabricius ) and other species pass the winter in regions where
this season is unfavorable to an active mode of life. It is, however,
fairly certain that they over- winter in the adult stage; undoubtedly
they seek out protected places to pass the winter. In addition to
Hungerford's study of G. ocidatus (Fabricius), Kevan (Froc. R.
Ent. Soc. London, ser. A, vol. 17. 1942, pp. 109-110.) has studied
the life history of Nerthra nepaeformis (Fabricius) in Trinidad.
Although he was unable to rear these insects through their com-
plete life cycle, he was successful in rearing captured fifth instar
nymphs through to adults. Adults were observed mating, but no
eggs were obtained. Except for captured nymphs of various
instars, which cannot be identified in the light of present knowledge,
nothing is known concerning the life cycles of the other species of
this family.

While all the species of the Gelastocorinae are found on the mud
or sand banks of streams, ponds, etc., this is apparently not entirely
true for all the species of the Nerthrinae. Melin (Zoologia Bidrag
Fran Uppsala, 1929*, Band 12, p. 194.) wrote: "Of the three
specimens of M. bipunctatus I collected in Peru, the male lived
in a decomposed trunk in a 'chacra' of bananas, the female on the
ground in the forest amongst dry leaves, and the larva on a trunk
in a clearing." Kevan (I.e.) states: "Af. nepaeformis has been taken
commonly, both by writer, and by Dr. H. B. Hynes, in a small
banana plantation attached to the Imperial College of Tropical
Agriculture, St. Augustine, Trinidad, during both wet and the dry
seasons of 1941-1942. There is neither standing nor running water
anywhere in the vicinity of this plantation, and, in the dry season,
the ground becomes parched and dry. The insects are most readily
found under debris, such as heaps of dry grass or old pawpaw
stumps, and seem to spend much of their time burrowing in the
soil, where they are common also, though less easily detected, while
nymphal exuviae have often been found underground. In the
laboratory, too, they will readily biuTOw in moderately loose soil.

* See page 288.

284 The University Science Bulletin

All their time, however, is not spent in this manner, for the writer
has swept the species from low vegetation on to which it had
climbed, presumably in search of prey." According to Torre-
Bueno (Ent. News, 1906, vol. 17, p. 54.), Biolley found a speci-
men of N. fuscipes (Guerin-Meneville) in a rotten log in Costa
Rica in December of 1904. Maxwell-Lefroy (Indian Insect
Life, Calcutta, 1909, p. 709.) writing about N. indica (Atkin-
son) states: ". . . it is found on grass paths, on the soil and
under stones, as well as on hard roads; it is not aquatic and is possi-
bly predaceous on small insects." In contrast to the above. La Riv-
ers (Wasmann Journ. of Biol., vol. 11, no. 1, 1953, pp. 83-84.) re-
ports some very interesting observations on the habitat of Mononyx
fuscipes Guerin-Meneville (probably N. martini Todd). In dis-
cussing the capture of specimens in Inyo County, California, he
states: "Most of the specimens were immatures taken at or near
the smface of the water. They readily crawl beneath the water
surface along plant stems while searching for prey. ' Concerning
two collections from Clark County, Nevada, he reports: "The
first Nevada record obtained, at Warm Springs, resulted in some
half-a-dozen mature and as many immature specimens, taken, for
the most part, from their hiding places on the undersurfaces of
pieces of wood partially submerged in a marshy tract of quiet,
shallow, warm water (83° F, pH 7.3). The bottom was fine, black
silt. Most of the specimens were clinging to the wood just below
the water surface." and "The Ash Meadows specimens were taken
from the undersurfaces of rocks bathed by swiftly flowing 90° F.
water (pH 7.3) and, like the Clark County individuals, were not
common; . . ." The observations of La Rivers brings to mind
a comment by Doctor R. H. Beamer pertaining to the capture of a
specimen of Nerthra rtigosa ( Desjardins ) . Doctor Beamer stated
that the specimen was first observed near some debris adjacent to
the surf on Matecumba Key, Florida, and that when it was disturbed
it jumped into the water among some floating vegetation and that
it clung there to a submerged f)lant until collected. This may
have been entirely accidental, but is of interest in light of La Rivers'
observations.

Apparently all species are predatory as evidenced by the rap-
torial front legs. Their prey must consist of small insects and
especially the larvae of those insects which are found in asso-
ciation with the Gelastocoridae. Kevan ( /. c. ) has reported that
in the laboratory N. nepaeformis (Fabricius) preferred termites as
food, while small moths and beetles were not received with favor.

The Family Gelastocoridae (Hkmtptera) 285

In regard to the burrowing habit of certain species, an observa-
tion of Doctor H. B. Hungerford on G. ocidatus (Fabricius) is of
interest. Doctor Hungerford states that he has seen broad, sandy,
barren flats where toad bugs hved become iniuidated by rapid
currents of water for a few hours; nevertheless, when the water
receded and the sun came again, the toad bugs were there as
before. Since there was neither vegetation nor sizable stones for
their anchorage, he supposes that they "dug in". Although G.
ocidatus (Fabricius) has well-developed wings, the writer has
never seen a specimen fly. When they are disturbed, they will
either hop away or crouch down against the mud or sand, appar-
ently depending upon their coloration to protect them from de-
tection. The facts that they burrow and that they apparently
do not often fly perhaps will explain, in part, the reduction of the
membrane of the hemelytra and of the entire hind wing in some
species. The fact that some species have the hemelytra united
may be due to the above habits. At least, it is the opinion of the
writer that any mutation toward the brachypterous condition
would not be harmful as far as their habits are concerned.

PHYLOGENY

Because of the habitat and the structure of the body, the family
Gelastocoridae has usually been considered as representing an
intermediate group in the progression from a terrestrial to an
aquatic mode of life. Whether this family represents an inde-
pendent line of development toward the semiaquatic mode of
life or whether they have arisen from a common ancestor with
existant aquatic forms, for example the Naucoridae, is not clear.
The morphological similarity between the subfamily Nerthrinae
and the Naucoridae is, in some respects very great. However,
Doctor R. L. Usinger, in a discussion with the writer, pointed out
that 8rd instar nymphs of Naucoridae possess abdominal scent
glands while the n\mphs of Nerthrinae apparently lack such glands.
He considers that this is a very fundamental difference. He further
stated that he does not believe that the absence of scent glands
in the Nerthrinae could be due to a reduction or a loss. The
similarities of the eggs, mouthparts, grasping forelegs, antennae,
and abdominal segmentation must therefore be considered as par-
allel developments. On the basis of our present knowledge we
must conclude that the Gelastocoridae represent an independent
line of development toward the semiaquatic mode of life. It has
generally been assumed that the approach was from terrestrial

286 The University Sc:ience Bulletin

forms to semiaqiiatic forms to truely aquatic forms, however, in
the Gelastocoridae the immediate ancestors may have Hved en-
tirely in the aquatic (semiaqiiatic) state and our present forms
may represent departure from the water to the terrestrial mode of
life. It is also possible that the Nerthrinae alone had aquatic an-
cestors and the Gelastocorinae did not enter the water. The hid-
den antennae of the Gelastocoridae could have been so developed
either as an adaptation to an aquatic mode of life or as an adap-
tation to the burrowing habit. Thus morphological evidence based
upon the antennae could be used to support either type of pro-
gression. At the present time only one species has been found
living in the water. The writer has not seen all the specimens
collected by La Rivers, but two males from the Warm Springs,
Clark Co., Nevada series collected by La Rivers and Johnson are
Nertliro mariini Todd. The species has also been collected in a
terrestrial habitat. On the other hand species of Nerthra have
been found in situations quite remote from water in India, Peru,
and Trinidad. The normal habitat of the majority of the species
of the Nerthrinae and of all the Gelastocorinae would seem to
be the moist terrestrial margins of streams and other bodies of
water. It seems best to consider the specimens collected by La
Rivers as representing the first record of a species entering the
water rather than the last species leaving the water. This con-
clusion is supported by the less specialized condition of the Gelas-
tocorinae. In the opinion of the writer, based on a comparative
study of the legs, nwuthparts, , abdominal segmentation and the
male genitalia, the Gelastocorinae is the more primitive of the
two subfamilies. Normally we would expect to find the more
primitive forms, not the more specialized forms, occupying a habi-
tat most closely resembling the habitat of their ancestors.

Fossil evidence would undoubtedly be helpful in the solution of
the problem of direction of progression of habitat and also such
evidence would certainly aid in the explanation of the current distri-
bution of the species of this family. Unfortunately fossils of the
Gelastocoridae are unknown. Limnochares antiquus Heyden which
was described as a mite and Necygontis rotimdatiis Scudder ( U. S.
Geological Survey of the Territories, 1890, pp. 347-348, pi. 7 fig. 8. )
have been placed in the family by Bertkau and by Scudder. In the
opinion of the writer they do not belong to the Gelastocoridae.
Fossils of aquatic hemiptera such as Nepidae, Naucoridae, Notonec-
tidae, etc. are known and some date back to Triassic and Jurassic
times. The absence of gelastocorid fossils may be an indication that

The Family Gelastoc:oridae ( Hemiptera ) 287

ancient members of this family were terrestrial and that they did not
inhabit areas which were immediately adjacent to water. It is true,
however, that many of the fossils of aquatic insects are from Euro-
pean deposits and gelastocorids do not occur iu Europe at the
present time and they may never have occurred there. Since the
Gelastocorinae are at present known only from the Western Hemi-
sphere,- we must assume either that they originated here or that
they once had a more widespread distribution. If the first assump-
tion is correct and the Gelastocoridae are the more primitive, then
the Nerthrinae must also have originated in the Western Hemi-
sphere. If this is true, then the spread of the species could have
been by way of Alaska into Asia and then toward Australia and
Africa. If, on the other hand, the Gelastocorinae had a more wide-
spread distribution in the past, then the spread of Nerthrinae could
have been in the reverse direction in which event, Nerthra grandi-
coJlis (Germar) could represent a relict species. The species of
Africa, China, India and Sumatra exhibit characteristics in common
witli both the Australian species and with the species from the
Western Hemisphere. This could be used as evidence for either
direction of distributional spread. In the absence of fossil evidence,
it is not possible to conclude at this time whether the distributional
spread was as suggested above. Perhaps future biological and
morphological studies will help solve this problem.

TAXONOMY OF THE GELASTOCORIDAE
Previous Studies of the Gelastocoridae

Before consideration of characteristics which have been used by
the writer, it seems best to present a brief review of the literature,
placing special emphasis upon the principal workers and upon the
dates and the authors of the original generic and subgeneric de-
scriptions.

In 1802, Latreille established the genus Galguhis for Naucoris
oculata Fabricius, 1798. Billberg, 1820, first used the name Gal-
gulidae (Galgulides). For nearly a century thereafter the family
was known bv this name. Thomas Sav, 1832, described a new
genus and species which he named Nerthra stygica. He apparently
was separating it from Naucoris and did not mention the Galgulidae.
One year later, in 1833,* Laporte described the genus Mononyx with
Naucoris raptoria Fabricius, 1803, as the type of the genus. He
placed this genus in the family Galgulidae. Nerthra nepaeformis

* Harris, H. M., Pan-Pacific Entomologist, vol. 18, no. 4, 1942, pp. 161-162, shows
that De Laporte's paper did not appear in 18'32 as usually given.

288 The University Science Bulletin

(Fabricius), which was described in 1775 as Naucoris nepaeformis,
was not inckided in this family until 1868. During 1843 and 1844,
Guerin-Meneville proposed, as a subgenus of Mononijx, the name
Peltoptertis. He based this subgenus upon Naucoris rugosa Des-
jardins. The genera Phintius, Matinus and Scylaecus were erected
in 1861 by Stal for Monomjx grandicollis Germar, Monoyx alati-
collis Stal and Galgtilus mucrothorax Montrouzier respectively. The
generic names Phintius and Scylaecus of Stal were placed in the
synonomy of Mononyx and Peltopterus by Montandon in 1899. The
latter author was the first worker to present a detailed species key.
It included seventeen species and was published in 1899 ( Bull. Soc.
Sci. de Bucarest-Roumanie, an. VIII, nos. 4 & 5, pp. 392-410.).
Furthermore, in 1900 (ibid., an. VIII, no. 6, pp. 1-9.) he presented a
key to five species of Matinus. This worker also described a dozen
or more new species in this family during the years 1895 to 1914.
Dumeril ( Mem. Acad. Sci. I'lnst. Imp. France, 1860, tom. XXXI, 2nd
partie, p. 1040.) was the first to recognize that Galgulus Fabricius
was preoccupied by Galgulus Pliny a genus of birds. It was not
until 1897 that the situation was altered when Kirkaldy (Entomolo-
gist, 1897, vol. 30, p. 258. ) renamed the genus Gelastocoris. In the
preparation of the Biologia Centrali-Americana, Rhynchota Heter-
optera, vol. 2, Champion used the manuscript names of Montandon
for two new species and he attempted to credit these species to
Montandon, however, the fact remains that Champion published
these names with his own descriptions and Montandon in 1910,
credited G. vicinus to Champion.

More recently, Blatchley (Ent. News, 1926, Vol. 36, pp. 49-52.)
described what he believed to be a new genus and species, Glos-
soaspis brunnea. Since the specimens on which this name was
founded were from Florida, Blatchley understandably did not con-
sider that they might be the same as a species from the Eastern
Hemisphere. A specimen from Pearl Island on the Pacific side
of Panama and the description of Naucoris rugosa Desjardins made
the present writer suspicious. A specimen from Florida was sent
to Doctor Eugene Seguy at the Paris Museum and a comparison
of this specimen with one of the cotypes of N. rugosa Desj. has
shown that the two are identical. In the process of identification
of specimens collected in northern South America, Douglas Melin
(Zool. Bidr. Fran Uppsala, 1929 *, Band 12, pp. 151-194.) undertook

* All soiirces list this date as 19.30, however, Melin's paper was printed separately in
December, 1928 and Dr. H. B. Hungerford has a copy which was mailed to him by the
author on February 1, 1929.

The Family Gelastocoridae (Hemiptera) 289

a revision of the Gelastocoridae of the Western Hemisphere. The
results were: 1 new genus, Montandonius, 17 new species and 3
new subspecies. His keys, which are based entirely upon external
characters, are fairly good and will permit the identification of a
majority of the species. In this work Melin refuses to accept the
rule of priority, but accepts as the author of the species, the first
worker whose description Melin considered sufficient to identify
the species. For two new species of Monomjx, he used the names
bipunctatiis and obscurus. Since both names had previously been
used in the genus, his names are homonyms and it has been neces-
sary to rename these species. Kevan (Ann. & Mag. Nat. Hist., ser.
11, vol. 14, 1948, p. 813. ) has used the name Monomjx terrestris for
Monomjx hipunctatus Melin. The other species is renamed in this
work. At the suggestion of Doctor H. B. Hungerford, Martin
(Univ. Kansas Sci. Bull., 1928*, vol. 18, no. 4, pp. 351-369) made
an exploratory survey of the genus Gelastocoris and was the first
to use male genitalia for the identification of the species. He fur-
ther discussed the value of the other characters for the identifica-
tion of the species of this genus. In the paper he described three
new species, but he did not make a key to the genus.

Taxonomic Value of Certain Characters

In this study, characters which were used by earlier workers and
a number of other characters were examined to determine their
value in the identification of species of Gelastocoridae.

Color, which was used almost entirely in the descriptions of
earlier workers, has been found to be too variable to be of much
value. In a few species the color of certain structures may be
characteristic. Such a case is seen in the white tubercles on the
frons of Nerthra tuherculata (Montandon). It is the opinion of
the writer that coloration should not be relied upon alone and that
other characters should be used whenever possible.

Size, alone, is not of specific value, except when used with other
characters.

In the Nerthrinae, the tubercles of the head have been studied.
The tubercles are variable to some extent, but are of value in sep-
arating the species into groups. Complete absence of these tuber-
cles may be used and also the number and position of the tubercles
may be of value. However, this character should not be relied

* The actual publication date of this paper was in 1929, probably sometime in the period
of June to August.

10—6730

290 The University Science Bulletin

upon alone, because these tubercles are rather variable within
some species and may be nearly obsolete in species normally having
prominent tubercles.

The apical margin of the head is more or less rounded and sim-
ilar for most of the species of Gelasfocoris, however, G. angulatiis
( Melin ) usually has the apical margin slightly concave.

The antennae, while somewhat different in the two subfamilies,
are not characteristic for a species. They are difficult to study and
are nearly identical for all the species of each group.

In the past, the presence of ocelli has been one of the key char-
acters in the separation of this family from the Naucoridae, how-
ever, I have found that six species lack ocelli. These six species
belong to three rather distantly related groups of the Nerthrinae
and undoubtedly ocelli have been lost independently. Although
the absence of ocelli does not show a phylogenetic relationship,
the presence or absence of ocelli is a useful key character and has
been so used by the writer.

The shape of the pronotum, especially at the lateral margin,
although slightly variable, seems to be constant for most species.
In some species, on the other hand, the degree of variation is quite
great and some specimens even show a remarkable difference in
the two sides. Also the general shape for many species, within
both subfamilies, is more or less similar. One species, G. angidatus
(Melin), has 6 to 8 distinct, short, longitudinal carinae on the
posterior portion of the pronotum. These carinae are lacking in
the other species of this genus.

The scutellum, while it differs some between the two subfamilies
and to some extent in some of the species of each, is not considered
to be of much value in determination of species. In what was for-
merly the subgenus Peltoptenis, the scutellum is somewhat smaller
than in the other Nerthrinae, but as reduction in size of scutellum
follows loss of normal flight wings, this is not considered important
and certainly the difference is not sufficient for generic or sub-
generic consideration. Furthermore, most specimens are pinned
through the scutellum, which makes it difficult to study this
structure.

The hemelytra may vary in length for the different species and
may be of some value. Reduction of the membrane occurs in both
subfamilies and in the Nerthrinae the hemelytra may be entirely
coriaceous and separate or fused togetlier. These characters are
of value in the identification of species, but as with the absence of

The Family Gelastocoridae (Hemiptera) 291

ocelli, have arisen independently and do not imply a close phylo-
genetic relationship. The embolium exhibits characters that are
specific. A few species show a strongly dilated lateral margin.
This is considered as a generic characteristic by Melin. The shape
of the lateral margin of the embolium of the other species is of some
value when used with other characters.

Bristles found on different parts of the insects, especially in
Nerthrinae, may prove to be of value upon further investigation.
This writer has used bristles, along with other characters, to identify
several species. The bristles tend to form groups on the tubercles
of the scutellum and upon the hemelytra and they vary in both
size and shape for a given species. Also, they are too frequently
covered by the encrustation or are rubbed off to be a character of
positive identification in most species.

In the genus Gelastocoris, some of the granulations, especially
upon the hemelytra, may be enlarged and blisterlike or more
elevated and peglike. Both may be used with other characters
in the identification of species. The blisterlike granules are some-
what variable in number and size within a species, but can be
used to help separate a few species.

The legs of Gelastocoris are all more or less the same for the
different species. In a few species, the legs, especially the hind
pair, are a little longer in proportion to the length of the body,
but the difference was not great enough to be used. The size and
shape of the anterior dilation of the front femora will separate two
species in the Nerthrinae from the other species, but the femora
of the others are so similar as to be of no value. The trochanters
were studied and in particular small tubercles on the apices were
examined, but these were found to be too variable within a species
to be of value. Proportions of the intermediate and hind legs
were checked and results showed them to be more or less the
same for all species. Spines on the legs are not characteristic of a
species.

The mesosternal elevation was used by Montandon in identifica-
tion of some of the species of Nerthrinae from the Eastern Hemi-
sphere. This structure is fairly variable within a species and in the
opinion of the writer should only be used with other characters, if
at all, and only when there is no possibility of confusion with an-
other species, i. e. N. grandicollis (Germar). It is difficult to study
because it is so often covered by the fore and intermediate legs
and commonly destroyed by the insect pin passing through or
close to it.

292 The University Science Bulletin

The shape and size of the abdominal segments of the male are
relatively constant for a species and of specific value in some
cases. In the Nerthrinae, the seventh ventral abdominal segment
of the females from the Western Hemisphere, Asia and Africa have
the posterior margin deeply emarginate and possess elevations
and depressions or both, which are characteristic for the species.
The females from the Australian region and Pacific Islands have
the seventh ventral abdominal segment more or less projecting
posteriorly and covering the ovipositors.

The male genitalia were examined and found to be very charac-
teristic for a species. They should be used as a final check when
males are available to the worker. The ovipositors of the females
were examined and are of some value when used with other
characters and in a few instances are in themselves sufficient to
identify species. The ovipositors of some of the species are spined,
but the number and position of the spines was found to be too
variable to be of value as specific characters.

Various proportions of different parts of the insects were com-
pared and with a few exceptions were practically the same for
most of the species in each of the two subfamilies.

Techniques

In the measurement of the length of the specimens, the distance
from the anterior margin of the eyes to the posterior end of the
abdomen or to the apices of the hemelytra, whichever was the
greater, was used. In the measurement of the width of specimens,
both the greatest width of pronotum and width of the abdomen at
a level with the apex of the scutellum were recorded. In the
males of Nerthrinae, the distance between the caudolateral angles
of the sixth ventral abdominal segment was compared with one-
half the width of the posterior margin of the fourth ventral ab-
dominal segment. The right side of the fourth segment is used
and the distance is measured from the median notch along the
posterior margin to the caudolateral angle of this segment. For
all other measurements the greatest width or length was used,
unless otherwise stated.

The technique for extracting the male genitalia is as follows:
The male is determined by the structure of the ventral abdominal
segments, the labels may then be pushed down to about one fourth
of an inch from the point of the pin and the specimen then
immersed in 5% alcohol with the labels resting on the outside
of the lip of the relaxing dish, holding the specimen in place.

The Family Gelastocoridae (Hemiptera) 293

In this manner a fairly large number of specimens may be re-
laxed at a time, the number depending upon the speed at which
the worker can extract the genitalia. The length of time required
for relaxation will vary with the size, the age of the specimens
and to some degree, the extent to which the specimens are covered
bv encrustations. Approximately 15 to 20 minutes is sufficient for
Gdastocoris, while Nerthra, especially the larger species require
a longer period. The process can be speeded by using a warm
relaxing fluid, but the writer has found that the number of speci-
mens that could be handled at one time must be reduced or
the specimens may become too relaxed. This makes extraction of
the genitalia more difficult in that the last few segments may tear
off in the attempt to remove the genital capsule, unless extreme
care is taken in the dissection. A special relaxing fluid (Alcohol
757c, 106 cc; Distilled water, 98 cc; Benzol, 14 cc; Ethyl ace-
tate, 38 cc. ) may be used in place of 5% alcohol. When the
specimen is sufficiently relaxed, that is to say, when it is relaxed
enough to prevent breakage yet stiff enough so that the abdominal
segments may be moved only by applying considerable pressure,
the genitalia may be extracted. From this point the technique
varies according to the subfamily being studied. For the Gela-
stocorinae, the procedure is as follows: holding the specimen in
one hand between the thumb and one finger, insert a small dis-
secting needle a short distance into the left side of the aperture
of the posterior end of the abdomen. Then pushing to the right
with the point of the dissecting needle and pulling the needle out
at the same time the genital capsule will be extracted. The
genitalia may then be turned a quarter of a turn counterclock-
wise, in this position the genitalia will "lock" against the mem-
brane of the hemelytra and will be exposed at the end of the
abdomen, where they are available for study at any time. Occa-
sionally, in old specimens or in specimens which are not com-
pletely relaxed, the membranes and the muscles which hold the
genital capsule to the preceding segment may break causing the
genitalia to become completely separated from the specimen. In
such cases, the genital capsule may be placed in a small vial
(4x10 mm.) in a small amount of glycerin. These vials may
be kept with the specimen by running the pin through the
cork. When such a procedure is used, it is advisable to run the
pin through the cork at a slant and also to use as little glycerin
as is necessary to cover the genital capsule. The procedure for
the extraction of the genitalia of the Nerthrinae is as follows:

294 The University Science Bulletin

if the entire right clasper is to be studied it is necessary to remove
the last two abdominal segments. This portion of the insect
is then placed in a drop of glycerin on a slide, where the clasper
is removed by breaking the segments longitudinally, and cutting
the strong muscles at the base of the clasper. It is unfortunate
that it is necessary to destroy these segments, but no other manner
could be found for removing the entire clasper. An effort was
made to remove the clasper from the specimen without removal
of the eighth abdominal segment. It was found, however, that
the ninth segment could not be removed separately in most
species and that the danger of breaking the clasper was great
when such a procedure was attempted. A better procedure and
one completely satisfactory for identification is as follows: After
relaxation, with a dissecting needle, gently pull the segments to
the rear and to the left of the insect until the clasper is clear of
the abdomen. Then allow the segments to pull back to their
normal position, but direct the clasper with the needle to the
ventral side of the abdomen. It will remain in this position and
can be studied readily by simply inverting the specimen.

All the drawings of the complete right claspers of the Nerth-
rinae were made so that the worker will see the ventral surface
of the clasper. In this manner comparison may be made with
the drawing when the latter method of extracting the clasper
is used. In order to make such a comparison, the specimen should
be tilted slightly so that the last two segments will be horizontal,
thus assuring a true ventral view of the clasper. In emphasizing
the value of the second procedure of extraction of the claspers
of Nerthrinae, it may be pointed out that the most distinctive
part of the clasper in almost all species is the apical half, which
ordinarily is fully visible when this procedure is used.

As has been mentioned previously, many specimens, especially
in the Nerthrinae, may be covered with an encrustation of mud
or other debris. It often becomes necessary therefore, to re-
move this encrustation, in order to study certain structures. In
the past this has been a difficult task, which required much
brushing of completely relaxed specimens. Recently, Nelson
( Coleopterist's Bull, 1949, vol. 3, no. 6, pp. 89-92.), has discovered
a method of removing the encrustations from specimens of the
family Elmidae ( Coleoptera ) . This method is satisfactory for
cleaning Gelastocorinae and may be used for the Nerthrinae. For
insects collected dry or in alcohol, or those which have been
pinned in the uncleaned condition, the procedure is as follows:

The Family Gelastocoridae (Hemiptera) 295

place the specimen in 5 to 10% acetic acid for 20 to 30 minutes.
Then put the specimen in a dish of warm tap water, a Syracuse
dish is satisfactory for small species, then add a pinch of tri-
sodium-phosphate, known commercially as "T. S. P." The length
of time the specimen is left in this solution varies with the degree
of encrustation. When the encrustation has become loosened, it
is removed from the specimen by means of a brush with short
stiff bristles. Such a brush can be made by taking a small in-
expensive paint brush and cutting the bristles down to a length
of about 1 mm. and then bending the metal tip inwardly to crowd
the bristles together and thus making the brush stifiFer. When
collecting specimens, if they are placed in Pampel's Fluid, they
may be cleaned amazingly well when subjected to 20 or 30
minutes in T. S. P. Pampel's Fluid is composed of the follow-
ing materials: Distilled water, 30 parts; 95% Ethyl alcohol, 15
parts; Glacial acetic acid, 4 parts; Formaldehyde (40%), 6 parts.

Family Characteristics

Squat, medium sized bugs, head triangular in front view,
fitting into concavity between the anterior angles of the pronotum.
Rostrum four-segmented. Eyes large, reniform and projecting
dorsolaterad. Ocelli usually present. Antennae short, four-seg-
mented, inserted beneath tlie eyes and hidden betvveen the head
and prothorax. Pronotum large, much wider than the head,
disk elevated. Scutellum, usually rather large, triangular and
elevated. Hemelytra with clavus, corium, embolium and mem-
brane, or with membrane reduced or entirely lacking and in a
few species with the hemelytra united. Anterior legs raptorial.
Abdominal segments of the male more or less asymmetrical. The
nymphs of the species of the entire family possess ocelli as far
as is known. This family may be separated from the Ochteridae
and Saldidae by the hidden antennae and from the Naucoridae
by presence of ocelH in most cases and by the fact that the
head is not flattened dorsoventrally.

Key to the Subfamilies and Genera

OF Gelastocoridae

A. Fore tarsus not fused to tibia, articulate; two well-developed tarsal
claws on foreleg in adult; rostrum arising from apex of head, stout,

recurved posteriorly ( Gelastocorinae ) Gelastocoris Kirkaldy.

AA. Fore tarsus fused to tibia, not articulate; one well-developed
tarsal claw on foreleg in adult; rostrum appearing to arise
on ventral surface of head, slender, projecting anteriorly or
ventrally ( Nerthrinae) Nerthra Say.

296 The University Science Bulletin

Gelastocorinae Champion

Gelastocorids with the rostrum arising from apex of head, stout,
recurved posteriorly. Anterior leg with a single, articulate, tarsal
segment which bears two well-developed tarsal claws. Ninth
ventral abdominal segment of the male completely invaginated
within the body cavity, not visible externally. The male genitalia
with posterior process (keel) and right clasper greatly modified,
the two structures guiding the simple, tubular aedaegus; left
clasper small, simple. Ventrally the abdominal segments of female
nearly symmetrical.

The Genus Gelastocoris Kirkaldy

(Type of Genus, Naiicoris oculata Fabricius)

1897. Kirkaldy, G. W., Entomologist, vol. XXX, p. 258. (New name for

Galgulus Latrielle. )
1901. Champion, G. C., Biologia Centrali- Americana, Rhynchota-Heteroptera,

vol. II, p. 347.
1906. Kirkaldv, G. W., Trans. Am. Ent. Soc., vol. XXXII, p. 149.

1909. Kirkaldy, G. W. and J. R. de la Torre-Bueno, Proc. Ent. Soc. Washing-
ton, vol. X, p. 180.

1910. Montandon, A. L., Annuario del Museo Zoologico della R. Universita di
Napoli (Nouva Serie), vol. 3, no. 10, pp. 1-4.

1910. Banks, N., Cat. Nearctic Hemip.-Heteroptera, Sep. pub. Am. Ento-
mological Soc, pp. 10-11.

1913. Torre-Bueno, J. R. de la. Can. Ent. vol. 45, no. 2, p. 60.

1917. Van Duzee, E. P., Catalogue of Hemiptera, Univ. California Publ., p.
472.

1919. Hungerford, H. B., Univ. Kansas Sci. Bull., vol. XI, p. 46.

1929. iMelin, D., Zcologiska Bidrag Fran Uppsala, Band 12, pp. 153-154.

1929. Martin, C. H., Univ. Kansas Sci. Bull., vol. XVIII, p. 352.

1951. Adams, P. A. and C. Don MacNeill, Pan-Pacific Ent., vol. 27, no. 2,
p. 71.

1952. Ellis, L. L., Am. Midland Nat, vol. 48, no. 2, p. 310.

References to this genus under Galgulus Latreille:

1802. Latreille, P. A., Histoire naturelle, generale et particuliere des Crustaces

et des Insectes, vol. Ill, p. 253. (Naucoris oculata Fabricius, only

species. )
1804. Latreille, P. A., Histoire naturelle, generale et particuliere des Crustaces

et des Insectes, vol. XII, p. 286.
1807. Latreille, P. A., Genera Crustaceorum et Insectorum, etc., vol. Ill,

p. 143.
1810. Latreille, P. A., Considerations generales sur I'ordre naturel des animaux,

etc., pp. 260, 434.

1815. Leach, W. E., Brewster's Edinburgh Encyclopedia, vol. IX, p. 123.

1816. Lamarck, C. de, Histoire naturelle des animaux sans vertcbres, vol. Ill,
p. 509.

1825. Latreille, P. A., Families Naturelles du Regne Animal, pp. 424-425.
1825. Le Peletier, A. L. M. and A. Serville, Encyclopedic Methodique, vol. X,

p. 270.
1833. Laporte, F. L. de, Essai d'une Classification Systematique de L'order

des Hemipteres, pp. 15-16.
1835. Burmeister, H. C. C, Handbuch der Entomologie, Band 2, Ab. 1, p. 201.
1835. Brulle, G. A., Histoire Naturelle des Insectes, vol IX, p. 274.

The Family Gelastocoridae (Hemiptera) 297

1837. Spinola, M. M., Essai sur les Insects Hemipteres, etc., Genes, p. 62.

(Another issue, Paris, 1840.)
(1839)1840. Herrich-Schaffer, G. A. W., Die WanzenurUj;- • Insecten, vol. V,

p. 87.
1840. Blanchard, E., Historic Naturelle des Animaux Atricules, Band 3, p. 93.
1843. Annot, C. J. B. and A. Serville, Histoire Naturelle des Insectes.

Hemipteres, p. 424.

1850. Spinola, M. M., Tavola Sinottica, etc., 25, 1 p. 49.

1851. Fieber, F. X., Genera Hydrocoridum {from Abhandlungcn der Konig.
Bohmischen Gescllschaft der Wissenschaften in Prague), p. 13.

1853. Hcrrich-Schiiffer, G. A. W., Die Wanzenartigen Insecten, vol. IX, pp.
19, 24.

1859. Dohm, F. A., Catalogus Hemipteroruni ( Herausgegeben von dem
Stettin Entomologischer Verein), p. 53.

1860. Dumeril, A. M. C., Mem. Acad. Sci. Inst. Imp. de France, tom. 31, 2nd
partie, p. 1040.

1873. Walker, F., Catalogue of Hemiptera in the British Museum, part VIII,

p. 170.
1876. Stal, C., Kongliga Svenska Vetenskaps-Akademiens Handlingar, Band

14, No. 4, p. 137.
1879. Berg, C., Hemiptera Argentina enumeravit speciesque novas descripsit,

p. 185.
1884. Uhler, P. R., Standard Natural Histor>', vol. II, p. 263.

Small to medium sized bugs, extremely variable in color, but
usually more or less variegated. Rostrum arising from apex of
head, stout, recurved posteriorly. Anterior leg with a single, articu-
late, tarsal segment which bears two well-developed tarsal claws.
Ocelli present, though rather small in some species. Ninth ventral
abdominal segment of the male completely invaginated within the
body cavity, not visible externally, the posterior process (keel)
and right clasper greatly modified, left clasper small, simple. Ven-
tral abdominal segments of female nearly symmetrical, occasionally
with the incision of the posterior margin of the last segment bent
to the left of the specimen.

Key to the Species of Gelastocoris*

1. Basal portion of pronotum wiih 6 to 8 longitudinal

carinae . G. angulatus (Melin) p. 339

Basal portion of pronotum lacking carinae 2

2. ( 1 ) Lateral margin of pronotum straight or nearly so 3

Lateral margin of pronotum notched 4

3. (2) Lateral angle of pronotum projecting beyond base of em-

bolium, not bent ventrad, connexivum quite

visible G. bufo (Herrich-Schaffer) p. 322

Lateral margin of pronotum not or scarcely projecting be-
yond base of embolium, bent ventrad, connexivum com-
pletely or nearly covered by
hemelytra G. rotundatus Champion p. 314

* Gelastocoris apureensis Melin is not included. ( See page 342. )

298 The University Science Bulletin

4. (2) Membrane of hemelytron reduced, its length subequal to or

less than the width of hind femur 5

Membrane of hemelytron well developed, its length usually

about twice or more the width of the hind fenuir 8

5. (4) Pan of male genitaha bilobed G. maior Montandon p. 327

Pan of male genitalia not bilobed 6

6. (5) Pan wider than long, more or less rounded 7

Pan not wider than long, rather

elongate G. peruensis Melin p. 329

7. (6) Tumescence of right clasper adnate to base of clasper; keel

hook very long and recurved, extending completely back

across base of pan G. nebulosus ( Guerin-Meneville ) p. 331

Tumescence of right clasper not adnate to base of clasper;

keel hook not recurved back over pan . . G. viridis n. sp. p. 338

8. ( 4 ) Tumescence of right clasper adnate to base of clasper; keel

hook recurved completely back over base of

pan G. nebulosus (Guerin-Meneville) p. 331

Tumescence of right clasper not adnate to base of clasper,

keel hook not recurved completely back over base of pan P

9. ( 8 ) Pan of male genitalia very thin, elongate, narrow and twisted

apically, turned completely over G. amazonensis Melin p. 325

Pan of male genitalia not as above, usually thick, when thin,

not elongate or twisted completely o\'er 10

10. (9) Keel hood located to the right side of the

keel G. fuscus Martin jd. 333

Keel hood, not as above, located either to the left side of

the keel or covering the entire apex H

11.(10) Fringe of keel hood nearly transverse, simply concave; sec-
ondary keel hood not prominent 12

Fringe of keel hood bent toward apex at middle, then
obliquely concave to right margin of apex of keel; sec-
ondary keel hood prominent G. Jiungerfordi Melin p. 316
12.(11) Pan rather flattened, wider than long, in an oblique plane
in relation to the horizontal plane of the genital
capsule G. viciniis Champion p. 319

Pan more or less thickened, elongate, not in a distinct oblique
plane in relation to the horizontal plane of the genital

capsule 13

13.(12) Posterior portion of lateral margin of pronotum usually ex-
tending more or less posterolaterad toward lateral angle;
color quite variable G. oculatus (Fabricius) p. 298

Posterior portion of lateral margin of pronotum transverse
or nearly so; white coloration more
abundant G. oculatus variegatus (Guerin-Meneville) p. 311

Gelastocoris oculatus oculatus (Fabricius)

(PI. I, figs. 1, 2; PI. Ill, figs. 5, 8; PL IV, fig. 11; PI. V, figs. 27, 28.)

1798. Naucoris ocidata Fabricius, Supplementum Entomologiae Systematicae,

p. 525.
1802. Galgulus oculatus Latreille, Histoire naturelle generale et particuliere

des Crustaces et des Insects, etc., Ill, p. 2.54.

The Family Gelastocoridae (Hemiptera) 299

1803. Naucoris oculatus Fabricius, Systema Rhyngotorum, p. 111.

1804. Galgulus oculatus Latreille, Histoire naturelle generale et particuliere
des Cnistaces et des Insects, etc., XII, p. 287.

1815. G. oculatus Leach, Brewster's Edinburgh Encyclopedia, IX, p. 123.
1825. G. oculatus Le Peletier and Serville, Encyclopedia Methodique, vol. X,

p. 270.
1835. G. oculatus Burmeister, Handbuch der Entomologie, Band 2, Abteil. I,

p. 202.
1843. G. oculatus Amyot and Serville, Histoire Naturelle des Insectes. Hemip-

teres, p. 425.
1859. G. oculatus Dohm, Catalogus Hemipterorum, p. 53.
1868. G. oculatus Stal, Hemiptera Fabriciana, I, p. 133.
1873. G. oculatus Walker, Catalogue of Hemiptera in British Museum, part

VIII, p. 170.
1876. G. oculatus Uhler, Bull. U. S. Geo!. Surv., vol. I, no. 5, p. 336, pi. XXI,

fig. 34.
1876. G. oculatus Stal, Kongl. Svenska Vetenskaps-Akademiens Handlingar,

Band 14, no. 4, p. 137.
1878. G. oculatus Uhler, Wheeler's Kept, to Chief Engineer for 1877, p. 1331.
1878. G. oculatus Uhler, Proc. Boston Soc. Nat. Hist., vol. XIX, p. 440.
1884. G. oculatus Uhler, Standard Natural History, vol. II, p. 263.
1886. G. oculatus Uhler, Bull. Brooklyn Ent. Soc, p. 27.
1891. G. oculatus Summers, Bull. Agric. Exp. Sta., Univ. Tennessee, vol. IV,

no. 3, p. 84.

1893. G. oculatus Osbom, Proc. Iowa Acad. Sci., vol. I, part IV, p. 123.

1894. G. oculatus Uhler, Proc. CaUfomia Acad. Sci., ser. 2, vol. IV, p. 290.

1895. G. oculatus Pittier and Biolley, Inst, fisico geografico Nacional (Costa
Rica), p. 23.

1895. G. oculatus Gillette and Baker, Colorado Agric. Exp. Station Bull., no.

31, ser. 1, p. 62.
1897. Gelastocoris oculatus Kirkaldy, Entomologist, XXX, p. 258. (n. n. for

Galgulus Latreille ) .
1901. G. oculatus Champion, Biologia Centrali-Americana, Rhynchota Het-

eroptera, vol. II, p. 348.

1905. G. oculatus Torre-Bueno, Jour. New York Ent. Soc., vol. XIII, p. 43.

1906. G. oculatus Kirkaldy, Trans. Am. Ent. Soc, vol. XXXII, p. 149.

1907. G. oculatus Torre-Bueno and Brimley, Ent. News, vol. XVIII, p. 433.
1909. G. oculatus Kirkaldy and Torre-Bueno, Proc. Ent. Soc. Washington, vol.

X, p. 180.

1913. G. oculatus Torre-Bueno, Canadian Ent., vol. XLV, no. 2, p. 60.

1914. G. oculatus Barber, Bull. Am. Mus. Nat. Hist., vol. XXXIII, p. 498.
1917. G. oculatus Van Duzee, Catalogue of Hemiptera, Univ. California Publ.

pp. 472-473.
1919. G. oculatus Hungerford, Univ. Kansas Sci. Bull., vol. XI, pp. 46-51
pis. Ill, V, IX.

1922. G. oculatus Hungerford, Univ. Kansas Sci. Bull., vol. XIV, pp. 145-171
pis. XIII, XIV.

1926. G. oculatus Torre-Bueno, Bull. Brooklyn Ent. Soc, vol. 21, no. 5, pp

190-191.
1926. G. oculatus Payne, Jour. Morphology and Physiology, vol. 43, no. 2, pp

299-345, 8 pis.
1929. G. oculatus Melin, Zoologiska Bidrag Fran Uppsala, Band 12, pp

164-167, figs. 6, 8, 9, 23-25.
1929. G. oculatus Martin, Univ. Kansas Sci. Bull. vol. XVIII, no. 4, pp. 357-

360, pi. .58, figs. 9, 10, 11, 12, 13; pi. 59, figs. 1-3, 18, 19c, and 20.
1938. G. oculatus Johnson, Jour. Morph., vol. 62, no. 1, pp. 113-134, 2 pis.
1942. G. oculatus Kevan, Proc. Roval Ent. Soc, London, (a) 17, pts. 10-12,

pp. 109-110.
1952. G. oculatus Ellis, Am. Midland Nat., vol. 48, no. 2, p. 310.

Also referring to this species:

1923. Gelastocoris barberi Torre-Bueno, Connecticut Nat. Hist. Surv. Bull., no
34, p. 393, fig. 38.

300 The University Science Bulletin

1926. Gelastocoris subdmilis Blatchley, Heteroptera of Eastern North America,

p. 1025.
1929. Gelastocoris califomiensis Melin, Zoologiska Bidrag Fran Uppsala, Band

12, p. 167, fig. 26.

Size. Male: Length, 6.1 to 8.0 mm.; width of pronotum, 4.0 to
5.3 mm.; width of abdomen, 3.9 to 5.2 mm. Female: Length, 6.7 to

9.3 mm.; width of pronotum, 4.3 to 5.9 mm.; width of abdomen,

4.4 to 6.0 mm.

Color. Extremely variable both as to color and pattern. Colors
present are black, brown, red, orange, yellow, white and green.
Legs are ringed with brown. It would be an endless and useless
task to try to describe all of the combinations of colors and patterns.
In general, the specimens from the southeastern United States are
smaller and darker, while those from the rest of the United States
tend to be larger and lighter, but both extremely dark and extremely
light forms as well as all intermediates may be found in either area.

Structural characteristics. Apex of head rounded or weakly
pointed; front moderately rough, provided with low, broad eleva-
tions which vary in size with the specimen; ocelli rather small,
equal to or slightly larger than the blisterlike granules of the
hemelytra. Pronotum with the disc moderately rough, though the
degree of roughness varies with the specimen; lateral margin quite
variable, usually with anterior portion subparallel for a distance
equal to two-thirds the length of the ocellocular space, slightly
divergent, then lateral margin extending diagonally posterolaterad
to the lateral angle, in some specimens this portion of the lateral
margin may be nearly transversal, usually this portion of the lateral
margin is convex, more rounded toward the lateral angle, lateral
margin weakly serrate; posterolateral margin extending diagonally
posteromerad to base of embolium, serrate; lateral angle projecting
laterad beyond the base of embolium; width of protonum subequa!
to width of abdomen; base of pronotum lacking short, longitudinal
carinae. Hemelytra extending beyond the end of abdomen; mem-
brane well-developed; basal half of lateral margin of embolium
expanded, width of expansion about three times as wide as largest
blisterlike granule, expansion forming an obtuse angle posteriorly.
Connexivum prominent. The blisterlike granules few in number,
slightly less in width than base of hind tarsus. Spatulate right side
of seventh ventral abdominal segment of male about half covered
by the preceding segment, not extending to the lateral margin
of the abdomen; left side projecting posteriorly only slightly.
Female ventral abdominal segments nearly symmetrical, the incision

The Family Gelastocoridae ( Hemiptera ) 301

may be symmetrical or bent to the left. Keel hood of male geni-
talia broadly triangular, apex blunt, an oval concavity near apex
on the ventral surface, fringe nearly transversal, somewhat con-
cave in center; keel hook present, rather stout, recurved and apex
extending partially over pan; pan with apex thick, but narrow,
about one-half width of base, with a slight depression on ventral
surface, base more flattened, pan as v/hole in plane at right angle
to the dorso\entral axis of the insect; right clasper hook quite
variable, usually apical half digitiform, usually rather thin, trans-
versal portion of base variable as to thickness, there may be a
lateral projection present at the base of digitiform process, this
also is variable in size; tumescence of right clasper not adnate
to basal portion of clasper; anterolateral projection of keel absent.
Location of type. In the Kiel Museum, Kiel, Germany. This
museum was destroyed during the recent war, but the Fabrician
Collection was removed and saved. Doctor H. B. Hungerford
visited the museum in 1928, and he reports that the type has
been partially destroyed by dermestids and that only the hemelytra
and scutellum remain. The type is labeled "Carolina".

Distribiitionol data. This species has been reported from the
United States to Brazil by various workers, but this wo'ker has seen
only three specimens from outside the limits of the United States:
one from northern Mexico, the other t\vo from Ontario. Since
many of the species in this group are very similar in external
appearance, it seems likely that reports of the typical subspecies
from Central and South America are probably misidentifications.
The writer has studied 1500 specimens or more from the follow-
ing localities:

Alabama: Burnsville, July 20, 1930, R. H. Beamer, 1 female;
Coatopa, July 18, 1930, P. W. Oman, 1 male; same place and
date, R. H. Beamer, 2 females; Decatur, July 6, 1939, J. D.
Beamer, 1 male and 1 female; Marion Jet., July 16, 1930, P. W.
Oman, 2 males and 2 females; Mt. Meigs, July 21, 1930, R. H.
Beamer, 1 female; same place and date, L. D. Tuthill, 1 male
and 1 female; Tuskegee, July 22, 1930, P. W. Oman, 1 male
and 1 female.

Arizona: Bill Williams Fork, Aug., F. H. Snow, 6 males and
8 females; Box Canyon, Nhiy 9, 1938, F. Piirker, 1 male; Chiricahua
Mts., Sept. 16, 1935, F. H. Parker, 1 female; Clifton, July 14,
1933, F. Parker, 2 males; Coconino Co., July 1, 1929, L. D. Ander-
son, 1 female; Congress Jet., July, F. H. Snow, 1 female; Doug-

302 The University Science Bulletin

las, Aug., F. H. Snow, 1 female; Fort Yuma, Aug. 4, Hubbard,
1 female; Grand Canyon, Aug. 11, 1927, L. A. Anderson, 2 males
and 1 female; Humphrey's Peak, Aug., F. H. Snow, 1 female;
Maricopa Co., July 1, 1929, L. A. Anderson, 1 female; same place
and date, R. H. Beamer, 1 female; Oak Creek Canon, Aug.,
F. H. Snow, 1 female; same place, July 9, 1941, B. Hodgden, 5
males and 4 females; Prescott, 1 male and 4 females; Sabina
Canyon, Sta. Catalina Mts., July 24, 1940, 2 males; same place,
July 23, 1917, P. A. Munz, 1 male and 1 female; Sierra Ancha,
July 19, 1927, 1 female; Tuscon, F. H. Snow, 28 males and 58
females; Southern Arizona, 2 females.

Arkansas: Fayetteville, Apr. 20, 1895, 1 male and 1 female;
same place, Aug. 1923, 2 females; same place, Apr. 22, 1930,
H, H. Schwardt, 1 male; same place, Apr. 23, H. H. Schwardt, 1
male and 1 female; Lawrence Co., nr. Imboden, June, 1925, B.
C. Marshall, 1 male and 1 female (USNM); Ark., 1 male.

California: Alma, Santa Cruz Mts., Apr. 22, 1931, C. H. Martin,
1 female; Bakersfield, July 14, 1929, P. W. Oman, 2 females;
same place, July 24, 1929, R. H. Beamer, 2 males and 1 female;
Barrett, Cottonwood Creek, Apr. 13, 1930, C. H. Martin, 2 males
and 2 females; Bautista Canyon, July 5, 1931, C. H. Martin, 1
female; Berkeley, April 26, 1933, Jean Linsdale, 1 male and 1
female; Cajon Pass, Aug. 6, 1936, R. H. Beamer, 1 female; Cal-
exico, Aug. 1, 1931, H. W. Capps, 1 female; Claremont, 1916,
M. H. Hatch, 2 males and 1 female; Cypress Rdg., Marion Co.,
March 29, 1921, C. T. Dodds, 1 female; Dulzura, Aug. 9, 1935,
J. Beamer, 1 male and 2 females; Indio, July 24, 1929, P. W.
Oman, 1 female; Lake Tahoe, Aug. 11, 1940, L. C. Kuitert, 2 males
and 3 females; Los Angeles, W. T. Davis, 1 female; Los An-
geles River, Oct. 24, 1915, 1 male; same place, Aug. 15, 1916,
V. Duran, 1 female; Los Gatos Cr., Sept. 24, 1910, 1 female; Los
Penasquitos Creek, Poway V., Apr. 10, 1930, C. H. Martin, 1
male and 4 females; same place, Apr. 11, 1930, C. H. Martin, 1
male; Lucerne, July 17, 1935, R. H. Beamer, 1 male; Marin Co.,
Aug. 3, 1929, L. D. Anderson, 2 males and 2 females; same place,
no date or collector, 1 female; Mount Shasta, no date or collector,
1 male and 2 females; Palo Alto, Apr. 13, 1893, no collector, 1
female; same place, Apr. 24, 1893, no collector, 1 female; same
place, Apr. 27, 1893, no collector, 2 males and 1 female; Pasa-
dena, no date or collector, 1 male and 4 females; Pine Valley,
July 27, 1938, R. I. Sailer, 1 male; Rio Hondo, Los Angeles Co.,

The Family Gelastocoridae (Hemiptera) 303

Mar. 21, 1930, C. H. Martin, 1 female; same place, Mar. 23,
1930, C. H. Martin, 1 male and 1 female; same place. Mar. 30,
1930, C. H. Martin, 1 male and 2 females; Rosamond, July 23,
1940, L. C. Kuitert, 1 male and 3 females; Salinas River, San
Ardo, Apr. 25, 1930, C. H. Martin, 1 male and 1 female; San
Diego, no date, Drake, 1 male; San Diego Co., Lake Hodges,
Aug. 5, 1937, F. W. Furry, 1 female; San Gabriel River, March,
1930, C. H. Martin, 1 male and 1 female; San Jacinto Mts.,
July 21, 1929, R. H. Reamer, 1 female; same place and date,
P. W. Oman, 1 male; San I.uis Rey River, Apr. 15, 1930, C. H.
Martin, 1 male and 2 females; San Mateo Creek, May 8, 1930,
C. H. Martin, 1 male; San Mateo Co., no date, Heidemann, 1
female; San Pablo, Alameda Co., Apr. 30, 1921, C. T. Dodds,
2 males; S. Pablo Val, May 9, 1921, C. T. Dodds, 1 male and
1 female; Santa Ana Canyon, June 8, 1930, D. W. Martin, 2
males and 1 female; Santa Margarita Creek, May 8, 1930, C.
H. Martin, 1 male and 1 female; Santa Margarita River, Apr.
8, 1930, C. H. Martin, 3 females; Santa Monica, Mar. 26, 1879,
1 male (Milw. Mus.); Sonoma Co., no date or collector, 1
male and 1 female; Stanford U., Sept. 12, 1909, no collector, 1
male; same place, Oct. 9, 1909, no collector, 1 male and 1 fe-
male; same place, Nov., 1910, no collector, 1 male; Sunol, May
8, 1921, C. T. Dodds, 1 male and 1 female; Sweetwater River,
Descanso Jet., Apr. 13, 1930, C. H. Martin, 2 males and 1 female;
Sweetwater River, San Diego Co., no date, W. J. Chamberlin, 2
males (Mich. Coll.); Warm Spring Creek, Temecula, April 18,
1930, C. H. Martin, 1 male and 1 female; Warner Sprgs., July
28, 1938, R. I. Sailer, 1 male and 1 female; Winters, Aug. 6,
1929, R. H. Reamer, 5 males and 7 females; same place and
date, P. W. Oman, 2 males and 1 female; Vina, May 15, 1921,
C. T. Dodds, 2 males; California, no date or collector, 1 female;
Tehama Co., Redblulf, Aug. 28, 1926, C. L. Hubbs, 1 female.

Colorado: Colorado Springs, Aug., E. S. Tucker, 1 female; Fort
Colhns, Aug. 12, 1931, M. W. Sanderson, 1 male and 2 females; same
place, July 16, 1900, no collector, 2 males.

District of Columbia: Brightwood, Apr. 30, 1901, Heidemann, 1
male; same place. May 20, Heidemann, 1 male; Rock Creek, June 17,
1893, Heidemann, 1 female; same place, July 8, 1893, Heidemann, 1
male; same place, June 19, 1902, Heidemann, 1 female; Washington,
June, 1893, Heidemann, 1 female; same place. Mar. 2, 1894, Heide-
mann, 1 male; same place, June 8, 1894, Heidemann, 1 male; same

304 The University Science Bulletin

place, June 29, Heidemann, 1 male; same place, July 8, Heidemann,
2 females.

Florida: Biscayne, May 24, 1887, Heidemann, 1 male; Branford,
July 31, 1930, R. H. Beamer, 1 male and 1 female; Chokolookee, no
date or collector, 1 male; Crescent City, Apr. 1908, Van Duzee, 1
male and 1 female; Farmdale, June 27, 1948, E. L. Todd, 4 males
and 4 females; Floral City, July 7, 1948, E. L. Todd, 8 males and 6
females; Ft. Pierce, Aug. 7, 1930, R. H. Beamer, 1 male and 1 female;
Fort Walton, June 27, 1948, E. L. Todd, 1 female; Gulf Port, no
date, A. G. Reynolds, 6 males and 2 females; Hilliard, Aug. 31,
1930, R. H. Beamer, 1 male and 1 female; Ind. River, no date
or collector, 1 female (Milw. Mus. ); Inglis, July 7, 1948, E. L. Todd,
6 males and 3 females; Labelle, Apr. 19, D. M. DeLong, 1 male;
same place, July 16, 1939, R. H. Beamer, 2 females; same place,
July 19, 1948, B. T. McDermott, 10 males and 8 females; same place
and date, H. W. Crowder, 1 male and 1 female; same place and
date, E. L. Todd, 54 males and 43 females; same place, Apr. 19,
J. N. Knull, 5 females; same place and date, D. M. DeLong, 2 males
and 2 females; Lacoochee, Aug. 18, 1930, P. W. Oman, 2 males
and 1 female; same place, July 7, 1948, E. L. Todd, 16 males and

14 females; Lake Placid, Feb. 13, 1943, M. Cazier, 1 female; same
place, July 13, 1948, B. T. McDermott, 3 males and 4 females; same
place and date, H. W. Crowder, 7 males and 3 females; same place
and date, E. L. Todd, 24 males and 16 females; Leon Co., Lake
Jackson, June 22, 1922, J. S. Alexander, 1 male and 1 female; Liberty
Co., Apr. 16, 1938, F. E. Lutz, 1 female; Lake Worth, Feb. 6, 1924,
C. S. Bromley, 1 female; Liberty Co., Rock Bluff, July 19, 1930,
H. Spieth, 1 female; M'Glamery Co., Apr. 2, 1927, C. O. Bare, 18
males and 13 females; Moore Haven, Apr. 17, D. M. DeLong, 2
males; same place and date, J. N. Knull, 1 male and 3 females; Otter
Creek, July 6, 1948, B. T. McDermott, 7 males and 6 females; same
place and date, R. H. Beamer, 2 females; same place and date, H.
W. Crowder, 6 males and 2 females; same place and date, E. L.
Todd, 26 males and 23 females; Paradise, Ky., Apr. 10, J. N. Knull,
4 males and 2 females; Pensacola, June 26, 1948, R. H. Beamer,
1 male and 2 females; Perrine, no date or collector, 22 males and

15 females; Royal Palm Pk., Sept. 18, 1930, no collector, 1 male
and 2 females; same place, Oct. 18, 1930, no collector, 1 male and
1 female; Suwannee Sprgs., July 3, 1948, B. T. McDermott, 1 female;
Tavures, July 18, 1894, Heidemann, 2 females; Torreya Ravine,
Liberty Co., Apr. 15, 1938, F. E. Lutz, 1 male and 1 female;

The Family Gelastocoridae (Hemiftera) 305

Wakulla Sprgs., July 14, 1934, R. H. Beamer, 1 male; Wakulla, June

29, 1948, R. H. Beamer, 3 males and 4 females; same place and
date, B. T. McDermott, 7 males and 13 females; same place and
date, H. W. Crowder, 6 males and 5 females; same place, June 30,
1948, E. L. Todd, 12 males and 8 females; Winter Park, Mar. 21,
1938, F. E. Lutz, 1 female; Fla., no date or collector, 1 male and
1 female.

Georgia: Atlanta, May 14, 1911, J. C. Bradley, 1 male; Clayton,
May 18-26, 1911, J. C. Bradley, 1 male and 1 female; Floyd Co.,
Rome, July 13, 1930, C. F. Byers, 1 female; Okefenokee Swamp,
Aug. 3, 1934, R. H. Beamer, 1 female; same place, July 25, 1939,
D. E. Hardy, 1 female; same place and date, J. D. Beamer, 5 males
and 5 females; same place, Aug. 1, 1939, R. H. Beamer, 2 males;
Spring Creek, Decatur Co., June 7-23, 1911, J. C. Bradley, 2 males
and 4 females; Thomasville, July 31, 1927, C. H. Martin, 1 female;
same place, Aug. 21, 1927, C. H. Martin, 1 male and 1 female;
same place, Aug. 28, 1927, C. H. Martin, 1 female; same place, June

30, 1948, B. T. McDermott, 1 female; same place and date, E. L.
Todd, 3 females; same place. Mar. 26, 1903, no collector, 1 male.

Idaho: Parma, Sept. 4, 1934, C. H. Martin, 1 male; Juliaetta, no
date or collector, 1 male and 1 female; Id., no date or collector, 1
male.

Illinois: Villa Ridge, June 7, 1892, no collector, 1 male.

Indiana: Jefferson Co., Hanover, Aug. 28, 1922, T. H. Hubbell,
1 female (Mich. Coll.); Kosciusko Co., May 13, 1932, G. E. Gould,
1 male and 1 female; Lake Co., Gary, no date, H. Ramstadt, 1
female (Mich. Coll.); Warsaw, Apr. 19, 1932, G. E. Gould, 1 male
and 1 female.

Kansas: Bourbon Co., 1915, R. H. Beamer, 1 male; Chanute, no
date, G. Wiley, 3 males; same place, Dec. 20, 1919, H. B. Hunger-
ford, 1 male and 1 female; Cherokee Co., Aug. 1920, H. B. Hunger-
ford and R. H. Beamer, 7 males and 10 females; same place, Aug.
29, 1926, R. H. Beamer, 3 males and 3 females; Cold water, Sept. 7,
1923, C. O. Bare, 3 males and 1 female; same place, Apr. 1925,
R. H. Beamer and C. O. Bare, 5 males and 4 females; same place,
June 19, 1927, R. H. Beamer, 1 male and 1 female; Comanche
Co., Aug. 1922, C. O. Bare, 2 males and 2 females; same place,
Aug. 23, 1924, C. O. Bare, 5 males and 5 females; Douglas Co., Oct.
23, 1926, H. B. Hungerford, 1 male and 1 female; same place, Oct.
8, 1946, B. Hodgden, 2 males; same place, Apr. 4, 1948, G. T.
Brooks, 1 male and 3 females; same place, Apr. 8, 1950, W. Smith,

306 The University Science Bulletin

3 males and 1 female; Ellsworth Co., July 3, 1923, C. H. Martin,

1 male; Graham Co., Aug. 16, 1912, F. X. Williams, 5 males and 2
females; Great Bend, Oct., 1920, Cavanaugh, 5 females; Lake View,
Oct., 1925, H. B. Hungerford, 1 male and 1 female; Logan Co., no
date, F. X. Williams, 3 males and 4 females; Meade Co., 13 mi.
SW Meade, Aug., 1939, C. W. Hubbard, 1 female (Mich. Coll.);
same place, no date or collector, 1 male; Montgomery Co., Elk City,
1927, H. B. Hungerford, 2 males and 1 female; Morton Co., July 20,
1924, C. O. Bare, 21 males and 15 females; Neosho Co., 1919,
G. Wiley, 5 males and 5 females; Norton Co., Aug. 24, 1912, F. X.
Williams, 1 male; Onaga, July 13, 1925, R. H. Beamer, 1 female;
Pawnee Co., July 26-Aug. 2, 1917, no collector, 7 males and 3
females; same place, no date, H. B. Hungerford, 7 males and 10
females; Reno Co., Oct. 4, Lantz, 1 male; Rice Co., July 3, 1923,
R. H. Beamer, 6 males; same place, July, 1923, C. H. Martin, 1
female; same place and date, L. C. Woodruff, 1 female; Riley Co.,
Apr. 22, 1924, R. H. Beamer, 1 female; same place, Sept. 20, J. B.
Norton, 17 males and 3 females; Saline Co., July 14, 1923, W. B.
Whitlow, 1 male and 3 females; Shawnee Co., Apr. 1, 1945, A. M.
Egbert, 1 female; Wallace Co., no date, F. X. Williams, 1 male and
3 females; same place, no date or collector, 1 female; Yates Center,
July 30, 1923, R. H. Beamer and P. B. Lawson, 1 male and 1 female.

Kentucky: Pineville, July 20, 1924, G. P. Engelhardt, 2 males and

2 females.

Louisiana: Aloha, Dec. 23, 1931, R. H. Beamer, 1 female; Coving-
ton, June 23, 1948, R. H. Beamer, 3 males and 2 females; same place
and date, E. L. Todd, 14 males and 15 females; Creole, June 17,
1948, R. H. Beamer, 2 males; same place, June 18, 1948, L. D.
Beamer, 1 male; same place and date, E. L. Todd, 7 males and
9 females; Denham Sprgs., June 20, 1948, E. L. Todd, 1 female;
Madison Parish, Allig. bayou, July 4, 1930, R. Bunn, 2 males and
1 female; Mandeville, June 24, 1948, R. H. Beamer, 1 male; same
place and date, B. T. McDermott, 1 female; same place, June 25,
1948, E. L. Todd, 2 males and 3 females; Mound (No. 276), May
25, 1918, no collector, 2 males and 4 females; same place (No. 301),
June 11, 1918, no collector, 2 males; Natchitoches Parish, Apr. 30,
1915, K. P. Schmidt, 1 male and 2 females.

Maryland: Glen Echo, June 9, O. Heidemann, 1 female; Grt. Falls,
May 13, 1890, Heidemann, 1 male; Plum Island, July 18, 1909, G. P.
Engelhardt, 1 female; Md., July 4, 1886, Heidemann, 1 female.

Michigan: Ann Arbor, May 20, 1932, no collector, 1 male ( Mich.

The Family Gelastocoridae (Hemiptera) 307

Coll.); same place, May 30, 1929, W. Clanton, 1 male and 1
female (Mich. Coll.); Berrien Co., New Buffalo, Sept. 2, 1919,
R. F. Hussey, 10 males and 7 females (Mich. Coll.); Berrien Co.,
E. K. Warren Preserv., Sand Dunes, Aug. 31, 1919, R. F. Hussey,
25 males and 18 females (Mich. Coll.); Berrien Co., E. K.
Warren Preserv., Warren Woods, Sept. 1, 1919, R. F. Hussey, 3
males and 2 females (Mich. Coll.); Huron Co., Port Austin,
Aug. 28, 1924, F. M. Gaige, 21 males and 13 females (Mich.
Coll.); Huron Co., Sand Point, July 30, 1934, I. J. Cantrall, 1
female (Mich. Coll.); same place, Sept. 13, 1927, F. M. Gaige,
1 male (Mich. Coll.); Midland Co., Apr. 4, 1932, R. R. Dreisbach,
1 male (Mich. Coll.); Midland Co., Chippewa R., 8 mi. W
Midland, July 15, 1935, A. Olson-L. K. Gloyd, 1 female (Mich.
Coll.); Monroe Co., 3 mi. W Temperance, Aug. 12, 1941, I. J.
Cantrall, 5 females (Mich. Coll.); Montcalm Co., no date or
collector, 2 females (Mich. Coll.); Ottawa Co., Sept. 5, 1940,
G. Orton, 1 male and 1 female (Mich. Coll.); Sanilac Co., 36
mi. S Port Sanilac, June 29, 1941, A. Watson, 1 male and 1
female (Mich. Coll.); Walnut Lake, June 26, 1906, no collector,
3 females; Washtenaw Co., Apr. 28, 1932, no collector, 1 male
(Mich, Coll.); same place, May 18, 1933, no collector, 1 male
(Mich. Coll.); same place, May 23, 1933, no collector, 2 males
(Mich, Coll.); same place, Apr. 23, 1935, no collector, 1 female
(Mich. Coll.); Washtenaw Co., Ann Arbor, May 18, 1933, R.
M, Bailey, 1 male (Mich. Coll.); same place. May 24, 1932,
R. M. Bailey, 1 female (Mich. Coll.); same place. May 8, 1937,
J, J. Friauf, 3 males and 3 females (Mich. Coll.); Washtenaw
Co., Irish Hills, May 15, 1932, R. M, Bailey, 1 female (Mich.
Coll.); Washtenaw Co., North Lake, Sept. 22, 1935, L. J. Can-
trall, 1 female (Mich. Coll.); Washtenaw Co., Sylvan Twp.,
Crooked Lake, May 8, 1937, D. L, and I. J. Cantrall, 6 males
and 2 females (Mich. Coll.); Washtenaw Co., Third Sister
Lake, Sept. 1, 1935, L, K. Gloyd, 3 males (Mich. Coll.); Wash-
tenaw Co., Whitmore Lake, May 22, 1932, R. M. Bailey, 1
female (Mich. Coll.); Cheboygan Co., Maple River, Douglas L,
and Black Lake, H. B. Hungerford, not common,

Mississippi: Hamilton, July 15, 1930, R. H. Reamer, 1 male
and 2 females; luka, July 14, 1930, R, H, Reamer, 1 male and
1 female; Pass Christian, June 25, 1948, R. H. Reamer, 2 males
and 3 females; same place and date, E. L. Todd, 6 males and
10 females; Smithville, July 15, 1930, R, H. Reamer, 1 female;
Tuskegee, July 22, 1930, R. H. Reamer, 3 females.

308 The University Science Bulletin

Missouri: lilack R., Clark Nat. Forest, July 22, 1937, C. P.
Brown, 1 female (Mich. Coll.); Blackwell, May, 1910, J. F.
Abbot, 1 female; Hollister, July 28, 1929, no collector, 1 male
and 1 female; Postosi, June 28, 1923, W. J. Clench, 1 female;
St. Louis, Creul-Coenir Lake, May, 1910, J. F. Abbot, 1 male;
Van Buren, Ozark Mts., June 7, 1930, E. A. Pence, 2 females
(Mich. Coll.); same place, June 16, 1930, E. A. Pence, 3 males
and 1 female (Mich. Coll.); same place, June 12, 1930, E. A.
Pence, 2 males (Mich. Coll.); same place, June 20, 1930, E. A.
Pence, 1 male (Mich. Coll.); same place, July 9, 1930, E. A.
Pence, 1 female (Mich. Coll.).

Nebraska: Jefferson Co., Apr. 20, 1924, H. B. Hungerford
and R. H. Beamer, 2 females.

Nevada: Carson City, Aug. 9, 1929, R. H. Beamer, 1 female;
same place, July, 1930, Wickham, 1 female; Reno, Sept. 14,
1940, LaR., 1 female; same place, Apr. 19, 1941, LaR., 1 female.

New Jersey: Anglesea, May 30, no collector, 1 male; Lake-
hurst, Sept. 4, 1911, F. M. Schott, 2 females; Rancoca, Aug. 29,
1927, E. M. Becton, 4 males and 4 females; Trenton, Aug. 8, no
collector, 1 male.

New Mexico: Catron Co., Aug. 15, 1938, J. Hildalgo Jr., 5
males and 4 females; Monzano, June 26, 1941, B. Hodgden, 1
male; State College, Dona Ana Co., Oct. 5, 1936, no collector,

1 female; Valencia Co., McCartys, Sept. 19, 1926, C. L. Hubbs,

2 females ( Mich. Coll. )

North Carolina: Charlotte, July 20, 1936, C. D. Lyman, 1
male (Mich. Coll.); Haywood Co., Crestmont, July 27, 1922, T.
H. Hubbell, 3 females (Mich. Coll.); Hertford, no date, W. R.
Walton, 1 male (USNM); Lake Ellis, May 8, 1906, no collector,
1 male; Roanoke Is., July 25, no collector, 2 females; Raleigh,
Mar. 21, 1905, no collector, 3 males; same place. Mar. 24, 1905,
no collector, 2 males and 3 females; same place. Mar. 25, 1905,
no collector, 1 male; same place. Mar. 31, 1906, C. S. Brimley,
1 male and 2 females; same place, Apr. 7, 1906, C. S. Brimley, 1
male and 1 female; same place, Apr. 13, 1906, C. S. Brimley, 3
males and 3 females; same place. May 25, no collector, 1 male
and 1 female; Southern Pines, May 15, 1919, A. H. Manee, 3 males;
Whitney, Apr. 22, 1907, no collector, 1 female; North Carolina,
no date or collector, 1 female.

Ohio: Columbus, no date or collector, 1 female; Hooking Co.,
Good Hope Twp., Sept. 17, 1931, R. M. Bailey, 1 female (Mich.

The Family Gelastocoridae (Hemiptera) 309

Coll.); Lucas Co., Spencer Twp., Sept. 24, 1938, I. J. Cantrall,
12 males and 13 females ( Mich. Coll. ) ; Lucas Co., 6 mi. W Toledo,
Sept. 24, 1938, I. J. Cantrall, 2 males and 2 females (Mich. Coll.).
Oklahoma: Comanche Co., Wichita Natl. Forest, June 11,
1926, T. H. Hubbell 1 male (Mich. Coll.); McCurtain Co.,
Glover, June 16, 1938, W. F. Blair, 1 male.

Oregon: Boardman, July 15, 1931, L. D. Anderson, 1 male;
Corvallis, Sept. 16, 1897, no collector, 1 male; same place, Sept.
28, 1899, no collector, 1 female; same place, 1906, Woods, 1
female; same place, Aug. 1907, no collector, 1 female; same
place, Sept. 21, 1907, no collector, 1 female; Dilley, Aug. 20,
1902, no collector, 8 males and 9 females; Dixie, July 9, 1931,
L. D. Anderson, 1 male; Eugene, July 15, 1941, Malkin, 1 female;
Freewater, Sept. 8, 1914, G. F. Moznette, 1 male; same place.
Sept. 9, 1914, G. F. Moznette, 1 male; same place, Sept. 21.
1914, G. F. Moznette, 1 female; Grant's Pass, Rogue R., Sept.
14, 1930, F. H. C, 1 male and 1 female; same place, Sept. 24,
F. R. C, 1 female; Hood R., July 17, 1931, L. D. Anderson, 1
female; Monroe, June 18, 1930, M. H. Hatch, 1 female; Philo-
math, 1905, no collector, 1 female; St. Angel, no date, Heidemann,
1 male.

South Carolina: Cheraw, Apr. 24, 1938, F. E. Lutz, 1 male
(AMNH); same place, Apr. 14, 1938, F. E. Lutz, 2 females
( AMNH); same place, Apr. 7, 1938, F. E. Lutz, 1 female (AMNH).
Tennessee: Clarksville, July 3, 1939, E. G. Wegenek, 1 female;
Coal Creek, June 28, 1916, W. S. Adkins, 2 males; Cumberland
Co., Grassy Cove, July 14, 1922, T. H. Hubbell, 1 male (Mich.
Coll.); same place, July 7, 1922, T. H. Hubbell, 1 female (Mich.
Coll.); Elkmont, May 4, 1913, no collector, 1 female; Kenton,
Aug. 4, 1948, B. T. McDermott, 3 males; Knoxville, June 6, 1890,
no collector, 2 males and 2 females; same place, Apr. 25, 1911, no
collector, 3 males and 2 females; same place, Apr. 11, 1914, no
collector, 1 female; same place. May, 1914, no collector, 1 male and
1 female; same place, July 15, 1919, no collector, 1 male; same
place, June, 190.^, no collector, 1 male; Obion Co., Reelfoot Lake,
Sept. 8, 1919, F. M. Gaige, 6 males and 5 females (Mich. Coll.);
same place, Sept. 6, 1919, F. M. Gaige, 3 males and 5 females (Mich.
Coll.); Perryville, July 20, 1926, T. E. White, 1 female; Scott Co.,
Big South Fork of Cumberland River, July 24, 1924, T. H. Hubbell,
1 female (Mich. Coll.); Thunderhead Mt, Blount Co., July 19,
1927, G. Steyskal, 2 males (Mich. Coll.).

310 The University Science Bulletin

Texas: Austin, Oct. 12, 1923, R. K. Nabors, 6 males; same place,
Oct. 1899, no collector, 1 female; same place, Oct. 1900, A. L. Me-
lander, 3 males and 1 female; same place, Nov. 18, 1899, no col-
lector, 1 male; Burton, June 24, 1938, L. W. Hepner, 2 females;
Cisco, June 19, 1947, R. Olmsted, 1 male; same place and date,
L. D. Beamer, 1 female; Colorado Co., Apr. 3, 1922, G. Wiley, 17
males and 11 females; same place, Apr, 7, 1922, G. Wiley, 2 males;
same place, Apr. 11, 1922, G. Wiley, 13 males and 11 females; same
place, Apr. 14, 1922, G. Wiley, 1 female; same place, Apr. 24, 1922,
G. Wiley, 3 males; same place. May 6, 1922, G. Wiley, 1 male;
same place. May 11, 1922, G. Wiley, 1 male and 1 female; same
place, May 15, 1922, G. Wiley, 2 males; same place. May 16, 1922,
G. Wiley, 9 males and 3 females; Comal Co., no date or collector,
1 male and 3 females; Concan, July 6, 1936, R. H. Beamer, 1
female; Dallas Co., Apr. 14, 1938, Cole, 1 female; same place. Mar,
9, 1939, D. D. Millspaugh, 3 males and 1 female; same place,
summer, 1931, J. K. G. Silvey, 3 females (Mich. Coll.); Eastland
Co., May 6, 1921, G, Wiley, 1 female; same place. May 12, 1921,
G. Wiley, 1 male; same place, May 17, 1922, G. Wiley, 1 male and
1 female; Guadalupe Co., 6 mi. N New Braunfels, Feb. 22,
1946, I. J. Cantrall, 3 males and 1 female (Mich. Coll.); Peeler,
June 22, 1938, D. W. Craik, 1 male and 1 female; Sequin, June
26, 1938, R. I. Sailer, 1 female; Tarrant Co., summer 1931, J. K. G.
Silvey, 1 female (Mich. Coll.); Webb Co., summer 1931, J, K, G,
Silvey, 1 male (Mich. Coll.); Willis, May, 1903, Bridwell, 1 female;
Texas, no date or collector, 3 males and 2 females; Texas, summer
1931, J. K. G. Silvey, 2 males and 1 female.

Utah: Beav. Cany., no date or collector, 1 male and 1 female.

Washington: Dayton, Apr. 17, 1934, C. H. Martin, 1 male and
1 female; Kalama R., July 21, 1931, L. D. Anderson, 3 males; same
place and date, J. Nottingham, 1 female; Mason Co., Lake Cush-
man, June 24, 1919, F. M. Gaige, 1 female (Mich. Coll.); Pasco,
Apr. 17, 1934, C. H. Martin, 2 males and 1 female; Pullman, no
date, C. V. Piper, 2 females.

Wisconsin: Milw. Co., Sept. 1, 1900, C. E. Brown, 1 female
(Milw. Mus.).

Ontario: Lambton Co., Stag Isl., Sept. 26, 1927, S. Moore, 1
male and 1 female (Mich. Coll.).

Mexico: Rio Corona, 18 mi. NE Victoria, Tamaulipas, Dec. 17,
1941, Cantrall-Friauf, 1 female (Mich. Coll.),
Africa: Cape of Good Hope, no date or collector, 2 females.

The Family Gelastocoridae (Hemiptera) 311

These specimens quite obviously have been mislabeled, they may
have come from Good Hope Twp., Ohio, a locality from which
the writer has seen other specimens.

Comparative notes. With the exception of G. rottindatus Cham-
pion, this is the only species the writer has seen from within the
limits of the United States. It may be separated from the closely
related species by the shape of the lateral margin of the pronotum
and/or male genitalia. G. californiensis Melin is in the opinion of
the writer a synonym of G. oculatiis oculatiis ( Fabr. ) and is so
treated. The characters utilized in the separation of this species
fall within the range of variation of G. oculatus oculatus ( Fabr. ) and
are therefore not valid.

Gelastocoris oculatus variegatus (Guerin-Meneville)

(PL IV, fig. 16; PI. VI, fig. 31)

1844. Galgulus variegatus Guerin-Meneville, Iconographie du Regne Animal

de B. Cuvier., part 7, p. 352.
1859. G. variegatus Dohm, Catalogue Hemipterorum, p. 53.
1873. G. variegatus Uhler, Bull. U. S. Geol. Survey, vol. I, no. 5, pp. 336-337.
1876. G. variegatus Stal, Kongl. Svenska Vetenskaps-Akademiens Handlingar,

Band 14, no. 4, p. 137.
1878. G. variegatus Uhler, Wheeler's Rept. to Chief of Engineers for 1877,

p. 1331.
1886. G. variegatus Uhler, Bull. Brooklyn Ent. Soc, p. 27.
1894. G. variegatus Uhler, Proc. California Acad. Sci., ser. 2, vol. IV., p. 290.
1901. Gelastocoris variegatus (Guerin-Meneville); Champion, Biologia Cen-

trah-Americana, Rhynchota Heteroptera, vol. II, pp. 349-350, pi. XX,

figs. 19, 20.
1906. G. variegatus Baker, Brooklyn Inst., Sci. Bull., vol. I, p. 288.

1909. G. variegatus Kirkaldy and Torre-Bueno, Proc. Ent. Soc. Washington,
vol. X, p. 181.

1910. G. variegatus Montandon, Annuario del Museo Zoologico della R. Uni-
versita di Napoli (nuova serie), vol. 3, no. 10, p. 2.

1910. G. variegatus Banks, Cat. Nearctic Hemip-Heteroptera, p. 11.

1914. G. variegatus Van Duzee, Trans. San Diego Soc. Nat. Hist., II, p. 33.

1917. G. variegatus Van Duzee, Catalogue of Hemiptera, Univ. California

Publ, p. 473.
1929. G. variegatus Melin, Zoologiska Bidrag Fran Uppsala, Band 12, pp.

162-163, figs. 7, 21.
1929. G. variegatus Martin, Univ. Kans. Sci. Bull., vol. XVIII, no. 4, p. 362,

pi. 58, fig. 8; pi. 59, fig. 10.
1939. G. variegatus Millspaugh, Field and Laboratory, vol. VII, no. 2, p. 82.
1952. G. variegatus EUis, Am. Midland Nat., vol. 48, no. 2, p. 310.

Also referring to this subspecies:

1854. Galgulus pulcher Stal, Ofversigt af Kongl. Vetenskaps-Akademiens For-

handlingar. Band XI, no. 3, p. 239.
1859. G. pulcher Dohrn, Catalogus Hemipterorum, p. 53.
1873. G. pulcher Walker, Catalogue of Hemiptera in British Museum, part

VIII, p. 171.
1876. G. pulcher Stal, Kongl. Svenska Vetenskaps-Akademiens Handlingar,

Band 14, no. 4, p. 137.
1909. Gelastocoris pulcher (Stal); Kirkaldy and Torre-Bueno, Proc. Ent. Soc.

Washington, vol. X, p. 180.
1929. G. pulcher Martin, Univ. Kansas Sci. Bull., vol. XVIII, no. 4, p. 362.

312 The University Science Bulletin

Size. Male: Length, 5.7 to 6.5 mm.; width of pronotum, 3.4 to

4.2 mm,; width of abdomen, 3.5 to 4.2 mm. Female: Length, 6.6
to 7.5 mm.; width of pronotum, 4.3 to 4.7 mm.; width of abdomen,

4.3 to 4.8 mm.

Color. Variegated, in general appearance usually light brown
with patches of white and brown or black. Some specimens with
areas of red, the amount varying with the individual specimen.
Below, with the abdominal segments lighter laterally; legs ringed
with light brown.

Structural characteristics. Apex of head rounded or slightly
pointed; front of head with moderate, broad, elevations; ocelli
rather small, slightly smaller than the largest blisterlike granules.
Disc of pronotum not very rough; lateral margin with anterior
portion subparallel to longitudinal axis of body, for a distance
equal to two thirds the length of the ocellocular space, then lateral
margin extending almost directly laterad to the lateral angle, pos-
terolateral margin strongly projecting for anterior two thirds, pos-
terior third converging from the projection to the posterior angle
of pronotum, anterior two thirds of posterolateral margin very
strongly serrate; lateral angle projecting beyond base of embolium;
width of pronotum subequal to width of abdomen; base of pro-
notum lacking short, longitudinal carinae. Hemelytra extending
to or beyond end of abdomen; membrane well developed; basal
half of lateral margin of embolium expanded, about two times as
wide as largest blisterlike granules, lateral margin serrate, expan-
sion angulate posteriorly. Connexivum visible in both sexes, but
a little more evident in the females than in the males. The blister-
like granules not very numerous, usually five or six in number on
each hemelytron, largest granule about as wide as base of hind
tarsus. Seventh ventral abdominal segment of male with the
spatulate right side partially covered by preceding segment, spat-
ulate right side short not reaching lateral margin, broad, quite
rounded and constricted at base of spatula; left side projecting pos-
teriorly only slightly. Female ventral abdominal segments nearly
symmetrical, incision bent to the left. Male genitalia practically
identical to those of G. oculatus oculatus (Fabricius).

Location of type. Unknown. Guerin-Meneville's types were in
the Museo Zoologico della R. Universita di Napoli, Italy. This
museum building was destroyed during World War II and I have
not been able to determine whether the collections had been re-
moved or whether they were lost.

The Family Gelastoc:oridae (Hemiptera) 313

Distributional data. Described from the Bay of Campeche, Mex-
ico, this subspecies seems to be restricted to Central America and
Mexico. The writer has seen authenic G. o. variegatus (Guerin-
Mcneville) from Brownsville, Texas, and it is possible that this
species might be found in other semitropical areas of the south-
western United States but the writer has yet to see such specimens.
Champion gives the distribution of this species to be southern and
southwestern United States to Argentina. The writer has not seen
this species from any of the South American countries and it seems
quite unlikely that the species occurs in any of the southern states
of the United States. Specimens from the following localities have
been examined:

Texas: Brownsville, Oct. 13, 1944, R. L. Usinger, 1 male (Usinger
Coll.). Other specimens from Brownsville are in the collections
of the U. S. National Museum.

Mexico: San Bernardo, Rio Mayo, Sonora, Oct. 14, 1934, H. S.
Gentry, 2 females; Oaxaca, Oax., Sept. 19, 1923, E. G. Smyth, 1
female (USNM); same place, Aug. 20, 1937, M. Embury, 3 males;
Vera Cruz, V. C, no date, Wickham, 2 males and 1 female.

Guatemala: Gualan, Jan. 14, 1905, no collector, 6 males and 2
females; same place and date, Drake, 1 male; Los Amates, Jan.
16, 1905, no collector, 4 males and 2 females; same place, no date
or collector, 1 male and 2 females (MCZH).

Honduras: Progreso, Nov. 29, 192.3, T. H. Hubbell, 1 male and
1 female.

British Honduras: Rio Grande, Jan. 1932, J. J. White, 17 males
and 11 females.

Tanamci: Progreso, Chiriqui Prov., Apr. 18, 1923, F. M. Gaige, 4
males and 1 female.

Comparative notes. The original description was based entirely
upon color. Because of this numerous specimens of G. ocidatus
ocidatus (Fabr.) and other species have been identified as G.
variegatus (Guerin-Meneville). Champion pointed out that the
middle portion of the lateral margin extends transversely at nearly
right angles to the longitudinal axis of the body. Champion speaks
as if this were well known, so he may have received this informa-
tion from Montandon and assuming that it had previously been
published. On this character alone, it is possible to recognize the
subspecies, though some specimens of G. oculatus oculatus ( Fabr. )
approach this subspecies in the shape of the lateral margin of the
pronotum, but to my knowledge there is no cline for this character.

314 The University Science Bulletin

The genitalia, as mentioned above, are practically identical to those
of the typical subspecies, differing no more from this subspecies
than the individual variation within that subspecies. In consider-
ation of the similarity of the male genitalia, it seems best for the
present to place G. voriegatus (Guerin-Meneville) as a subspecies
of G. ocidatus (Fabricius).

Gelastocoris rotiindatiis Champion

(PI. IV, fig. 18; PL VI, fig. 30)

1901. Gelastocoris rotundatus Champion, Biologia Centrali-Americana,

Rhynchota Heteroptera, vol. II, pp. 347-348, pi. 20, fig. 18.
1909. G. rotundatus Kirkaldy and Torre-Bueno, Proc. Ent. Soc. Washington,

vol. X, p. 180.
1929. G. rotundatus Melin, Zoologiska Bidrag Fran Uppsala, Band 12, pp.

155-156, fig. 10.
1929. G. rotundatus Martin, Univ. Kansas Sci. Bull., vol. XVIII, no. 4, p. 363,

PI. 58, fig. 16, PI. 59, figs. 8, 19e and 19f.
1939. G. rotundatus Millspaugh, Field and Laboratory, vol. VII, no. 2, p. 83.
1951. G. rotundatus Adams and McNeill, Pan-Pacific Entomologist, vol.

XXVII, no. 2, p. 71.

Size. Male: Length, 6.4 to 7.8 mm.; width of pronotum, 4.1 to
5.0 mm.; width of abdomen, 4.5 to 5.5 mm. Female: Length, 7.1
to 8.7 mm.; width of pronotum, 4.5 to 5.3 mm.; width of abdomen,
5.0 to 5.8 mm.

Color: Quite variable as in other species, mostly light yellowish-
brown, others decidedly reddish, grey or black. Specimens may
be nearly concolorous or with distinct patterns, i. e., scutellum and
disc of pronotum black, rest light yellow or yellowish brown.

Structural characteristics. Body almost oval due to peculiar
form of pronotum. Apex of head rounded; front not especially
rough; ocelh moderate in size, slightly larger than largest blister-
like granules. Pronotum with disc rather smooth; lateral margin
of pronotum very slightly concave anteriorly then extending postero-
laterad to lateral angle, usually slightly convex; lateral angle bent
ventrad, not at all visible from above, only very slightly projecting
laterad beyond base of embolium; abdomen wider than pronotum;
base of pronotum lacking short, longitudinal carinae. Hemelytra
extending to or beyond the end of the abdomen; membrane well
developed; basal half of lateral margin of embolium scarcely
expanded laterally, entire lateral margin of embolium nearly smooth
and rounded. Connexivum completely or nearly completely cov-
ered by the hemelytra. Blisterlike granules few, small, the largest
less than width of hind tarsus in size, all granules are small and
rather uniform. Spatulate right side of seventh ventral abdominal

The Family Gelastocoridae (Hemiptera) 315

segment of male short, rounded, not extending to the lateral margin
of the abdomen; left side not or only slightly projecting posteriorly.
Female ventral abdominal segments nearly symmetrical, incision
of the posterior margin of last segment slightly bent to the left.
Keel hood of male genitalia covering apex of keel, fringe more or
less concave in middle and bent rather abruptly anterolaterad on
left side; keel hook present, recurved back toward base of pan;
pan not very large, apex flattened at right angles to the base, round
and with a slight crescent-shaped depression on ventral half; right
clasper hook reduced, apical portion digitiform, short, basal trans-
verse portion widest at base, slight lateral projection present at
base of digitiform apex; tumescence of right clasper not adnate
to base of clasper; anterolateral projection of keel absent.
Location of type. In the British Museum at London, England.

Distributional data. Champion gives the distribution of this
species as Mexico and Guatemala. Martin reported it from Ari-
zona and Millspaugh has reported it from Texas. Adams and
MacNeill have recently reported this species from San Bernardino
Co., California. The writer has examined specimens from the follow-
ing localities:

Texas: Glenn Spring, Brewster Co., June 21, 1928, F, M.
Gaige, 3 males and 8 females, (Mich. Coll.); Kerr Co., Apr.
9, 1939, D. Millspaugh, 2 females.

Arizona: Castle Hot Springs, Aug. 5, 1941, R. H. Beamer, 7
males and 13 females; same place and date B. Hodgden, 25
males and 29 females; same place and date, L. H. Banker, 4
males and 3 females; same place and date, E. L. Todd, 1
male; Douglas, Aug., F. H. Snow, 2 females.

Mexico: Corriente, Aug. 24, 1927, H. D. Thomas, 1 male and
3 females; Colima Vulcano, no date, L. Conrad, 1 male; San
Luis de Barbarocos, Chihuahua, Dec. 30, 1934, H. S. Gentry,
1 female; Garrapatas, Gro., kil. 369 S Mex. City, Oct. 31, 1936,
H, D. Thomas, 1 female; La Sabana, Guerrero, kil. 226 S Mex.
City, Oct. 10, 1936, H. D. Thomas, 9 males and 9 females; Rio
Balsas, jet. Acapulco Hwy., Guerrero, June 24, 1932, H. Smith,
1 female; Tecolotlan, Jahsco, July 16, 1938, H. D. Thomas, 2
females; same place, Sept. 14, 1938, L. J. Lipovsky, 2 males and
1 female; Union de Tula, Jahsco, Sept. 16, 1938, H. D. Thomas,
13 males and 7 females; El Sabino, Uruapan, July 20, 1936, H. D.
Thomas, 5 males and 1 female; Near Chinapa, Michoacan, Sept.
5, 1938, H. D. Thomas, 1 male and 2 females; Tancitaro,

316 The University Science Bulletin

Michoacan, July 29, 1940, H. Hoogstraal, 4 males and 3 females;
Alpuyeca, Morelos, 102 km. S Mexico (city), Oct. 15, 1936, H.
D. Thomas, 1 male; Cuernavaca, Morelos, Aug. 3, 193S, L. J.
Lipovsky, 1 female; Rio Amacuza, Morelos, 133 km. S Mexico
(city), Oct. 14, 1936, H. D. Thomas, 1 male and 1 female; Palmar
Dist., Alamos, Sonora, Oct. 27, 1934, H. S. Gentry, 4 males and
6 females; Arroyo de los Mascales, Rio Mayo, Sonora, Feb. 16,
1935, H. S. Gentry, 2 females; San Bernardo, Rio Mayo, Sonora,
October 14, 1934, H. S. Gentry, 8 males and 6 females; Tejupilco,
Dist. of Temascaltepec, June-July, 1933, H. E. Hinton, 8 males
and 9 females; near Tarandacuao, Aug. 6, 1932, H. Smith, 2
males and 1 female; "Mex.", no date and no collector, 1 female
(MCZH).

Guatemala: Gualan, July 14, 1905, no collector, 1 male; Sonar-
ate, no date, Kellerman, 1 female (MCZH); "Guatemala", no
date or collector, 5 males and 3 females (MCZH), 1 female
compared with type by Doctor H. B. Hungerford.

Honduras: Copan, Feb. 16, 1937, C. Roys, 1 male.

Comparative notes. The oval shape of the body and the
shape of the pan of the male genitalia will separate this species
from G. oculatus (Fabricius) to which it is apparently closely
related.

Gelastocoris hungerfordi Melin

(PI. V, fig. 19; PI. VI, fig. 33)

1929. Gelastocoris hungerfordi Melin, Zoologiska Bidrag Fran Uppsala, Band
12, p. 168, figs. 5, 27-31.

Also referring to this species:

1929. Gelastocoris ctictiUatits Martin, Univ. Kans. Sci. Bull., vol. XVIII, no.

4, pp. 363-364, pi. 58, fig. 6, pi. 59, figs. 11, 12, 13.
1939. G. cucuUatus Millspaugh, Field and Laboratory, vol. VII, no. 2, pp.

82-83.

Size. Male: Length, 5.9 to 6.5 mm.; width of pronotum, 3.5 to

4.1 mm.; width of abdomen, 3.6 to 4.2 mm. Female: Length,

6.2 to 7.4 mm.; width of pronotum 3.9 to 4.5 mm.; width of
abdomen, 4.0 to 4.9 mm.

Color. In general appearance ranging from almost white to
a very dark brown. Both color and pattern extremely variable.
Colors present are red, brown, yellow, white, black, and green.
Below, abdomen variable, from pale yellow to dark brown, but
most specimens have the median area of abdomen darker than
rest of specimen. Legs ringed with brown, front femora fre-

The Family Gelastocoridae (Hemiptera) 317

(juently with rings expanded, fnsed and covering almost the
basal two thirds of the femur, the fusion is not complete and
there are spots of paler coloration present.

Structural characteristics. Apex of head rounded; front not
at all rough; ocelli fairly large, larger than the blisterlike granules
of the hemelytra. Disc of pronotum not very rough; lateral margin
of pronotum subparallel for a distance about equal to ocellocular
space, then extending diagonally posterolaterad to the lateral
angle, this portion of lateral margin slightly convex; posterolateral
margin nearly straight and converging to humeral angle of pro-
notum; lateral angle projecting beyond base of embolium; ab-
domen slightly wider than pronotum; base of pronotum lacking
short, longitudinal carinae. Hemelytra extending beyond end
of abdomen; membrane well developed; basal half of lateral
margin of embolium slightly expanded, expansion equal to one
fourth or one fifth of the ocellocular space, expansion rounded
or obtusely angulate posteriorly. Connexivum usually slightly
exposed, some males have the connexivum completely covered
by the hemelytra. Blisterlike granules of the hemelytra few,
small, less than width of base of hind tarsus. Spatulate right
side of seventh ventral abdominal segment of male short, not
extending to the lateral margin of the abdomen, partially covered
by the preceding segment; left side only slightly projecting
posteriorly. Female ventral abdominal segments symmetrical
except for incision of posterior margin of last segment, which is
strongly bent to the left. Keel hood of male genitalia rather
long, mainly to the left side of keel, but covering most of apex;
membranous secondary keel hood present and nearly as large
as keel hood; fringe of keel hood with left side nearly transverse
and slightly concave, right half extending almost anteriorly to
right margin of keel; keel hook present, slender, recurved to
base of pan; pan small, thick, rounded apically and apex pro-
vided with slight circular depression; right clasper hook, like
that of G. oculatus (Fabricius) and G. rotundatus Champion,
apical portion digitiform, transverse basal portion much wider,
with distinct lateral projections at base of digitiform apex; tumes-
cence of right clasper not adnate to basal portion of clasper;
anterolateral projection of keel absent.

Location of type. In the Museum of Paris, Paris, France,
Distributional Data. The types are from Venezuela. Melin
also examined specimens from Mexico, Guatemala, Nicaragua,

318 The University Science Bulletin

Costa Rica, Panama and Colombia. Millspaugh reports this
species from Texas, but the writer has not seen specimens from
that state. Specimens available for study v^ere from the follow-
ing localities:

Mexico: Acapulco, Guerrero, Aug. 16-30, 1938, L. J. Lipovsky,
1 male; same place, Aug. 28, 1938, L. J. Lipovsky, 1 female;
10 mi. S Chilpancingo, Guerrero, June 23, 1932, H. Smith, 1
female; La Sabana, Guerrero, 226 km. S Mexico (city), Oct.
20, 1936, H. D. Thomas, 27 males and 25 females; Mexcala,
Guerrero, Rio Balsas, Dec, 14, 1928, Dampf., 5 males and 3 fe-
males; Rincon, Guerrero, 260 km. S Mex. City, Oct. 31, 1936,
H. D. Thomas, 2 males and 2 females; Rio Agua, Guerrero,
437 km. S Mexico (city), Oct. 31, 1936, H. D. Thomas, 16 males
and 11 females; Guerrero, 438 km. S Mex. City, Nov. 1, 1936,
H. D. Thomas, 3 males and 3 females; Sacacoyuca, Gro., 216
km. S Mexico (city), Oct. 31, 1936, H. D. Thomas, 3 males
and 2 females; Tecolotlan, Jalisco, Sept. 14, 1938, L. J. Lipovsky,
1 female; Alpuyeca, Morelos, 102 km. S Mex. City, Oct. 15,
1936, H. D. Thomas, 19 males and 26 females; Rio Amacuya,
Morelos, 133 km. S Mexico (city), Oct. 14, 1936, H. D. Thomas,
7 males and 9 females; 5 mi S Valles, San Luis Potosi, June
13, 1932, H. Smith, 2 males and 3 females; Rio Mayo, San
Bernardo, Sonora, Oct. 14, 1934, H. S. Gentry, 2 females; Teju-
pilco, Dist. of Temascaltepec, June-July, 1933, H. E. Hinton, 2
males; Minatitlan, Vera Cruz, Sept. 22, 1936, H. D. Thomas,

1 male.

Guatemala: El Salto Escuintla, 1934, F. X. Williams, 1 male
and 1 female; Gualan, Jan. 14, 1905, no collector (holotype of
G. cucullatus Martin); same place, Jan. 4, 1905, no collector,

2 males and 2 females; Guatemala Cit}% no date. Champion (1
male paratype of G. cucullatus); Los Amates, Jan. 16, 1905, no
collector, 2 males and 1 female (paratypes of G. cucullatus);
same place, no date, Kellerman (4 male paratypes of G. cucullatus).

Honduras: Copan, Feb. 18, 1937, C. Roys, 3 males and 2
females; Progreso, Feb. 23, 1936, J. Deal, 5 males and 4 females.

Nicaragua: "Nicaragua", O. Sacken., 1 male (AMNH); ?. O.
Sacken. (1 male paratype of G. cucullatus) (AMNH).

Costa Rica: Rio Torres, Feb. 10, 1932, H. Schmidt, 3 females;
Rio Virilla, Dec. 26, 1931, H. Schmidt, 55 males and 48 females;
San Jose, June and July, 1931, H. Schmidt, 2 males and 2 fe-
males; San Jose, 1932, H. Schmidt, 16 males and 40 females.

The Family Gelastocoridae (Hemiptera) 319

Panama: Barro Colorado, C. Z., July 13, 1924, N. Banks (1
female paratype of G. cuciilJatus) (MCZH); same place, Dec. 22,

1928, C. H. Curran, 1 male (AMNH); Caldera, Feb. 28, 1935,
D. V. Brown, 38 males and 33 females; Cochea River, Mar. 6,
1935, D. V. Brown, 4 males and 3 females; Gamboa, C. Z.,
Apr. 2, 1921, J. G. Saunders, 4 males and 8 females; Progreso,
Chiriqui Prov., Apr. 18, 1923, F. M. Gaige, 1 female; Pehonoma,
1926, H. Brown (allotype female of G. cucuUatiis Martin).

Colombia: Arroyo de Arenas, Sta. Marta Mts., July 25, 1920,
F. M. Gaige, 1 male and 2 females (Mich. Coll.); Cuyaima
Tolima, Nov. 9, 1944, R. A. Stirton, 1 female; Villavieja, 1944,
R. A. Stirton, 3 females.

Comparative notes. Externally, very similar to G. oculatus
(Fabricius), but considerably smaller than most specimens of
the latter species. Easily separated by the presence of a large,
membranous secondary keel hood on the male genitalia.

Gelastocoris vicinus Champion

(PI. IV, fig. 17; PI. VI, fig. 32)

1901. Gelastocoris vicinus Champion, Biologia Centrali-Americana, Rhynchota

Heteroptera, vol. II, p. 349.
1906. G. vicinus Torre-Bueno, Ent. News, vol. XVII, p. 54.

1909. G. vicinus Kirkaldy and Torre-Bueno, Proc. Ent. Soc. Washington, vol.
X, p. 181.

1910. G. vicinus Montandon, Annuario del Museo Zoologico della R. Univer-
sita di Napoli (nuova serie), vol. 3, no. 10, p. 2.

1910. G. vicinus Banks, Catalogue Nearctic Hemip. -Heteroptera, Am. Ent.

Soc. (sep. publ. ), p. 11.
1917. G. vicinus Van Duzee, Catalogue of Hemiptera N. America, Univ.

California Publ., p. 473.

1929. G. vicinus Melin, Zoologiska Bidrag Fran Uppsala, Band 12, pp. 163-164,
fig. 22.

1929. G. vicinus Martin, Univ. Kansas Sci. Bull. vol. XVIII, no. 4, pp. 362-363,

pi. 58, fig. 7, pi. 59, fig. 9.
1939. G. vicinus Millspaugh, Field and Laboratory, vol. VII, no. 2, p. 88.

Size. Male: Length, 6.2 to 7.1 mm.; width of pronotum, 3.9 to
4.3 mm.; width of abdomen, 5.0 to 4.3 mm. Female: Length, 7.1
to 8.0 mm.; width of pronotum, 4.3 to 4.8 mm.; width of abdomen,
4.3 to 5.0 mm.

Color. In general appearance usually some shade of brown. Of
approximately one hundred specimens, there is one that is light green
and only three that are light brown or yellowish-brown. The black
spot of the corium at the middle of the claval suture, as well as
other black spots of the hemelytra, are conspicuous on most speci-
mens. The color and patterns are variable to some extent, but not
to the degree as in some of the other species. Colors present are

320 The University Science Bulletin

red, brown, black, yellow, green, and white, although the last two
are not commonly found. Below, abdomen usually with lateral
portions lighter than median portion, but abdomen as a whole lighter
than upper surface of body, occasionally suflFused with red. Legs
ringed with dark brown, front femora indistinctly ringed, but this
may vary to almost completely fused rings, the intensity of the
brown also variable.

Structural characteristics. Apex of head rounded; front quite
rough; ocelli rather large, one and one-half times size of largest
blisterlike granules. Disc of pronotum rough; lateral margin sub-
parallel for a distance equal to one and one-fourth times the width
of the ocellocular space, slightly divergent posteriorly, then ex-
tending posterolaterad to lateral angle; posterolateral margin con-
cave; width of pronotum subequal to width of abdomen; distinctly
projecting laterad beyond the base of embolium; base of pronotum
lacking short, longitudinal carinae. Hemelytra extending to or
beyond the end of abdomen in both sexes; membrane well-
developed; basal half of lateral margin of embolium expanded
laterad, width of the expansion equal to one half the ocellocular
space, expansion acute or obtusely angulate posteriorly. Connexi-
vum visible in both sexes, but not too exposed. Blisterlike granules
numerous, not very large, about equal in size to base of hind
tarsus, many of the smaller ones more elevated, becoming peglike.
Spatulate right side of seventh ventral abdominal segment of male
short, not extending to lateral margin of the abdomen, about one-
half covered by the preceding segment; left side moderately pro-
jecting posteriorly. Female ventral abdominal segments more or
less symmetrical except that the incision of the posterior margin of
the last segment is slightly bent to the left. Keel hood of male
genitalia triangular, covering apex, fringe nearly transverse, follow-
ing margin basad on left side of keel; keel hook present, fairly
stout, not very recurved, but reaching margin of pan; pan short,
more or less flattened and in an oblique plane in respect to the hori-
zontal plane of the genital capsule; right clasper hook with digiti-
form apex, transverse basal portion very wide at base and narrowing
rapidly to base of digitiform apex; tumescence of right clasper not
adnate to basal portion of clasper; anterolateral projection of keel
absent.

Location of type. In the British Museum at London, England.
Distributional data. In the original description Champion gives
for the distribution of this species the following: "Habitat: North

The Family Gelastocoridae (Hemiptera) 321

America, southern and western United States; Mexico; Panama;
South America to Brazil." This seems too great a range to the
present writer, as he has not seen specimens from the United States
or from South America. It seems Hkely that the records of speci-
mens from these areas have been based upon misidentifications. The
writer has studied specimens from the following localities:

Mexico: Colima, Vulcano, no date, L. Conrad, 1 female, com-
pared with type by Doctor H. B. Hungerford; La Sabana, Guerrero,
kil. 226 S Mex. City, October 20, 1936, H. D. Thomas, 1 female;
Chiapas, Juarez, March 19, 1939, P. Brodkorb, 1 male (Mich. Coll.);
Cuautla, Morelos, June 27, 1948, P. P. Dowhng, 1 male (Mich. Coll);
Tejupilco, Dist. of Temascaltepec, June-July, 1933, H. E. Hinton,
4 males; Nayarit, Tepic, March, 1923, no collector, 2 females
(USNM); 5 mi. S Valles, San Luis Potosi, June 13, 1932, H. Smith,
1 female.

Guatemala: Los Amates, January 16, 1905, no collector, 2 females.

Honduras: Aguan R. valley, Copete farm, April 10, 1923, T. H.
Hubbell, 1 female; Copan, February 18, 1937, C. Roys, 1 male;
Tela, Guaimas Dist., May 10, 1923, T. H. Hubbell, 2 females.

Costa Rica: San Jose, June-July, 1931, H. Schmidt, 4 males and
6 females; same place, purchased 6, 1931, H. Schmidt, 2 females;
same place, 1932, H. Schmidt, 5 males and 2 females; same place,
no date, Heidemann, 2 males; R. Surubres, San Mateo, February,
1905, P. Biolley, 35 males and 29 females; Rio Torres, February
10, 1932, H. Schmidt, 1 female; Rio Virilla, December 26, 1931, H.
Schmidt, 32 males and 18 females; Surrubres, no date, A. Heyne,
3 males and 5 females.

Panama: Barro Colorado, July 31, 1924, N. Banks, 2 females
(MCZH); Boquete, Chiriqui Prov., Mar. 2, 1923, F. XL Gaige, 1
female; Cochea River, Mar. 6, 1935, D. V. Brown, 1 female.

Isla de Finos: Jan. 15, 1923, C. H. Ballou, 1 female.

Comparative notes. Similar to G. oculatus (Fabricius), but
with the posterior portion of the lateral margin of the pronotum
straighter and the lateral angle more angulate. Although the geni-
talia are similar to the genitalia of G. oculatus (Fabricius), there
is one constant difference. The pan of this species is always in
an oblique plane in respect to the horizontal plane of the genital
capsule and the pan is more flattened. In his 1910 paper, Mon-
tandon places G. vicinus Champion in the synonomy of G. quadri-
maculatus (Guerin-Meneville). The writer cannot accept Mon-

11—6730

322 The University Science Bulletin

tandon's action in this matter. Montandon states that the type of
G. quadrimactdattis (Guerin-Mcneville) does not agree in colora-
tion with the specimens he had previously identified as G. vicimis
Champion. Coloration in itself is not generally considered of
much value in the Gelastocorinae, yet in this case, it may be an
indication that the type of G. qiiadrimactilattis (Guerin-Meneville)
is not the same as G. vicinus Champion. The writer has not seen
the pattern, as described by Montandon, in G. vicinus Champion.
It is a common pattern in several other species, for example G.
oculatus (Fabr. ) and G. nehtilosiis (Guerin-Meneville). Mon-
tandon doubts the correctness of the locality label (Bolivia) of the
type of G. quadrimaculatus (Guerin-Meneville). He states that
the type agrees in form with specimens he had labelled G. vicinus
Champion and which were from ". . . Cost Rica, Mexique,
Perou (Mus. Nat. Hung. Budapest), Venezuela, Bresil (Mus.
Senckenberg Frankfurt), Chulumani Bolivie (coll. Schouteden ) ."
Montandon does not indicate that there was a similarity of a par-
ticular morphological structure, so we must assume that a general
similarity of habitus prompted his action in placing G. vicinus
Champion in the synonomy of G. quadrimacidatus (Guerin-
Meneville). Further there is no evidence that he had material
with him which he could have compared with the type of the
latter species. The writer has seen no specimens of G. vicinus
Champion from South America and he does not believe it occurs
there, particularly in Bolivia. In this paper G. quadrimaculatus
(Guerin-Meneville) is treated as a synonym of G. nebulosus
(Guerin-Meneville). Those specimens which were from South
America and which Montandon had formerly labelled G. vicinus
Champion were probably a mixed series of several species. G.
fuscus Martin, G. amazonensis Melin and G. nebulosus (Guerin-
Meneville) may have been represented. The first two species both
resemble G. vicinus Champion in general habitus. It is quite pos-
sible that G. quadrimacidatus (Guerin-Meneville) will prove to be
an older name for G. fuscus Martin, provided that the type of the
former was not destroyed during World War II and provided it
can be located and examined.

Gelastocoris bufo (Herrich-Schaffer)

(PI. IV, fig. 15; PI. VI, fig. 34)

(1839) 1840. GaJgulus bufo Herrich-Schaffer, Die Wanzenartigen Insecten,

V, p. 88, pi. 174, fig. 536.
1873. G. bufo Walker, Catalogue of Hemiptera in the British Museum, part

VllI, p. 171.

The Family Gelastocoridae (Hemiptera) 323

1876. G. bufo Uhler, Bull. U. S. Geol. Surv., vol. I, no. 5. p 337.

1S76. G. bufo Stal, Kongl. Svenska Vetenskaps-Akademiens Handlingar, Band

14, no. 4, p. 137.
1895. G. bufo Pittier and Biolley, Institute fisico Geografico Nacional (Costa

Rica), p. 23.
1901. Gelastocoris bufo (Herrich-Schaffer); Champion, Biologia Centrali-

Americana, Rh>'nchota Heteroptera, vol. II, p. 348, pi. XX, fig. 16.
1906. G. bufo Torre-Bueno, Ent. News, vol. XVII, p. 54.
1909. G. bufo Kirkaldy and Torre-Bueno, Proc. Ent. Soc. Washington, vol. X,

p. 180.
1929. G. bufo Melin, Zoologiska Bidrag Fran Uppsala, Band 12, pp. 156-158,

fig. 11.
1929. G. bufo Martin, Univ. Kansas Sci. Bull., vol. XVIII, no. 4, p. 360, pi. 58,

fig. 14.

Size. Male: Length, 7.0 to 8.0 mm.; width of pronotum, 4.1 to 4.6
mm.; width of abdomen, 4.4 to 4.9 mm. Female: Length, 8.1 to 8.8
mm.; width of pronotum, 4.7 to 5.3 mm.; width of abdomen, 4.8 to 5.5
mm.

Color. \"ariable, from pale yellow to black in general appearance.
The main colors present are brown, red, yellow and black; a few
specimens show some green coloration, but white is rarely found,
except on some specimens, where it is limited to the blisterlike
granules of the hemelytra. The pattern is as variable as the colors.
Below, with the abdomen dark browTi, legs annulated with dark
broum, front femora with the basal tsvo thirds, except for a small
yellow spot on the anterior margin at the middle, entirely dark
brown.

Structural characterhiics. Apex of head rounded; front slightly
elevated and moderately rough; ocelli small, subequal to the blister-
like granules of the hemel>i:ra in size, located on very small tubercles
between the eyes. Pronotum with the disc moderately rough; lateral
margin of pronotum slightly concave, extending posterolaterad to
the lateral angle; posterolateral margin nearly straight, extending
posteromesad to humeral angle, forming a right angle with the
lateral margin; not quite as wide as abdomen; lateral angle pro-
jecting beyond the base of embolium; base of pronotum lacking
short, longitudinal carinae. Hemel\i:ra extending beyond the end
of the abdomen in both sexes; membrane well developed; basal half
of lateral margin of embolium slightly expanded, width of expansion
equal to one third of ocellocular space, expansion posteriorly very
obliquely rounded, tapering into margin of posterior half of em-
bolium. Conne.xi\-um visible in both sexes. BlisterHke granules
ver\' small, fairly numerous, two thirds width of base of hind tarsus
in size. Spatulate right side of seventh ventral abdominal segment
of male only slightly covered by the preceding segment, not extend-

324 The University Science Bulletin

ing to the lateral margin of the abdomen; left side projecting pos-
teriorly only slightly. Female ventral abdominal segments nearly
symmetrical. Keel hood of male genitalia broadly triangular, one-
third to one-fourth as long as wide, fringe transverse, slightly con-
cave; keel, just basad of keel hood, broadly expanded to the right;
keel hook present, rather large, not recurved and apex not extending
over any portion of pan; pan rather broad, cup-shaped due to
large depression; right clasper hook similar to that of G. octilatus
(Fabricius), apical half, thin, digitiform, transversal portion much
thicker and increasing in width toward base; tumescence of right
clasper not adnate to basal portion of right clasper; anterolateral
projection of keel absent.

Location of type. Unknown. Other Hemipterous insects of
the collection of Herrich-Schiiffer are located in the Museum of
Miinchen, Miinchen, Germany.

Distribntionol data. The original description merely states:
"Aus Amerika." Champion and Martin examined specimens from
Mexico, Guatemala and Costa Rica. Melin studied specimens from
these localities and from Panama. This worker has seen specimens
from the following localities:

Mexico: Cocula, Jalisco, November, 1923, no collector, 1 female
(USNM); El Sabino, Uruapan, Mich., July 20, 1936, H. D. Thomas,
2 females; same place, August 2, 1936, H. D. Thomas, 1 female;
Las Barrancas, March, 1923, W. M. Mann, 1 female (USNM);
San Antonio, Nr. El Salto, June 10, 1937, M. Embury, 18 males and
16 females; Real de Arriba, Dist. of Temascaltepec, May-June, 1933,
H. E. Hinton, 25 males and 29 females; Tejupilco, Dist. of Tema-
scaltepec, June-July, 1933, H. E. Hinton, 3 males and 4 females.

Guatemala: El Salto Escuintla, 1934, F. X. Wihiams, 1 male;
Guatemala City, no date. Champion, 1 male; Alta Vera Paz, Dept.
Baja, no date or collector, 1 female (MCZH); "Guatemala," no
date or collector, 1 female (MCZH).

Costa Rica: Rio Virilla, December 26, 1931, H. Schmidt, 2 males
and 5 females; San Jose, June and July, 1931, H. Schmidt, 1 female;
same place, purchased June, 1931, H. Schmidt, 1 male; same place,
1932, H. Schmidt, 1 female.

Panama: Cochea River, March 6, 1933, D. V. Brown, 1 male.

Comparative notes. This species is distinguished from most
species by the lateral margin of the pronotum being nearly straight.
It is easily separated from G. angulatus (Melin) since this species

The Family Gelastocoridae (Hemiptera) 325

lacks the short, longitudinal carinae of the pronotum and the geni-
talia are quite different. The male genitalia are rather similar to
those of G. oculatus (Fabricius), but the pan is more flattened
and shorter than in that species.

Two references have not been included in the bibliography of
this species because it is not possible to determine whether they
actually refer to this species or not. The first of these papers is
that of Spinola (Essai sur les Insects Hemipteres, etc., Genes, 1837,
p. 62). Spinola uses the name Galguhis biifo Lefebvre and adds
this statement: "Nouvelle espece qui m'a ete envoyee par M.
Lefebvre." I have been unable to find a paper by Lefebvre in
which this species is described. The name apparently was a manu-
script name of Lefebvre's and it is a nomen nudum in Spinola, since
he did not describe or figure the species. Herrich-Schiiffer also
credits the species to Lefebvre, but it is quite possible that he was
following Spinola. He states: "Da ich diese einzige Art dieser
Gattung vor mir habe, so kann ich nicht mit Bestimmtheit be-
haupten, ob es wirklich Lefebvre's Bufo ist." It is therefore possible
that Herrich-Schiiffer did not have the same species before him
as that before Spinola. Herrich-Schaffer did not describe the
species, but he did figure it and therefore becomes the author.
The second paper omitted from the bibliography of this species is
that of Dohrn (Catalogus Hemipterorum, 1859, p. 53). Dohrn
also lists this name with Lefebvre as the author. He may have
been following either Spinola or Herrich-Schaffer.

GeJastocoris amazonensis Melin

(PI. V, fig. 22; PI. VI, fig. 36)

1929. Glastocoris amazonensis Melin, Zoologiska Bidrag Fran Uppsala, Band
12, p. 158, figs. 1. 3, 12.

Size. Male: Length, 7.9 to 8.2 mm.; width of pronotum, 4.6 to
4.9 mm.; width of abdomen, 4.6 to 4.9 mm. Female: Length, 7.5
to 8.6 mm.; width of pronotum, 4.8 to 5.0 mm.; width of abdomen,
5.0 to 5.1 mm.

Color. In general appearance brown to reddish-brown, nearly
concolorous, the lateral portions of the pronotum slightly lighter
than the disc, occasionally with darker areas along the lateral mar-
gins of the hemelytra. Colors present are mainly brown, red and
black. Below, dark brown, legs lighter, ringed with dark brown.
Blisterlike granules of the hemelytra are very pale green in two
specimens, light or dark brown in the others.

Structural characteristics. Apex of head rounded; front rather

326 The University Science Bulletin

rough; ocelli large, nearly twice the size of the blisterlike granules.
Disc of pronotum rather rough; lateral margin approaching that
of G. bttfo ( Herrich-Schiiffer ) , but a little more notched, margin
extending diagonally, more posteriorly than laterally, for a distance
equal to one and one-half times the ocellocular space, then proceed-
ing posterolaterad to the lateral angle, usually there is a small tri-
angular projection on the anterior one third of the lateral margin,
this projection is smaller than a similar projection in G. ftiscus Mar-
tin; posterolateral margin straight slightly concave, extending from
the lateral margin at almost a right angle, the posterolateral margin
is more convergent than in G. fuscus Martin; pronotum and ab-
domen about equal in width, the body rather elongate in appear-
ance; lateral angle projecting laterad beyond base of embolium;
base of pronotum lacking short, longitudinal carinae. Hemelytra
extending to or beyond the end of the abdomen; membrane well-
developed; basal half of lateral margin of embolium slightly ex-
panded, the expansion equal to one third the width of the ocelloc-
ular space, margin weakly serrate, expansion acutely angulate
posteriorly. Connexivum visible in both sexes, moderately expanded
laterally, blisterlike granules of hemelytra small, fairly abundant,
rather elevated, subconate, about one-half the width of base of
hind tarsus in size. Spatulate right side of seventh ventral abdominal
segment of male short, not extending to the lateral margin of the
abdomen, oval in shape, nearly covered by the preceding segment;
left side moderately projecting posteriorly. Female ventral ab-
dominal segments symmetrical, incision of the posterior margin
of the last segment rather broad. Keel hood of male genitalia
crescent-shaped, fringe nearly transverse, concave; keel hook pres-
ent, rather slender and not very recurved; pan flat with elongate
apex which is twisted completely around; right clasper hook, simple,
more or less sickle-shaped; tumescence of right clasper not adnate
to basal portion of clasper; anterolateral projection of keel absent.

Location of type. In the Museum of Stockholm, Stockholm,
Sweden.

Distributional data. Known only from Amazonas, Brazil. Melin
studied specimens from Rio Autaz, Amazonas, Oct., A. Roman,
holotype male, allotype female and 1 male paratype; Rio Negro,
Umarituba, Amazonas, Apr. 28, A. Roman, 3 males and 3 females
(paratypes); and Tefe, Amazonas, 1 male and 1 female para-
types. This writer has seen all the specimens from the first locality
and 3 female paratypes from the second locality.

The Family Gelastocoridae (Hemiptera) 327

Comparative notes. Resembling G. fusctis Martin and G. hiifo
Herrich-Schiiffer, but a more slender species than either of these.
The worker has examined the genitalia of the holotype and of the
male paratype from Rio Autaz. They are identical and extremely
different from all other species. The pan is elongate, thin and
twisted completely over.

Gelastocoris major Montandon

(PI. V, fig. 21; PL VI, fig. 39)

1910. Gelastocoris major Montandon, Annuario del Museo della R. Universita

di Napoli ( Nuova Serie), vol. 3, no. 10, p. 2.
1952. G. major Hussey, Florida Entomologist, vol. XXXV, no. 2, pp. 69-70.

Also referring to this species:

1929. Gelastocoris andinus Melin, Zoologiska Bidrag Fran Uppsala, Band 12,

pp. 159-160, fig. 14.
1929. Gelastocoris ditplicatus Martin, Univ. Kansas Sci. Bull., vol. XVIII, no.

4, p. 364, pi. 58, fig. 17, pi. 59, fig. 7.

Size. Male: Length, 6.8 to 8.0 mm.; width of pronotum, 3.8 to
4.7 mm; wide of abdomen, 4.4 to 5.1. mm. Female: Length, 7.4
to 8.8 mm.; width of pronotum, 4.4 to 4.9 mm.; width of abdomen,
5.0 to 5.8 mm.

Color. In general appearance ranging from light brown to black,
both color and pattern are quite variable. Colors present are
yellow, orange, brown, black and green, but the latter is in the
blisterlike granulations of the hemelytra. Below, with the abdomi-
nal segments lighter laterally; legs ringed with brown, the intensity
depending directly upon the darkness of the individual specimen.

Structural characteristics. Apex of the head rounded; or slightly
pointed; front of head with moderate elevations; ocelli small, about
one-half to three-fourths the size of the largest blisterlike granules.
Pronotum with the disc moderately rough; lateral margin with an-
terior portion subparallel, but convex, for a distance equal to two
thirds the length of the ocellocular space, then lateral margin pre-
ceding posterolaterad to the lateral angle, lateral margin very
faintly serrate to smooth; posterolateral margin converging very
slightly to the humeral angle of the pronotum, serrate; lateral angle
projecting laterad only slightly beyond base of embolium; width
of pronotinn less than width of abdomen; base of pronotum lacking
short, longitudinal carinae. Hemelytra of males extending to end
of abdomen, in females hemelytra may reach end of abdomen or
may be slightly shorter; membrane greatly reduced, hmited to a
narrow strip around the apex and extending along inner margin to

328 The University Science Bulletin

claval suture; basal half of lateral margin of embolium expanded
slightly, about the width of blisterlike granule, expansion posteriorly
may be either angulate or broadly rounded. Connexivum barely
visible in the males, more evident but not prominent in the females.
The blisterlike granules fairly numerous, largest subequal in width
to width of hind tarsi. Seventh ventral abdominal segment of
male with spatulate right side nearly covered by sixth segment and
not reaching the lateral margin of the abdomen; left side projecting
posteriorly only slightly. Female ventral abdominal segments
nearly symmetrical, the incision may be bent slightly to the left.
Male genitalia very distinctive; keel hood rounded, broad, crescent-
shaped, fringe of long hairs, extending transversely then basally
along right margin of keel; keel hook absent, replaced by two very
heavily sclerotized, broadly triangular projections; pan very dis-
tinctive, bilobed; right clasper hook short, sickle-shaped, very
broad at base, but rapidly narrowing; tumescence of right clasper
not adnate to basal portion of clasper; anterolateral projection
curved mesad, bluntly claw-shaped.

Location of type. Unknown. The matter of a type of this species
requires a discussion of the description of the species. Montandon
had misidentified this species in routine identification as G. favtis
(Guerin-Meneville). When he examined the type of the latter
species he realized his error and described "G. MAJOR Montand.
nov. sp. = G. flaviis Montand. olim, (nee Guerin) et Champion
Biol. Cent. Am. 2, p. 349, . . ." Montandon did not indicate a
type in the description. In the discussion of "GeJastocoris nehulosus
Guerin loc cit. = G. flaviis Guer. loc. cit." he does list some of the
specimens which he had labelled G. fioviis (Guerin-Meneville)
and mentions where they were located. In this discussion he states
that part of those he had labelled G. flaviis (Guerin-Meneville) were
from Chandra Maja, Perou and were in his personal collection. He
also states that he had distributed many specimens from the above
locality to the Museum of Naples and to many correspondents. If he
did mark a specimen as a type, it may be in the British Museum,
or it may have been in the collection of the University of Naples.
There are four specimens in the collection of the United States Na-
tional Museum which bear the label "Gelastocoris major Montd.,
Det. Montd. 1909." These specimens are the same species as G.
andinns Melin, so it would appear that the latter specific name will
fall as a synonym of G. 7najor Montandon. Montandon places the
G. flavus of Champion as this species, but it could also be the species

The Family Gelastocoridae (Hemiptera) 329

herein described as Gelastocoris viridis n. sp. Contrary to Hussey's
statement, Melin did not propose Gelastocoris andimis to replace
G. fiavus Champion, 1901, ncc Guerin-Meneville.

Distributional data. It would seem that if Montandon actually
had material before him when he described this species it was prob-
ably specimens from Chandra Maja, Perou. The four specimens
(2 males and 2 females) which Montandon labelled G. major Montd.
are from Cachabe (Cachabi), Ecuador. In addition to the four
specimens mentioned above, the writer has seen specimens from
the following localities:

Ecuador: Naranjapata, Dec. 14, 1933, F. X. Williams, holotype
male and allotype female (G. duplicatus Martin); Cachabi, no date,
Rosenberg, 1 male,

Colombia: Bogota, no date, Lindig, holotype male, allot>'pe fe-
male, 2 male and 3 female paratypes (G. andinus Melin); Mt. San
Lorenzo, Santa Marta Mts., June 8, 1920, F. M. Gaige, 6 males and
2 females (Mich. Coll.); same place, June 12, 1920, F. M. Gaige,
1 male and 1 female, (Mich. Coll.); same place, June 29, 1920,
F. M. Gaige, 1 male and 4 females, (Mich. Coll.); same place, Aug.
17, 1920, F. xM. Gaige, 1 male, (Mich. Coll.); Sasaima, Dec.,' 1936,
A. Maria, 1 male and 1 female; Valle de Tabacal, Santa Marta
Mts., Sept. 2, 1920, F. M. Gaige, 1 female, (Mich. Coll.).

Panama: Gamboa, C. Z., Feb. 14, 1921, J. G. Sanders, 1 male;
Ft. Clayton, C. Z., 1933, Cpt. R. F. Edwards, 1 female; Canal Zone,
Mar. 19, 1945, Cpl. G. S. Ryan, 1 male and 2 females.

Comparative notes. Easily separated from all other species by
means of the bilobed pan of the male genital capsule.

Gelastocoris peruensis Melin

(PI. V, fig. 26; PI. VI, fig. 38)

1929. Gelastocoris andinus peruensis Melin, Zoologiska Bidrag Fran Upnsala
Band 12, pp. 160-161, figs. 4, 15, '

Size. Male: Length, 7.7 to 8.4 mm.; width of pronotum, 4.4 to
4.8.; width of abdomen, 5.1 to 5.6 mm. Female: Length, 8.6 to 10.0
mm.; width of pronotum, 4.7 to 5.5 mm.; width of abdomen, 5.7 to
6.4 mm.

Color. Nearly concolorous, in general appearance various shades
of brown, usually reddish-brown. Legs usually sHghtly hghter than
body, and faintly ringed with brown, the dorsal surface of the legs
particularly, often suffused with red.

Structural characteristics. Apex of head rounded to slightly

330 The University Science Bulletin

pointed; front not very rough, nearly flat; ocelli on the anterolateral
surface of small elevations, very small, about three fourths width
of largest blisterlike granules in size. Pronotum with the disc not
very rough; lateral margin of pronotum subparallel for a distance
equal to one and one fourth the ocellocular space, slightly diverging
posteriorly, then extending obliquely posterolaterad for a short dis-
tance to lateral angle; posterolateral margins more or less subparallel,
usually very slightly converging posteriorly or with entire margin
convex; lateral angle scarcely projecting laterad beyond the base
of the embolium; abdomen distinctly wider than the pronotum;
base of pronotum lacking short, longitudinal carinae. Hemelytra
extending to end of the abdomen; membrane greatly reduced, lim-
ited to thin strip around apex and extending along inner margin to
claval suture; basal half of lateral margin of embolium scarcely
expanded, width of expansion equal to one eighth of ocellocular
space, expansion posteriorly usually running smoothly into margin
of posterior half of embolium or very weakly obtuse. Connexivum
almost completely hidden in the males, a little more obvious in
females, not at all prominent. Blisterlike granules rather small, but
numerous, about equal in width to base of hind tarsus. Spatulate
righ side of seventh ventral abdominal segment of male short,
rounded, only slightly covered by preceding segment, not extending
to lateral margin of abdomen; left side only slightly projecting pos-
teriorly. Female ventral abdominal segments more or less symmetri-
cal, incision of posterior margin of last segment broad. Keel hood
of male genitalia semicircular, broad, sclerotized overfold not very
long, fringe long and extending completely across apex and down
the right side of keel; keel hook short, not prominent, greatly modi-
fied, forming a heavily sclerotized roof over the base of the pan;
pan elongate, rather flat; tumescence of right clasper not adnate to
basal portion of clasper; right clasper hook curved, wide basally
and tapering to a thin point apically, apical portion sickle-shaped;
anterolateral projection of keel present, slender, clavate and bent
mesad.

Location of type. In the Museum of Stockholm, Stockholm,
Sweden.

Distributional data. Apparently restricted to Peru. The writer
has examined specimens from the following localities in Peru:
Roque, nr. Moyobamba, March 31, D. Melin, holotype female, 4
male and 1 female paratypes (Mus. of Stockholm); same place,
Sept. 8, D. Mehn, 1 male paratype (Mus. of Stockholm); Chandra

The Family Gelastocoridae (Hemiptera) 331

Maya, no date or collector, 2 males and 1 female; Prov. La Mar.
Sivia, Dept. Ayacucho, June 16-20, 1941, F. Woytkowski, 22 males
and 17 females; Rio Perene and Rio Chanchosmayo, Dept. Junin,
May, 1934, F. Woytkowski, 6 males and 6 females; Loc Shapajilla,
Dept. Huanuco, July 29 to August 10, 1938, F. Woytkowski, 1
female; Marcopata, no date or collector, 8 males and 10 females;
Rio Paucartambo, Quiroz, Dept. Junin, January, 1934, G. Nelson,
3 females; Tambo, Enenas Cam. del Pichis, July 3, 1920, Corn.
Univ. Ex., 1 male; Rioja, Dept. San Martin, September 9 to
October 3, 1936, F. Woytkowski, 2 males and 4 females; San Pedro,
May 15-19, 1935, F. Woytkowski, 3 males; same place. May 29,
1935, F. Woytkowski, 8 males and 5 females; Sani Beni, River
Sani Beni, September 5, 1935, F. Woytkowski, 3 males; "Peru,"
1937, F. Woytkowski, 22 males and 25 females.

Comporative notes. Similar to G. nebulosus (Guerin-Meneville)
but generally larger and easily distinguished by the male genitalia.
The genitalia show the close relationship to G. major Montandon
(G. andimis Melin). The writer has removed the name of G.
peruensis Melin from the position of a subspecies of the latter
species and has raised it to specific status.

Gelastocoris nebulosus (Guerin-Meneville)

(PI. V, fig. 24; PI. VI. fig. 35)

1844. Galguhis nebulosus Guerin-Meneville, Iconographie du Regne Animal

de B. Cuvier, pt. 7, pp. 351-352.
1859. G. nebulosus Dohm, Catalogus Hemipterorum, p. 53.
1876. G. nebulosus Stal, Kongl. Svenska Vetenskaps-Akademiens Handlingar,

Band 14, no. 4, p. 137.
1879. G. nebulosus Berg, Hemiptera Argentina, p. 184.

1909. Gelastocoris nebulosus (Guerin-Meneville); Kirkaldy and Torre-Bueno,
Proc. Ent. Soc. Washington, vol. X, p. 180.

1910. G. nebulosus Montandon, Annuario del Museo Zoologico della R. Uni-
versita di Napoli (nuova serie), vol. 3, no. 10, p. 2.

Also referring to this species:

1844. Galgulus flavus Guerin-Meneville, Iconographie du Regne Animal de

B. Cuvier, pt. 7, p. 351, pi. 57, figs. 4, 4a.
1859. G. flavus Dohrn, Catalogus Hemipterorum, p. 53.
1909. Gelastocoris flavus (Guerin-Meneville); Kirkaldy and Torre-Bueno,

Proc. Ent. Soc. Washington, vol. X, p. 180.
1929. G. flavus Mehn, Zoologiska Bidrag Fran Uppsala, Band 12, pp. 161-162,

figs. 2, 16-20.
1954. G. flavus DeCarlo, Mision de Estudios de Patologia Regional Argentina,

ano XXIV, numeros 83-84, pp. 88-90, figs. 1, 7.
1876. Galgulus nebulosus var. b. Stal, Kongl. Svenska Vetenskaps-Akademiens

Handlingar, Band 14, no. 4, p. 137.
1909. Gelastocoris stali Torre-Bueno, in Kirkaldy and Torre-Bueno, Proc. Ent.

Soc. Washington, vol. X, p. 180 (named Stal's var. b).
1929. G. quadrimaculatus Martin, Univ. Kans. Sci. Bull., vol. XVIII, no. 4,

pp. 361-362, pi. 58, figs. 1-5; pi. 59, figs. 14-16.

332 The University Science Bulletin

1954. Gelastocoris paraguayensis DeCarlo, Mision de Estudios de Patologia
Regional Argentina, ano XXIV, nunieros 83-84, pp. 90-92, figs. 2, 8.

1954. Gelastocoris bolivicmits DeCarlo, Mision de Estudios de Patologia Re-
gional Argentina, ano XXIV, numeros 83-84, pp. 92-94, figs. 3, 9.

1954. Gelastocoris vianai DeCarlo, Mision de Estudios de Patologia Regional
Argentina, ano XXIV, numeros 83-84, pp. 94-95, figs. 4, 10.

1954. Gelastocoris bergi DeCarlo, Mision de Estudios de Patologia Regional
Argentina, ano XXIV, numeros 83-84, pp. 96-97, figs. 5, 11.

Possibly referring to this species:

1844. Galgulus quadrimaculatus Guerin-Meneville, Iconographie du Regne

Animal de G. Cuvier, part 7, p. 351.
1859. G. quadrimaculatus Dohm, Catalogus Hemipteronmi, p. 53.
1876. G. quadrimaculatus Stal, Kongl. Svenska Vetenskaps-Akademiens Hand-

lingar, Band 14, no. 4, p. 137.

1909. Gelastocoris quadrimaculatus Kirkaldy and Torre-Bueno, Proc. Ent. See,
Washington, vol. X, p. 180.

1910. G. quadrimacidatus Montandon, Annuario del Museo Zoologico della
R. Universita di Napoli (nuova serie), vol. 3, no. 10, p. 2.

Size. Male: Length, 5.5 to 8.0 mm.; width of pronotum, 3.7
to 4.6 mm.; width of abdomen, 4.2 to 5.3 mm. Female: Length,
6.4 to 8.8 mm.; width of pronotum, 4.0 to 5.4 mm.; width of abdo-
men, 4.3 to 6.3 mm.

Color. Quite variable, ranging from light brown to black. Both
colors and patterns extremely variable. Colors present may be
brown, black, orange, green, yellow and red. Blisterlike tubercles
of the hemelytra variable in color, orange, white, green, brown and
black. Legs pale green to reddish-brown, ringed with brown,
although the distinctness of the rings varies with the specimen.

Structural characteristics. Apex of head pointed or very narrowly
rounded; front slightly to moderately rough; ocelli fairly large,
subequal to width of blisterlike granules in size. Disc of pronotum
rather rough; lateral margin subparallel for a distance equal to
two and one-half times the ocellocular space, then extending pos-
terolaterad for a short distance to lateral angle; posterolateral
margin straight, short, either subparallel or converging posteriorly;
abdomen in specimens from Peru and Ecuador distinctly wider
than pronotum, those from Brazil have the pronotum a little wider,
but still less than width of abdomen; lateral angle scarcely project-
ing laterad of base of embolium; base of pronotum lacking short,
longitudinal carinae. Hemelytra extending to or beyond end of
abdomen in males, extending to end of abdomen in females; mem-
brane well developed or reduced depending upon where the speci-
men was taken, those from Peru and Bolivia have the membrane
reduced to about one half its normal length; basal half of lateral
margin of embolium distinctly expanded laterally, the expansion
about one half the width of the ocellocular space, expansion obtusely

The Family Gelastocoridae (Hemiptera) 333

angulate posteriorly. Connexivum not very visible, but not com-
pletely covered by the hemelytra. Blisterlike granules numerous,
equal to or slightly larger than width of base of hind tarsus. Spatu-
late right side of seventh ventral abdominal segment of male large,
but not extending to the lateral margin of the abdomen, only
slightly covered by the preceding segment; left side only slightly
projecting posteriorly. Female ventral abdominal segments sym-
metrical, incision of the posterior margin of the last segment very
broad. Keel hood of male genitalia short, crescent-shaped, fringe
transverse, concave; keel hook present, greatly recurved, extending
completely back across base of pan; pan more or less flat, wider than
long and truncate or slightly rounded apically; right clasper hook
large, with rather large lateral projection which gives the clasper
a footlike appearance; tumescence of right clasper adnate to base
of clasper; anterolateral projection of keel absent.

Location of type. Unknown. The type was in the Museo Zo-
ologico della R. Universita di Napoli, Italy, which was destroyed
during World War II. The writer is unable to state whether the
type survived the war.

Distributional data. Guerin-Meneville's type was from Brazil.
This species has also been reported from Peru, Ecuador, Venezuela,
Bolivia and Argentina. The writer has seen specimens from the
following localities:

Brazil: Abama, no date, Mann, 1 female (MCZH); Alto Parana,
Mar. 1929, F. Schade, 1 female; Ariro Angra dos Reis, Estado do
Rio, Mar. 28, 1948, A. L. de Carvalho, 6 males and 7 females; same
place, May 29, 1948, A. L. de Carvalho, 17 males and 18 females;
Campinas, Estado do Sao Paulo, Mar. 10, 1924, F. X. Williams, 1
female; Campo Ode., Sao Paulo, no date or collector, 1 male and
1 female; Espirito Santo, no date or collector, 1 male and 3 females;
Kabelstation, Sept. 25, 1938, Geyskes, 1 female; Lassance, Minas
Ger., Nov. 9-19, 1919, R. G. Harris, 1 male and 1 female; Nova
Teutonia, Bauru, May, 1947, F. Plaumann, 19 males and 9 females;
Nova Teutonia, Marilia, Apr., 1947, F. Plaumann, 1 male and 1
female; Nova Teutonia, Tupan L. 1, Apr. 1947, F. Plaumann, 31
males and 23 females; Nova Teutonia, Tupan L. 2, May 1947, F.
Plaumann, 6 males and 9 females; Para, Aug. 4, 1915, H. S. Parish,
1 female; Rezendo, Estado do Rio, Feb. 29, 1924, F. X. Williams,
1 female; Salfada, Cruzes, Oct. 5, 1908, no collector, 1 male (Car.
Mus.); Sao Paulo, Oct., 1947, F. Plaumann, 1 female; Zanderij, I,
Boven Para Dist., Apr. 23, 1927, no collector, 1 female.

334 The University Science Bulletin

Paraguay: Albovena, Srojozuas, Nov. 16, 1926, F. Schade, 4 males
and 7 females.

Venezuela: S. Esteban, Carbobo, Feb. 5, J. and E. B. Williamson,

1 male and 1 female (Mich. Coll.).

Ecuador: Tena, Apr. 13, 1923, F. X. Williams, 1 female.

Peru: Aguaitia, Dept. Loreto, Sept., 1946, F. Woytkowski, 1
male; Boqueron de Padre, Abad. Cordillera, Dept. Loreto, Aug. 3,
1946, F. Woytkowski, 1 male and 2 females; same place, Aug. 31,
1946, F. Woytkowski, 3 males and 1 female; Callanga, no date or
collector, 1 male and 2 females; Prov. La Mar. Sivia, Dept. Aya-
cucho, June 16-20, 1941, P*; Woytkowski, 1 male and 3 females;
Loc Shapajilla, Dept. Huanuco, July 29 to Aug. 10, 1938, F. Woyt-
kowski, 1 female; Rio Perene, El Campamiento, June 18, 1920,
Corn. Univ. Ex., 1 male and 1 female; Marcopata, no date or col-
lector, 1 male; Puerto Bermudez, Rio Pichis, July 13-19, 1920, Corn.
Univ. Ex., 1 male and 3 females; Rio Paucartambo, Quiroz, Dist.
Junin, Jan. 1934, G. Nelson, 4 males and 3 females; Rio Perene and
Chanchamago, Dept. Junin, May 1934, F. Woytkowski, 4 males;
Tumaturmari, Aug., 1913, B, C, 3 males (AMNH); Satipo, Dec,
1942, P. Papryzki, 1 female; same place, July, 1942, P. Papryski, 1
male; Rioja, Dept. San Martin, Sept. 9 to Oct. 3, 1936, F. Woytkow-
ski, 1 female; Rio Negro, Nov. 4, 1935, F. Woytkowski, 1 female;
Sani Beni, Rio Sani Beni, Aug. 5, 1935, F. Woytkowski, 1 male and

2 females; same place, Sept. 5, 1935, F. Woytkowski, 6 females;
San Pedro, May 15-19, 1935, F. Woytkowski, 9 males and 11
females; same place. May 29, 1935, F. Woytkowski, 4 males and
9 females; "Peru," 1937, F. Woytkowski, 2 males and 1 female.

Bolivia: Santa Cruz, no date, J. Steinbach, 42 males and 44
females; Sud Yungas, Dept. de La Paz, Sept., 1938, A. M. Olalla,
1 male; "Bolivia," no date or collector, 1 female.

Chile: Santiago, Aug. 26, 1949, L. E. Pena, 6 males.

Comparative notes. Similar to G. peruensis Melin, especially
in the case of specimens with the membrane of the hemelytra re-
duced. It may be separated from the latter species by the charac-
teristic genital capsule, which has the keel hook recurved completely
back across the base of the pan and which has the tumescence of
the right clasper adnate to the base of the clasper. The shape of
the lateral margin of the pronotum will distinguish it from most
other species.

Galgulus flavus, G. quadrimaculatus and G. nehulosus were de-

The Family Gelastocoridae (Hemiptera) 335

scribed in that order by Guerin-Meneville. The descriptions were
based entirely upon coloration and only the first named species was
figured. Martin, 1929, used Gelastocoris flaviis (Guerin-Meneville)
for the species known as Gelastocoris angiikitus (Melin). He ap-
parently thought that G. faviis (Guerin-Meneville) had the lateral
margin of the pronotum like that of G. hufo ( Herr.-Schiiff . ) but
the figure of the species in Guerin-Mcneville's paper shows that
this is not the case. Martin used the name Gelastocoris quaclri-
macnlatiis (Guerin-Meneville) for this species. Stal, 1876, used
G. nehulosus (Guerin-Meneville) in spite of the facts that G. fiaviis
(Guerin-Meneville) has line priority and that it is the only species
figured. Montandon, 1910, also used G. nehulosus (Guerin-
Meneville ) but probably because he had misidentified so many spec-
imens under the name of G. flavus ( Guerin-Meneville ) . Melin, to re-
vise the genus, used the latter name. The present writer cons'derf.'d
Melin to be "the first reviser" and in past years has determined many
specimens under this name. However, the International Committee
on Zoological Nomenclature has defined "the first reviser" and
under this definition Stal, not Melin, becomes the first reviser and
the name G. nehulosus (Guerin-Meneville) should be used.

Gelastocoris quadritnaculatus (Guerin-Meneville) possibly should
not be placed in the synonymy of this species (see the discussion
under G. vicinus Champion).

It has been necessary to place Gelastocoris hergi DeCarlo, G.
vianai DeCarlo, G. holivianus DeCarlo and G. paraguayensis
DeCarlo in the synonymy of Gelastocoris nehulosus (Guerin-
Meneville ) . DeCarlo based these names on slight variations of the
lateral margins of the pronotum, of the granulations of the corium
and of the male genitalia. This writer believes that these differ-
ences are only individual variations. The illustrations of the ma'e
genitalia of the species described by DeCarlo are all those of G
nehulosus (Guerin-Meneville). The illustrations show the foot-
shaped right clasper hook and the adnate condition of the trmes-
cence of the right clasper and the base of the right clasper. The
details of the keel are not shown and therefore the very character-
istic keel hook may not be identified. If DeCarlo had had before
him all the species of the genus, more material of this species from
other South American localities and all the literature pertaining to
the genus, particularly the paper by Martin, 1928, perhaps he would
not have proposed the above names.

336 The University Science Bulletin

Gelastocoris fuscus Martin

(PI. V, fig. 20; PI. VI, fig. 40)

1929. Gelastocoris fuscus Martin, Univ. Kansas Sci. Bull., Vol XVIII, No. 4,
p. 364, PI. 58, fig. 15, Pi. 59, fig. 17, 19a, 19b.

Also referring to this species:

1954. Gelastocoris martinezi DeCarlo, Mision de Estiidios de Patologia Re-
gional Argentina, ano XXIV, numeros 83-84, pp. 97-98, figs. 6, 12.

Size. Male: Length, 7.3 to 8.2 mm.; width of pronotum, 4.6 to
5.0 mm.; width of abdomen, 4.7 to 5.3 mm. Female: Length, 8.4
to 9.5 mm.; width of pronotum, 5.1 to 5.8 mm.; width of abdomen,
5.3 to 6.0 mm.

Color. In general appearance from pale to dark brown. Colors
present are yellow, brown, black and green. The ground color is
usually some shade of brown and there are patches of green and
black on the hemelytra, but these patches are usually indistinct.
In three specimens there is a yellow transverse band on the posterior
portion of the pronotum, but this is not characteristic of this species
as several others often exhibit this pattern of coloration. In about
one half of the specimens studied the tibiae and tarsi were green
while the basal portions of the legs were yellow or yellow-brown
in color.

Structural characteristics. Apex of head rounded; front mod-
erately rough; ocelli small, less than width of the largest blister-
like granules in size. Disc of pronotum not very rough; lateral
margin of pronotum subparallel for a distance equal to the ocellocu-
lar space, this portion has a small triangular lateral projection, then
the margin extends posterolaterad to lateral angle; the postero-
lateral margin straight or slightly concave, forming a right angle
with the lateral margin; lateral angle projecting laterad beyond
the base of the embolium; abdomen slightly wider than the pro-
notum; base of the pronotum lacking short, longitudinal carinae.
Hemelytra extending to or beyond the end of the abdomen; mem-
brane well developed; basal half of the lateral margin of the
embolium only slightly expanded laterally, the expansion equal
to one-sixth the width of the ocellocular space, expansion obtusely
angulate or rounded posteriorly. Connexivum visible in both sexes,
but more so in the females than in the males. Blisterlike granules
of the hemelytra moderate in size, not over two thirds the width of
the base of the hind tarsus, not very numerous. Spatulate right side
of seventh ventral abdominal segment of male short, more or less
ovate, not extending to the lateral margin of the abdomen, nearly

The Family Gelastocoridae (Hemiptera) 337

hidden by the preceding segment; left side moderately projecting
posteriorly. Female ventral abdominal segments nearly symmetri-
cal, the incision of the posterior margin of the last segment rather
broad. Keel hood located to the right side of the keel, fringe ex-
tending from near apex of keel to a point near the tip of the keel
hook, keel hook present, rather stout, scarcely recurved; pan longer
than wide, rounded apically, rather stout; right clasper hook,
slender, fairly long, simple and sickle-shaped; tumescence of right
clasper not adnate to base of clasper; anterolateral projection of
keel absent.

Location of type. In the Francis Huntington Snow Entomological
Collection at the University of Kansas, Lawrence, Kansas.

Distributional data. Martin had specimens from Ecuador and
Brazil before him, when he described this species. This worker has
examined specimens from the following localities:

Ecuador: Barios, Jan. 22, 1923, F. X. Williams, allotype female;
Near Napo, Feb. 14, 1923, F. X. Williams, 1 female paratype; Rio
Upano, Macas, Jan. 25, 1939, F. M. Brown, 1 female (AMNH);
Tena, Feb. 28, 1923, F. X. Williams, holotype male; same place,
Feb. 23, 1923, F. X. Williams, 2 male paratypes; same place, Apr.
12, 1923, F. X. Williams, 1 male and 1 female paratypes.

Peru: Prov. La Mar. Sivia, Dept. Ayacucho, June 16-20, 1941,
F. Woytkowski, 1 male; Santa Elena Roqueron, Padro Abad., Dept.
Loreto, Aug. 8, 1946, F. Woytkowski, 1 female; Satipo, July, 1942,
P. Paprzycki, 1 female; 11 km. NE. Tingo Maria, Loc. Shapajilla,
Dept. Huanuco, May 9, 1939, F. Woytkowski, 1 male.

Chile: Santiago, Nov., 1949, L. E. Peiia, 1 female.

Comparative notes: In general appearance, quite similar to G.
amazonensis Melin, G. hufo (Herrich-Schaffer), and to some ex-
tent G. vicinus Champion. It is larger than the latter species,
not as slender as G. amazonensis Melin, and does not have the
lateral margin of the pronotum as straight as is the case in G.
hufo (Herrich-Schaffer). It may be separated from all three by
the shape of the genital capsule of the male. This may be G. quadri-
maculatus ( Guerin-Meneville ) . See the discussion under G. vicinus
Champion.

The illustration of the genitalia of the male, the nature of the
lateral margin of the pronotum and the distribution clearly show
that GeJastocoris martinezi DeCarlo is a synonym of this species.
DeCarlo obviously was not aware of the description of G. fuscus
Martin,

338 The University Science Bulletin

GeJastocoris viridis n. sp.

(PI. IV, fig. 13; PI. V, fig. 23; PI. VI, fig. 37)

Probably referring to this species:

1901. Gelastocoris jlavus Champion, Biologia Centrali- Americana, Rhynchota
Heteroptera, vol. II, p. 349.

Size. Male: Length, 6.6 to 6.8 mm.; width of pronotum, 4.1 to
4.2 mm.; width of abdomen, 4.4 to 4.5 mm. Female: Length, 6.9
to 7.8 mm.; width of pronotum, 4.0 to 4.7 mm.; width of abdomen,
4.7 to 5.2 mm.

Color: In general apperance from light brown to dark greyish-
brown. Usually with the lateral portions of the pronotum lighter
than the disc. Colors present are black, brown, yellow, orange, red,
green and white. The blisterlike granules are green in all but one
specimen. In this specimen they are white. Below, mostly dark,
especially the abdomen, which is dark brown. The legs are lighter,
ringed with dark brown, dorsal surface suffused with red.

Structural characteristics. Apex of head rounded; front mod-
erately rough; ocelli extremely small, scarcely visible, one-third
width of largest blisterlike granules of the hemelytra in size. Pro-
notum with the disc rather rough; lateral margin of pronotum
subparallel for a distance equal to one and one-fourth times the
ocellocular space, diverging slightly posteriorly, then extending
posterolaterad to lateral angle; posterolateral margin with anterior
half convex, posterior half concave; lateral angle projecing laterad
beyond base of embolium; width of pronotum slightly less than
width of abdomen; base of pronotum lacking short, longitudinal
carinae. Hemelytra extending to or beyond the end of the abdomen;
membrane is slightly reduced; basal half of lateral margin of em-
bolium expanded, width of expansion equal to one third the
ocellocular space, expansion posteriorly not angulate, rounding
into margin of posterior portion of embolium. Connexivum com-
pletely covered or nearly so by the hemelytra in the males, more
expanded and visible in the females. Blisterlike granules of the
hemelytra and of the other parts very large, numerous, one and
one-half the width of base of hind tarsus in size. Spatulate right
side of seventh ventral abdominal segment of male about half
covered by preceding segment; not extending to lateral margin
of abdomen; left side only slightly projecting posteriorly. Female
ventral abdominal segments nearly symmetrical, incision of posterior
margin of last segment broad, symmetrical or slightly bent to left.
Keel hood of male genitalia triangular, covering apex, fringe trans-

The Family Gelastocoridae (Hemiptera) 339

verse, hairs of fringe not very long; keel hook present, short, very
stout, only slightly curved, not covering any portion of pan; pan
wider than long, swollen into rather thick ridge around apex, basad
of this is a deep transverse, crescent-shaped depression; tumescence
of right clasper not adnate to basal portion of clasper; right clasper
hook nearly sickle-shaped, moderately long, thin and with a very
slight lateral projection at the middle; anterolateral projection of
keel absent.

Location of type. Holotype male, allotype female and one male
paratype, Mt. Obando Queit, Chiapas, Mexico, Apr. 15, 1940,
H. M. Smith; 1 male and 3 female paratypes. El Salto Escuintla,
Guatemala, 1934, F. X. Williams in the Francis Huntington Snow
Entomological Collection at the University of Kansas.

Distributional data. As given above.

Comparative notes. This small species resembles G. nebulosus
(Guerin-Mcneville) somewhat in external appearance, but it may
be readily separated from this species by comparing the male
genitalia. G. viridis does not have the keel hook recurved com-
pletely back over the base of the pan and the tumescence of the
right clasper is not adnate to the basal portion of the clasper. This
species may be separated from the other species by the shape of
the pan, size of the blisterlike granules of the hemelytra and by
the shape of the lateral margin of the pronotum.

Gelastocoris angidatus (Melin)

(PI. IV, fig. 14; PI. V, figs. 25, 29)

1929. Montandonius angulatus Melin, Zoologiska Bidrag Fran Uppsala, Band
12, p. 169, figs. 32, 33.

Also referring to this species:

1929. Gelastocoris jlavus Martin, Univ. Kansas Sci. Bull., vol. XVIII, no. 4,
p. 364, pi. 58, fig. 18, pi. 59, fig. 7.

1954. Montandonius willineri DeCarlo, Mision de Estudios de Patologia Re-
gional Argentina, ano XXIV, numeros 83-84, pp. 99-100, figs. 13-16.

1954. Moyttandoniits mansosotoi DeCarlo, Mision de Estudios de Patologia
Regional Argentina, ano XXIV, numeros 83-84, pp. 100-101, figs. 17, 18.

1954. Montandonius bridarolli DeCarlo, Mision de Estudios de Patologia
Regional Argentina, ano XXIV, numeros 83-84, pp. 101-102, figs. 19-20.

Size. Male: Length, 7.5 to 8.8 mm.; width of pronotum, 4.7 to
5.4 mm.; width of abdomen, 4.8 to 5.7 mm. Female: Length,
8.1 to 10.6 mm.; width of pronotum, 5.2 to 6.2 mm.; width of
abdomen, 5.0 to 6.5 mm.

Color. Yellowish-brown to dark reddish-brown, usually more or
less concolorous. Scutellum a little darker than rest of body, one

340 The University Science Bulletin

specimen also has disc of pronotum darker than remainder of
body. In most of the specimens examined, there are small black-
spots along the lateral margins of the pronotum and the expanded
basal portion of the embolium. The connexivum has the segments
darker along the bases. The large blisterlike and peglike granules
are yellowish-brown, brown, reddish-brown or black. Legs faintly
or darkly ringed with brown depending upon the coloration of the
specimen as a whole.

Structural characteristics. Apex of head slightly concave or
straight; front very rough; ocelli small, two fifths width of largest
blisterlike granules of the hemelytra in size, located on small
tubercles between the eyes. Pronotum with the disc rough; lateral
margin of pronotum, nearly straight, usually faintly concave;
lateral angle when present very obtuse, usually this portion of
the pronotum is more or less rounded; projecting laterad beyond
base of embolium; base of pronotum provided with six to eight
short, longitudinal carinae. Hemelytra extending to or beyond end
of abdomen, in males, usually shorter than abdomen in females,
one small female has the hemelytra extending well beyond the
end of the abdomen; membrane varies in size, but it is rather
reduced in most specimens, the small female has a very well-
developed membrane; basal half of lateral margin of embolium
greatly expanded, width of expansion equal to one half ocellocular
space, expansion posteriorly not angulate, rounded, expansion as
a whole rectangular. Connexivum, broad, extending from ex-
pansion of embolium all around hemelytra in females, quite visible
in males. Large granules of hemelytra and other parts more
elevated, peglike, numerous, largest granules oval, about two
times as long as width of base of hind tarsus. Spatulate right
side of seventh ventral abdominal segment of male elongate, ex-
tending to lateral margin of abdomen; hardly covered by preceding
segment; left side greatly projecting posteriorly. Female ventral
abdominal segments nearly symmetrical, the incision of the pos-
terior margin of last segment broad, and bent slightly to the left.
Keel hood of male genitalia on the right side of keel, greatly
reduced, fringe with hairs extremely long and extending from left
side, around apex and down right side of keel; keel hook absent;
pan extremely large, rather oval in shape, as wide as shaft of
keel level with right clasper hook; right clasper hook very charac-
teristic, extremely long, shaped more or less like an S made
backwards; tumescence of right clasper not adnate to base of

The Family Gelastocoridae (Hemiptera) 341

right clasper, devoid of black peglike spines; anterolateral pro-
jection of keel absent.

Location of type. In the British Museum, London, England.

Distributional data. This species was described by Melin from
specimens from Brazil. The writer has examined specimens from
the following localities:

Bolivia: Coroico, no date, purchased from Dr. Staudinger, 1 male;
Santa Cruz, no date, J. Steinbach, 1 male.

Brazil: Chapada, Aug., no collector, 1 male; same place, July, no
collector, 1 male; same place. May, no collector, 1 male; same
place, June, no collector, 1 female (AMNH); same place, July, no
collector, 1 male (AMNH); same place, Aug., no collector, 2 males
and 1 female (AMNH).

Paraguay: Caravene, Jan. 12, 1925, F. Schade, 1 female;
Moliuasque, Dept. Caruga, Dec, 1925, F. Schade, 1 female; Villar-
rica, Serro Pelado, Jan 28, 1929, F. Schade, 1 male; same place,
Nov. 28, 1929, F. Schade, 1 female; Serro Pelado, Cordillero de la
Villarica, Dec, 1926, F. Schade, 1 female; Villarrica, Mar., 1926, F.
Schade, 1 female.

Comparative notes. Differing considerably from all other species.
In shape of the lateral margin of the pronotum, rather similar to
G. bufo ( Herrich-Schliff er ) , but with the lateral angle rounded and
not angulate as in the latter species. The 6 to 8 short, longitudinal
carinae on the base of the pronotum; straight or concave margin
of the apex of the head; peglike nature of the granules of the
hemelytra, etc.; wide expansion of the basal half of the embolia]
margin; and the male genitalia make the identification of this
species a simple matter.

DeCarlo, 1954, has proposed three new species in the genus
Montandonius Melin, namely M. willineri DeCarlo, M. mansosotoi
DeCarlo and M. bridarolli DeCarlo. All three are considered to
be synonyms of Gelastocoris angulatus (Melin) by this writer.
DeCarlo states that the genus Montandonius differs mainly from
Gelastocoris by the presence of a clavus in the latter. G. angulatus
(Melin), contrary to the statement by DeCarlo, has a clavus. It
is true that some specimens do not appear to have a claval suture
on the dorsal surface, however, a glance at the ventral surface
show a distinct clavus and claval suture. Most specimens do show
the claval suture dorsally. This writer can see no difference be-
tween the illusti-ations of M. willineri DeCarlo and G. angulatus

342 The University Science Bulletin

( Melin ) in regard to the shape of the pronotum and the embohum.
The other two DeCarlo names, based on unique females, are sup-
posed to differ from M. willineri DeCarlo by differences in the
granulations, scutellum more elevated with shallow furrows (M.
mansosotoi DeCarlo), shorter hemelytra (M. bridaroUi DeCarlo)
and by the "characteristic valves of the genitalia." This writer
considers these to be individual variations.

Gelastocoris apureensis Melin

1929. Gelastocoris apureensis Melin, Zoologiska Bidrag Fran. Uppsala, Band
12, p. 159, fig. 13.

Original descriptio7i: "Front of caput somewhat rough, between
ocelli a small knoblike protuberance, laterally slightly sinuated, and
apically rounded; fringe usually stout, of medial length. Structure
of pronotum fairly marked; side without distinctly flattened margin,
level with transversal furrow, projecting to an almost right, some-
what rounded angle, anterior part of edge slightly angularly bent,
in front somewhat crenulated; posterior part somewhat dentated;
anterior corner fairly stout, posterior one not very prominent;
posterior margin of pronotum fairly deeply sinuated before scu-
tellum. Membrane fairly narrow (exposing a broad connexivum)
with fairly big granules, terminally not very hyaline; embolium,
laterally, narrowly shelved. Mesosternal process short, broad, den-
tated. The 6th abdominal sternite of female slightly restricted.

"Coloration, greyish-brown; legs indistinctly ringed.

"Length 6.5, breadth 4 mm.

"Museum of Paris 3 sp. (types): 1?, 2 f. S. Fernando de Apure,
Venezuela ( M aindron ) .

"This species somewhat resembles G. amazonensis but with the
exception of a smaller size may be distinguished by the side of the
pronotum lacking a distinctly flattened margin and the process
being less prominent. The nature of apical incision of the 6th ab-
dominal segment of female could not be observed on collected
specimens."

The writer has not seen the type specimens and has in the col-
lections before him only two specimens from Venezuela. These
two specimens, one male and one female, seem to agree fairly
closely with the original description of G. apureensis Melin, but
the genitalia of the male shows that it is typical G. nebtilosus
(Guerin-Meneville). Since the size and the shape of the lateral
angle of the pronotum are quite variable in the latter species, it is

The Family Gelastocoridae (Hemiptera) 343

the opinion of the writer that Mehn's species will probably prove
to be a synonym of G. nchidosus (Guerin-Meneville). For the
present, however, it seems best to retain this species imtil the types
can be examined.

Nerthrinae Kirkaldy

Gelastocorids with the rostrum appearing to arise from the ventral
surface of the head, not especially stout, apical portion often pro-
jecting anteriorly. Anterior leg with the tarsus fused to the tibia,
not articulate, with only one well-developed tarsal claw in adults.
Ninth ventral abdominal segment of the male quite visible exter-
nally, not completely invaginated within body cavity; the posterior
process only slightly modified, right clasper greatly developed and
folded longitudinally to act as a guide to the simple, tubular
aedeagus; in the normal position, right clasper resting in a depres-
sion of the right side of the dorsal surface of the abdomen; left
clasper absent. Ventral abdominal segments of female symmetrica I
or asymmetrical, depending upon the species.

The Genus Nerthra Say
(Type of the genus, Nerthra stygica Say)

1832. Say, T., Heteropterus Hemiptera of North America, New Harmony,
Indiana, p. 37 {stygica only species).

1859. Leconte, J. L., "Complete Writings of Thomas Say," vol. I, p. 364.
1876. Stal, C, Kongl. Svenska Vetenskaps-Akademiens. HandHngar, Band 14,

No. 4, p. 139.
1898. Kirkaldy, G. W., Entomologist, vol. XXXI, p. 2.

1905. Torre-Bueno, J. R. de la, Ohio Nat., vol. V, p. 287.

1906. Kirkaldy, G. W., Trans. Am. Ent. Soc, vol. XXXII, p. 150.

1909. Kirkaldy, G. W. and Torre-Bueno, J. R. de la, Proc. Ent. Soc. Washing-
ton, vol. X, p. 182.

1910. Banks, N., Catalogue Nearctic Hemip-Heteroptera, Sep. Publ., Am.
Ent. Soc, Phila., p. 11.

1925. Blatchley, W. S., Ent. News, vol. 36, pp. 49-52.
1954. Todd, E. L., Pan-Pacific Ent., vol. 30, no. 2, p. 13.

References to this genus under Mononyx Laporte:

1833. Laporte, F. L. de, Essai d'une Classification Systematique de L'Order des
Hemipteres, p. 16.

1835. Bunneister, H. C. C, Handbuch der Entomologia, Bd. 2, Abt. 1, p. 201.

1836. Brulle, G. A., Histoire Naturelle des Insectes, ed. 9, pp. 274-275.
1840. Spinola, M. M., Essai sur les Insectes Hemipteres, p. 63.

1840. Blanchard, E., Histoire Naturelle des Animaux Articules, vol. 3, p. 93.

1842. Erickson, W. F., Archiv FUr Naturgeschichte, VIII, (1), p. 285.

1843. Amyot, C. J. B. and Sersille, A., Histoire Naturelle des Insectes.
Hemipteres, p. 425.

1850. Spinola, M. M., Tavola sinattica dei generi spettanti alia classi degli
Insetti arthrodignati Hemiptera, 25, 1, p. 49.

1851. Fieber, F. X., Genera Hydrocoridum secundum ordinem naturalem in
familias di sposita. in: Sep. Pragae, Calve, 4, p. 12.

1859. Dohm, F. A., Catalogus Hemipterorum, ( Herausgegeben von dem
Entomologischen Verein zu Stettin), p. 53.

344 The University Science Bulletin

1861. Stal, C, Ofversigt af Kongl. Vetenskaps-Akademiens Forhandlingar,

Arg. 18, p. 201.
1863. Stal, C, Berlin Entomologische Zeitschrift, vol. XXIV, p. 40.5.
1865. Stal, C, Hemiptcra Africana, vol. Ill, p. 171.
1876. Stal, C, Kongl. Svenska Vetenskaps-Akademiens. Handlingar, band

14, no. 4, p. 138.
1884. Uhler, P. R., Standard Natural History, voL II, p. 264.

1899. Montandon, A. L., Bulletin de la Societe des Sciences de Bucarest-
Roumanie, an. VIII, no. 4 et 5, p. 392.

1901. Champion, G. C, Biologia Centrali-Americana, Rhynchota Heteroptera,

vol. II, p. 350.
1906. Distant, W. L., Fauna British India, Rhynchota, vol. Ill, p. 14.
1906. Kirkaldy, G. W., Trans. Am. Ent. Soc, vol. XXXII, p. 149.

1909. Kirkaldy, G. W. and J. R. de la Torre-Bueno, Proc. Ent. Soc. Washing-
ton, vol. X, p. 181.

1910. Banks, N., Cat. Nearctic Hemip. -Heteroptera, sep. publ. Am. Ent. Soc,
Phila., p. 11.

1910. Montandon, A. L., Annuario del Museo Zoologico della R. Universita

di Napoli (nuova serie), vol. 3, no. 10, p. 2.
1917. Van Duzee, E. P., Cat. Hemiptera, Univ. Cahfomia publ., pp. 473-474.
1925. Blatchley, W. S., Ent. News, vol. 36, pp. 49-52.
1929. Melin, D., Zoologiska Bidrag Fran Uppsala, Band 12, pp. 171-194.

References to this genus under Phintius Stal:

1861. Stal, C, Ofversigt af Kongl. Vetenskaps-Akademiens Forhandlingar,

Arg. 18, p. 201.
1863. Stal, C, Berlin Entomologische Zeitschrift, vol. XXIV, p. 407.
1865. Stal, C, Hemiptera Africana, vol. Ill, p. 172.
1876. Stal, C, Kongl. Svenska Vetenskaps-Akademiens Handlingar, Band 14,

no. 4, p. 139.
1929. Mehn, D., Zoologiska Bidrag Fran Uppsala, Band 12, pp. 192-193.

References to this genus under Matinus Stal:

1861. Stal, C, Ofversigt af Kongl. Vetenskaps-Akademiens Forhandlingar,

Arg. 18, p. 201.
1863. Stal, C, Berhn Entomologische Zeitschrift, vol. XXIV, p. 407.
1865. Stal, C, Hemiptera Africana, vol. Ill, p. 172.
1876. Stal, C, Kongl. Svenska Vetenskaps-Akademiens Handlingar, Band

14, no. 4, p. 139.

1900. Montandon, A. L., Bulletin de la Societe des Sciences de Bucarest-
Roumanie, an. VIII, no. 6, pp. 1-7.

References to this genus under Peltopterus Guerin-Meneville:

1843. Guerin-Meneville, F. E., Revue Zoologique Travaux Inedits, p. 113,

as subgenus of (A/ononyx Laporte)
1865. Stal, C, Hemiptera Africana, vol. Ill, p. 173.
1876. Stal, C, Kongl. Svenska Vetenskaps-Akademiens. HandHngar, Band

14, no. 4, p. 140.
1900. Montandon, A. L., Bulletin de la Societe des Sciences de Bucarest-

Roumanie, an. VIII, no. 6, pp. 8-9.
1910. Montandon, A. L., Annuario del Museo Zoologico della R. Universita

de Napoli (nuova serie), vol. 3, no. 10, p. 3.

References to this genus under Scylaecus Stal:

1861. Stal, C, Ofversigt af Kongl. Vetenskaps-Akademiens. Forhandlingar,

Arg. 18, p. 201.
1865. Stal, C, Hemiptera Africana, vol. Ill, p. 173.
1876. Stal, C, Kongl. Svenska Vetenskaps-Akademiens. Handlingar, Band

14, no. 4, p. 139.

The Family Gelastocoridae (Hemiptera) 345

References to this genus under Glossoaspis Blatchley:
1925. Blatchley, W. S., Ent. News, vol. 36, pp. 49-52.

Small to large bugs; usually of some shade of brown, nearly con-
colorous in most species. Front of head usually provided with tooth-
like tubercles; ocelli usually present, but absent in some species;
rostrum appearing to arise from the ventral surface of the head, not
stout, apical portion projecting ventrally or anteriorly. Anterior leg
with the tarsus fused to the tibia, not articulate, with only one well-
developed tarsal claw in the adults. Ninth abdominal segment of
the male quite visible externally, not completely invaginated within
the body cavity; the posterior process only slightly modified; right
clasper greatly developed and folded longitudinally as a guide to
the simple, tubular aedeagus; in the normal position, right clasper
resting in a depression of the right side of the dorsal surface of the
abdomen; left clasper absent. Abdominal sternites of female either
symmetrical or asymmetrical. Hemelytra usually with well-devel-
oped membrane, but membrane reduced or entirely lacking in
some species; hemelytra fused together in some species, but usually
separate.

At the time of the writing of this paper the species of this genus
were placed in six genera, Nerthra Say, Monomjx Laporte, Phintius
Stal, Matintts Stal, Peltopterus Guerin-Meneville and Glossoaspis
Blatchley. The number of species in each was 1, 31, 1, 6, 3, and 1
respectively. The genus Phintius Stal (type of genus, Monomjx
grandicoUis Germar) was placed in the synonomy of Mononyx
Laporte (type of genus, Naucoris raptoria Fabricius) by Montan-
don in 1899. Melin, 1929, raised this name back to generic status
for a specimen of questionable origin. Most of the genera were
characterized by the nature of their hemelytra. Mononyx Laporte
had the hemelytra separate with well-developed membranes. In
Nerthra Say (type of genus, Nerthra stygica Say) the membrane
was lacking and the hemelytra were fused together. Matinus Stal
(type of genus, Mononyx alaticollis Stal) had the membrane re-
duced. Peltopterus Guerin-Meneville (type of genus, Naucoris
rugosa Desjardins) had the membrane lacking, hemelytra fused
together, scutellum slightly reduced, and although not mentioned,
the ocelli are absent. Glossoaspis Blatchley (type of genus, Glos-
soaspis brunnea Blatchley) was characterized in the same manner as
Peltopterus Guerin-Meneville. A recent comparison of a specimen
of Glossoaspis brunnea Blatchley and a type of Peltopterus rugosus
(Desjardins ) has proved that these two are identical.

346

The University Science Bulletin

Since brachypterous forms occur rather commonly in closely re-
lated families, it seems impractical to consider reduction of mem-
brane and fusion of hemelytra as generic characters, especially when
it is quite apparent that in most cases these characters have de-
veloped independently in each of the species possessing them.

Ordinarily the presence or absence of ocelli would seem to be a
rather good character, but the loss of ocelli also has apparently oc-
curred independently. This writer has found that six species lack
ocelli. They all have the hemelytra entirely coriaceous and fused
together, but when the structure of the head, the ventral abdominal
segments of both male and female, and the male claspers are studied
it can be shown that they are more closely related to other forms
occurring in the areas where they are found than they are to each
other.

When our knowledge of the species of this subfamily is more
fully developed, it is quite possible that some of the generic names
herein placed in the synonomy of Nerthra Say may be used at least
at the subgeneric level. The writer has tentatively divided the
species into eight species groups as follows:

I Stijgica Group

Nerthra stygica Say

Nerthra mexicana (Melin)

Nerthra martini Todd

Nerthra usingeri Todd

II Raptoria Group

Nerthra raptoria (Fabricius)

Nerthra buenoi n. sp.

Nerthra ranina ( Herrich-Schaf-

fer)
Nerthra nepaeformis (Fabricius)
Nerthra terrestris ( Kevan )
Nerthra horealis (Melin)
Nerthra tenehrosa n. n.
Nerthra parvula ( Sign. )
Nerthra unicornis (Melin)
Nerthra peruviana (Montandon)
Nerthra montandoni (Melin)
Nerthra quinquedentata (Melin)
Nerthra americana ( Montandon )
Nerthra williamsi n. sp.

Ill Grandicollis Group

Nerthra grandicollis (Germar)
Nerthra indica (Atkinson)
Nerthra lobata ( Montandon )
Nerthra serrata (Montandon)
Nerthra asiatica ( Horvath )
Nerthra spissa (Distant)

IV Rugosa Group

Nerthra rugosa ( Desjardins )
Nerthra macrothorax (Montrou-

zier)
Nerthra nervosa (Montandon)

V Fuscipes Group

Nerthra fuscipes ( Guerin-Mene-

ville)
Nerthra manni n. sp.
Nerthra hungerfordi n. sp.
Nerthra hracchialis n. sp.

VI Amplicollis Group

Nerthra amplicollis (Stal)
Nerthra ecuadorensis (Melin)
Nerthra ater ( Melin )
Nerthra lota (Montandon)
Nerthra rudis ( Melin )
Nerthra corudis n. sp.

VII Alaticollis Group

Nerthra alaticollis (Stal)
Nerthra adspersa (Stal)
Nerthra stali ( Montandon )
Nerthra elongata ( Montandon )
Nerthra hirsuta n. sp.
Nerthra tasmaniensis n. sp.
Nerthra tuberculata (Montan-
don)

The Family Gelastocortoae (Hemiptera) 347

VIII LaticoUis Group

Nerthra laticollis (Guerin-Mene- Nerthra ampliata (Montandon)

ville) Nerthra macrostyla n. sp.

Nerthra gurneiji n. sp. Nerthra robusta n. sp.

Nerthra mixta (Montandon) Nerthra hitcovaria (Distant)

Nerthra omani n. sp. Nerthra sinuosa n. sp.

Nerthra femoralis (Montandon) Nerthra grandis (Montandon)

Nerthra nudata n. sp. Nerthra tcalkeri n. sp.

The writer has been unable to recognize N. turgidula (Distant),
N. anmilipes (Horvath), N. planifrons (Mehn) and IV. suberosa
( Erichson ) . Therefore these species have not been inckided in the
species groups given above. It is possible that IV. tuherculata
(Montandon), which is included in the alaficoUis group, should be
placed in a group to itself. Nerthra riidis (Melin) and N. corudis
n. sp. are placed in the amplicollis group, but they seem to be inter-
mediate between this group and the fuscipes group. The females
of the rugosa, laticollis and alaticollis groups have the last visible
abdominal sternite more or less projecting posteriorly, not emargi-
nate as in the other species groups, and many species have spined
ovipositors. The terminal (VI to IX) abdominal sternites of the
males of these three groups are usually rather small, the width of
the posterior margin of the sixth segment less than one half the
width of the posterior margin of the fourth segment; in the males
of the other species groups the posterior margin of the sixth segment
is equal to or greater than one half the width of the posterior margin
of the fourth abdominal sternite. The name Peltopterus (type of
genus, Naticoris rugosa Desjardins) would be available in either a
generic or subgeneric sense if it were decided to remove these three
species groups from Nerthra Say or to place them in an atypical sub-
genus. The writer does not believe that we have suflBcient knowl-
edge at this time to take this action and therefore he has not done so.

Key to the Species of Nerthra *

1. Hemelytron with membrane reduced or entirely coriaceous ... 2
Hemelytron with membrane well developed 15

2. ( 1 ) Ocelli present 8

Ocelli absent 3

3. (2) Lateral margin of hemelytron from nodal furrow to apex,

bent up almost at a right angle to rest of hemelytron,

N. hirsuta n. sp. p. 417
Lateral margin of hemelytron not as above 4

* Does not include N. elongate (Montandon), (p. 436); N. turgidula (Distant), (p.
406); N. serrata (Montandon) (p. 409); N. annulipes (Horvath), (p 426); N. planifrons
(Melin), (p. 388); N. suberosus (Erichson), (p. 438).

348 The University Science Bulletin

4. (3) Apex and lateral margins of head provided widi sharp-

pointed tubercles; scutellum lacking groups of black

bristles on lateral portions 5

Apex and lateral margins of head lacking sharp-pointed
tubercles; scutellum with a group of bristles on lateral
portions 6

5. (4) Large species, over 8 mm.; last three abdominal sternites

of male small, right side of seventh abdominal stemite
spatulate, partially covered by preceding stemite; clasper
of male simple, sickle-shaped; bristles spiniform,

N. williamsi n. sp. p. 383
Smaller species, under 8 mm.; last three abdominal
sternites of male rather large, right side of seventh ab-
dominal stemite entirely covered by preceding sternite;
clasper of the male with a large dorsomedial process;
bristles clavate N. americana (Montandon) p. 381

6. (4) Hemelytra with large longitudinal carinae 7

Hemelytra without such carinae, nearly smooth, with a
broad, rounded elevation in the middle of each hemel-
ytron, this elevation usually covered with a group of
bristles N. rugosa (Desjardins) p. 412

7. ( 6 ) Pronotum widest at posterior angle; posterior angle of

pronotum acute; tubercles on front of head very large,
densely covered with short, clavate bristles,

N. macrothorax ( Montrouzier ) p. 414
Pronotum widest at middle of lateral margin; posterior
angle obtuse; tubercles on front of head rather small, not
densely covered with bristles; longitudinal carinae of
hemelytra very prominent... N. nervosa (Montandon) p. 416

8. ( 2 ) Membrane limited to a very narrow strip extending

along inner margin from apex of hemelytron to scutellum,

N. sintiosa n. sp. p. 440

Membrane lacking, hemelytron entirely coriaceous 9

9. (8) Hemelytra fused together 10

Hemelytra separate, not fused together 11

10. (9) Temiinal abdominal stemites of male large, width of
posterior margin of sixth abdominal sternite (measured
from one caudolateral angle to the other) greater than
one half the width of the posterior margin of the fourth
abdominal sternite (measured along posterior margin
from the median notch to the lateral margin of the right
side); posterior margin of last visible abdominal stemite
of the female emarginate N. stygica Say p. 354

Terminal abdominal sternites of male small, width of pos-
terior margin of sixth abdominal sternite less than one
half the width of the posterior margin of the fourth
abdominal sternite; posterior margin of last visible
abdominal sternite of the female not emarginate,

N. walkeri n. sp. p. 439

The Family Gelastocoridae (Hemiptera) 349

11. (9) Apex and lateral margin ot head with five large tooth-
like tubercles; base of scutelluni medially strongly
depressed; hemelytra, scutcllum and pronotum with
groups of jet black, rather large, clavate bristles,

N. grandis (Montandon) p. 440
Apex and lateral margin of head lacking toothlike
tubercles or if present, very small; base of scutelluni not
strongly depressed; hemelytra, scutelluni and pronotum

lacking groups of large, black, clavate bristles 12

12.( 11 ) Posterior angle of pronotum angulate 13

Posterior angle of pronotum rounded, not angulate,

N. ahticoUis (Stal) p. 432
13.(12) Lateral margin of head with a tubercle directly below
inner margin of eye; apex of head depressed, base of
depression wider than distance between ocelli,

xV. adspersa (Stal) p. 434
Lateral margin of head lacking a tubercle below the in-
ner margin of eye; apex of head depressed, base of de-
pression less in width than distance between ocelli 14

14.(13) Connexivaim very prominent from dorsal view (at least 4
segments exposed); clasper of male recurved and bluntly

knobbed apically N. stall ( Montandon ) p. 435

Connexivum not as above (only two segments exposed);
clasper of male apically neither recurved nor bluntly
knobbed, straight, tapering to a sharp point

A/, tasmaniensis n. sp. p. 437
15. ( 1 ) Basal half of embolium expanded laterally ( except N.
asiatica Horvath); ovipositor lobes of female quite dis-
tinctly projecting posteriorly beyond rest of abdomen 16

Basal half of embolium not expanded; ovipositor lobes of
female usually not projecting posteriorly beyond rest of

abdomen 19

16.(15) Mesosternal elevation apically greatly expanded laterally,
two times as wide as width of mid-femur, apex depressed
in middle, directed anteriorly as a shelf,

N. grandicoUis (Germar) p. 401
Mesosternal process apically not very expanded laterally,
about as wide as width of mid-femur, apex convex, not

directed anteriorly as a shelf 17

17.(16) Embolium not expanded laterally; females large over
10.5 mm.; lateral edge of pronotum more or less rounded,

N. asiatica (Horvath) p. 410
Embolium expanded laterally; females medium-sized,
not over 10.5 mm.; lateral edge of pronotum with several

sinuosities 18

18.(17) Ovipositor of female strongly lobed and projecting pos-
teriorly beyond rest of abdomen; expansion of embolium
more or less triangular, very salient; clasper of male with
right side of base of swollen apical half broadly angulate;

350 The University Science Bulletin

last visible abdominal stemite of female with lateral, sub-
marginal tumescences N. lohata ( Montandon ) p. 407

Ovipositor of female slightly projecting posteriorly be-
yond rest of abdomen; expansion of embolium more
elongated, but not so salient; clasper of male with right
side of base of swollen apical half more or less rounded;
last visible abdominal stemite of female without lateral,

submarginal tumescences N. indica (Atkinson) p. 405

19.(15) Females with posterior margin of last visible abdominal
stemite conspicuously emarginated; ovipositor lobes com-
pletely exposed, fitting into the emargination; males with
the greatest width of posterior margin of sixth abdominal
stemite (measured from one caudolateral angle to the
other) subequal to or greater than one half the width of
the fourth abdominal stemite (measured along the pos-
terior margin, from the median notch to the lateral
margin of the right side) [except N. nepaeformis

(Fabricius)] 31

Females with posterior margin of last visible abdominal
stemite not emarginate, usually more or less projecting
posteriorly, lobes of ovipositor usually only slightly
exposed or entirely covered by the last visible abdominal
stemite; males with width of the posterior margin of the
sixth abdominal stemite less than one half the width
of the posterior margin of the fourth abdominal ster-
nite [except IV. spissa (Distant), in which case the

wddths are subequal] 20

20. ( 19 ) Front of head with rows of white tubercles which fomi

an inverted V N. tuberculata (Montandon) p. 431

Front of head lacking such white tubercles 21

21.(20) Apex of head without toothlike tubercles 22

Apex of head with toothlike tubercles present 23

22.(21) Head entirely lacking toothlike tubercles N. ntidata n. sp. p. 425
Head with two small, broad, superapical, toothlike tuber-
cles present N. spissa (Distant) p. 412

23.(21) Anterolateral and lateral margins of pronotum concave,

N. laticollis (Guerin-Meneville) p. 418

Not with both anterolateral and lateral margins concave 24

24.(23) Pronotum distinctly wider than greatest width of abdomen,
abdomen less than 90% of width of pronotum,

N. inixta ( Montandon ) p. 420
Pronotum about equal to or only slightly wider than

abdomen, abdomen more than 90% of width of pronotum ... 25
25.(24) Basal portion of lateral margin of embolium convex; front
femora usually suflFused with red,

N. femaralis (Montandon) p. 423
Basal portion of lateral margin of embolium more or less

concave; front femora not suflFused with red 26

26.(25) Small species, under 9 mm.; basal portion of lateral margin
of embolium very slightly concave or straight; last visible

The Family Gelastocoridae (Hemiptera) 351

abdominal sternite of female with the posterior margin
angularly projecting on each side of apex, which is
slightly concave; clasper of male with basal portion of
apical half broadly expanded, flattened dorsoventrally,
small under 2.5 mm N. luteovaria (Distant) p. 430

Larger species, over 9 mm.; basal portion of lateral
margin of embohum concave; last visible abdominal
sternite of female with posterior margin laterad of apex
more or less rounded; clasper of male not as above,
not flattened dorsoventrally, either with apical half of
clasper expanded or clasper with the apical half not

expanded 27

27.(26) Clasper of male with the greatest portion of apical half

greatly expanded 28

Clasper of male not greatly expanded apically 30

*28.(27) Clasper with the dilation laterally angularly projecting,

forming abnost a right angle N. robusta n. sp. p. 429

Clasper with the dilation laterally not angularly project-
ing, more or less lobed, concavely depressed basad of

of the broadest part of the dilation 29

29.(28) Clasper large, over 4 mm. in length; lobe on right side very

prominent, constricted at base N. macrostyla n. sp. p. 428

Clasper smaller, under 3 mm. in length, indistinctly lobed,

lobe not constricted at base N. ampliata (Montandon) p. 426
30.(27) Lateral margin of embolium strongly concave at basal
third; clasper nearly straight, bent laterad at apex,

N. omani n. sp. p. 422

Lateral margin or embolium not as concave at basal third;
clasper more or less curved, not distinctly bent laterad

at apex N. gurneyi n. sp. p. 419

3L(19) Edge of anterior dilation of front femur nearly forming

a right angle with posterior side of femur 32

Edge of anterior dilation or front femur forming an acute

angle with posterior side of femur 33

32.(31 ) Anterior basal angle of anterior dilation of front femur more
or less rounded; last abdominal segment of female ven-
trally with a tumescence on either side of the emargina-
tion of posterior margin N. raptoria (Fabricius) p. 366

Anterior basal angle of anterior dilation of front femur
pointed; last abdominal segment of female ventrally
without a tumescence on either side of emargination of

posterior margin N. buenoi n. sp. p. 365

33.(31) Eighth abdominal sternite of male equal to or less than
length of ninth abdominal segment; ovipositor of female
longer than wide 34

* The writer also has been unable to find key characters for females of N. rohusta n. sp.,
N. macrostyla n. sp., N. ampliata (Montandon), N. omani n. sp., and N. gurneyi n. sp.
For the present it is necessary to use distribution, size and association with male to identify
these females. Fortunately, these are insular species and may be separated on distributional
data in most cases.

352 The University Science Bulletin

Eighth abdominal sternite of male longer than (usually
at least twice as long) ninth abdominal sternite; oviposi-
tor of female small, as wide as or wider than long 57

34.(33) Prominent patches of black bristles on scutellum 35

Without such patches of black bristles on scutellum 39

35. ( 34 ) Bristles on scutellum in narrow longitudinal bands 36

Bristles not in narrow longitudinal bands, usually in rather

oval patches 38

36. ( 35 ) Posterior part of abdomen strongly dilated laterally 37

Posterior part of abdomen not strongly dilated laterally,

N. unicornis ( Melin ) p. 378
37.(36) Bristles of scutellum short, clavate . N. montandoni (Melin) p. 380
Bristles of scutellum long, scarcely or not at all clavate,

N. peruviana (Montandon) p. 379
38.(35) Apex of the head concavely excavated,

A/, ranina ( Herrich-Schalfer ) p. 368
Apex of the head rather pointedly projecting; tubercles at

the apex fused basally N. unicornis ( Melin ) p. 378

39.(34) Apex of head pointedly projecting, terminating in one or

more apical tubercles 40

Apex of head not projecting pointedly, rather concavely

excavated 44

40.(39) Superapical tubercles present 51

Superapical tubercles absent 41

41.(40) Last abnominal sternite of male very large, approximately
two-fifths as wide as fifth abdominal sternite; last visible
abdominal sternite of female with anterior part of emar-

gination forming an acute angle 42

Last abdominal sternite of male smaller, about one-fourth
as wide as fifth abdominal sternite; last visible abdominal
sternite of female with anterior part of emargination

rounded 43

42.(41) Sides of pronotum at the level of the transverse furrow dis-
tinctly projecting beyond the base of embolium; sides
of pronotum noticeably converging anteriorly,

N. terrestris ( Kevan ) p. 373
Sides of pronotum at the level of the transverse furrow
very slightly projecting beyond base of embolium; sides
of pronotum more or less parallel JV. borealis ( Melin ) p. 375
43.(41) Basal third of embolium dilated, edge rounded; body flat-
tened; greatest width of seventh abdominal sternite of the
male less than one-half tlie width of the posterior margin
of the fourth abdominal sternite,

N. nepaeformis (Fabricius) p. 371
Basal third of embolium narrowed, not rounded, usually
slightly sinuous; body not so flattened; greatest width of
seventh abdominal sternite of male about equal to one-
half the width of the posterior margin of the fourth ab-
dominal sternite N. tenebrosa n. n. p. 376

The Family Gelastocoridae (Hemiptera) 353

44.(39) Males 45

Females 49

45. ( 44 ) Apical half of clasper sword- or sickle-shaped, more or less

simple 46

Apical half of clasper not as above 47

46. ( 45 ) Sides of pronotum rounded and projecting,

N. ranina (Herrich-Schaffer) p. 368
Sides of pronotum straight, slightly sinuous,

N. parvula (Signoret) p. 362
47.(45) Apical part of clasper bifurcate, processes of approximate

equal size and pointed N. ampIicoUis (Stal) p. 384

Not conspicuously bifurcate, but with a dorsomedial process,

this process much smaller than apical portion of clasper 48

48.(47) Dorsomedial process, oval, flattened in tangential plane to

longitudinal axis of clasper IV. ater (Melin) p. 389

Dorsomedial process, cylindrical or sometimes flattened in
radial plane to longitudinal axis of clasper,

N. ecuadorensis ( Melin ) p. 387
49.(44) Emargination of last visible abdominal stemite continuing
as a long curved groove to the right, forming a distinct

lobe or flap N. ranina ( Herrich-Schaffer ) p. 368

Emargination not as above 50

50.(49) Anterior end of left ovipositor lobe twisted to the right and
covering the anterior end of the right ovipositor lobe,

N. riidis (Melin) p. 390
Anterior end of left ovipositor lobe not twisted to cover an-
terior end of right ovipositor lobe,
N. amplicollis (Stal) p. 384 and N. ecuadorensis (Melin) p. 387

51.(40) Males 52

Females 54

52.(51) Apex of clasper of male very broad, flattened in an

oblique dorsoventral plane N. iisingeri Todd p. 360

Apex of clasper swollen but not as broad and not flattened

as above 53

53.(52) Clasper with slight processes on the lateral and dorsal

surfaces N. rruirtini Todd p. 358

Clasper with a prominent process on the dorsomedial

surface IV. mexicana ( MeUn ) p. 356

54.(51) Last abdominal stemite of female without obvious de-
pressions or tumescences, nearly symmetrical,

IV. lata (Montandon) p. 389
Last abdominal stemite of female with obvious depres-
sions or tumescences or both, asymmetrical 55

55. ( 54 ) Head with two apical tubercles 56

Head with only one or without apical tubercles, "N. parvula

Signoret p. 362 and N. quinquedentata (Melin) p. 363

56. ( 55 ) Last abdominal stemite more or less pointedly emarginated,

N. nwrtini Todd p. 358
Last abdominal stemite with a rounded emargination,

N. mexicana (Melin) p. 356 and N. usingeri Todd p. 360

12—6730

354 The University Science Bulletin

*57.(33) Ninth abdominal sternite moderately large, about one-half
as wide as eighth abdominal sternite; right side of pos-
terior margin of fifth abdominal sternite strongly convex,

N. conidis n. sp. p. 392
Ninth abdominal sternite small, approximately one-third
as wide as eighth abdominal sternite; right side of poste-
rior margin of fifth abdominal sternite nearly straight,

slightly concave 58

58.(57) Eighth abdominal sternite of male over one-half as wide

as fifth abdominal sternite N. hungerfordi n. sp. p. 398

Eighth abdominal sternite of male about one-half as wide

as fifth abdominal sternite 59

59.(58) Clasper with two projections on tlie medial surface,

N. fuscipes ( Guerin-Meneville ) p. 393

Clasper with one projection on the medial surface 60

60.(59) Mediobasal angle of medial projection of the clasper

rounded N. bracchialis n. sp. p. 400

Mediobasal angle of medial projection of the clasper

deeply sinuated iV. manni n. sp. p. 396

Nerthra stygica Say

(PI. XII, fig. 110; PI. XIII, fig. 117; PI. XIV, fig. 130)

1832. Nerthra stygica Say, Heteropterous Hemiptera of North America, New

Hamiony, Indiana, p. 37.
1859. N. stygica Leconte, Complete Writings of Thomas Say, vol. I, p. 364.
1876. A^. stygica Stal, Kongl. Svenska Vetenskaps-Akademiens Handlingar,

Band 14, no. 4, p. 139.

1905. N. stygica Torre-Bueno, Ohio Nat., vol. V, p. 288, figs. 1, 2.

1906. N. stygica Kirkaldy, Trans. Am. Ent. Soc, vol. XXXII, p. 149.

1910. N. stygica Banks, Catalogue of the Nearctic Hemiptera-Heteroptera,

separate publ. of Am. Ent. Soc, p. 11.
1914. N. stygica Barber, Bull. Am. Mus. Nat. Hist., vol. XXXIII, p. 498.
1917. iV. stygica Van Duzee, Catalogue of Hemiptera, Univ. California Publ.,

vol. I, p. 474.
1925. N. stygica Blatchley, Ent. News, vol. 36, pp. 49-52.

Also referring to this species:

1863. Mononyx stygica Stal, Berl. Ent. Zeitsch., vol. VII, p. 406.
1886. M. stygica Uhler, Check List of Hemiptera-Heteroptera of North
America, p. 27.

Size. Male: Length, 6.1 to 6.6 mm., width of pronotum, 4.4
mm., width of abdomen, 4.4 to 4.5 mm. Female: Length, 7.3 to
7.8 mm., width of pronotum, 5.0 to 5.2 mm., width of abdomen, 5.2
to 5.4 mm.

Color. Above: yellowish-brown to dark brown or even black,
usually with the anterior two thirds of each segment of the connexi-
vum, the scutellum, the disc of the pronotum and the head, darker.
Below: forelegs dark brown, others lighter, femora ringed with two

* Remaining couplets refer to males only. Females are unknown or do not appear to
possess key characters. Identification of the females must therefore be based upon size,
distribution and association with males.

The Family Gelastocoridae (Hemiptera) 355

brown bands; abdomen brown, posterolateral area of each segment
yellowish-brown, median portion of abdomen with a U-shaped
series of yellow or yellowish-brown spots.

Structural characteristics. Front of head projecting, apex with
small, rather sharp-pointed tubercles, superapical tubercles present,
broad, not sharp-pointed, a lateral toothlike tubercle present be-
tween the superapical tubercle and the eye, the lateral tubercle
small, irregular in shape; ocelli present, rather small, often diffi-
cult to locate in light colored specimens. Lateral margins of
pronotum nearly straight but slightly convergent anteriorly, an-
terior one fourth more abruptly converging; posterolateral mar-
gin rounded; posterior margin nearly straight, slightly sinuous
before scutellum; pronotum widest at transverse furrow, equal
to or slightly less than abdomen in width; disc elevated, lateral
expansions nearly flat. Scutellum moderately large, flat, de-
pressed at base and inclined to apex which is slightly elevated.
Hemelytra entirely coriaceous, fused together, extending posteriorly
as far as end of abdomen; basal half of embolium expanded lat-
erally, nearly as wide as pronotum. Connexivum expanded, but
more so in the female than the male. Bristles moderately long,
usually dark brown or black, clavate, those on the hemelytra ar-
ranged to some extent in faint rows following or paralleling the
sutures. Abdominal sternites of female asymmetrical; posterior
margin of last visible sternite deeply and triangularly emargi-
nate, keeled, keel anteriorly running obliquely to the right, a
small submarginal tumescence on the right side, a larger tumes-
cence to the left side, also a very prominent depression antero-
laterad of the left side of the emargination. Nearly identical
in the shape of the abdominal sternites to N. martini Todd or N.
mexicana (Melin). Abdominal sternites of male asymmetrical;
ninth sternite oval, wider than long, 32:15, fairly large, about one
fourth width of abdomen, subequal in length to eighth sternite, but
longer than seventh. Clasper of male very similar to that of the
N. mexicana (Melin) and A/, martini Todd. Apical half swollen
slightly and slightly recurved. Apex narrowed, rather abruptly
into short process which curves in toward the middle of the body,
a projection arising from the dorsomedial surface at the base of
the swollen portion of the clasper.

Location of type. Unknown. Some of Say's types are supposedly
in the collection of the Philadelphia Academy of Science, Philadel-
phia, Pennsylvania.

356 The University Science Bulletin

Distributional data. This species is known from Florida and
Georgia. The writer has studied specimens from the following
localities in Florida:

Capon, Apr. 18, no collector, 1 female, (USNM); Ch. Hbr.
(Charlotte Harbor?), no date, Mrs. A. T. Slosson, 1 female,
(AMNH); Dmiedin, March 20, 1923, W. S. Blatchley, 1 female,
(Pm-due Coll.); same place, Feb. 21, 1926, W. S. Blatchley, 1
female, (Purdue Coll.); Enterprise, May 13, no collector, 1 male,
(USNM); Fort Myers, Nov. 15, 1911, no collector, 1 male and
1 female, (AMNH); Jacksonville, no date, Ashmead, 1 female;
Pensacola, Oct. 11-14, 1914, no collector, 1 female, (AMNH).

Comparative notes. This species is quite close to N. mexicana
(Melin) and N. martini Todd. It differs from these two in that
the hemelytra are entirely coriaceous and fused together. If this
form had normal wings it would be quite difficult to separate it
from the two previously mentioned species. The pronotum is
slightly different also, but this may be due to the brachypterous
condition. This species can be separated from N. americana ( Mon-
tandon) and N. ivilJiamsi n. sp., which also have entirely coriaceous
and united hemelytra, by the presence of ocelli, shape of the abdom-
inal sternites of the female, and by the male clasper.

Nerthra mexicana (Melin)

(PI. Vll, fig. 43; PI. IX, figs. 69, 75; PI. X, fig. 94)

1929. Monomjx mexicanus Melin, Zoologiska Bidrag Fran Uppsala, Band 12,
pp. ] 87-188, figs. 80-83.

Size. Male: Length, 6.8 to 7.4 mm.; width of pronotum, 4.3 to
5.0 mm.; width of abdomen, 4.2 to 4.9 mm. Female: Length, 8.0
to 8.1 mm.; width of pronotum, 5.3 mm.; width of abdomen, 5.4 mm.

Color. Dark brown to black, ventral side slightly lighter than
dorsal surface, last four segments of male abdomen yellowish-
brown with dark brown punctations. Last abdominal segment and
anal flaps of female pale yellow. Posterior edge of segments of
connexivum with a thin, pale yellow area. Intermediate and hind
legs with two dark rings at middle of femur and three dark rings
on tibia, one basal, one apical and one medial. Front legs light brown
with dilation of femur and the trochanters lighter than the rest.

Structural characteristics. Apex of head more or less pointedly
projecting, apical tubercles rather slender, usually sharply pointed,
appearing between superapical tubercles when viewing the speci-
men from above; a few small tubercles laterad of superapical

The Family Gelastocoridae (Hemiptera) 357

tubercles. Pronotum widest at transverse furrow, not quite as wide
as abdomen in the female, slightly wider than abdomen in the
male; lateral margin of pronotum more or less rounded, anterior
two thirds to three fourths nearly straight, converging anteriorly,
posterior fourth of lateral margin nearly straight, weakly oblique.
Scutellum broad, rather large, without prominent tumescences.
Hemelytra extending beyond tip of abdomen; embolium narrow,
lateral margin nearly straight. Connexivum not very prominent,
almost covered by the hemelytra, especially in the male. Bristles
small, sparsely distributed on body on the specimens before the
writer. Abdominal sternites of the male asymmetrical; ninth ab-
dominal sternite small, oval, wider than long, slightly longer than
eighth abdominal sternite; seventh abdominal sternite roundly
spatulate on right side; fifth abdominal sternite very short medially.
Abdominal sternites of the female asymmetrical; posterior margin
of last visible abdominal sternite roundly emarginated; a moderate
tumescence on the left side, a faint tumescence on the right side
of the last visible abdominal sternite; a deep, elongate, transverse
depression extending along the anterior margin of the last visible
abdominal sternite from the median line of the abdomen to the
tumescence on the left side. Clasper of male swollen apically, a
rather prominent projection at the middle of the medial surface,
aedeagal furrow visible on apical half of ventral surface.

Location of type. In the Museum of Stockholm, Stockholm,
Sweden.

Distributional data. Melin listed four specimens in his descrip-
tion of this species. One is from Guatemala City, Guatemala, and is
a female. The other three are labeled "Mexico," "Salle" and com-
prise the type series. The writer has examined the type and also a
series of six specimens, three males and three females, from Teju-
pilco, Dist. of Temascaltepec, Mexico, June-July, 1933, H. E. Hinton.

Comparative notes. Closely related to N. martini Todd but differ-
ing in the shape of the clasper of the male and having the emargina-
tion of the posterior margin of the last visible abdominal sternite
of the female more rounded anteriorly than is the case in N. martini
Todd, and also differing in that the pronotum in this species is
usually wider than or subequal to the width of the abdomen, while
in N. martini Todd the pronotum is not as wide as the abdomen.

358 The University Science Bulletin

Nerthra martini Todd

(PI. VII, fig. 42; PI. IX, fig. 70; PL X, fig. 91)

1954. Nerthra martini Todd, Pan-Pacific Entomologist, XXX, No. 2, pp.
113-116, figs. 1 and 5.

Referring to this species:

1876. Mononyx badius Uhler, Bull. U. S. Geol. Surv., pp. 336-337.

1894. Mononyx stygicus Uhler, Proc. California Acad. Sci., ser. 2, vol. IV,
pp. 290-291.

Size. Male: Length, 7.0 to 7.6 mm.; width of pronotum, 4.5 to
4.8 mm.; width of abdomen, 4.5 to 5.0 mm. Female: Length, 7.8
to 8.8 mm.; width of pronotum, 4.8 to 5.3 mm.; width of abdomen,
5.0 to 5.8 mm.

Color. Varying from a yellowish-brown to black, generally a
brown or reddish-brown. A whitish spot on each hemelytron mesad
of the junction of the embolial and nodal sutures. In the black speci-
mens the posterior third of the pronotum is generally spotted with
pale yellow or white. Ventrally slightly lighter than dorsally; front
femur uniform brown, intermediate legs light brown ringed with
dark brown areas which may be faint in some specimens; last three
abdominal segments lighter than rest of abdomen; connexivum
with posterior third of each segment yellowish, anterior two thirds
dark. A few specimens examined were uniformly reddish-brown
above and with a lighter marginal band ventrally which extended
the entire length of the abdomen.

Structural characteristics. Apex of the head pointedly projecting
with a pair of rather sharp pointed tubercles at the tip, sometimes
these tubercles are rather rudimentary; superapical tubercles pres-
ent. Pronotum not very dilated laterally; lateral edge nearly straight,
occasionally weakly sinuous at middle; lateral edge conspicuously
converging anteriorly; posterior margin sinuated before scutellum.
Pronotum at greatest width not as wide as abdomen in either
sex. Scutellum broad, rather large with slight tumescence on either
side, mesad of which are two faint rounded depressions. Body
sparsely covered with very small bristles, those on the scutellum
being slightly clavate. Hemelytra well developed extending beyond
the apex of the abdomen. Lateral margin of embolium nearly
straight, not dilated. Connexivum barely visible from a dorsal view
in the male, quite prominent in the female because the posterior
abdominal segments are greatly dilated laterally. Ninth abdominal
sternite of the male rather large, over half as wide as eighth ab-
dominal sternite, slightly longer than seventh or eighth abdominal

The Family Gelastocoridae (Hemiptera) 359

sternites, asymmetrical. Abdominal sternites of female asymmetri-
cal; posterior margin of last visible abdominal sternite more or
less pointedly notched; a moderate tumescence on the left side, a
faint tumescence on the right side of the last visible abdominal
sternite; a deep, elongate, transverse depression extending along
the anterior margin of the last visible abdominal sternite from the
median line of the abdomen to the tumescence on the left side.
Clasper of male swollen apically; a weak process on the lateral sur-
face and another on the dorsal surface; aedeagal furrow visible on
the apical half of the ventral surface.

Location of type. Holotype male, allotype female, and 24 male
and 2 female paratypes, Los Pensaguitos Cr., San Diego Co.,
California, Apr. 8, 1930, C. H. Martin, in the Francis Huntington
Snow Entomological Collection at the University of Kansas. One
male and one female paratypes, San Felipe Cr., California, Apr. 14,
1935, C. E. Norland, (Usinger Coll.). One male and one female
paratypes, Los Angeles Co., California, no date, Coquillet, (USNM).
One male and one female paratypes. Riverside, California, March 2,
1927, T. Craig, (CAS).

Distributional data. In addition to the type series, the writer has
seen specimens from the following localities:

California. Afton Canyon, San Bernardino Co., July 17, 1931,
R. E. Blackwelder, 1 male, (CAS); Ardeou, Dec. 25, 1915, J. O.
Martin, 1 female, ( CAS ) ; Los Angeles, no date, P. R. Uhler, 1 male,
(USNM); Palm Canyon, Apr. 15, 1916, J. O. Martin, 1 male,
(CAS); Palm Springs, Riverside Co., Apr. 3, 1925, E. C. Van Dyke,
1 male, (CAS); same place, Jan. 3, 1929, Van Dyke, 1 female;
same place, Aug. 2, Hubbard, 1 female, (USNM); Panamint Vy.,
Apr. 1891, C. V. Riley, 2 females, (USNM); Pico, Mar. 7, 1916, J. O.
Martin, 1 male and 1 female, (CAS); Pine Valley, July 27, 1938,
R. I. Sailer, 1 female; Santa Ana Canyon, July 12, 1931, C. H.
Martin, 7 males and 3 females; Sierra Madre, June 7, 1930, C. H.
Hicks, 2 males, (J. C. Lutz Coll.); Sierra Nevada Mts., no date,
P. R. Uhler, 1 male, (USNM).

Baja California: Big Cyn., Sierra Laguna, Oct. 13, 1941, Ross
and Bohart, 2 males and 2 females, (CAS); Catavina, June 19,
1938, Michelbacher and Ross, 1 male, (CAS); Las Animas, Sierra
Laguna, Oct. 12, 1941, Ross and Bohart, 1 male and 2 females,
(CAS); Las Parras, Oct., 1925, W. M. Mann, 1 male, (USNM);
Loreto, Feb., 1923, W. M. Mann, 2 males and 3 females, (USNM);
19 mi. E. Rosario, June 17, 1938, Michelbacher and Ross, 3 males,

360 The University Science Bulletin

(CAS); "L. Cal," no date, P. R. Uhler, 1 male, (USNM).

Arizona: "Ariz.," no date, P. R. Uhler, 1 male (USNM); "Ariz.,"
no date, C. F. Baker, 1 male and 1 female, (USNM).

Georgia: "Dayton," July 9, 1926, J. R. Delaney, 2 females, (PAS),
J. C. Lutz Coll.*

Comparative notes. This species is very closely related to IV.
mexicana Melin and to N. stijgica Say. It is separated from the
former species by the narrow pronotum, slightly larger size, the
notch of the posterior margin of the last visible abdominal sternite
of the female being rather pointed at the anterior end and by the
shape of the clasper of the male. It differs from N. stijgica Say in
that it has separate hemelytra with well-developed membranes, the
emargination of the last visible abdominal sternite of the female
slightly more pointed, and in the shape of the male genitalia. It is
interesting to note that most specimens of N. stygica Say have
been taken during the winter or early spring. If the specimens
collected by J. R. Delaney are from Georgia, then there is a possi-
bility that Nerthra stygica Say is a winter form of this species.
During the summer of 1948, the writer attempted to locate this
species in the Gulf States, but he was not successful. Perhaps
future collecting and the development of our knowledge of the
biology of these species will reveal the true relationships of N.
stygica Say, N. mexicana (Melin) and N. martini Todd.

Nerthra usingeri Todd

(PL XV, fig. 141)

1954. Nerthra usingeri Todd, Pan-Pacific Entomologist, XXX, No. 2, pp.
116-117, fig. 2.

Size. Male: Length, 6.4 to 6.8 mm.; width of pronotum, 4.5 mm.;
width of abdomen, 4.4 to 4.5 mm. Female: Length, 7.1 to 7.7
mm.; width of pronotum, 4.8 to 5.0 mm.; width of abdomen, 4.7
to 5.0 mm.

Color. General over-all color yellowish-brown to dark brown
nearly black. Some specimens have the dorsal surface mottled
with black and yellowish-brown. Ventrally slightly lighter in
color than on the dorsal surface; front femora brown, darkest
basally, intermediate and hind legs yellowish-brown with dark
brown rings. Abdominal segments both above and below have
light yellowish spots at the latero-caudal angles of each segment.

* In the original description erroneously credited to the collection of the Philadelphia
Academy of Sciences.

The Family Gelastocoridae (Hemiptera) 361

Structural characteristics. Apex of head pointedly projecting,
usually with a pair of sharp-pointed tubercles at the tip, some speci-
mens have three and some only one apical tubercle, all are variable
in size. Pronotum with the lateral margins more or less rounded,
converging anteriorly and posteriorly; widest level with the trans-
verse furrow; posterior margin deeply sinuated at the middle
before the scutellum. Scutellum large, not strongly elevated.
Hemelytra with well-developed membranes, extending to or beyond
the end of the abdomen; lateral margins of the embolia nearly
straight for the basal half then gently curving medially. Connexi-
vum visible in both sexes, but not prominent; width of abdomen
equal to or subequal to the width of pronotum. Bristles very small,
short, peg-like, sparsely distributed over the body. Abdominal
sternites of the male asymmetrical; ninth sternite small, oval wider
than long, slightly longer than eighth sternite; seventh abdominal
sternite spatulate on right side. Abdominal sternites of female
asymmetrical; posterior margin of last visible abdominal sternite
roundly emarginated; a moderate tumescence on the left side, a
faint tumescence on the right side of the last visible abdominal
sternite; a deep, elongate, transverse depression extending along
the anterior margin of the last visible abdominal sternite from the
median line of the abdomen to the tumescence on the left side.
Clasper of male very distinctive, broad and flattened apically, widest
near apex; aedeagal furrow visible on apical half of ventral surface,
terminating before reaching apex; clasper flattened in an oblique
dorsoventral plane.

Location of type. Holotype male, Nr. Parker Dam, Cahfornia,
Apr. 12, 1952, R. L. Usinger and allotype female. Wash 3.5 mi. N.
Cross Roads, California, April 12, 1952, J. D. Lattin, in the collection
of the California Academy of Science at San Francisco. One male
and five female paratypes. Wash 3.5 mi. N. Cross Roads, California,
April 12, 1952, J. D. Lattin, in the R. L. Usinger Collection at the
University of California. One male and one female paratypes, same
data, in the personal collection of Mr. Lattin.

Distribution. Known at this time only from the type locality.
The difference in the locality of the holotype and the remainder of
the specimens examined is merely a difference in labelling the
specimens on the part of the two collectors. These data represent
the same locality.

Comparative notes. Closely related to Nerthra sttjgica Say, N.
mexicana (Melin) and N. martini Todd. This species may be sep-

362 The University Science Bulletin

arated from N. stijgica Say by the fact that the hemelytra are normal
with well-developed membranes. The distinctive clasper of the
male will separate this species from the males of N. mexicana
(Melin) and N. martini Todd. The females of this species differ
from the females of N. martini Todd in that the emargination of
the last visible abdominal sternite is broadly rounded basally. To
the knowledge of the writer the females of this species may be sep-
arated from females of N. mexicana (Melin) only by distribution.

Nerthra parviila ( Signoret )

(PI. VII, fig. 44; PI. VIII, fig. 67)

1864. Mononyx parvultis Signoret, Annales de la Societe des Entomologique

de France, XXXIII, p. 588.
1873. M. porvulus Walker, Catalogue of Hemiptera in British Museum, Part

VIII, p. 172.
1899. M. parvulus Montandon, Bulletin de la Societe des Societe des Sciences

de I3ucarest-Roumanie, An. VIII, Nos. 4 & 5, p. 402.
1909. M. parvulus Kirkaldy and Torre-Bueno, Proc. Ent. Soc. Washington,

Vol. X, p. 182.
1929. M. parvulus Mehn, Zoologiska Bidrag Fran Uppsala, Band 12, pp.

188-189, figs. 88-90, 110.
1935. M. parvulus De Carlo, Revista Chilena de Historia Natural, Afio

XXXIX, pp. 106-107.

Size. Male: Length, 5.8 to 6.1 mm.; width of pronotum, 3.7 to
3.8 mm.; width of abdomen 3.6 to 3.9 mm. Female: Length, 7.0 to
7.5 mm.; width of pronotum, 4.3 to 4.9 mm.; width of abdomen,
5.0 to 5.4 mm.

Color. Yellowish-brown, scutellum darker; a dark brown spot
at distal end of embolial suture; a small, median, blackish spot on
the posterior margin of the head; segments of the connexivum en-
tirely dark brown or darker on the basal portion; front legs brown;
intermediate legs, hind legs and abdomen pale yellowish-brown,
ringed and spotted with brown respectively.

Structural characteristics. Apex of the head weakly pointedly
projecting, terminating in a small blunt knob; small indistinct
superapical tubercles present (larger in females); lateral tubercles
small, indistinct (more prominent in females). Pronotum widest at
at level with the transverse furrow, barely projecting beyond base
of embolium, about equal to abdomen in width in males, distinctly
less in width than abdomen in females; lateral margins nearly
straight for anterior three-fourths, slightly sinuated at middle;
posterior margin sinuated before scutellum. Scutellum rather large,
with a moderate tumescence on either side and a weak median
longitudinal carination. Hemelytra extending to or beyond the end
of the abdomen; membrane well developed; embolium elongate, lat-

The Family Gelastocoridae (Hemiptera) 363

eral margin straight or weakly convex. Connexivum nearly covered
by the hemlytra in males, very prominent in females. Abdominal
sternites of female similar to those of N. quinquedentata (MeHn)
except, there are no lateral tumescences on the last visible ab-
dominal sternite and the depressions of that sternite are generally
broader and not so deep as in the aforementioned species. Ab-
dominal sternites of the male asymmetrical; ninth abdominal ster-
nite oval, wider than long, about equal in length to eighth sternite.
Clasper of the male simple, sickle-shaped, rather slender, aedeagal
furrow visible on the apical half of the medioventral surface.

Location of type. In the British Museum at London, England.

Distributional data. This species is known only from Chile. The
writer has seen specimens from the following localities in Chile:
Las Brisas, Aug. 12-14, 1951, Luis Peiia, 4 males and 1 female;
Toulemo, Nov. 15-27, 1951, Luis Peiia, 1 male and 1 nymph; El
Canelo, Dec. 1950, Luis Peiia, 1 male and 1 female; Guayacan, Jan.
25, 1951, Luis Peiia, 1 female; El Manzano, Nov. 1951, Luis Pena,
2 females; "Sonora," no date, P. R. Uhler, 1 male, (USNM); "Chile,"
no date, Porter, 1 male, (USNM).

Comparative notes. This small species may be separated from
N. ranina ( Herrich-SchaflFer ) by the shape of the head, the pro-
notum not so produced laterally and by the shape of the clasper of
the male and by the shape and depressions of the abdominal ster-
nites of the female. This species is apparently very closely related
to Nerthra quinquedentata (Melin) and the latter may only be a
larger form, but there appear to be diflFerences in the females avail-
able to this worker and it would seem best to consider them as dis-
tinct species for the present.

Nerthra quinquedentata (Melin)

(PI. X, fig. 84)

1929. Monomjx quinquedentatus Melin, Zoologiska Bidrag Fran Uppsala,
Band 12, p. 188, figs. 84-87.

Size. Female: Length, 7.5 to 8.2 mm.; width of pronotum, 4.9
to 5.4 mm.; width of abdomen, 5.0 to 5.8 mm.

Color. Light yellowish brown to black, scutellum and median
part of pronotum darker; a small dark brown spot at the distal
end of the embolial suture; anterior part of segments of the con-
nexivum, brown or black, posterior part pale yellow; front legs
light brown, intermediate and hind legs pale yellow ringed with
brown; abdominal segments dark brown with yellowish-brown
punctations.

364 The University Science Bulletin

Structural characteristics. Apex of head pointedly projecting,
terminating in one apical tubercle; a pair of superapical tubercles
present; one lateral tubercle present on either side of the head,
as large as the superapical tubercles. Pronotum widest at a level
with the transverse furrow, not as wide as the abdomen; lateral
margin with the anterior three fourths nearly straight to weakly
rounded, distinctly converging anteriorly; posterior margin sinu-
ated before scutellum. Scutellum fairly large with moderate
lateral tumescences and a weak longitudinal carination; a slight
depression between the lateral tumescence and carination; tu-
mescences and carination covered rather thickly with brown bristles
which are slightly clavate. Hemelytra extending beyond the end
of the abdomen. Connexivum moderately prominent. Abdominal
sternites of the female asymmetrical; last visible abdominal sternite
with the posterior margin medially deeply emarginated, anterior
end of emargination twisted slightly to the right; a tumescence near
the lateral margin on each side, the one on the left side much
larger than that on the right side; three depressions on the last
sternite, one on either side of the emargination and one medial
depression, the lateral depression on the left side large and extend-
ing from a point slightly to the left of the median line along the
anterior margin of the last visible sternite to the base of the large
tumescence on the left side.

Location of type. In the Museum of Copenhagen, Copenhagen,
Denmark.

Distributional data. Melin studied two specimens from Argen-
tina. This worker has examined specimens from the following
localities:

Argentina: "Argentina," no date, Wehseregt, 1 female, (Mus.
Prague); Mendoza, no date, G. S. Reed, 1 female.

Brazil: Nova Teutonia, Dec, 1946, Fritz Plaumann, 1 female.

Chile: "Chile," Dec, 1923, no collector, 4 females.

Comparative notes. This species is easily identified by the pres-
ence of five tubercles on the front of the head and by the shape
of the abdominal sternites of the female. It is very closely related
to N. parvula (Signoret), but the last visible abdominal sternite has
lateral tumescences which are lacking in the latter species and the
depressions are narrower and deeper. In many ways this species
shows affinities to the stygica group even though its range is remote
from those species.

The Family Gelastocoridae (Hemiptera) 365

The four females from Chile are not typical. The toothlike
tubercles of the head are almost completely obsolete and the
abdominal sternites vary slightly from those of the typical female
in that the apex of the emargination of the last visible sternite is
not bent to the right and in that the tumescence on the left side
of the last visible sternite is a little larger than in the typical female.

Nerthra biienoi n. sp.

(PI. Ill, fig. 10; PI. X, fig. 82)

Size. Female: Length, 4.2 mm.; width of f)ronotum, 2.6 mm.;
width of abdomen, 2.7 mm.

Color. Uniformly brown above, except the posterior half of
each segment of the connexivum and the tip of the membrane of
the hemelytra which are pale yellowish-brown. Light yellowish-
brown ventrally with brown punctations on the abdominal seg-
ments, tarsal claw of foreleg dark brown, intermediate and hind
legs with the femora ringed medially by two faint, slightly darker
areas.

Structural characteristics. Front of head with apex concavely
excavated, a stout tubercle on either edge of the excavation; lateral
tubercles small forming into a group between the eye and the
excavation of the apex. Lateral margin of the pronotum with the
anterior two thirds faintly rounded, posterior third straight, ob-
liquely converging to base of embolium; posterior margin sinuated
before scutellum. Pronotum widest at transverse furrow, not quite
as wide as abdomen. Scutellum fairly large with weak tumescences
on either side. Hemelytra extending beyond tip of abdomen; em-
bolium narrow, elongate, lateral margin weakly curved. Body
covered sparsely with short clavate bristles. Anterior margin of front
femur forming approximately a right angle with posterior margin
of femur; anterior dilation projecting into a rather sharp point at
the anterior basal angle. Abdominal sternites of the female nearly
symmetrical; emargination of the posterior margin of the last visible
abdominal sternite rounded; lateral areas on either side of emargina-
tion flattened, without a trace of tumescences.

Location of type. Holotype, female, from Bnito Prov., Pernam-
buco, Brazil, Feb., '83, no collector, in the Francis Huntington Snow
Entomological Museum at the University of Kansas.

Distributional data. As given for the type.

Comparative notes. This species though very closely related to
N. raptoria (Fabricius) seems sufficiently different to constitute

366 The University Science Bulletin

another species. It differs from the preceding species by the more
pointedly projecting anterior dilation of the front femur, by the
absence of tumescences on the last visible abdominal stemite and
by its smaller size.

Nerthra raptoria (Fabricius)

(PI. HI, fig. 7; PI. VII, fig. 46; PI. VIII, fig. 64; PI. X, fig. 88)

1803. Naucoris raptoria Fabricius, Systema Eleutheratorum, vol, 3, Systema

Rhyngotarum, p. 111.
1833. Mononyx raptoria Laporte, Essai d'une Classification Systematique de

L'order des Hemipteres, p. 16.
1835. M. raptoria BruUe, Histoire Naturelle des Insectes, vol. IX, p. 275.
1840. M. raptoria Spinola, Essai sur les Insectes Hemipteres, Rhyngotes ou

Heteropteres, p. 63.
1840. M. raptoria Blanchard, Histoire Naturelle des Animaux Articules, vol. 3,

p. 93.
1843. M. raptorius Guerin-Meneville, Revue Zoologique Travaux Inedits,

p. 114.
1859. M. raptorius Dohm, Catalogus Hemipterorum, Herausgegeben von dem

Entomologischen Verein zu Stettin, p. 53.
1863. M. raptorius Stal, Berliner Entomologische Zeitschrift, Bd. VII, p. 405.
1865. M. raptorius Mayr, Zoologischer Theil, Band II, Abt. 1, p. 181.
1868. M. raptorius Stal, Hemiptera Fabriciana, I, p. 134.
1876. M. raptorius Stal, Kongl. Svenska Vetenskaps-Akademiens. Handlingar,

Band 14, no. 4, p. 139.
1879. M. raptorius Berg. Hemiptera Argentina, p. 186.
1886. M. raptorius Uhler, Bull. Brooklyn Ent. Soc, p. 27.
1895. M. raptorius Montandon, BoUettino dei Musei di Zoologia ed Anatomia

comparata della R. Universita di Torino, no. 219, vol. X, p. 8.
1899. M. raptorius Montandon, Bulletin de la Societe des Sciences de Bucarest-

Roumanie, an. VIII, nos. 4 et 5, p. 402.
1901. M. raptorius Champion, Biologia Centrali- Americana, Rhynchota Heter-

optera, vol. II, p. 352, pi. 20, figs. 27, 27a.
1906. M. raptorius Distant, Fauna British India, vol. Ill, p. 14.
1906. M. raptorius Kirkaldy, Trans. Am. Ent. Soc, XXXII, p. 149.
1909. M. raptorius Kirkaldy and Torre-Bueno, Proc. Ent. Soc. Washington,

vol. X, p. 182.
1929. M. raptorius Melin, Zoologiska Bidrag Fran Uppsala, Band 12, pp.
191-192, figs. 96-99, 105, 111.

Also referring to this species:

1858. M. fusco-conspersus Stal, Kongl. Svenska Vetenskaps-Akademiens

Handlingar, 2, p. 82.
1863. M. fusco-conspersus Stal, Berliner Entomologische Zeitschrift, VII,

p. 406.
1873. M. fusco-conspersus Walker, Catalogue of Hemiptera in the British

Museum, pt. VIII, p. 172.

Size. Males: Length, 5.0 to 5.5 mm.; width of pronotum, 3.1 to
3.4 mm.; width of abdomen, 3.0 to 3.35 mm. Female: Length, 5.4
to 5.5 mm.; width of pronotum, 3.55 to 3.6 mm.; width of abdomen,
3.55 to 3.6 mm.

Color. Light to dark brown above; scutellum darkest, occa-
sionally black. Anterior dilation of front femora, intermediate and
hind legs pale yellowish. Tibiae and tarsi of intermediate and hind

The Family Gelastocoridae (Hemiptera) 367

legs with two indistinct dark rings. Venter of abdomen, brown with
darker punctations. Margin of abdominal segments with the
anterior half dark brown, the posterior half pale yellow.

Structural characteristics. Apex of head concavely excavated
with a stout tubercle on either side of the excavation; lateral tuber-
cles irregular, small, grouped together to form a small raised knob
between eye and apex of head; ocelli rather large. Margins of pro-
notum more or less rounded, occasionally with sides straighter,
forming two obtuse lateral angles; slightly wider than abdomen
in the males, about the same width as abdomen in the females;
posterior margin with area before scutellum deeply sinuous. Scutel-
lum with lateral tumescences prominent; a weak, median longitu-
dinal carination present; tumescences and carina covered very
sparsely with slightly clavate bristles. Hemelytra extending beyond
tip of abdomen; a rather large spot which is dark and smooth at
the medial angle of embolium; two groups of bristles mesad and
one group of bristles caudad of this spot; bristles present along
embolial suture, external edge of embolium and edge of corium;
embolium narrow, elongate and slightly convex at external edge.
Anterior edge of front femur greatly dilated, forming an almost
right angle with the posterior edge. Abdominal sternites of the
male asymmetrical; ninth sternite slightly longer than eighth ster-
nite, wider than long. Abdominal sternites of the female nearly
symmetrical; last visible abdominal sternite deeply emarginated;
ovipositor lobes nearly as long as last segment of abdomen; a
rather small tumescence on either side of emargination near the
lateral margins, the one on the left side being slightly the larger.
Clasper of the male small, sickle-shaped, apex faintly hooked,
aedeagal furrow visible on the distal half.

Location of type. The type specimen, a male, is located in the
Museum of Copenhagen, Copenhagen, Denmark.

Distributional data. The literature lists this species from Mexico
through Central and South America to Argentina. This writer has
examined specimens from the following localities:

Fanamd: Paraiso, C. Z., Jan. 16, 1911, E. A. Schwarz, 1 female,
(USNM); "Pan.," no date, P. R. Uhler, 1 female, (USNM).

Brazil: Santarem, no date or collector, 1 male, compared with
type by Doctor H. B. Hungerford in 1928; Rio Caiari-Uaupes,
Amazonas, Sept. 1906, H. Schmidt, 1 male, (AMNH); West border
of Matto Grosso, May, 1931, R. C. Shannon, 1 female, (USNM).

368 The University Science Bulletin

Comparative notes: This species in the past was easily separated
from all others by the shape of the anterior dilation of the front
femur. It can be separated from N. buenoi n. sp. by its larger size,
the presence of tumescences on the last visible abdominal sternite
of the female and by the rounded anterior basal angle of the
anterior dilation of the front femur.

Nerthra ranina (Herrich-Schaffer)

(PI. II, figs. 3, 4; PI. Ill, figs. 6, 9; PI. IV, fig. 12; PI. VII, fig. 41; PI. X, fig. 81)

1853. Mononyx raninus Herrich-Schaffer, Die Wanzenartigen Insecten, vol.

IX, p. 28, fig. 896.
1859. M. raninus Dohrn, Catalogus Hemipterorum, Herausgegeben von

dem Entomologischen Verein zu Stettin, p. 53.
1863. M. raninus Stal, Berliner Entomologische Zeitschrift, vol. VII, p. 406.
1899. M. raninus Montandon, Bulletin de la Societe des Sciences de Bucarest-

Roumanie, an. VIII, nos. 4 and 5, p. 401.
1909. M. raninus Kirkaldy and Torre-Bueno, Proc. Ent. Soc. Washington,

vol. X, p. 182.
1929. M. raninus Melin, Zoologiska Bidrag Fran Uppsala, Band 12, pp.

189-191, figs. 91-94.

Also referring to this species:

1929. Mononyx niger Melin, Zoologiska Bidrag Fran Uppsala, Band 12,
pp. 182-184, figs. 63, 64.

Size. Male: Length, 6.2 to 7.0 mm.; width of pronotum, 4.2 to
4.9 mm.; width of abdomen, 4.0 to 4.9 mm. Female: Length, 5.8
to 8.5 mm.; width of pronotimi, 3.8 to 6.0 mm.; width of abdomen,
3.8 to 6.0 mm.

Color. As with size this species is very variable in coloration.
The color ranges from pale yellow or light grey to black. The
writer has examined some specimens showing a combination of
pale yellow with a dorsal median stripe of black.

Structvral characteristics. Apex of head concavely excavated,
with a broad tubercle on either side of excavation; lateral tubercles
rudimentary, generally forming an indistinct toothlike protuber-
ance before the lateral depression of the head, but occasionally sep-
arate and conspicuous. Pronotum with greatest width at transverse
furrow, lateral margins usually rounded; posterior margin sinuated
before the scutellum. Scutellum fairly large with a tumescence
frequently covered with bristles, especially in small specimens
from the southern part of its range. Hemelytra extending beyond
end of abdomen. Bristles sparsely scattered over body, worn off
in most specimens, slightly clavate in larger specimens, distinctly
clavate in small specimens. Connexivum prominent, especially in
the female; posterior half of each segment light, anterior half dark.

The Family Gelastocoridae (Hemiptera) 369

Abdominal sternites of both male and female asymmetrical. Ninth
abdominal sternite of male small, approximately equal in length to
eighth abdominal sternite. Seventh abdominal sternite at its
greatest length as long as eighth abdominal sternite or occasionally
longer. Abdominal sternites of female very characteristic. Notch
of posterior margin of last visible abdominal sternite continued as
a long, narrow curved groove to the right, forming a lobe on that
side; an elongate depression continuing from the groove posteriorly
and a circular depression at the median part of the anterior margin.
A large tumescence on the left side extending along the margin of
the last two abdominal sternites, occasionally a very small tumes-
cence on the right side of the last visible abdominal sternite near the
lateral margin. The clasper of the male small and sickle-shaped,
twisted, aedeagal furrow visible on apical half viewing the speci-
men ventrally.

Location of type. Unknown. Other Hemipterous insects of the
collection of Herrich-Schaffer are located in the Museum of
Munchen, Munchen, Germany.

Distributional data. The literature lists this species from Argen-
tina, Brazil and Paraguay. The writer has studied specimens from
the following localities:

Peru: Callango, no date, G. W. Kirkaldy, 5 males and 1 female;
same place, no date, Haglund, 1 female, (Mus. Stockholm), type of
Mononyx niger Melin; Guayabamba, Andes, Dept. Amazonas, Aug.
14-19, 1936, F. Woytkowski, 1 male; Huanuco, Andes, Dept.
Huanuco, Sept. 6, 1937, F. Woytkowski, 18 males and 16 females;
Marcapata, no date or collector, 1 male and 9 females; Pacasmayo,
May 19-20, 1936, F. Woytkowski, 1 male; Puerto Bermudez, Rio
Pichis, July 13-19, 1920, no collector, 1 male; Recardo Palma, 44
kil. E. Lima, Nov. 27-29, 1934, F. Woytkowski, 1 female; Rio Perene
and Chanchamayo, Dept. Junin, May, 1934, F. Woytkowski, 2 males
and 4 females; San Beni, Dept. Junin, Sept. 5, 1935, F. Woytkowski,
2 males and 2 females; San Ildefonso, Rio Utcubamba, Dept. Ama-
zonas, July 29, 1936, F. Woytkowski, 1 male and 1 female; San
Miguel, Prov. La. Mar., Dept. Ayacucho, July 9-12, 1941, F. Woyt-
kowski, 12 males and 8 females; San Pedro, May 15-29, 1935, F.
Woytkowski, 1 male; Santa Clara, River Rimac, Sept. 22-25,
1934, F. Woytkowski, 7 males and 4 females; Tinccochaca, Aug.,
1911, no collector, 1 female; Yanamonte, Prov. La. Mar., Dept.
Ayacucho, Sept. 28-30, 1941, F. Woytkowski, 2 males and 1 female.

370 The University Science Bulletin

Chile: Santiago,* Aug. 26, 1949, L. E. Pena, 1 female.

Bolivia: Coroico, no date, purchased from Dr. O. Staudinger, 1
female; Espia, 1921-22, W. M. Mann, 1 male and 1 female,
(USNM); Mojos, no date, N. Holingren, 2 males, (Mus. Stock-
holm); San Fernando Rapids, 1921-22, W. M. Mann, 1 female,
(USNM); Sta. Helene, 1921, W. M. Mann, 1 female, (USNM).

Paraguay: "Albovena Srojo guasi," Nov. 16, 1926, F. Schade, 3
females; Alboveno, May 1, 1925, F. Schade, 1 female; Asuncion,
July, no collector, 3 females, (Car. Mus.); same place, Nov., no
collector, 1 female, (Car. Mus.); Horgueta, Aug. 11, 1933, A.
Schultze, 1 male and 1 female, ( Usinger Coll. ) ; same place, 45
mi. E., July 6, 1933, A. Schulze, 1 female, (J. C. Lutz Coll.); same
place, Aug. 7, 1933, A. Schulze, 1 male and 1 female, (J. C. Lutz
Coll.); same place, Aug. 10, 1933, A. Schulze, 3 females (J. C.
Lutz Coll.); Villarrica, Jan. 6, 1923, F. Schade, 1 female; same
place, May 9, 1923, F. Schade, 1 female, Villarica Serro Pelado,
Nov. 28, 1929, F. Schade, 1 female.

Argentina: Catamarca, Jan. 24, 1924, Weiser, 1 female; same
place, Corral Quemado, Mar., 1924, Weiser, 1 male and 1 female;
same place, Apr. 1924, Walters, 1 female; same place, Fuerte Que-
mado, Feb. 1924, F. Walters, 1 male and 1 female; same place,
Valle de Santa Maria, no date, F. Walters, 1 male; Chafiiian, Valde
9, Jan., 1922, Weiser, 1 male; Potrerillos, Mar. 16-19, 1920, no col-
lector, 1 female; "Argentina," no date, Jensen-Haarup, 1 female;
"Argentina," no date, Weiser, 1 male, (Mus. Prague).

Brazil: Nova Teutonia, Jan. 8, 1949, F. Plaumann, 4 males and
16 females; same place, Jan., 1948, F. Plaumann, 5 males and 4
females; same place, Apr. 1948, F. Plaumann, 4 males and 7 fe-
males; Nova Teutonia, Santa Catharina, Apr. 27, 1948, F. Plaumann,
5 males and 3 females; same place. Mar. 5, 1948, F. Plaumann,
1 female; Nova Teutonia, Tupan L. 2, May, 1947, F. Plaumann, 12
males and 5 females; Para, July, no collector, 1 female, (Car. Mus.);
Sao Paulo, no date, Mraz., 1 male, (Mus. Prague).

Comparative notes. This species can be easily separated from
the other species of the genus by the shape of the male clasper
and by the characteristics of the abdominal sternites of the female.
The writer, after examining the type of Mononyx niger Melin, is
of the opinion that the latter species is a synonym of Nerthra
ranina ( Herrich-Schaffer ) . There is considerable variation within

* This locality may be incorrect, however, the specimen is from Chile and extends the
range of this species to that country.

The Family Gelastocoridae (Hemiptera) 371

this species as to color and size. The specimens from Peru and
Bolivia are on the whole darker and somewhat larger than those
from Brazil, Paraguay and Argentina. The abdominal stemites of
the female and the male clasper as well as the other structures of
the body are identical.

Nerthra nepaeformis ( Fabricius )

(PI. VII, fig. 49; PI. IX, figs. 72, 74; PI. X, fig. 93)

1775. Naucoris nepaeformis Fabricius, Systema Entomologiae, 2, p. 693.

1803. N. nepaeformis Fabricius, Systema Eleutheratorum, vol. 3, Systema
Rhyngotarum, p. 111.

1868. Mononyx nepaeformis Stal, Hemiptera Fabriciana, I, p. 134.

1873. M. nepaeformis Walker, Catalogue of Hemiptera in the British Mu-
seum, pt. VIII, p. 171.

1876. M. nepaeformis Stal, Kongl. Svenska Vetenskaps-Akademiens Hand-
lingar, vol. V, p. 138.

1879. M. nepaeformis Berg, Hemiptera Argentina, pp. 185-186.

1895. M. nepaeformis Pittier and BioUey, Instituto Fisico Geografico Nacional
(Costa Rica), p. 23.

1895. M. nepaeformis Montandon, Bollettino dei Musei di Zoologia ed
Anatomia comparata deUa R. Universita di Torino, no. 219, vol. X,
p. 8.

1899. M. nepaeformis Montandon, Bulletin de la Societe des Sciences de
Bucarest-Roumanie, an. VIII, nos. 4 et 5, p. 401.

1901. M. nepaeformis Van Duzee, Trans. Am. Ent. Soc, vol. XXXVII, p. 352.

1901. M. nepaeformis Champion, Biologia Centrali-Americana, Rhynchota
Heteroptera, vol. II, p. 351.

1906. M. nepaeformis Torre-Bueno, Ent. News, vol. 17, p. 54.

1909. M. nepaeformis Kirkaldy and Torre-Bueno, Proc. Ent. Soc. Washing-
ton, vol. 10, p. 182.

1925. M. nepaeformis Blatchley, Ent. News, vol. 36, p. 52.

1929. M. nepaeformis Melin, Zoologiska Bidrag Fran Uppsala, Band 12, pp.
174-177, figs. 36-38, 103, 107.

1935. M. nepaeformis Carlo, Revista Chilena de Historia Natural, ano XXXIX,
pp. 106-107.

1942. M. nepaeformis Kevan, Proc. Royal Ent. Soc. London, (a) 17, pts. 10-12,
p. 110.

Also referring to this species:

1835. Mononyx raptorius, Burmeister, Handbuch der Entomologie, Bd. 2,
Abteil. 1, p. 201. (nee. Fabricius)

1853. M. raptorius Herrich-Schiiffer, Die Wanzenartigen Insecten, IX, p. 27,
t. 291, fig. 895.

1843. Mononyx raptorius, Amyot et Serville, Histoire Naturelle des Insectes,
Hemipteres, p. 425. (nee. Fabricius)

1854. Mononyx bipunctatus Stal, Ofversigt af Kongl. Vetenskaps-Akademiens
Fordhandlingar, no. 3, XI, p. 239.

1858. M. bipunctatus Stul, Kongl. Svenska Vetenskaps-Akademiens Hand-
lingar, 2, p. 82.

1859. M. bipunctatus Dolirn, Catalogus Hemipterorum, Herausgegeben von
dem Entomologischen Verein zu Stettin, p. 53.

1863. M. bipunctatus Stal, Berliner Entomologische Zeitschrift, VII, p. 405.

Size. Male: Length, 7.6 to 7.9 mm.; width of pronotum, 5.0 to 5.3
mm.; width of abdomen, 5.3 to 5.4 mm. Female: Length, 7.9 to 9.1
mm.; width of pronotum, 5.5 to 6.1 mm.; width of abdomen 5.5 to
6.2 mm.

372 The University Science Bulletin

Color. Yellowish-brown to reddish-brown, usually the former;
a dark brown spot at distal end of embolial suture, a black spot
medially at posterior margin of head; segments of connexivum with
a thin dark area at the anterior margin; front femora yellowish-
brown, darker on the posterior third; intermediate and hind legs
yellowish ringed with brown; abdominal segments brown with spots
of yellow, especially along the lateral margins.

Structural characteristics. Apex of head more or less pointedly
projecting, terminating in a pair of apical tubercles; lateral tubercles
of the head smaller, separate and four or five in number on each side
of the head. Pronotum widest at a level of the transverse furrow,
about as wide as abdomen, projecting beyond base of embolium;
lateral margins of the pronotum irregular in shape, usually with the
median part straight or faintly rounded, the anterior and posterior
parts straight or slightly sinuous and converging sharply toward
the eye and base of embolium respectively; posterior margin nearly
straight, weakly sinuated before scutellum. Scutellum without no-
ticeable tumescences and a slight depression on either side of the
median line of the body. Hemelytra extending to the end of the
abdomen; embolium rather dilated, lateral margin more or less
rounded. Connexivum prominent in both sexes, but especially so
in the female. Ninth abdominal sternite of the male small, oval,
two-thirds as long as wide, longer than eighth abdominal sternite;
abdominal sternites asymmetrical. Abdominal sternites of the fe-
male almost symmetrical; last visible abdominal sternite medially
roundly emarginated, a shallow depression on either side of emargi-
nation laterad of which is a small tumescence. Ovipositor lobes
large, elongate. Clasper of the male sickle-shaped, simple, a shght
constriction present just before apex making the apex bluntly spatu-
late; aedeagal furrow visible on the ventral surface for only a short
distance at the apex.

Location of type. Unknown. Doctor H. B. Hungerford examined
a specimen of this species in the Kiel Museum, Kiel, Germany in
1928. He states in his notes that the specimen is a female "with
Fabr. label."

Distributional data. The literature gives this species a rather
extensive range. Specimens from Mexico to Chile have been identi-
fied as this species. All the specimens examined by Melin, with the
exception of one nymph which was questionably identified as be-
longing to this species, were from Brazil. This worker has examined
specimens from the following localities:

The Family Gelastocoridae (Hemiptera) 373

Brazil: Bahia, no date, Rolle, 1 female, (CAS); Vic. Joao Pessoa
(Sao Phelipe), River Junui, July 10 to Sept. 20, 1936, A. M. Olalla,
1 female; Jurujuba, Rio de Janeiro, Jan. 6, 1920, E. G. Holt, 1 female;
Nova Friburgo, May, 1935, D. M. Cochran, 1 female, (USNM);
Nova Teutonia, May 15, 1935, F. Plaumann, 1 male and 1 female,
(Usinger Coll.); Rio de Janeiro, Mar. 4, 1935, P. Sandig, 1 female,
(USNM); Rio, no date, P. R. Uhler, 1 female, (USNM); Sao Paulo,
no date, Marz., 7 males and 12 females, (Mus. Prague); "Braz.,"
no date, P. R. Uhler, 1 male, (USNM).

Paraguay: Horquaeta, 45 mi. E., Aug. 7, 1933, A. Schulze, 1 fe-
male, (J. C. Lutz Coll.); same place, July 10, 1934, A. Schulze, 1
female, (J. C. Lutz Coll.); Santa Barbara, Dec. 8, 1924, F. Schade,
1 female; Villarrica, Aug. 30, 1923, F. Schade, 1 male.

Argentina: "Argentina," no date, Weiser, 1 male, (Mus. Prague).

Comparative notes. This species resembles N. tcrrestris ( Kevan )
and IV. tenehrosa n. n. It can be separated from the former by the
small size of the ninth abdominal sternite of the male and by the
rounded emargination of the posterior margin of the last visible
abdominal sternite of the female. It can be separated from the
latter species by the dilation of the embolium and by the more sym-
metrical abdominal sternites of the female.

Nerthra terrestris (Kevan)

(PI. VII, fig. 48; PI. VIII, fig. 62; PI. X, fig. 87)

1948. Monomjx terrestris Kevan, Annals and Magazine Nat. Hist., 11th series,
vol. 14, no. 119, p. 813. (n. n. for M. hipunctatus Melin).

Referring to this species:

1929. Monomjx hipunctatus Melin, Zoologiska Bidrag Fran Uppsala, Band 12,

p. 177, figs. 39-42, 108.
1942. M. hipunctatus, Kevan, Proc. Royal Ent. Soc. London (a) 17, pts. 10-12,

pp. 109-110.

Size. Male: Length, 8.0 to 10.8 mm.; width of pronotimi 5.2 to

6.7 mm.; width of abdomen, 5.1 to 6.9 mm. Female: Length, 8.5
to 11.1 mm.; widtli of pronotum, 5.4 to 7.0 mm.; width of abdomen,

5.8 to 7.5 mm.

Color. A yellowish-brown to reddish-brown, median portion of
pronotum and scutellum darkest; a blackish spot at the medial end
of nodal furrow of the hemelytra; segments of the connexivum dark
brown at the anterior half, yellow or yellowish-brown at the pos-
terior half; base of femur and trochanter of fore leg light; femora
of intermediate and hind legs Hght with darker rings, the basal ring
of the femur of the hind leg often very faint; tibiae and tarsi of

374 The University Science Bulletin

intermediate and hind legs dark brown; abdominal segments dark
brown mottled with yellowish-brown; anal flaps of female yellowish-
brown.

Structural characteristics. Apex of the head more or less pointedly
projecting, terminating in a pair of sharp tubercles which are
sometimes blunt or even rudimentary; lateral tubercles small, sep-
arate and four or five in number. Pronotum laterally projecting at
a level with the transverse furrow; lateral margin often strongly
crenulated, anterior part usually not sinuated but rounded, poste-
rior part often sinuated, usually forming an obtuse angle with
base of embolium; posterior margin before scutellum slightly sinu-
ated. Scutellum large, sides sinuated at the base and near the
apex; without noticeable tumescences, but a weak depression on
either side of median line. Hemelytra extending beyond tip of
abdomen in the male, about the same length as the abdomen in
the female; embolium with lateral margin sometimes sinuated at
the base, usually straight to slightly rounded; nodal suture longer
in proportion to the length of the embolium in the female than in
the male; connexivum barely visible in the male, prominent in the
female. Ninth abdominal sternite of the male very large and
broad, nearly twice as long as the eighth abdominal sternite, about
two-fifths as wide as fifth abdominal sternite. Last visible ab-
dominal sternite of the female with posterior margin, medially,
pointedly notched, nearly symmetrical. Clasper of male sickle-
shaped, simple, tapering to a rather blunt point; aedeagal furrow
wide, visible on the apical half of the ventral surface.

Location of type. In the Museum of Stockholm, Stockholm,
Sweden.

Distributional data. Melin examined specimens from Peru,
Bolivia, Brazil, French Guiana, Trinidad, and Mt. Roraima, (Vene-
zuela ?). The writer has seen specimens from the following locali-
ties:

Peru: Roque near Moyobamba, Aug. 31, 1923-1925, D. Melin,
1 male and 1 female (types) (Mus. Stockholm).
Bolivia: "Bolivien," no date or collector, 1 male.

Colombia: Meta Villavicencio, May 11, 1946, E. A. Chapin, 1
male, (USNM).

Brazil: Corumba, April, no collector, 2 females, (Car. Mus.);
Manaos, Aug. 1924, Begnaert, 1 female; Sao Paulo, no date, A. A.
BarbieDini, 1 female, (USNM); "Paramaribo," Oct. 6, 1938, D. G.
Geiskes, 1 male; "Brazil," no date or collector, 1 female.

The Family Gelastocoridae (Hemiptera) 375

British Guiana: Near New Amsterdam, July 30, 1923, F. X.
Williams, 1 male; Demerara, June 25, 1901, R. J. Crew, 1 male,

(CAS).

French Guiana: St. Jean, no date, W, Schaus, 1 male, (USNM).

Trinidad: St. Augustine, Sept. 20, 1942, D. K. McE. Kevan, 2
males and 2 females.

Comparative notes. This species is very similar to the following
species, N. borealis (Melin), from which it may be separated by
the more laterally projecting pronotum. The writer has examined
the types of both species and it is his opinion that the two forms
probably represent the same species. The claspers of the males are
very similar, that of N. borealis ( Melin ) being sHghtly more slender
apically. However, until such time as we are in possession of the
details of the Hfe histories, ecology, etc., it seems best to treat
N. terrestris (Kevan) and N. borealis (Melin) as distinct species.

The name Mononyx bipunctatus Melin is a homonym of Mononijx
bipunctatus Stal, a synonym of Nerthra nepaeformis (Fabricius),
accordingly, Kevan, 1948, renamed the species Mononyx terrestris
n. n.

Nerthra borealis ( Melin )

(PI. Vm, fig. 59)

1929. Mononyx borealis Melin, Zoologiska Bidrag Fran Uppsala, Band 12,
p. 179, figs. 47-49.

Size. Male: Length, 7.2 mm.; width of pronotum, 4.4 mm.; width
of abdomen, 4.7 mm. These are the measurements of the type, they
vary slightly from the measurements given for the type by Melin.

Color. Dark yellowish-brown, pronotum and scutellum darker
than hemelytra; a dark brown spot on the embolial suture at the
junction with the nodal suture; posterior margin of the pronotum
with thin, longitudinal, black marks; posterior margin of front
femur dark brown, the rest yellowish-brown; intermediate and
hind legs yellowish with darker rings on the femora, two rings
on the femur of the intermediate leg, one on the femur of the
hind leg; last three abdominal segments yellowish-brown, the rest
of abdomen dark brown; segments of the connexivum mostly dark
brown with indistinct light spots near the posterior margins of the
segments.

Structural characteristics. Apex of the head more or less pointedly
projecting, terminating in two small apical tubercles; lateral tuber-
cles of the head irregularly spaced between apical tubercles and
eyes. Pronotum not very dilated laterally, less than the abdomen

376 The University Science Bulletin

in width; lateral margin nearly straight, slightly sinuated at the
middle; posterior margin before scutellum moderately sinuated.
Scutellum large, without tumescences and a very faint depression
on either side of the median line. Hemelytra extending to end of
abdomen; embolium slightly dilated, lateral margin weakly rounded.
Abdominal sternites of male like those of N. terrestris (Kevan);
last abdominal sternite large, longer than eighth abdominal sternite
and about two-fifths as wide as the fifth abdominal sternite.

Location of type. In the Museum of Stockholm. Stockholm,
Sweden.

Distributional data. This unique specimen is from Brazil, no date,
Boucord (Mus. Stockholm).

Comparative notes. Very near N. terrestris (Kevan), but differ-
ing by having the lateral dilation of the pronotum weak, the lateral
margin of the pronotum nearly straight.

Nerthra tenebrosa n. n.

(PI. VII, fig. 50; PI. VIII, fig. 61; PI. X, fig. 92)

1929. Mononyx obscurus Melin, Zoologiska Bidrag Fran Uppsala, Band 12,
p. 178, figs. 43-46.

Size. Male: Length, 8.3 to 8.5 mm.; width of pronotum, 5.5 to
5.6 mm.; width of abdomen, 5.3 to 5.8 mm. Female: Length, 8.7
to 9.7 mm.; width of pronotum, 5.8 to 6.2 mm.; width of abdomen,
5.9 to 6.5 mm.

Color. Variable as in the other species from a yellowish-brown
to reddish-brown; a dark brown spot at the distal end of the em-
bolial suture; a small, medial, black spot on the posterior margin
of the head; posterior margin of the pronotum with small irregular
spots of brown; segments of the connexivum dark brown or with the
posterior third yellowish; trochanter and base of femur of fore
leg, femora of intermediate and hind legs yellowish; femora of
intermediate and hind legs ringed with brown; abdominal seg-
ments either entirely dark brown or spotted with yellowish-brown.

Structural characteristics. Apex of head more or less pointedly
projecting, terminating in a pair of apical tubercles; lateral tuber-
cles of the head small, separate, four or five in number. Pronotum
widest at a level with the transverse furrow; lateral margin project-
ing beyond base of embolium; anterior three fourths of lateral
margin nearly straight, usually slightly convex; posterior fourth
straight, extending obliquely to base of embolium; posterior margin
sinuated before scutellum. Scutellum large, without lateral tumes-

The Family Gelastocoridae (Hemiptera) 377

cences; a faint depression on either side of median line of the
body. Hemelytra extending to or beyond the end of the abdomen
in the males and some females, but shorter than abdomen in some
female specimens; embolium sinnated at basal third. Abdominal
sternites of male asymmetrical; ninth sternite small, slightly wider
than long, longer than eighth sternite. Abdominal sternites of female
asymmetrical; posterior margin of last visible abdominal sternite
deeply and roundly notched medially; last visible abdominal ster-
nite very long, nearly as long as wide; area about notch pinched
up as a result of the narrowing; a prominent depression on either
side of the notch, laterad of which is a moderate tumescence, the
one on the left slightly the larger. Clasper of the male sickle-shaped,
simple, rather abruptly curved and pointed at the apex; aedeagal
furrow visible on the apical half of the ventral surface.

Location of type. In the Museum of Stockholm, Stockholm,
Sweden.

Distributional data. Melin examined specimens from Mexico,
Colombia and Bolivia. The writer has seen specimens from the fol-
lowing localities:

Colombia: Bogota, not date, Lindig., typus male and allotypus
female, (Mus. Stockholm); Cali, no date, W. F. H. Rosenberg, 2
females, (USNM); Gallegos, 1937, no collector, 1 female, (USNM);
Medellin, 1938, no collector, 1 female, (USNM); Minca, May, 1919,
no collector, 1 female, (Car. Mus.); Muzo, no date, H. Apolimar-
Maria, 1 male, (CAS); Rio Dagua, no date, W. F. H. Rosenberg,
1 male, (USNM); Rio Frio, Feb., 1924, W. M. Mann, 1 male and
4 females, (USNM).

Ecuador: Santa Inez, no date, R. Haensch, 1 male, (CAS);
Tena, Feb. 23, 1923, F. X. Williams, 1 male and 2 females.

Panama: Colon, no date, P. R. Uhler, 1 male, (USNM); same
place, no date, C. F. Baker, 1 male, (USNM).

British Honduras: Punta Gorda, Feb., 1931, J. J. White, 1 male,
(J. C. Lutz Coll.).

Comparative notes. This species may be separated from N.
nepaeformis (Fabricius) by the embolium being sinuous at the
basal third, by the clasper of the male, and by the asymmetry of the
last visible abdominal sternite of the female.

The name Mononyx obscurus Melin is a homonym of Mononyx
obscurus Stal and must be renamed. I propose the new name
Nerthra tenebrosa for this species.

378 The University Science Bulletin

Nerthra unicornis (Melin)

(PI. VIII, figs. 57, 57a; PI. IX, fig. 77)

1929. Mononyx unicornis Melin, Zoologiska Bidrag Fran Uppsala, Band 12,
pp. 179-180, figs. 50-53.

Size. Male: Length, 7.1 to 7.2 mm.; width of pronotum, 4.5 to
4.6 mm.; width of abdomen, 4.5 to 4.6 mm.

Color. Brown to reddish-brown; a dark brown spot at the distal
end of the embolial suture; posterior margins of the connexivum,
intermediate legs, hind legs and abdominal segments with yellowish-
brown areas.

Structural characteristics. Apex of head pointedly projecting,
terminating in a pair of sharp apical tubercles, fused medially, the
degree of fusion variable in the different specimens; lateral tubercles
small, irregular, forming an indistinct toothlike protuberance before
the eye. Pronotum very slightly projecting laterally beyond the
base of embolium, widest at a level with the transverse furrow;
lateral margins nearly straight for the anterior three-fourths, slightly
sinuated before scutellum. Scutellum moderately large, a weak tu-
mescence on either side and a slight median longitudinal carina-
tion present. Hemelytra extending to or beyond end of abdomen;
embolium elongate, lateral margins nearly straight or slightly con-
vex. Connexivum almost covered by the hemelytra. Body covered
with slightly clavate bristles, a group of black bristles on the tumes-
cences of the scutellum either in oval patches or short irregular
longitudinal bands. Abdominal sternites of male asymmetrical;
ninth sternite small, almost identical with the abdominal sternites
of N. nepaeformis (Fabricius). Clasper of male also resembling
that of N. nepaeformis (Fabricius).

Distributional data. Melin studied specimens from French Guiana
and Brazil. The writer has examined the type, a male from Sao
Leopoldo, Brazil, no date or collector, (Mus. Stockholm) and an-
other male from Para, Brazil, no date or collector, (Car. Mus.).

Comparative notes. N. unicornis Melin seems to the writer to be
very near N. nepaeformis (Fabricius). It differs from the latter
species by the fusion of the apical tubercles, by the less dilated pro-
notum and by a slight difference in the shape of the male clasper.
The writer unfortunately has not been able to examine a female of
this species.

The Family Gelastocoridae (Hemiptera) 379

Nerthra peruviana (Montandon)

(PI. VII, fig. 47; PI. VIII, fig. 65; PI. X, fig. 83)

1905. Mononyx peruvianus Montandon, Annales Musei Nationalis Hungarici,
vol. Ill, p. 403.

1909. M. peruvianus Kirkaldy and Torre-Bueno, Proc. Ent. Soc. Washington,
vol. X, p. 182.

1929. M. peruvianus Melin, Zoologiska Bidrag Fran Uppsala, Band 12, p.
180, figs. 54-57.

1931. M. peruvianus De Carlo, Revista de la Sociedad Entomologica Ar-
gentina, III, no. 17, p. 329.

Size. Male: Length, 6.9 to 7.3 mm.; width of pronotum, 5.0 to
5.3 mm.; width of abdomen, 5.0 to 5.3 mm. Female: Length, 7.4
to 8.3 mm.; width of pronotum, 5.4 to 6.0 mm.; width of abdomen,
5.6 to 6.8 mm.

Color. Uniformly yellowish-brown to reddish-brown above with
a dark spot at junction of embolial and nodal sutures. Ventrally
slighty darker. Front femur pale yellow to light brown, lighter than
the trochanter. Intermediate and hind legs mostly dark brown,
with a lighter area at the base of the femora, occasionally the legs
are light with dark rings.

Structural characteristics. Apex of the head more or less project-
ing with a fairly large pair of sharp-pointed tubercles at apex; lateral
tubercles smaller, separate, four or five in number. Pronotum with
lateral margins rounded, occasionally with anterior third and
posterior fourth straight to slightly concave. Pronotum of male as
wide as abdomen, not as wide as abdomen in females. Scutellum
with two longitudinal bands of long narrow black bristles. Bristles
of the body numerous, small, light brown, not swollen apically.
Hemelytra well developed, extending to or beyond end of ab-
domen. Abdomen of female prominently expanded laterally. Ab-
dominal sternites of male asymmetrical; ninth sternite rounded in
shape, longer than seventh or eight sternite. Abdominal sternites
of female nearly symmetrical. Posterior margin of last visible ad-
dominal sternite medially emarginated with a slightly lateral de-
pression on either side, laterad of which is a moderate tumescence,
the one on the left slightly larger than the one on the right.

Location of type. In the Budapest Museum, Budapest, Hungary.

Distributional data. This species has been reported from Peru
and Argentina. Specimens from the following localities have been
examined by this worker:

Peru: Callango, no date or collector, 9 males and 11 females,
one compared with the type by Doctor H. B. Hungerford, in 1928;
Chanchamayo, no date, W. F. H. Rosenberg, 1 female, (USNM):

380 The University Science Bulletin

Rioja, Dcpt. San Martin, Sept. 9 to Oct. 3, 1936, F. Woytkowski,
1 male and 1 female; Rio Perene and Chanchamayo, Dept. Jimin,
May, 1934, F. Woytkowski, 1 male and 1 female; Sani Beni, Dept.
Junin, Nov. 9, 1935, F. Woytkowski, 1 female; San Pedro, May 29,
1935, F. Woytkowski, 4 males and 1 female; "Peru," no date or
collector, 1 female, (Mus. Prague).

Ecuador: Tena, Apr. 13, 1923, F. X. Williams, 1 male and 1
female.

Bolivia: Huachi, Rio Beni, Sept. 1921, W. M. Mann, 1 male,
(USNM).

Comparative notes. This species is easily recognized by the
longitudinal bands of long, narrow black bristles on the scutellum,
the abdominal sternites of the female and the clasper of the male.
Of the specimens examined only one specimen lacked the bristles
on the scutellum, apparently having been rubbed off in pinning the
specimen.

Nerthra montando7n (Melin)

(PI. Vlll, figs. 55, 55a; PI. X, fig. 85)

1929. Mononijx montandoni Melin, Zoologiska Bidrag Fran Uppsala, Band

12, p. 195, fig. 95.
1942. M. montandoni Kevan, Proc. Royal Ent. Soc. London (a), 17, pts. 10-

12, p. 110.

Size. Male: Length, 5.5 to 5.8 mm.; width of pronotum, 3.7 to
4.1 mm.; width of abdomen, 3.7 to 4.1 mm. Female: Length, 5.5
to 6.1 mm.; width of pronotum, 3.8 to 4.1 mm.; width of abdomen,
3.9 to 4.3 mm. .

Color. Yellowish-brown, spotted with small brown markings;
a dark brown spot at the distal end of the embolial suture; a black-
ish spot at median part of posterior margin of head; segments of
connexivum dark brown anteriorly, pale yellowish posteriorly; ven-
tral surface more or less light yellow; front femora, rings on inter-
mediate and hind femora and spots on the abdomen brown.

Structural characteristics. Apex of the head concavely excavated
with a blunt tubercle on either side of the excavation; lateral tuber-
cles of the head small, blunt and irregularly spaced. Pronotum
about as wide as abdomen, less than width of abdomen in some
specimens; lateral margins more or less rounded very slightly pro-
jecting beyond the base of the embolium; posterior margin prom-
inently sinuated before scutellum. Scutellum fairly large, some-
what elevated; a moderate tumescence on either side and a weak
median longitudinal carination present. Hemelytra extending to or

The Family Gelastocoridae (Hemiptera) 381

beyond end of abdomen; embolium elongate, narrow with lateral
margin slightly rounded. Rather large, clavate bristles scattered
over body, more or less clumped; a narrow longitudinal band of
black clavate bristles on tumescences of the scutellum. Abdomen
posteriorly broadly dilated laterally, rather truncate. Abdominal
sternites of male asymmetrical; ninth sternite oval, slightly wider
than long, about the same length as eighth sternite or slightly
longer. Abdominal sternites of female slightly asymmetrical; pos-
terior margin of last visible abdominal sternite emarginated me-
dially, notch more or less triangular in shape; a slight depression on
either side of notch laterad of which is a moderate tumescence, the
one on the left the larger. Clasper of male simple, apical part
more or less wedge-shaped, directed mesad.

Location of ttjpe. Holotype, female, in the Museum of Stock-
holm, Sweden. Allotype, male, Icta, Trinidad, XI- 19- 1941, N.
Hynes, in the Francis Huntington Snow Entomological Museum at
the University of Kansas.

Distributional data. Melin examined two females of this species
from Venezuela. The writer has before him three specimens from
Trinidad. One male and one female from Icta, Nov. 19, 1941, N.
Hynes; the other a male, merely labeled "Trinidad, W. I.," Dec. 14,
1902, Chipman, (CAS).

Comparative notes. Melin said this species was near N. ranina
( Herrich-Schiiffer ) , but the abdominal sternites of the female are
entirely different. The species may be separated from N. peru-
viana ( Montandon ) by the concavely excavated apex of the head.
The male can be separated from both species by the shape of the
clasper.

Melin had only the females of this species before him when he
described this species, therefore, I designate the male specimen
from Icta, Trinidad, Nov. 19, 1941, N. Hynes, as the allotype of
this species.

Nerthra americana (Montandon)

(PL XII, fig. 115; PL XIII, fig. 120)

1905. Matinus americanus Montandon, Annales Musei Nationalis Hungarici,
vol. Ill, pp. 404-405.

1906. M. americanus Torre-Bueno, Proc. Ent. Soc. Washington, vol. VIII, nos.
1 and 2, p. 51.

1931. M. americanus Carlo, J. A. de and Gemignani, E. V., Revista de la
Sociedad Entomologica Argentina, vol. 3, no. 6, pp. 329-330.

Size. Male: Length, 6.3 to 6.6 mm., width of pronotum, 4.3 to 4.4
mm., width of abdomen, 4.4 mm. Female: Length, 6.6 to 7.1 mm.,
width of pronotum, 4.5 to 4.9 mm., width of abdomen, 4.6 to 5.1 mm.

382 The University Science Bulletin

Color. Above, yellowish-brown to brown, scutellum and disc of
pronotum usually darker; basal two thirds of each segment of the
connexivum dark brown; hemelytra and lateral expanses of pronotum
are sometimes spotted with dark brown. Below, legs light with dark
brown rings, front femora irregularly spotted with brown; abdominal
segments of female dark, but may be spotted with yellowish-brown;
ovipositor lobes of female and last two abdominal segments of the
male, lighter than rest of abdominal segments.

Structural characteristics. Apex of head pointedly projecting,
anterior margin with numerous small, sharp-pointed tubercles;
ocelli absent. Lateral margins of pronotum subparallel for the pos-
terior two thirds, straight or very faintly concave and slightly con-
verging anteriorly; anterolateral margin converging abruptly toward
the eye, rather concave; posterolateral margin doubly concave;
posterior margin nearly straight, with three broad, very shallow,
concavities; disc elevated, lateral expanses nearly flat, slightly ele-
vated medially next to disc; pronotum and abdomen subequal in
width in males, abdomen slightly wider than pronotum in female.
Scutellum not at all elevated, no higher than hemelytra and de-
pressed slightly at base, the apex being the most elevated portion,
Hemelytra usually extending as far as or beyond end of abdomen,
entirely coriaceous and fused together; embolium with basal portion
expanded laterally, and bent up for about one half its length. Con-
nexivum visible, more expanded laterally in females. Bristles of
dorsal part of body, moderately long and clavate, those of the heme-
lytra darker than rest. Abdominal sternites of female nearly sym-
metrical; emargination of posterior margin of last visible sternite
nearly triangular, slightly rounded apically; ovipositor lobes triangu-
lar, slightly longer than wide; caudolateral angles of last visible
sternite weakly projecting posteriorly. Ninth abdominal sternite of
male oval, with oblique furrow from left to right, wider than long,
33:19, longer than seventh or eighth sternites; right side of seventh
sternite completely covered by the sixth sternite. Clasper of male
distinctive, sickle-shaped, but with a large tliumblike projection aris-
ing on the dorsolateral surface.

Location of type. Unknown. It may be in the National Museum
of Hungary at Budapest, some of the specimens before the author
at the time of the description were from this museum. The rest of
the specimens were from Montandon's own collection, but Dr. H.
B. Hungerford did not find this type in the British or Paris Museums.

Distributional data. Montandon's specimens were from Espirito

The Family Gelastocoridae (Hemiptera) 383

Santo, Brazil. Specimens studied by this worker are from Sao
Paulo, Brazil, no date, Mraz. Lgt., 2 males and 13 females. The
specimens belong to the Museum of Prague, Prague, Czechoslovakia.
A male and female from this series has been retained and are now
in the Francis Huntington Snow Entomological Collection at the
University of Kansas.

Comparative notes. This species may be separated from IV. ivil-
Uainsi n. sp. by the right side of the seventh abdominal sternite of
the male being covered by the sixth segment, by the smaller size, by
the presence of clavate bristles and by the shape of the clasper of
the male. It can be separated from the other American species by
the absence of ocelli, and by the entirely coriaceous hemelytra
which are fused together.

Nerthra williamsi n. sp.

(PI. XII, fig. 112)

Size. Male: Length, 8.0 to 8.6 mm., width of pronotum, 5.8 to
6.1 mm., width of abdomen, 5.7 to 6.0 mm. Female: Length, 9.1 to
9.6 mm., width of pronotum, 6.4 to 6.8 mm., width of abdomen, 7.0
to 7.1 mm.

Color. Above from light yellowish-brown to brown, base of
scutellum, disc of pronotum, front of head, and antero-lateral angles
of segments of connexivum, darker. Below, head dark, front legs
mostly dark brown or black, irregularly marked with yellow and
yellowish-brown, femora of middle and hind legs yellowish-brown,
ringed with two bands of dark brown, abdominal segments brown
to black, caudolateral angles of each segment lighter in color.

Structural characteristics. Front of head with sharp-pointed tuber-
cles, apical ones ventrad and slightly caudad of others, the latter, of
which there are about five on each side, are irregular as to size and
position, usually along the anterolateral margin of the head. Apical
tubercles variable as to number and size, usually one large median
apical tubercle; ocelli absent. Width of pronotum about equal to
width of abdomen in males, less than width of abdomen in females;
lateral margin of pronotum nearly straight, barely concave, for
the posterior three fourths of the margin of the pronotum, abruptly
but roundly converging to the eye anteriorly, posterolateral margin
rounded; posterior margin of pronotum relatively straight, very
weakly concave before scutellum, slight convexities before the
basal angles of the scutellum and before base of embolial suture
of hemelytron; disc elevated, lateral expanses of pronotum nearly

384 The University Science Bulletin

flat. Scutellum, not especially elevated, except at apex; base sunken,
inclined posteriorly to apex. Hemelytra entirely coriaceous, fused
together. Connexivum expanded laterally in both sexes, wider
than embolium, but more so in females. Body covered with slender,
brown or black bristles which are slightly curved near apex, pointed,
not at all clavate. Abdominal sternites of female with last sternite
fairly deeply emarginated, apex of emargination more or less
rounded; sternites nearly symmetrical; lobes of ovipositor rather
large, triangular, slightly longer than wide and each with a rather
deep depression. Ninth abdominal sternite of male small, oval,
wider than long, 20:11, about equal to or a little longer than
eighth sternite, longer than seventh sternite. Spatulate right side
of seventh abdominal sternite, elongate and narrowed. Clasper of
male sickle-shaped, quite similar to the clasper of N. peruviana
( Montandon ) and N. nepaeformis (Fabricius).

Location of type. Holotype male, allotype female and 3 male
and 3 female paratypes, Sao Paulo, Brazil, no date, Mraz Lgt., in
the Museum of Prague, Prague, Czechoslovakia. One male and
one female paratype, same data, in the Francis Huntington Snow
Entomological Collections at the University of Kansas.

Distributional data. As given for type series.

Comparative notes. This form is closely related to N. americana
(Montandon), if one considers the hemelytra, it may be separated
from this species by the clasper of the male being simple, without
a large process on the left side, bristles not clavate, right side of
seventh abdominal sternite of male not covered completely by
sixth sternite and by the presence of non-clavate bristles. It is the
opinion of the writer, that this species is actually closely related to
N. peruviana (Montandon) as is indicated by the similarities of
head, pronotum, abdominal segments, male claspers, non-clavate
bristles, etc. It may be distinguished from the latter species by
the fused, entirely coriaceous hemelytra and absence of ocelli.

This species is named in honor of F. X. Williams, who has col-
lected considerable material of this family from various regions
of the world.

Nerthra amplicollis ( Stal )

(PI. VII, fig. 53; PL VIII, fig. 68; PI. X, fig. 95)

1854. Mononyx amplicollis Stal, Ofversigt af Kongl. Vetenskaps-Akademiens

Fordhandlingar, Band XI, no. 3, p. 239.
1859. M. amplicollis Dohrn, Catalogus Hemipterorum, Herausgegeben von

dem Entomologischen Verein zu Stettin, p. 53.
1863. M. amplicollis Stal, Berliner Entomologische Zeitschrift, VII, p. 406.

The Family Gelastocoridae (Hemiptera) 385

1873. M. amplicoUis Walker, Catalogue of Hemiptera in British Museum,

part VIII, p. 172.
1876. M. ampicnllis Stal, Kongl. Svenska Vetenskaps-Akademiens Hand-

lingar. Band 14, no. 4, p. 138.
1899. M. amplicoUis Montandon, Bulletin de la Societe des Sciences de

Bucarest-Roumaine, an. VIII, nos. 4 and 5, p. 400.
1901. M. amplicoUis Champion, Biologia Centrali-Americana, Rhynchota

Heteroptera, vol. II, p. 350, pi. XX, fig. 26.
1909. M. ampUcoUis Kirkaldy and Torre-Bueno, Proc. Ent. Soc. Washington

vol. X, p. 180.
1929. M. amplicoUis Melin, Zoologiska Bidrag Fran Uppsala, Band 12, pp.

184-185, figs. 65-68, 104, 109.

Size. Male: Length, 10.8 to 12.7 mm.; width of pronotiim, 6.8
to 7.7 mm.; width of abdomen, 6.7 to 7.5 mm. Female: Length, 10.5
to 14.7 mm.; width of pronotum, 6.3 to 9.3 mm.; width of abdomen,
6.7 to 9.1 mm.

Color. Brown to black, frequently reddish brown; connexivum
entirely dark or with thin pale areas at the posterior margins of
the segments; apex of head, front trochanter, anterior and basal
part of front femur and a spot near apex of hind femur yellow
or orange; posterior margin of front femur, front tibia-tarsus, in-
termediate and hind legs and abdomen dark brown.

Structural characteristics. Apex of the head broadly concavely
excavated; a short broad tubercle on each side of the excavation;
lateral tubercles indistinct, forming a slight toothlike protuberance.
Pronotum as wide at the anterior third as at the level of the trans-
verse furrow; projecting beyond the base of the embolium, but not
as wide as the abdomen; lateral margin with the median third,
straight and parallel with the longitudinal axis of the body, anterior
third straight obliquely converging to the eye, posterior third
straight converging obliquely to the base of the embolium; posterior
margin sinuated before the scutellum. Scutellum large, elevated
except at the basal angles. Hemelytra extending beyond the end
of the abdomen; embolium slightly dilated; lateral margin concave
at the base, slightly convex the remainder of its length. Connexivum
almost completely covered by the hemelytra in the male, moderately
prominent in the female. Abdominal sternites of male asymmetri-
cal; ninth sternite twice as wide as long, longer than eighth sternite
and about the same length as the seventh sternite; seventh sternite
not visible on the lateral two thirds of the right side. Female
abdominal sternites nearly symmetrical; posterior margin of last
visible abdominal sternite deeply and pointedly notched. Lobes
of ovipositor elongate and rather large. Clasper of the male more
or less bifurcate.

13—6730

386 The University Science Bulletin

Location of type. In the Museum of Stockholm at Stockholm,
Sweden.

Distribiitio7ial data. Kirkaldy and Torre-Bueno list this species
from Costa Bica, Colombia, Venezuela, Ecuador and Bolivia. This
worker has examined specimens from the following localities:

Costa Rica: San Jose, no date, M. Valerio, 1 male, (USNM).

Venezuela: Merida, no date, W. Robinson, 5 males and 3 females,
(USNM); same place, no date or collector, 1 female, (AMNH);
"Venez.," no date or collector, 1 male and 7 females, (USNM).

Bolivia: "Bolivia," no date or collector, 1 male and 1 female.

Colombia: Cincinnati, Feb. 1924, W. M. Mann, 1 male, (USNM);
Fusagasuga, June, 1917, Apolinar Maria, 1 female; Villavicencio,
Oct., 1916, A. Maria, 1 male, (CAS); "Colombia," no date, F. C.
Nicholas, 1 female, (AMNH); same place, no date, C. F. Baker,
4 females, ( USNM ) ; same place, no date or collector, 1 female.

Ecuador: Balzapamba, no date, R. Haensch, 1 male; Rio Mapoto,
Mar. 24, 1939, F. M. Brown, 1 female, (AMNH); Santa Inez, no date,
R. Haensch, 1 male and 1 female; same place, no date, R. Haensch,
3 males and 2 females, ( CAS ) .

Peru: Lima, no date, Soukup, 1 male, (USNM); same place, no
date or collector, 1 female; "Peru," no date, P. R. Uhler, 1 male,
(USNM); "Peru," no date or collector, 1 female, (AMNH).

Comparative notes. This is the largest American species, how-
ever, it varies considerably in size and thus size alone is not always
sufficient to seperate this species from the others. The deep tri-
angular emargination and the large elongate ovipositor lobes which
do not overlap anteriorly will separate the females from all other
species except N. ecuadorensis (Melin) which is smaller, which
has the abdomen more broadly expanded laterally than most females
of N. amplicollis (Stal) and which has the emargination of the
last abdominal sternite more rounded than in N. ampIicoUis ( Stal ) .
The males of this species have the clasper more or less bifurcate,
the two parts of this bifurcation are of approximately the same size
and both are sharp-pointed. In this respect it differs from N. ecua-
dorensis (Melin) and N. ater (Melin), both of which havf the
dorsal process greatly reduced and not at all sharp-poin*"ed.

The Family Gelastocoridae (Hemiptera) 387

Nerthra ecuadorensis ( Melin )

(PI. VIII, fig. 58; PI. IX. fig. 73; PI. X. fig. 86)

1929. Mononyx amplicoUis var. ecuadorensis Melin, Zoologiska Bidrag Fran
Uppsala, Band 12, p. 185, figs. 69, 70.

Size. Male: Length, 8.1 to 10.2 mm.; width of pronotum, 5.0
to 6.2 mm.; width of abdomen, 5.2 to 6.2 mm. Female: Length, 8.8
to 10.2 mm.; width of pronotum, 5.6 to 7.0 mm.; width of abdomen,
5.9 to 6.8 mm.

Color. As in Nerthra amplicoUis (Stal).

Structural characteristics. This species is very similar to the
preceding species and differs from that species only in a few
respects. It is smaller than IV. amplicoUis (Stal); in the case of the
females, the abdomen usually is broader in proportion to the length
of the body than in the latter species, however, this is not always
true and in the opinion of the writer, it is not a very good character;
the emargination of the posterior margin of the last visible ab-
dominal sternite of the female is more rounded than in N. amplicoUis
(Stal); the males are smaller than in the latter species and the
clasper differs in that the dorsal process is reduced, it varies slightly
in that the process may be more or less cylindrical while in other
specimens it is flattened in a radial plane to the longitudinal axis
of the clasper.

Location of type. In the Museum of Helsingfors, (Helsinki),
Finland.

Distributional data. Melin described this species as a subspecies
of IV. amplicoUis (Stal), from two female specimens from Ecuador.
This worker has studied specimens from the following localities:

Ecuador: Macas, no date, A. Heyne, 1 male; same place, no date
or collector, 2 females; "Normandia," no date or collector, 1 female.

Colombia: "Bet. Queremal and Buenaventura," Feb. 3, 1935, H. F.
Schwarz, 1 male, (AMNH).

Peru: Aquaitia, Dept. Loreto, Sept., 1946, F. Woytkowski, 1
male; Vic. Guayabamba, Andes, Dept. Amazonas, Aug. 14-19, 1936,
F. Woytkowski, 1 female; Rio Perene and Chanchamayo, Dept.
Junin, May, 1934, F. Woytkowski, 1 male; Vic. San Pedro, May
29, 1935, F. Woytkowski, 2 females; Tarma, Dept. Junin, March
1-15, 1948, F. Woytkowski, 5 males and 2 females; Tinccochaca,
Aug. 8, 1911, Yale Peruv. Exp., 1 male and 1 female, (USNM).

Panama: Boquete, Chiriqui Prov., March 6, 1923, F. M. Gaige,
1 male.

388 The University Science Bulletin

Comparative notes. This species has been raised from subspecific
rank to that of a full species on the fact that the males are consistent
in the reduction of the dorsal process of the clasper and in being
of a smaller size, and because at the present time there is no evi-
dence that the two forms do not occur in the same localities. It
should be pointed out, however, that there is no definite proof that
the males before this worker actually are the same species as the
females. In the series of specimens a pair was taken at Tinccochaca,
Peru, on the same day and 5 males and 2 females were taken at
Tarma, Peru, in a fifteen day period. While such evidence is not
positive, it is sufficient to consider the two sexes as one species,
in view of their morphological similarities.

Nerthra planifrons (Melin)

1929. Mononyx planifrons Melin, Zoologiska Bidrag Fran Uppsala, Band 12,
p. 186, figs. 74-76.

The writer has not seen Nerthra planifrons (Melin), or at least
has been unable to identify tlie species. The original description
of the species follows.

"Front of caput fairly even, without medial ridge or knob; medial
processes knoblike, well separated; some small, apical knobs ap-
pearing between them; lateral ones forming an irregular, toothlike
knob before depression, which is rather slightly marked. Structure
of pronotum somewhat but not strongly marked; side anteriorly,
strongly projecting beyond base of hemelytrae; edge rounded and
crenulated, somewhat sinuated in the middle; posterior part not bent
angularly; anterior corner forming an obtuse angle; posterior margin
before scutellum less sinuated than in ampUcolUs. Scutellum with
distinct traces of protuberances. Embolium dilated. Abdomen of
female dilated terminally, apically somewhat truncated; 6th seg-
ment of female jammed, at posterior margin deeply notched to an
acute, somewhat rounded angle; lamellae broadly wedge-shaped.
Bristles fine and fairly short.

Coloration blackish-brown; segments of connexivum posteriorly
with indistinct light spots; abdomen, ventrally, darkish-brown with
posterior corner of segments light; legs dark; femora of 2nd and
3rd pair lighter with traces of dark rings.

Length 10, breadth 6 mm.

Museum of Kobenhavn: 1 female, Parzudaki, Mexico, (type)."

The Family Gelastocoridae (Hemiptera) 389

Nerthra ater (Melin)

(PL VIII, fins. 56, 56a)

1929. Monomjx ater Melin, Zoologiska Bidrag Fran Uppsala, Band 12, pp. 185-
186, figs. 71-75.

Size. Male: Length, 9.1 to 9.2 mm.; width of pronotiim, 5.6 to
5.8 mm.; width of abdomen, 5.7 to 5.8 mm.

Color. Black above, posterior margin of the pronotum and the
vertex of the head light brown; much lighter below; legs mainly
light yellowish brown; abdomen slightly paler.

Structural characteristics. The writer has examined the type of
this species and has found it to be very similar to Nerthra ecuado-
rensis (Melin). It differs from the latter species in color, by the
more rounded anterior and posterior portions of the lateral margin
of the pronotum and by the shape of the clasper of the male. The
clasper of N. ater (Melin) has a thin concave plate-like process
which is flattened in a tangential plane to the longitudinal axis of
the clasper and it is located at the same point of origin as the dorsal
arm of the bifurcate clasper of N. ecuadorensis (Melin). The ab-
dominal sternites are identical with the abdominal sternites of the
latter species.

Location of type. In the Museum of Stockholm, Stockholm,
Sweden.

Distributional data. The type is from Bogota, Colombia. An-
other specimen, in the Museum of Paris, was questionably placed
in this species by Melin. It is from Costa Rica.

Comparative notes. It is the opinion of this writer that this may
possibly be only an aberrant specimen of N. ecuadorensis (Melin).
However, until the other specimen in the Museum of Paris can be
studied I prefer to retain this species. There is also a possibility
that this species could be the male of Nerthra rudis (Melin). If
such should prove to be the case, N. ater (Melin) would fall as a
synonym of the latter species.

Nerthra lata (Montandon)

1899. Monontjx latus Montandon, Bulletin de la Societe des Sciences de

Bucarest-Roumanie, An. VIII, nos. 4 and 5, p. 399.
1909. M. latus Kirkaldy and Torre-Bueno, Proc. Ent. Soc. Washington, Vol.

X, p. 181.
1929. M. latus Melin, Zoologiska Bidrag Fran Uppsala, Band 12, pp. 186-187,

figs. 77-79.

Size. Female: Length, 10.5 mm.; width of pronotum, 6.4 mm.;
width of abdomen, 6.5 mm.

390 The University Science Bulletin

Color. Brown, except for the femora of the legs which are pale
yellow.

Structural characteristics. Apex of head slightly projecting ven-
trally, concavely excavated above this projecting part; apex terminat-
ing in two small tubercles which are barely visible in a dorsal view.
Pronotum not as wide as abdomen; lateral margin more or less
rounded, anterior part slightly concave, margin faintly crenulated;
posterior margin sinuated before scutellum. Scutellum moderate,
with slight lateral tumescences and a weak median longitudinal
carination. Hemelytra extending beyond end of the abdomen; em-
bolium concave and narrow at base; a constriction at the nodal
suture. Connexivum slightly exposed. Abdominal sternites of the
female nearly symmetrical, rather broad; last visible abdominal
sternite with posterior margin medially emarginated, anterior part
of notch more or less rounded.

Location of type. In the British Museum, London, England.

Distributional data. The type is labeled "Nanegal-Equateur."
Melin examined two specimens from Ambato, Ecuador, and Antio-
chia, Colombia. The writer has seen only the latter specimen. It
has, in addition to the locality, the label marked "Schh.," but has
no date. It is located in the Museum of Stockholm.

Comparative notes. This specimen seems to fit Montandon's
description fairly well, although there are only two small apical
tubercles of the head instead of three, and in that the abdomen is
not so widely expanded that it can be readily separated from some
of the other American species, as Montandon states. Except for
the two very small apical tubercles and a slight difference in the
shape of the lateral margin of the pronotum, the specimen studied
by the writer looks like N. amplicollis (Stal) or more especially
like N. ecuadorensis (Melin). This name must be retained until
the type can be examined and compared with the other species.

Nerthra rudis (Melin)

(PI. X, fig. 90)

Referring to this species:

1929. Mononyx fuscipes var. rudis Melin, Zoologiska Bidrag Fran Uppsala,
Band 12, p. 182, fig. 62.

Size. Female: Length, 8.9 to 10.6 mm.; width of pronotum, 5.3
to 6.4 mm.; width of abdomen, 5.4 to 6.1 mm.

Color. Yellowish-brown to black; scutellum dark brown or black;
a small yellow spot at the posterior angle of each segment of the

The Family Gelastocoridae (Hemiptera) 391

connexivum; ventral surface light brown or brown with femora of
legs, median portion of abdomen and anal flaps yellow.

Structural characteristics. Apex of the head concavely excavated;
lateral tubercles forming a moderate toothlike protuberance be-
tween the eye and apex of the head. Pronotum widest at a level
with the transverse furrow, about as wide as abdomen; lateral
margin with the anterior three fourths nearly straight, slightly
con\'ex then weakly concave; posterior margin sinuated before
scutellum. Scutellum slightly elevated, a depression at each basal
angle. Hemelytra extending beyond tip of abdomen; lateral margin
of embolium slightly convex. Connexivum moderately exposed
dorsally. Abdominal sternite of the female nearly symmetrical;
posterior margin of the last visible abdominal sternite medially,
deeply and pointedly emarginated; a slight depression on either
side of the emargination. Ovipositor lobes large, elongate, the
anterior end of the left lobe twisted to the right and covering the
anterior end of the right lobe.

Location of type. In the Museum of Helsingfors, Helsingfors
(Helsinki), Finland.

Distrihiitional data. The type is from Vera Cruz, Mexico. An-
other specimen studied by Melin is from Guatemala. This worker
has seen specimens from the following localities:

Costa Rica: Rio Virilla, Dec. 26, 1931, H. Schmidt, 4 females.

Panama: Bohio, C. Z., Feb. 7, 1911, E. A. Schwarz, 1 female,
(USNM); Buenaventura, Mar. 10, 1911, A. Busck, 1 female,
(USNM); XX Plantation, 6 mi. E. Portobelo, Feb. 16, 1930, T. O.
Zschokke, 1 female, (CAS).

Ecuador: "Bucay," no date, Campos, 1 female.

Comparative notes. The writer has not seen the type of this
species, but the description and the drawings of this species made
by Melin could not possibly belong to N. fuscipes ( Guerin-Mene-
ville ) . The front of the head, the general shape and especially the
shape and size of the ovipositor lobes connect this species to the
amplicollis group. I take this opportunity to remove this name as
a subspecies of N. fuscipes (Guerin-Meneville) and elevate it to
specific rank. Further study may prove that this is only a subspecies
of N. amplicollis Stal, but the ovipositor lobes seem to me to be
sufficiently different to constitute a good species.

392 The University Science Bulletin

Nerthra corudis n. sp.

(PI. XV, figs, 144, 145)

Size. Male: Length, 10.0 to 10.4 mm.; width of pronotum, 6.0 to
6.5 mm.; width of abdomen, 6.0 to 6.5 mm.

Color. General coloration above yellowish-brown; head, pro-
notum and scutellum heavily spotted with darker brown; hemelytra
with finer maculations and therefore lighter; segments of the con-
nexivum, yellowish-brown on the posterior two thirds and with
apical one third nearly black. Below, nearly uniformly dark brown,
nearly black, except caudolateral angles of the abdominal ster-
nites; the eighth sternite has the lighter spot only on the right side.

Structural characteristics. Head with two broad, blunt, super-
apical tubercles and a broad rather indistinct lateral tubercle on
either side. Ocelli present. Pronotum equal to width of abdomen;
lateral margin with apical one third roundly converging toward
the eyes, the middle one third with the two sides subparallel, slightly
convergent anteriorly; pronotum widest at a level with the trans-
verse furrow. Hemelytra with well-developed membranes, extend-
ing to end of abdomen; embolium not expanded basally, lateral
margin straight or very slightly concave near base, convex at
middle. Scutellum broad, not strongly elevated. Bristles very short,
slightly clavate, curved, rather sparse. Ninth abdominal sternite
oval, twice as wide as long, shorter than eighth sternite; abdomen
very distinctly asymmetrical, seventh sternite almost entirely to
left side; posterior margins of sternites five and six broadly emargi-
nate to the left side of the abdomen. Clasper of male very character-
istic (Seefig. 145, PI. XV).

Location of type. Holotype male, Panzos, Guatemala, no date
or collector, in the Rijksmuseum van Natuurlijke Historic, Leiden,
Netherlands. Paratype male, same data in the Francis Huntington
Snow Entomological Collection, at the University of Kansas, Law-
rence, Kansas.

Distributional data. As given above for the type series.

Comparative notes. This species superficially resembles the
fuscipes group, the abdominal sternites are more like those of the
amplicollis group. There is a possibility that these specimens could
represent the male sex of N. rudis (Melin), but the necessary evi-
dence of proof is lacking, so for the present these specimens are
described as a distinct species.

The Family Gelastocoridae (Hemiptera) 393

Nerthra fuscipes (Guerin-Meneville)

(PI. VII, fig. 52; PI. VIII, fig. 63; PI. IX, fig. 76; PI. X, fig. 89)

1843. Monomjx fuscipes Guerin-Meneville, Revue Zoologique Travaux Inedits,

p. 114.
1859. M. fuscipes Dohrn, Catalogus Hemipterorum, Herausgegeben von dem

Entomologischen Verein zu Stettin, p. 53.
1863. M. fuscipes Stdl, Berliner Entomologische Zeitschrift, VII, p. 406.
1876. M. fuscipes Stal, Kongl. Svenska Vetenskaps-Akademiens Handlingar,

Band 14, no. 4, p. 138.
1886. M. fuscipes Uhler, Bull. Brooklyn Ent. Soc, p. 27.
1895. M. fuscipes Pittier and Biolley, Inst, fisico geografico Nacional (Costa

Rica), p. 23.
1899. M. fuscipes Montandon, Bulletin de la Societe des Sciences de Bucarest-

Roumanie, an. VIII. no. 4 et 5, p. 392.
1901. M. fuscipes Champion, Biologia Centrali-Americana, Rhynchota Heterop-

tera, vol. II, p. 348.
1906. M. fuscipes Torre-Bueno, Ent. News, vol. 17, p. 54.

1909. M. fuscipes Kirkaldy and Torre-Bueno, Proc. Ent. Soc. Washington, vol.
X, p. 180.

1910. M. fuscipes Montandon, Annuario del Museo Zoologico della R. Univer-
sita di Napoli ( Nuova Serie) vol. 3, no. 10, p. 2.

1917. M. fuscipes Van Duzee, Catalogue of Hemiptera, Univ. Cahfomia Publ.,

p. 474.
1925. M. fuscipes Blatchley, Ent. News, vol. 36, p. 52.
1929. M. fuscipes Melin, Zoologiska Bidrag Fran Uppsala, Band 12, p. 181,

figs. 58-61.

Also referring to this species:

1853. Mononyx badius Herrich-Schaffer, Die Wanzenartigen Insecten, IX, p.
28, fig. 896.

1859. M. badius Dohrn, Catalogus Hemipterorum, Herausgegeben von dem

Entomologischem Verein zu Stettin, p. 53.
1862. M. badius Stal, Stettiner Entomologische Zeitschrift, XXXII, p. 459.
1873. M. badius Walker, Catalogue of Hemiptera in British Museum, Part

VIII, p. 172.
1876. M. badius Uhler, Bull. U. S. Geol. Sur\-., vol. 1, no. 5, p. 337.

1854. Motwnyx obscurus Stal, Ofversigt af Kongl. Vetenskaps-Akademiens
Fordhandlingar, Arg. 11, p. 239.

1859. M. obscurus Dohrn, Catalogus Hemipteromm, Herausgegeben von dem

Entomologischen Verein zu Stettin, p. 54.
1873. M. obscurus Walker, Catalogue of Hemiptera in the British Museum,

Part VIII, p. 172.

Size. Male: Length, 8.2 to 10.0 mm.; width of pronotum, 5.2 to
6.4 mm.; width of abdomen, 5.2 to 6.5 mm. Female: length, 8.8
to 11.0 mm.; width of p)ronotum, 5.6 to 6.6 mm.; width of abdomen,
5.7 to 6.8 mm.

Color. Light brown to dark blackish-brown; posterior margins
of the segments of the connexivum pale yellowish-brown; two tliin
blackish-brown spots on the sinuated part of the posterior margin
of the pronotum. Ventral surface usually entirely dark brown or
with the median part of abdominal segments, posterior lateral angles
of tlie abdominal segments and the apex of the intermediate and
hind femora pale yellowish-brown.

394 The University Science Bulletin

Structural characteristics. Apex of the head more or less con-
cavely excavated, a rather broad tubercle on either side of the
excavation; lateral tubercles small, generally forming a toothlike
protuberance between the eye and the apex of the head. Pronotum
widest at the level of the transverse furrow, not quite as wide as the
abdomen; lateral margins rather variable, median portion may be
rounded, concave or straight, most specimens are of the latter type;
posterior margin sinuated before the scutellum. Scutellum rather
large, slightly elevated, depressed at the basal angles. Hemelytra
extending beyond the end of the abdomen; lateral margin of the
embolium usually weakly convex. Connexivum barely visible in the
males, moderately prominent in the females. Abdominal sternites
of the female nearly symmetrical; last visible abdominal sternite
with the posterior margin medianly rather shallowly notched. Lobes
of ovipositor small, about as wide as long. Abdominal sternites of
the male asymmetrical; ninth sternite small, oval in shape; eighth
sternite twice as long as the ninth sternite. Clasper of the male
very characteristic, two processes extending mesad from tlie medial
surface.

Location of type. The type was in the Zoological Museum of the
University of Naples, Naples, Italy. It is not known to the writer
whether the type survived the last World War.

Distributional data. This species was described from a specimen
or specimens from Colombia. To present the distribution as given
in the literature would be useless in consideration of the fact that no
less than five species have been identified as this species by previous
workers. Specimens of N. fuscipes ( Guerin-Meneville ) before this
worker are from the following localities:

Mexico: Chichen Itza, Yucatan, Aug. 29, 1936, H. D. Thomas,
1 male; Motzorongo, Vera Cruz, Jan., 1892, H. Osborn, 1 male,
(USNM); Orizaba, Jan. 17, 1908, F. Knab, 4 males, (USNM);
Orizaba, Vera Cruz, Jan. 9-16, 1892, H. Osborn, 1 male, (USNM);
Otozac, Nov. 1887, L. Bruner, 1 male, (USNM); "Mexico," no
date, Storkan, 1 male, (Mus. Prague).

Guatemala: Coban, Alta Vera Paz, May 22, 1926, J. M. Aldrich,
1 male, (USNM); Morales, Dec, 1929, J. J. White, 3 males, (J. C.
Lutz Coll. ) ; Secanquin, Alta Vera Paz, Jan. 4, 1905, A. Mac Laklan,
1 male, (USNM).

Costa Rica: La Carpintera, Apr., 1924, W. M. Mann, 1 male,
(USNM); San Carlos, no date, Schild Bugdorf, 1 male, (USNM).

The Family Gelastocoridae (Hemiptera) 395

Honduras: Progresso, Mar. 8, 1923, T. H. Hubbell, 2 females;
San Pedro, Sula, no date, W. M. Mann, 1 male, (USNM).

Salvador: La Libertad, Jan. 24, 1930, T. O. Zschokke, 1 female,
(CAS).

Panama: Albrook Field, Canal Zone, Nov, 5, 1937, R. Bliss, 1
male and 1 female, (J. C. Lutz Coll.); Ancon, Canal Zone, May 15-
28, 1918, J. Zetek, 1 male, (USNM); Barro Colorado I., Canal
Zone, Nov. 22, 1944, K. E. Frick, 1 male, (CAS); same place, Dec.,
1946 to Feb., 1947, J. Zetek, 1 male, (USNM); Boquete, Chiriqui
Prov., Mar. 7, 1923, F. M. Gaige, 1 female; Canal Zone, Feb., 1923,
C. V. Riley, 1 male, (USNM); Cano Saddle, Gatiin L., Aug. 6,
1923, R. C. Shannon, 1 male, (USNM); Chilibrillo Caves, Apr. 30,
1945, K. E. Frick, 1 female, (CAS); Colom (Colon ?), "Casey
bequest 1925," 1 male, (USNM); Corozal, May 12, 1937, R. Bliss, 1
male, (J. C. Lutz Coll.); Fort Clayton, Canal Zone, Feb., 1945,
K. E. Frick, 1 male, (CAS); Fort Kobbe Rd., Canal Zone, May
19, 1937, R. Bliss, 1 female, (J. C. Lutz Coll.); Gatun, Canal Zone,
Mar., 1930, T. O. Zschokke, (CAS); same place, no date, A. H.
Jennings, 1 male, (USNM); Gatun Dam, C. Z., Apr. 13, 1930, T. O.
Zschokke, 2 males and 1 female, (CAS); Gatun Lake, Aug. 16, 1931,
T. O. Zschokke, 2 males, (CAS); Paraiso, C. Z., June 5, 1911, A.
Busck, 1 male, (USNM); same place, no date, A. H. Jennings, 1
male, (USNM); "Pan.," no date, P. R. Uhler, 1 male, (USNM).

Colombia: Palmira, May, 1943, B. Losada S., 1 male, (USNM).

Brasil: "Rio de Jan." Oct., no collector, 1 male, (Car. Mus.).

Puerto Rico: Ponce, July 10, 1928, L. F. Con., 1 male and 2
females, (J. C. Lutz Coll.).

Comparative notes. The original description of this species is
not suflBcient to determine accurately this species, however, the de-
scription by Montandon, 1899, his confirmation of that description
after studying the type, 1910, and specimens determined as this
species by Montandon, led the writer to accept Montandon's con-
cept of the species. However, an examination of the male genitalia
soon revealed that what Montandon and other workers had con-
sidered one species was in reality a complex of at least four closely
related species. The question therefore arises as to which of these
four species should the name N. fuscipes ( Guerin-Meneville ) be
assigned. Since the location of the type is unknown at present, the
writer has been forced to base his decision upon geographical dis-
tribution. Of tlie four species, only one species was represented, in

396 The University Science Bulletin

the material studied, from areas as far south as Colombia. This
species I am presently calling N. ftiscipes ( Guerin-Meneville ) .

This species is most closely related to N. manni n. sp. and can
be separated from that species only by the male genitalia. While the
later species appears to occur mainly in Western Mexico, the
ranges of the two appear to overlap at the Isthmus of Tehuantepec.
Because of this overlap, females from Eastern Mexico, Southern
Mexico and Guatemala were not included in the distributional data
of either species.

Nerthra manni n. sp.

(PI. VII, fig. 54; PI. VIII, fig. 66; PI. IX, fig. 80)

Size. Male: Length, 9.1 to 10.2 mm.; width of pronotum, 5.6
to 6.1 mm.; width of abdomen, 5.6 to 6.3 mm. Female: Length, 8.8
to 11.0 mm.; width of pronotum, 5.6 to 6.6 mm.; width of abdomen,
5.7 to 6.8 mm.

Color. Light brown to dark blackish-brown; posterior margins
of the segments of the connexivum pale yellowish-brown; darker
irregular spots on the posterior margin of the pronotum; ventral
surface usually entirely dark brown or with the median part of
abdominal segments, posterior angles of the abdominal segments,
and the apex of the intermediate and hind femora pale yellowish-
brown.

Structural characteristics. Apex of the head more or less con-
cavely excavated, a rather broad tubercle which is variable in size
on either side of the excavation; lateral tubercles generally forming
a toothlike protuberance between the eye and the apex of the head,
but sometimes a few are separate between the protuberance and
the tubercle that is laterad of the apical excavation; ocelli present.
Pronotum widest at the level of the transverse furrow, not quite as
wide as abdomen, although this may vary with some specimens;
lateral margins variable, median part in particular, either slightly
rounded, straight or weakly concave; posterior margin sinuated
before the scutellum. Scutellum rather large, slightly elevated,
depressed at the basal angles. Hemelytra extending beyond the
end of the abdomen in normal specimens; lateral margin of the
embolium usually weakly convex. Connexivum barely visible in
the males, moderately prominent in the females. Abdominal
sternites of the female nearly symmetrical; last visible abdomi-
nal sternite with the posterior margin medially rather shallowly
notched. Lobes of ovipositor small, about as wide as long. Abdom-

The Family Gelastocoridae (Hemiptera) 397

inal sternites of the male asymmetrical; ninth sternite small, about
half as long as the eighth sternite. Clasper of the male with one
large process or projection on the medial surface, the medial basal
angle of the projection deeply sinuate.

Location of types. Holotype male and allotype female, Taxco,
Guerrero, Mexico, Oct., 1946, J. G. Shaw; 1 male paratype. La
Sabana, Guerrero, Mexico, kil. 226 S. Mexico City, Oct. 20, 1936,
H. D. Thomas; 3 male and 9 female paratypes, Tejupilco, District
of Temascaltepec, Mexico, June-July, 1933, H. E. Hinton in the
Francis Huntington Snow Entomological Collections at the Uni-
versity of Kansas. One male and 2 female paratypes, Tejupilco,
Dist. Temascaltepec, Mexico, June 24, 1933, R. L. Usinger and
H. E. Hinton; 1 male paratype, Bejucos, Dist. of Temascaltepec,
Mexico, July 3, 1933, H. E. Hinton and R. L. Usinger in the R. L.
Usinger Collection. One male paratype, Cuernavaca, Morales,
Mexico, July, 1945, N. L. H. Krauss; 9 male and 7 female paratypes,
Las Barrancas, Mexico, Mar. 1923, W. M. Mann; 13 male and 7
female paratypes. Above Tepic, Nayarit, Mexico, Mar., 1923, W. M.
Mann in the United States National Museum at Washington, D. C.
One male paratype, Cuernavaca, Morales, Mexico, Nov. 19, 1946,
E. S. Ross; 1 male paratype, 3 miles S. Acahuizotla, Guerrero,
Mexico, Nov. 17, 1946, E. S. Ross; 1 female paratype. Alamos,
Sonora, Mexico, Oct. 2, 1923, H. S. Gentry in the collections of the
California Academy of Science at San Francisco, California.

Distributional data. This species appears to be restricted to
Mexico, particularly to the western states. It may also possibly
occur in the extreme southern portions of California and Arizona
since I have seen a specimen from Sonora, Mexico (paratype in
the California Academy of Science's collection). I have also seen
a female specimen which was taken from the crop of a chicken
in Nogales, Arizona. There is no way of knowing where the
chicken came from, however, the state of preservation of the speci-
men would indicate that it had not been in the chicken's crop very
long. This specimen is in the Francis Huntington Snow Entomologi-
cal Collections at the University of Kansas.

Comparative notes. This species can be separated from N.
fuscipes ( Guerin-Meneville ) only by the male genitaha. In this
species there is a single large medial process while in the latter
species there are two smaller medial processes.

This species could have been Herrich-Schiiffer's Monomjx badiiis
or Stal's M. obscurus, however, since this species is apparently

398 The University Science Bulletin

rather restricted to western Mexico, I rather doubt that it was
either. The type of the former is apparently lost and the
type of the latter may be with other types of Stal in the Museum
of Stockholm, Stockholm, Sweden. If it is located there, Melin,
1929, does not mention it.

This species is named for W. M. Mann, Director of National
Zoological Park, Washington, D. C, who collected many specimens
not only of this species but of many other species in this family.

Nerthra hungerfordi n. sp.

(PI. VII, fig. 51; PI. VIII, fig. 60; PI. IX, fig. 78)

Size. Male: Length, 7.5 to 9.0 mm.; width of pronotum, 5.0 to
5.8 mm.; width of abdomen, 5.0 to 5.9 mm. Female: Length, 8.1
to 9.4 mm.; width of pronotum, 5.5 to 6.0 mm.; width of abdomen
5.8 to 6.4 mm.

Color. Brown to almost black; posterior margin of the pronotum
with indistinct alternating black and yellowish-brown spots; con-
nexivum either entirely dark or with a thin yellowish-brown or
orange area at the posterior margins of the segments; ventrally
mostly dark brown; the forelegs reddish-black; yellowish-brown
spots present on the apex of the intermediate and hind femora and
on the abdominal segments.

Structural characteristics. Apex of the head more or less con-
cavely excavated, with a tubercle on either side of the excavation;
lateral tubercles irregular in size and position, sometimes forming
a toothlike protuberance between the eye and the apex of the head.
Pronotum widest at the level of the transverse furrow, not as wide
as abdomen in the female, about as wide as the abdomen in the
male; lateral margins slightly variable in the specimens studied,
generally with the median part nearly straight and parallel to the
longitudinal axis of the body, anterior third roundly curved to the
eye, posterior fourth straight to slightly concave running mesad
in an oblique manner to the base of the embolium; posterior margin
sinuated before the scutellum. Scutellum rather large, sides and
apex slightly tumescent. Hemelytra extending beyond tlie end of
the abdomen in the males, in the females extending to the end or
slightly shorter than the abdomen; embolium with lateral margin
moderately convex. Connexivum barely visible in the males, moder-
ately so in the females. Body covered by very thin, rather long
black bristles. Abdominal sternites of the female nearly symmetri-
cal; posterior margin of the last visible sternite medially shallowly

The Family Gelastocoridae (Hemiptera) 399

emarginated. Lobes of ovipositor about as wide as long. Abdomi-
nal sternites of the male asymmetrical; ninth sternite small, about
half as long as the eighth sternite; eighth sternite over half as wide
as fifth abdominal sternite. Clasper of the male somewhat resem-
bling that of N. fuscipes (Guerin-Mcneville), but with the dorsal
surface more irregular, a small dorsal projection present near the
bend of the clasper and the medial projections closer to each other.
Location of type. Holotype male, allotype female, and 1 male and
8 female paratypes, Rio Virilla, Costa Rica, Dec. 26, 1931, H.
Schmidt; 8 male and 9 female paratypes, San Jose, Costa Rica,
1932, H. Schmidt; in the Francis Huntington Snow Entomological
Collection at the University of Kansas. In addition to the above
series, there is 1 male and 1 fem.ale paratypes, San Jose, Costa Rica,
1928, M. Valerdo, in the United States National Museum at Wash-
ington, D. C; 1 male parat>'pe, Finca, El Cipres, Suchitepequez
Prov., Guatemala, no date. J. R. Slevin, in the Collection of the
California Academy of Science at San Francisco, California; and,
1 male paratype, Boquete, Chiriqui Prov., Panama, Mar. 2, 1923,
F. M. Gaige, in the Collection of the University of Michigan at Ann
Arbor, Michigan.

Distributional data. Besides the types mentioned above, the
writer has studied specimens from the following localities:

Guatemala: Alta Vera Paz, Apr. 30, Schwarz and Barber, 1 male,
(USNM).
Nicaragua: "Nicarag.", no date, P. R. Uhler, 1 male, (USNM).

British Honduras: Belize River, Mar. 14, 1922, H. F. Loomis, 1
male, (USNM).

Honduras: Tela, May 23, 1923, T. H. Hubbell, 1 female.

Salvador: La Libertad, Jan. 24, 1930, T. D. Zschokke, 1 female,
(CAS); San Andres, July 25, 1944, E. J. Hambleton, 1 female,
(USNM); San Salvador, May 26, 1925, K. A. Salman, 2 males,
(USNM).

Costa Rica: Estrella, no date or collector, 1 male, (USNM); Rio
Virilla, Costa Rica, June to July, 1931, 1 male and 3 females; "Costa
Rica," no date, C. F. Baker, 1 female, (USNM).

Comparative notes: This species is closely related to N. fuscipes
(Guerin-Meneville) and N. manni n. sp., but it can be separated
from these two species in the case of the males by the shape of
the clasper, by the fact that the eighth abdominal sternite of the male
is more than half as wide as the fifth abdominal sternite, and by

400 The University Science Bulletin

the slightly smaller size. I name this species after Doctor H. B.
Hungerford, under whose supervision this investigation was made.

Nerthra bracchialis n. sp.

(PL VII, fig. 45; PI. IX, figs. 71, 79)

Size. Length 7.0 to 7.7 mm.; width of pronotum, 4.5 to 4.7 mm.:
width of abdomen, 4.8 mm. Female: Length, 8.0 mm.; width of
pronotum, 5.1 mm.; width of abdomen, 5.2 mm.

Color. Brown to blackish-brown; posterior part of the segments
of the connexivum in the female thinly yellow; ventrally mostly
dark brown with yellowish-brown spots on the intermediate and
hind legs and on the abdomen.

Structural characteristics. Apex of the head more or less con-
cavely excavated, a stout tubercle on each side of the excavation;
lateral tubercles of the head forming a toothlike protuberance be-
tween the eye and the apex of the head. Pronotum widest at a level
with the transverse furrow, slightly less in width than the abdomen;
lateral margins more or less rounded; posterior margin before
scutellum sinuated. Scutellum large, slightly elevated. Hemelytra
extending to or beyond the end of the abdomen; embolium with
the lateral margin slightly convex. Connexivum barely visible in
the male, prominent in the female. Abdominal sternites of male
asymmetrical; ninth sternite small, oval in shape, about half as
long as eighth sternite; eighth sternite about half as wide as fifth
sternite. Abdominal sternites of the female nearly symmetrical;
posterior margin of last visible sternite shallowly notched medially.
Lobes of ovipositor about as wide as long. Clasper of the male char-
acteristic; a large projection on the medial surface, medial basal
angle of the projection rounded.

Location of type. Holotype male, Mazatlan, Mex., P. R. Uhler
Coll. in the United States National Museum at Washington, D. C.
Allotype female, Real de Arriba, Dist. of Temascaltepec, Mexico,
VII- 6- 33, H. E. Hinton & R. L. Usinger; one male paratype.
Tejupilco, Dist. of Temascaltepec, Mexico, VI- 24- 33, H. E. Hinton
& R. L. Usinger in the R. L. Usinger Collection.

Distributional data. As given above for the type series.

Comparative notes. This species may be separated from the
other three species of this closely related group by the male clasper
and the smaller size.

The Family Gelastocoridae (Hemiptera) 401

Nerthra grandicolUs (Germar)

(PI. XI, fig. 96)

1837. Mononyx grandicolUs Germar, Silbermann's Revue Entomologique,

V, p. 122.
1859. M. grandicolUs Dohm, Catalogiis Hemipterorum, Herausgegeben von

dem Entomologischen Verein zu Stettin, p. 53, 54.
1863. M. grandicoUis Stal, Berlinger Entomologische Zeitschrift, VII, p. 407.
1873. M. grandicoUis Walker, Catalogue of Hemiptera in British Museum, pt.

VIII, p. 172.
1892. M. grandicoUis Gerstaecker, Jahrbuch der Hamburgischen Wissen-

schaftlichen Anstalten, Vol. IX, 2, p. 14.
1899. M. grandicoUis Montandon, Bulletin de la Societe des Sciences de

Bucarest-Roumanie, an. VIII, nos. 4 and 5, p. 396.
1899. M. grandicoUis Horvath, Termeszetrajzi Fuzetek, XXII, p. 268.
1908. M. grandicoUis Montandon, Herausgegeben von der Kongl. Schwe-

dischen Akademie der Wissenschaften, p. 20.
1914. M. grandicoUis Montandon, Insectes Hemipteres, II, p. 122.

1925. M. grandicoUis Singh-Pruthi, Trans. Royal Ent. Soc. London, p. 184.

1926. M. grandicoUis Jaczewski, Annalibus Zoologicis Musei Polonici Historiae
Naturalis, T. V., zesz. 2, pp. 72-74.

1934. M. grandicoUis Poisson, Bulletin de la Societe des Zoologique de

France, tome LIX, no. 1, p. 97.
1940. M. grandicoUis Poisson, Bulletin du Musee royal d'Histoire naturelle

de Belgique, tome XVI, no. 40, p. 13.

Also referring to this species:

1865. Phintius grandicoUis Stal, Hemiptera Africana, III, p. 172.

1876. P. grandicoUis Stal, Kongl. Svenska Vetenskaps-Akademiens Handlingar,

Band 14, no. 4, p. 139.
1892. P. grandicoUis Brancsik, Jahrbuche der naturqissenschaftlichen Ver-

eines des Trencsiner Comitates, XV, p. 251.
1853. Mononyx sordidus Herrich-Schaffer, Die Wanzenartigen Insecten, IX,

p. 26, fig. 893.

1859. M. sordidus Dohm, Catalogus Hemipterorum, Herausgegeben von dem
Entomologischen Verein zu Stettin, p. 54.

1860. M. sordidus Signoret, Annales de la Societe des Entomologique de
France, p. 969.

1855. Mononijx Umigenus Stal, Ofversigt af Kongliga Vetenskaps-Akademiens

Fordhandlingar, XII, p. 46.
1859. M. Umigenus Stal, Dohrn, Catalogus Hemipterorum, Herausgegeben

von dem Entomologischen Verein zu Stettin, p. 54.
1858. Mononyx rotundicoUis Signoret, in Thoms. Arch., 2, p. 329, fig. 628.
1929. Phintius staU Melin, Zoologiska Bidrag Fran Uppsala, Band 12, p. 193,

figs. 100-102, 106. (?)

Size. Male: Length, 7.5 to 9.3 mm.; width of pronotiim, 5.5 to
6.3 mm.; width of abdomen, 5.5 to 6.3 mm. Female: Length, 8.2 to
10.4 mm.; width of pronotum, 6.4 to 7.5 mm.; width of abdomen,
6.0 to 7.8 mm.

Color. Variable, from light yellowish-brown to dark brown;
connexivum with the posterior half of each segment yellowish-
brown, anterior half darker; ventrally mostly dark brown, dilation
of front femur, intermediate and hind femora and the abdominal
segments either dark or spotted with yellowish-brown.

Structural characteristics. Apex of the head more or less con-

402 The University Science Bulletin

cavely excavated; four tubercles present, three on the frons and
one on the apex, the latter is not visible from a dorsal view. Pro-
notum widest at a level with the transverse furrow, projecting
beyond the base of the embolium; lateral margins more or less
rounded, converging anteriorly; posterior margin sinuated before
scutellum, crossed by nine indistinct longitudinal carinations. Scu-
tellum elevated, tumescent laterally and at the apex. Hemelytra
extending to or beyond end of abdomen in the males, usually shorter
than abdomen in the females; embolium strongly dilated from the
base to the middle, lateral margin of the dilation nearly straight.
Connexivum moderately visible in the males, very prominent in
the females. Short clavate bristles in groups on the hemelytra, and
especially prominent on the tumescence of the scutellum and the
carinations of the pronotum. Front femur with anterior margin
wavy with toothlike microscopic spines. Abdominal sternites of
the female asymmetrical; posterior margin of the last visible
abdominal sternite medially deeply notched. Lobes of ovipositor
rather large, elongate, rounded posteriorly. Abdominal sternites
of the male asymmetrical; ninth sternite about as long as the
eighth sternite, twice as long as the seventh, slightly wider than
long. Clasper of the male twisted slightly mesad at apex, aedeagal
furrow visible for the entire length of the dorsal surface.

Location of type. Unknown. Germar's Hemipterous specimens
are located in the Museum of Halle and Dahlem Museum in Ger-
many and Museum of Lomberg in Poland. Doctor H. B. Hunger-
ford visited the Dahlem Museum in Berlin in 1928, and did not
find the type there.

Distributional data. This species apparently is the only species
of this family present in Africa, where it is found from east to west,
south of the Sahara. It also occurs on Madagascar. The writer
has studied specimens from the following localities:

Belgian Congo: Ava Kubi, Nov. 10-30, 1909, Lang and Chapin,
1 female, (AMNH); Faradje, Feb., 1911, Lang and Chapin, 1 male,
(AMNH); same place. Mar., 1911, Lang and Chapin, 1 female,
(AMNH); Garamba, May 3-11, 1912, Lang and Chapin, 1 female,
(AMNH); same f)lace, July, 1912, Lang and Chapin, 3 males,
(AMNH); Kinda Katana, no date or collector, 1 female; Luebo,
D. W. Snyder, no date, 2 females, (USNM); Medjie, May, 1914,
Lang and Chapin, 1 male, (AMNH); Niangara, Nov., 1910, Lang
and Chapin, 1 female, (AMNH); Parva, July, 1913, Lang and
Chapin, 1 male, (AMNH); Panga, Sept., 1914, Lang and Chapin,

The Family Gelastocoridae (Hemiptera) 403

1 female, (AMNH); Stanleyville, Mar., 1915, Lang and Chapin, 2
males, (AMNH); same place, Aug. 7, 1909, Lang and Chapin, 1
female, (AMNH); same place, Sept. 5, 1909, Lang and Chapin,
1 female, (AMNH); Vankerckhovenville, April, 1912, Lang and
Chapin, 1 female, (AMNH).

Cameroons: Batanga, Mar., 1914, F. H. Hope, 1 male and 1
female, (Car. Mus. ); same place, April, 1914, F. H. Hope, 1 male
and 1 female, (Car. Mus.); same place, June 15, 1920, F. H. Hope,

1 female, (Car. Mus.); Metet, May 13, 1919, A.I. Good, 1 female
(Car. Mus.); Victoria, no date or collector, 2 males and 1 female;
Yaunde, no date or collector, 1 male, (Car. Mus.); "N. Kamerun,
Johann-Albrechtshohe, L. Conradt, 96", 1 female, (USNM).

Ethiopia: Cheren, Erythraea, no date or collector, 1 male and

2 females.

French Equatorial Africa: Fort-Crampel, no date or collector, 2
females; Gabun, no date or collector, 1 male; Kangwe, no date
or collector, 1 male; Lambarene, May, 1892, A. C. Good, 1 male,
(Car. Mus); Ogove (Ogowe?) River, no date or collector, 1 male,
(Car. Mus.).

Kenya: Diani Dist., 20 mi. S. of Mombasa, Apr. 28, 1948, F. X.
Williams, 1 female.

Liberia: Bendija, 1940, W. M. Mann, 1 male, (USNM); Bromley,
1940, W. M. Mann, 1 female, (USNM); Mombo, 1940, W. M.
Mann, 1 female, (USNM); Mt. Coffee, Apr., 1897, R. P. Currie,
6 males and 2 females, (USNM); same place, 1897, Mrs. Sharp,
Imale, (USNM).

Mozambique: Delog. Bay (Delogoa Bay), no date or collector,
1 female, (USNM); Lowraulo Margues (Lourenco Margues), no
date or collector, 1 male, ( USNM ) .

Nigeria: Ibadan, Apr. 23-May 6, 1936, Van. Zwaluwenburg and
McCough, 1 male, (USNM).

Portuguese Guinea: Borma, June to Dec, 1899, L. Fea, 2 females,

(USNM).

Sierra Leone: "Sierra Leone," no date or collector, 1 male.

Tanganyika: Lindi, Apr., no collector, 1 male and 1 female; same
place, no date or collector, 1 male.

Transvaal: Ugogo, no date or collector, 1 male; Zontpansberg,
March, no collector, 1 female; "D. O. Afr., from Rolle," 1 male,
(CAS).

404 The University Science Bulletin

Colony Unknown: "Nliosamque, Feb. 1, 1910, Howard Coll.", 1
male and 3 females, ( USNM ) .

Madagascar: Diego-Suarez, 1893, Ch. Alluaud, 1 male, (Mus.
Paris); Great Oriental Forest, no date or collector, 2 males; Maro-
antsetra, no date or collector, 2 males and 1 female; same place,
no date or collector, 1 male and 2 females.

Comparative notes. Most closely related to N. indica (Atkinson)
from which it may be distinguished by the shape of the dilation of
the embolium, more rectangular, by the shape of the clasper of the
male, and of course by its peculiar distribution. It is the only species
of this genus known from Africa.

Melin, 1929, re-estabhshed the genus Phintiiis Stal and described
a new species from a single male specimen which is labeled "Mexico-
Liebmann." He named this species Phintiiis stali. While it is not
impossible, especially in consideration of the distribution of Nerthra
riigosa (Desjardins), that a species closely related to Nerthra grandi-
collis (Germar) could occur in the New World, I personally feel
that it is quite unlikely. It is my opinion that the label has been
changed on this specimen. Melin does not state how this specimen
differs from N. grandicolUs (Germar) and his drawings do not
show any differences. For these reasons I am temporarily placing
this species in the synonomy of N. grandicolUs (Germar). In the
event that an examination of the type proves that this a good
species, the name will have to be changed since it is a homonym
of N. staJi (Montandon) which was described much earlier. The
original description of Pliintiiis stali Melin follows:

"Edge of mouth with a toothlike process; front with s'^rongly
marked medial knob; medial processes fairly large, knoblike and
pointed; lateral ones forming a dentated ridge. Structure of pro-
notum fairly strongly marked with prominent ridges at transversal
furrow, but without stem of T-shaped part; side rounded, level
with the furrow, fairly strongly projecting beyond base of embolium;
anterior corner hookshaped; posterior margin before scutellum
broadly by not very deeply sinuated. Scutellum with traces of
protuberances, apical knob very prominent. Processes of femora
apically, obtuse-angled and rounded, anterior side wavy. Apical
segment in male fairly small, not very much longer than preceding
one. Bristles short, conspicuously clavate, densely grouped on
sides of scutellum.

"Coloration brownish; abdomen ventrally, darkish-brown with
posterior margin of anterior segments lighter; legs light brown;

The Family Gelastocoridae (Hemiptera) 405

femora of 1st pair posteriorly, darker; other femora indistinctly
ringed.

"Length 9, breadth 6 mm.

"Museum of Kobenhavn: 1 male, Mexico (Liebmann).

"As far as I know, no species of this genus has hitherto been
found in tlie New World. For this reason I thought the label of
the specimen might possibly have been changed, but I could not
find the type in any paper."

Nerthra indica ( Atkinson )

(PI. XI, fig. 98)

1888. Monomjx indicus Atkinson, Journal of the Asiatic Society of Bengal, vol.

LVII, pt. 2, p. 345.
1899. M. indicus Montandon, Bulletin de la Societe des Sciences de Bucarest-

Roumanie, an. VIII, nos. 4 and 5, p. 397.
1906. M. indicus Distant, Fauna British India, vol. Ill, p. 15.
1919. M. indicus Paiva, Records of the Indian Museum, vol. XVI, pt. 5, no. 23,

p. 372.

Also referring to this species:

1906. Mononyx projectus Distant, Fauna British India, vol. V, p. 310, fig. 174.

Size. Male: Length, 8.4 to 9.1 mm.; width of pronotum, 5.5 to
6.7 mm.; width of abdomen, 5.7 to 6.2 mm. Female: Length, 9.7
to 9.9 mm.; width of pronotum, 7.0 to 7.3 mm.; widtli of abdomen,
7.2 mm.

Color. Light brown to dark reddish-black; segments of connexi-
vum mostly uniform in color, occasionally with posterior half of
each segment lighter. Legs generally light yellowish-brown, ab-
domen, except for marginal spots, darker.

Structural characteristics. Front of head with four tubercles,
three on the frons and one at the apex which is not visible from a
dorsal view, the medial tubercle of the frons and the apical tubercle
sometimes rather indistinct. Pronotum about as wide at anterior
third as at the level with the transverse furrow; lateral margins
irregular in shape, usually sinuous, but variable even on the two
sides of the same specimen; posterior margin sinuated before scutel-
lum; posterior third of pronotum crossed by nine indistinct longi-
tudinal carinations. Scutellum elevated, tumescent laterally and
at apex, a curved ridge paralleling sinuosity of posterior margin of
pronotum. Hemelytra extending beyond end of abdomen in the
males, not quite reaching end of abdomen in the females; embolium
narrow at base, dilated before middle, anterior portion and apex
of dilation more or less rounded. Abdomen broadly dilated laterally
in the female. Bristles mostly short and clavate, groups of long

406 The University Science Bulletin

black bristles on basal tumescences and median part of pronotum.
Abdominal sternites of female nearly symmetrical; last visible ab-
dominal sternite medially emarginated, flattened on either side of
emargination. Lobes of ovipositor asymmetrical, slightly lobed
and projecting posteriorly. Abdominal sternites of the male asym-
metrical; ninth sternite wider than long, not quite as long as eighth
sternite but longer than seventh. Male clasper similar to that of N.
lohata (Montandon) but smaller, not as pointed apically.

Location of type. Unknown. It apparently is not in the British
Museum with the rest of Atkinson's specimens.

Distributional data. This species is apparently restricted to India.
This worker has seen specimens from the following localities:

Chabua, Assam, Aug. 2, 1943, D. E. Hardy, 1 male, (USNM);
Himalaya, 1900, Tulis, 1 female; Kouy-Tcheon, Reg. de Pin-Fa,
1908, Pere Cavalerie, 1 male, (Mus. Paris); Kurseong, no date, P.
Blaet, 1 male, (USNM); Sandhira, Sylket, no date, Heidemann, 1
female.

Comparative notes. Most closely related to N. lohata (Montan-
don ) from which it may be separated by the male genitalia, by the
smaller ovipositor lobes which are less projecting and by the lack
of lateral submarginal tumescences of the last visible abdominal
sternite in the case of the females. In both sexes the lateral dilation
of the embolial margin is not so triangular as in N. lohata.

A comparison of the male genitalia of the type of Mononyx pro-
jectiis Distant with the male genitalia of N. indica (Atkinson) as
identified in this study has shown that they are the same.

Nerthra turgidula (Distant)

1906. Mononijx turgidulus Distant, Fauna British India, vol. V, pp. 311-312,
fig. 175.

The writer has not seen this species or, at least, has not been
able to recognize it.

Original description: "Head, pronotum and scutellum more or
less dark greyishly pubescent; hemelytra pitchy black; connexivum
greenish ochraceous, with subquadrate piceous segmental spots;
head between eyes about twice as broad as long, anterior margin
shortly setose and with two short obscure spines at middle; eyes
a little recurved, brownish ochraceous, above interiorly fuscous;
pronotum with the anterior and posterior margins subequal in
breadth, the lateral margins moderately, subtruncately produced,
the anterior disk moderately gibbous, but not prominentiy nodulose,

The Family Gelastocoridae (Hemiptera) 407

finely granulose as seen between the pubescence, transversely im-
pressed at about one-third before basal margin, behind this im-
pression longitudinally carinate, the most prominent carinations
being six in number, the lateral areas are flatly ampliate, very
much less pubescent and more ochraceous in hue, their margins
shortly setose, prominently sinuate near anterior angles which are
shortly obtusely spinous; scutellum with a short uprightly setose
elevation on each side near base, its apex somewhat prominently
tuberculous; corium thickly, finely granulose; membrane not
reaching abdominal apex; connexivum broad, the posterior angles of
the segments moderately angulate, the segments also moderately
foveately depressed; body beneath palely fuscously pubescent, the
lateral areas of the prosternum more or less ochraceous; abdomen
with a lateral fascia on each side united subapically, black in ap-
pearance caused by the absence of the paler pubescence; anterior
legs fuscously pubescent, the tarsi castaneous at base and black
at apex; intermediate and posterior legs more or less olivaceous,
shortly setose.

"Length 9.5 millim.; breadth of pronotum 6.5 millim.

"Hab. Nepal; Gowchar (Coll. Dist. )."

This species may very easily be a synonym of N. inclica (Atkin-
son ) , however, it is the opinion of the writer that the species should
be retained until such time as the type may be studied and com-
pared to the other species from India.

Nerthra lobata (Montandon)

(PI. XI, fig. 97; PI. XV, fig. 138)

1899. Mononyx lohatus Montandon, Bulletin de la Societe des Sciences de
Bucarest-Roumanie, an. VIII, nos. 4 and 5, p. 397.

Size. Male: Length, 8.6 mm.; width of pronotum, 6.1 mm.;
width of abdomen, 6.2 mm. Female: Length, 10.0 to 10.5 mm.;
width of pronotum, 6.8 to 7.0 mm.; width of abdomen, 7.3 to 7,4 mm.

Color. Dorsally dark reddish-brown; segments of connexivum
posteriorly yellowish-brown, sometimes entirely dark; front femur
mostly dark, anterior dilation and area before apex yellowish-brown;
intermediate and hind femora light with a dark brown ring; joint
between tibia and femur of all legs black; lateral margins of the
abdominal segments yellowish-brown at posterior half.

Structural characteristics. Front of head more or less concavely
excavated; a pair of large pointed tubercles on frons, eyes notice-
ably projecting. Pronotum widest at anterior third; lateral margin

408 The University Science Bulletin

broadly sinuous, two sinuosities before and two behind anterior
third; posterior margin sinuated before scutelkim; posterior part
of the disc of the pronotum crossed by nine indistinct longitudinal
carinations. Scutellum moderate in size, elevated, a tumescence
at each basal angle and at apex. Hemelytra short not reaching end
of abdomen; embolium very narrow at base, suddenly dilated
laterally before middle, dilation more or less triangular. Abdomen
very prominently dilated laterally. Lobes of ovipositor large, lobed
and prominently projecting posteriorly. Bristles on body short
and clavate, except those on scutellum which are rather elongate,
not very clavate. Bristles on basal tumescences of scutellum very
dense and black in color, those on the apex also dense, but light
brown in color. Front femur with anterior margin irregular, cov-
ered with black toothlike miscroscopic spines. Abdominal sternites
of the female slightly asymmetrical, posterior margin of last visible
sternite notched medially with a slightly depressed area on each
side, laterad of which is a moderate tumescence, the one on the
left the larger. Abdominal sternites of the male asymmetrical;
ninth sternite rather oval, wider than long, slightly shorter than
eighth sternite; seventh sternite about half as long as eighth
sternite; fifth sternite very short medially. Male clasper swollen
apically, bent mesad, tapering to fairly acute point at apex.

Location of type. In the British Museum, at London, England.

Distributional data. When Montandon described this species
he had specimens from Sumatra and Java before him. All speci-
mens examined by the writer have been from Sumatra. They are
from the following localities:

Aek Coelangen, N. Harbisaran, Sept. 6, 1928, Meer Mohr, 1
male and 3 females; "Sumatra," no date, R. Weber, 1 female,
(AMNH).

Comparative notes. This species is mostly closely related to
Neiihra indica (Atkinson) but differs from that species in that
the abdomen of the female is wider, the ovipositor lobes much
more projecting posteriorly, lateral tumescences of the last visible
abdominal sternite are present, in the males the clasper while quite
similar is not dilated so abruptly as in N. indica (Atkinson) and is
more sinuous apically and in both sexes the lateral dilation of the
embolial margin is more triangular.

It may be that N. serrata (Montandon) will prove to be this
species in which case Nerthra lobata (Montandon) would fall as
a synonym since it was described two years after the former

The Family Gelastocoridae (Hemiptera) 409

species. Doctor R. L. Usinger examined the clasper of the type
of N. serrata ( Montandon ) and is of the opinion that it is the same
as my drawing of the clasper of A/, lobata (Montandon). Mon-
tandon's description of N. serrata states that the pronotum laterally
is very dilated and that the segments of the connexivum arc
concave. None of the specimens I have examined are of this
nature. For that reason and because both species were described
by Montandon, this writer prefers to retain both species until
more specimens can be examined. The original description o(
Nerthra serrata (Montandon) is given at another point in this
work.

Nerthra serrata (Montandon)

1897. Mononyx serrattis Montandon, Annals Musee Civique di St. Nat.

Genova, I, p. 365.
1899. M. scrratus Montandon, Bulletin de la Societe des Science de Bucarest-

Roumanie, an VIII, nos. 4 and 5, p. 396.
1906. M. serratus Distant, Fauna British India, vol. Ill, p. 15.

The writer has not seen this species or at least has not been able
to recognize it.

Original description: "2. Mononyx serratus nov. sp. — Carin
Cheba, Carin Ghecu. Collection du Musee Civique de Genes, du
Musee d'hist. Nat. de Hambourg et la mienne.

"Longueur 8.2 - 9.4 mill.; largeur max. pronotum 6-7 mill,,
abdominale 5.8 - 6.8 millimetres.

"Tete fortement tuberculeuse; en regardant I'insecte directement
en dessus on apergoit tres bien trois forts tubercules dentiformes
en avant, les deux de la partie anterieure de la tete laissent entre
eux une assez profonde echancrure au milieu de laquelle se presente
le sommet du troisieme dont la base est situee un peu en arriere des
premiers. Cotes lateraux de la tete faiblement sinues creneles; partie
superieure raboteuse, inegale.

"PronotLun fortement enlargi en demi cercle sur les cotes, le bord
externe avec de petites sinuosites formant des dents subarrondies,
inegales; depassant lateralement d'une fagon tres sensible le niveau
du bord externe des elytres et de I'abdomen; Tangle anterieur dirige
presque droit en avant, plus ou moins saillant, parfois assez aigu,
atteignant le niveau du milieu transversal de I'oeil.

"Ecusson assez bombe avec une petite carene longitudinale
mediane et deux forts tubercules lateraux longitudinaux, un peu
divergents en arriere, garnis d'epines dressees en brosse.

"Elytres finement granuleuses, fortement sinuees au bord externe,
tout pres de la base, cette sinuosite suivie d'un assez fort lobe

410 The University Science Bulletin

aplati, briisquement tronque en arriere au niveau du tiers basal
de I'elytre; subarrondies ensuite sur les deux tiers posterieurs. Con-
nexivum avec tous les segments legerement sinues, surtout les trois
derniers, les angles posterieurs un peu saillants et garnis d'une
rangee de petites soies blanchatres, courtes et raides.

"Tubercule du mesosternum en croix, tres attenue au sommet
qui est legerement arrondi, presque aigu. Femurs jaunatres pales
avec un anneau indecis, legerement brunatre vm peu au dela du
milieu, et I'extreme sommet etroitement noir brunatre; tibias et tarses
brunatres, un peu plus clairs vers leur base et plus fonces, presque
noirs vers leur extremite.

"Piece genitale ^ assez large, faiblement rejetee sur la gauche.
Piece genitale 5 un peu plus large que longue, s'enfongant dans
I'echancrure obtuse du dernier segment abdominal; les deux parties
de la piece genitale subtronquees posterieurement avec leur angle
externe peu saillant, subarrondi.

"La forme tres dilatee du pronotum, le lobe de la marge elytrale
et les sinuosites du connexivum permettront de reconnaitre facile-
ment cette espece."

A discussion of the possible relationship of this species with
Nerthra lobata (Montandon) will be found in the comparative
notes of that species.

Nerthra asiatica (Horvath)

(PI. XIV, fig. 128)

1892. Monontjx asiaticus Horvath, Termeszetrajzi Fuzetek, vol. XV, pt. 3,

p. 136.
1909. M. asiaticus Oshanin, Verzeichnis der Palaearktischen Hemipteren,

vol. I, p. 956.
1912. M. asiaticus Oshanin, Katalog der palaarktischen Hemipteren, p. 89.
1926. M. asiaticus Kirit., "Konowia," Zeitschrift fur systematische Insekten-

kunde unter Mitwirkung fuhrender Entomologen, Vol. 5, No. 3, p. 226.
1930. M. asiaticus Kirit., Annuaire du Musee Zoologique de TAcademie des

Sciences de I'URSS, p. 435.
1933. M. asiaticus Hoffmann, Lingnan Science Journal, vol. 12, Suppl., p. 250.
1935. M. asiaticus Wu, Catalogus Insectorum Sinensium, vol. 2, p. 559.
1941. M. asiaticus Hoffmann, Lingman Science Journal, vol. 20, no. 1, p. 44.

Also referring to this species:

1899. Mononyx grossus Montandon, Bulletin de la Societe des Sciences de

Bucarest-Roumanie, an. VIII, nos. 4 et 5, p. 398.
1906. M. grossus Distant, Fauna British India, vol. Ill, p. 16.

Size. Female: Length, 11.8 mm.; width of pronotum, 8.0 mm.;
width of abdomen, 8.5 mm.

Color. Above, more or less uniformly brown, scutellum slightly
darker than rest of dorsal surface. Below, mostly dark brown, a

The Family Gelastocoridae (Hemiptera) 411

few patches of yellowish-brown about the bases of the middle and
hind legs.

Structural characteristics. Apex of head more or less rounded,
apical tubercle absent, superapical and lateral .tubercles small, ir-
regular in shape, not sharp-pointed. Pronotum widest at transverse
furrow, not quite as wide as abdomen; lateral margins parallel or
nearly so, in median portion, anterior and posterior parts of the
margin distinctly converging mesad; disc elevated and rather
rough; posterior margin trisinuate. Scutellum elevated, apex slightly
lobed, with tumescences at the middle of the lateral margins. Hem-
elytra not extending to the end of the abdomen, membrane well de-
veloped; embolium with the basal half of the lateral margin nearly
straight or very slightly concave, not expanded laterad at middle.
Connexivum greatly expanded laterally, wider than width of pro-
notum or hemelytra. Bristles in rows and clumps on hemelytra and
in clumps on scutellum and pronotum, bristles short or moderately
long, clavate, slightly curved. Abdominal sternites of female nearly
symmetrical, lobes of ovipositor slightly projecting posteriorly; pos-
terior margin of last visible abdominal sternite triangularly emargi-
nate.

Location of type. Unknown. Perhaps in the Museum of Budapest,
although Doctor H. B. Hungerford does not mention it in the
notes he made during his visit to this museum in 1928.

Distributional data. This species has been reported from China,
India and Tibet. The type is from "Flumen Poi-ho," China (G. N.
Potanin). This worker has examined a paratype of Mononyx
grossus Montandon. This specimen is labeled Thibet (Mou-Pin),
1869-1870, (A. David). It was sent in exchange from the Museum
of Paris and is now in the Francis Huntington Snow Entomological
Collection at the University of Kansas.

Comparative notes. This species differs from N. indica (Atkin-
son ) by the shape of the tubercles of the head, by the lateral margin
of the pronotum and by the larger size. In the shape of the pronotum
it would seem to be cosely related to N. spissa (Distant), but the
worker has not seen a female of the latter species or a male of N.
asiatica ( Horvath ) , so no definite comparison can be made between
these two species at the present time.

412 The University Science Bulletin

Nerthra spissa (Distant)

(PI. XI, fig. 101)
1906. Monomjx spissus Distant, Fauna British India, vol. V, p. 313, fig. 176.

Size. Male: Length, 8.5 mm.; width of pronotum, 6.0 mm.;
width of abdomen, 6.2 mm.

Color. Light grayish-brown, scutellum and median portion of
pronotum darker; ventrally darker with faint yellowish-brown
spots on intermediate and hind femora and abdominal segments.

Structural characteristics. Front of head with two moderate
tubercles. Pronotum nearly rectangular, lateral margins mostly
straight, parallel to longitudinal axis of the body, anterior part
roundly converging to the eye; lateral margin posteriorly forming
almost a right angle with the posterior margin; posterior margin
slightly sinuated before scutellum; posterior third of pronotum
crossed by two moderate longitudinal carinations. Scutellum with
moderate tumescences at lateral sides and apex, median portion
depressed. Hemelytra extending beyond end of the abdomen;
embolium entirely dilated. Body covered with long erect bristles
which are moderately clavate. Abdominal sternites of the male
asymmetrical; ninth sternite wider than long, not as long as eighth
sternite. Male clasper rather large for the size of the insect,
nearly straight, cylindrical, abruptly narrowed to point at apex,
twisted, aedeagal furrow obliquely crossing basal half of clasper.

Location of type. In the British Museum at London, England.

Distributional data. Distant described this species from a speci-
men from Sibsagar, Assam, India. The writer has seen a single
male specimen labeled Misamari, Assam, India, June 26, 1943,
D. E. Hardy, (USNM).

Comparative notes. Shape of the pronotum, the long erect
bristles and the shape of the male clasper easily separates this
species from all others.

Nerthra rugosa (Desjardins)

(PI. XIII, fig. 121)

1837. Naucoris rugosa Desjardins, Annals de la Societe Entomologique de
France, vol. 6, p. 239.

1837. N. rugosa Serville, Annals de la Societe Entomologique de France, vol.
7, p. 243.

1840. N. rugosa Westwood, Introduction to the Modem Classification of In-
sects, vol. 2, p. 464.

1843. Peltopterus rugosus (Desjardins); Guerin-Meneville, Revue Zoologique
Travaux Inedits, pp. 112-113.

1863. F. rugosus Stal, Berliner Entomologische Zeitschrift, Band, VII, p. 407.

1865. P. rugosus Stal, Hemiptera Africana, Band. Ill, p. 173.

The Family Gelastocoridae (Hemiptera) 413

1876. P. nigosus Stal, Kongl. Svenska Vetenskaps-Akademiens Handlingar,

Band. 14. No. 4, p. 140.
1900. P. rugosus Montandon, Bulletin de la Socicte des Sciences de Bucarest-

Roumanie, an. VIII, no. 6, p. 9.
1906. P. rugosa Kirkaldy, Trans. Am. Ent. Soc, vol. XXXII, p. 149.
1910. P. rugosa Montandon, Annuario de Museo Zoologico della R. Universita

di Napoli (nuova serie) vol. 3, no. 10, p. 3.

Also referring to this species:

1859. Mononyx rugosus ( Desjardins ) ; Dohm, Catalogus Hemipterorum, Her-

ausgegeben von dem Entomologischen Verein zu Stettin, p. 53.
1925. Glossoospis brunnea Blatchley, Ent. Nevi's, vol. 36, pp. 49-52.

Possibly referring to this species:

1884. Mononijx sttjgicus Uhler, Standard Natural History, vol. II, p. 264, nee

Say.
1902. Nerthra stygica Say; Howard, The Insect Book, pi. XXXIX, fig. 16.

Size. Female: Length, 6.0 to 7.1 mm., width of pronotum, 4.1 to
5.0 mm., width of abdomen, 4.2 to 5.1 mm.

Color. Above; head, disc of pronotum and hemelytra, brown,
lateral expansions of pronotum, apical half of the scutellum and
apical halves of the segments of the connexivum, yellowish-brown,
the lateral portions of the pronotum may have some darker macula-
tions. Below; legs brown; abdomen brown with the caudolateral
portions of the segments paler, yellow or yellowish-brown.

Structural characteristics. Front of head with three rounded ele-
vations, apex smooth, rounded, lacking tubercles; ocelli absent.
Lateral margins of pronotum subparallel for posterior two thirds,
slightly rounded, anterior third roundly converging toward the
eyes; posterior angle rounded, not at all projecting beyond base of
embolium; posterior margin of pronotum trisinuate before the scutel-
lum. Scutellum rather small, apex rather elongate, tongue-shaped,
apex and basal angles slightly elevated, mesobasal portion depressed.
Hemelytra entirely coriaceous, fused together, extending to or be-
yond end of abdomen, smooth, with not a trace of longitudinal
carinae, but with a basal and medial tumescence on each hemely-
tron; embolium greatly expanded laterally at the basal one third,
expansion widest posteriorly and rather abruptly narrowing at that
point. Connexivum visible, variable as to the degree of expansion.
Bristles variable in shape, most of the body with short clavate
bristles, elevations with much longer clavate bristles which are very
slender at the base. Abdominal sternites of the female symmetrical;
last visible abdominal sternite not emarginate, but not covering the
ovipositor, usually slightly but broadly keeled. The trochanters of
the anterior legs have two apical tubercles in all the specimens
examined.

414 The University Science Bulletin

Location of type. In the Museum of Paris, Paris, France.

Distributional data. This small species has without a doubt the
most peculiar distribution of all the members of this family. The
type was described from "Isle of Maurice" (Mauritius), which is in
the Indian Ocean. Blatchley reported it from Florida (as Glosso-
aspis hrunnea ) . The writer has seen specimens from the following
localities :

Florida: Arch Cr., Mar. 21, 1911, W. S. Blatchley, 2 females, one
the type of Glossoaspis hrunnea Blatchley, ( Purdue Coll. ) ; Biscayne,
Apr. 5, 1887, Heidemann, 1 female; Matecumba Key, Mar. 14, 1947,
R. H. Beamer, 1 female.

Panama: San Jose Is., Pearl Is., Mar. 1, 1944, J. P. E. Morrison, 1
female, (USNM).

Comparative notes. A comparison by Doctor Eugene Seguy of
the specimen from Matecumba Key, Florida, has shown that Glosso-
aspis hrunnea Blatchley is a synonym of Peltopterus rugosus ( Desjar-
dins ) . This species may be separated from the other species, which
have the hemelytra entirely coriaceous and fused together and
which also have the ocelli absent, in the following ways: From N.
macrothorax ( Montrouzier ) and N. nervosa (Montandon) by the
absence of longitudinal carinae on the hemelytra. From N. williamsi
n. sp. and N. americana ( Montandon ) by the tubercles of the head
and the shape of the last visible abdominal sternite of the female.
From N. hirsuta n. sp. This species may be separated by the nature
of the edges of the hemelytra.

Nerthra macrothorax (Montrouzier)

(PI. Xll, fig. 113; PI. XIII, fig. 119; PL XIV, fig. 135.)

1855. Galgulus macrothorax Montrouzier, Annales des Sciences Physiques et

Naturelles, d' Agriculture et d'Industrie, Lyon., vol. 2, pp. 110-111.
1863. Peltopterus macrothorax Stal, Berliner Entomologische Zeitschrift, vol.

VII, p. 407.
1870. P. macrothorax Stal, Ofversigt af Kongl. Vetenskaps-Akademiens For-

handlingar, no. 7, p. 706.
1880. P. macrothorax Distant, Trans. Ent. Soc. London, pt. IV, p. 486.
1900. P. macrothorax Montandon, Bulletin de la Societe des Sciences de

Bucarest-Roumanie, an. VIII, no. 6, p. 8.
1906. P. macrothorax Kirkaldy, Trans. Am. Ent. Soc, vol. XXXII, p. 149.

Also referring to this species:

1861. Syclaecus macrothorax (Montrouzier); Stal, Ofversigt af. Kongl.

Vetenskaps-Akademiens Forhandlingar, Arg. 18, p. 201.
1865. S. macrothorax Stal, Hemiptera Africana, III, p. 173.
1876. S. macrothorax Stal, Kongl. Svenska Vetenskaps-Akademiens Hand-

lingar. Band 14, no. 4, p. 139.

Size. Male: Length, 7.9 mm., width of pronotum, 5.9 mm..

The Family Gelastocoridae (Hemiptera) 415

width of abdomen, 6.0 mm. Female: Length, 9.2 to 10.6 mm.,
width of pronotum, 6.8 to 8.2 mm., width of abdomen, 6.7 to 8.2 mm.
Color. Above, hght brown; the basal portions of the segments of
the connexivum, the carinae of the hemelytra and the disc of the
pronotum a little darker. Below, brown with a tinge of red.

Structural characteristics. Front of head provided with five large,
rounded tubercles, four of which are flattened on top and densely
covered with short clavate bristles; ocelli absent. Pronotum
greatly expanded laterally; lateral margins converging anteriorly,
subparallel for posterior one half; posterior angle projecting
obliquely posterolaterad, rather pointed; posterior margin with five
concavities. Scutellum, rather small, apex narrowed, basal portion
depressed, inchning to apex which is most elevated. Hemelytra
entirely coriaceous, fused together, extending slightly beyond end
of abdomen, large longitudinal carinae present; base of embolium
greatly expanded laterad. Connexivum broadly expanded laterally
in both sexes. Entire body covered with short, broadly clavate
bristles, bristles pale and especially dense on pronotum and on the
elevations of the head. Abdominal sternites of female nearly sym-
metrical except for posterior margin of last sternite, which is slightly
emarginated, but with apex slightly convex just below the lobes
of the ovipositor, the latter somewhat rounded and the left one
overlapping the right. Abdominal sternites of male rather small,
last visible abdominal sternite wider than long, 32:17, nearly twice
as long as seventh sternite, which has the right side elongate,
spatulate. Clasper of male rather sickle-shaped, but nearly straight,
very slightly enlarged at apex (apical third) then tapering to a
blunt point.

Location of type: In the Museum of Paris, Paris, France.

Distributional data. This species has been reported from the
following places in the East Indies and the Pacific: Philippines,
Aru Island, N. Borneo, Woodlark Island, Solomon Arch., and the
Mariannes. The writer has examined specimens from the following
localities :

New Guinea: Goodenough I., Aligabay Cr., Aug. 1, 1943, B. E.
Rees, 1 female, (USNM); Maffin Bay, Dutch N. Guinea, July 1,
1944, E. S. Ross, 1 female, (CAS); same place. May 25, 1944,
E. S. Ross, 1 female, (CAS).

Philippines: N. W. Panay, no date. Baker, 1 female, ( USNM ) ;
Biliran Isl., no date or collector, 1 female, ( USNM ) .

416 The University Science Bulletin

Solomons: Florida Island, Mar. 11, 1945, J. R. Stuntz, 1 male.

Comparative notes. The size, shape of the pronotum, longi-
tudinal carinae and clasper of the male will separate this species
from all the others. It is quite closely related to N. nervosa (Mon-
tandon), but differs in the shape of the pronotum.

Nerthra nervosa ( Montandon )

(PI. XIII, ag. 124)

1900. Peltopterus nervosus Montandon, Bulletin de la Societe des Sciences
de Bucarest-Roumanie, an. VIII, no. 6, pp. 8-9.

Size. Female: Length, 8.1 mm., width of pronotum, 5.7 mm.,
width of abdomen, 6.0 mm.

Color. Dark brown to reddish brown, more or less concolorous,
but with the disc of the pronotum, the carinae of the hemelytra,
and the scutellum very slightly darker.

Structural characteristics. Front of head with round, flattened
tubercles which are densely covered with short clavate bristles;
ocelli absent. Lateral margins of the pronotum rounded; pronotum
widest at the middle; disc elevated, lateral expansions nearly flat;
posterior margin of the pronotum nearly straight, not at all con-
cave before scutellum; pronotum less than abdomen in width.
Scutellum rather small, apex narrowed, basal portion depressed,
inclining to apex which is the most elevated portion. Hemelytra
entirely coriaceous, fused together, extending beyond end of ab-
domen, longitudinal carinae very large and conspicuous; base of
embolium greatly expanded laterally, exceeding or equal to
width of pronotum. Connexivum broadly expanded laterally. Bristles
of body short and clavate. Abdominal sternites of female nearly
symmetrical; posterior margin of last visible abdominal sternite
slightly emarginated, last visible sternite not covering the lobes
of the ovipositor.

Location of type. In the Museum of Paris, Paris, France.

Distributional data. Montandon described this species from a
single male from the Mariannes. This worker has studied a female
"Guam, 1895," (USNM), that was determined by Montandon in
1909.

Comparative notes. This species differs from N. macrothorax
(Montrouzier) in the shape of the lateral margin of pronotum being
more rounded, posterior margin nearly straight, smaller size, longi-
tudinal carinae of the hemelytra being somewhat larger and the
emargination of the posterior margin of the last visible abdominal

The Family Gelastocoridae (Hemiptera) 417

sternite smaller. To the knowledge of the writer, Montandon did
not designate this female as the allotype of the species, it may be
that he designated some other specimen as the allotype, although
I can find no reference to it in the literature. For this reason, it
seems best to refrain from designating this specimen as the allo-
type, at least until more material from some of the European
museums can be examined.

Nerthra hirsuta n. sp.

(PI. XIV, fig. 132)

Size. Female: Length, 6.2 mm., width of pronotum, 4.4 mm.,
width of abdomen, 4.4 mm.

Color. Brown to dark reddish-brown above, below, darker, except
trochanters, bases, and apices of femora of the middle and hind
legs, and the posterolateral angles of the ventral abdominal seg-
ments which are yellowish-brown.

Structural characteristics. Head with four rather broad toothlike
tubercles; ocelli, if present, very small and indistinguishable from
the granulation of the head. Lateral margins of pronotum sub-
parallel, but slightly converging anteriorly; anterolateral margin
converging rather abruptly toward eye; posterolateral margin
rounded. Scutellum moderate in size, depressed basally and ele-
vated laterally from middle to apex. Hemelytra parallel or nearly
so for three fifths their length, then converging posteriorly in a
broad semicircle; hemelytra thick, entirely coriaceous, fused to-
gether and with the costal margins from apex of embolium to apex
of hemelytron very distinctly bent up at about a 45 to 60 degree
angle to rest of hemelytra, notched at apex and not bent up. Con-
nexivum not visible. Entire upper surface and front of head pro-
vided with rather large broadly clavate bristles which are mostly
black in color and which are clumped together in spots on the hem-
elytra and on the elevations of the scutellum. The body as a whole
is rather truncate in appearance. Abdominal sternites of the female
similar to those of other species from the Australian region, last
visible abdominal sternite extending posteriorly and completely or
nearly completely covering lobes of ovipositor, broadly keeled and
slightly emarginate at apex.

Location of type. Holotype female. West Australia, Augusta,
Feb., W. S. Brooks, in the Museum of Comparative Zoology at
Harvard University.

Distributional data. As given above for the type.

14—6730

418 The University Science Bulletin

Comparative notes. This species may be separated from all others
with entirely coriaceous and fused hemelytra, by the small size,
by the turned-up edges of the hemelytra, and by the abundance of
broad, clavate bristles on the dorsal surface and the head.

Nerthra laticoUis ( Guerin-Meneville )

(PI. XII, fig. 108; PI. XV, fig. 139)

1843. Monontjx laticoUis Guerin-Meneville, Revue Zoologique Travaux Inedits,

pp. 112-114.
1863. M. laticoUis Stal, Berliner Entomologische Zeitschrift, VII, p. 406.
1899. M. laticoUis Montandon, Bulletin de la Societe des Sciences de Bucarest-

Roumanie, an. VIII, nos. 4 and 5, pp. 407-408.
1910. M. laticoUis Montandon, Annuario del Museo Zoologico della R. Univer-

sita di Napoli (nuova serie), vol. 3, no. 10, p. 2.

Size. Male: Length, 8.4 to 8.5 mm.; width of pronotum, 5.9 to
6.0 mm.; width of abdomen, 5.4 to 5.7 mm. Female: Length, 8.6
to 9.3 mm.; width of pronotum, 5.9 to 6.5 mm.; width of abdomen,
5.5 to 6.2 mm.

Color. Brown to blackish-brown; posterior half of each segment
of the connexivum dull yellowish-brown; ventrally mostly dark
brown; front femora pale yellowish-brown, femora of other legs
slightly darker; posterior half of each abdominal segment near
lateral margins with a dirty yellow spot.

Structural characteristics. Five tubercles on front of head, four
on frons, one on apex, the latter not visible from above, lateral
tubercles of frons smaller than others. Pronotum widest at a point
level with the transverse furrow, distinctly projecting beyond the
base of embolium, slightly wider than abdomen; lateral margin
sinuous, anterior and median portions distinctly concave, posterior
portion straight or weakly concave; posterior margin of pronotum
sinuated before scutellum; posterior one third of pronotiun crossed
by five indistinct longitudinal carinations. Scutellum elevated, a
moderate oval tumescence at each side and a more obscure elongate
median tumescence between them. Hemelytra extending poste-
riorly beyond end of abdomen, nearly covering abdomen completely
in the male, to a lesser extent in the female; embolium elongate,
basal one-third narrow and distinctly concave. Bristles short, cla-
vate, forming five groups on each hemelytron, a group on the lateral
tumescences of the scutellum and one medial group near anterior
margin of pronotum. Mesosternal process moderately pointed at
apex, lateral margins straight. Abdominal sternites of the female
symmetrical, more or less roundly projecting posteriorly, entirely
covering lobes of the ovipositor. Abdominal sternites of male

The Family Gelastocoridae (Hemiptera) 419

asymmetrical, ninth sternite much wider than long, slightly less in
length than eighth sternite, longer than seventh sternite. Male
clasper simple, slightly curved mesad, apical one half distinctly
narrowed, aedeagal furrow visible for a short distance at apex.

Location of type. Formerly in the Zoological Museum of the
University of Naples, Naples, Italy, the location of the type is now
unknown.

Distributional data. Guerin-Meneville in the original description
gives the distribution as New Guinea. Montandon also examined
specimens from that island. The specimens studied by the writer
have all been from New Guinea and are from the following
localities:

"Br. N. Guinea," no date or collector, 1 male and 1 female;
Finschhafen, Apr. 21, 1944, E. S. Ross, 1 female, (CAS); Lae, N. E.
New Guinea, June, 1944, F. E. Skinner, 2 males and 3 females,
(CAS); same place, Nov., 1944, Heifer, 1 male; MaflBn Bay,
"Dutch N. Guinea," Aug. 20, 1944, E. S. Ross, 1 female, (CAS);
same place, June 20, 1944, E. S. Ross, 1 female, (CAS); same
place, Sept. 1944, E. S. Ross, 1 female, (CAS); same place, July 1,
1944, E. S. Ross, 1 male and 1 female, (CAS); same place, Oct. 7,
1944, E. S. Ross, 1 male, (CAS); Nadyab, Markham R. val, June
1944, K. V. Krombein, 1 male and 2 females, (USNM); "Neth.
New Guinea," Dec. 10, 1944, T. Aarons, 2 females, (CAS); "New
Guinea," no date or collector, 3 females.

Comparative notes. Near N. giirneyi n. sp. from which it can be
separated by the sinuous lateral margins of the pronotum and by
the shape of the male clasper.

Nerthra gurneyi n. sp.

(PI. XII, fig. 109)

Size. Male: Length, 9.3 mm.; width of pronotum, 6.3 mm.;
width of abdomen, 6:15 mm. Female: Length, 9.7 mm.; width of
pronotum, 7.0 mm.; width of abdomen, 6.4 mm.

Color. Light to dark brown above; ventrally entirely dark reddish-
brown to blackish-brown.

Structural characteristics. Front of head with two large median
tubercles on the frons, laterad of which are a pair of indistinct
tubercles, apex with a small tubercle, not visible dorsally. Pronotum
widest at a level with the transverse furrow, prominently project-
ing beyond the base of the embolium, wider than the abdomen;
lateral margin more or less rounded, anterior portion slightly con-

420 The University Science Bulletin

cave, converging obliquely toward the eye, medial portion nearly
straight, posterior portion straight or very slightly concave, con-
verging obliquely to base of embolium; posterior margin sinuated
before scutellum; posterior third of pronotum crossed by two
moderate and five faint longitudinal carinations. Scutellum rather
large, elevated, sides medially tumescent, a faint longitudinal tu-
mescence medially. Hemelytra extending beyond end of abdomen
in male, not quite covering ovipositor lobes in the female; con-
nexivum only slightly visible dorsally; embolium narrow basally,
distinctly concave; hemelytra constricted at nodal suture. Groups
of clavate bristles present on pronotum, scutellum and hemelytra.
Apex of mesosternal process moderately pointed, lateral margins
nearly straight. Abdominal sternites of the female symmetrical;
last visible abdominal sternite more or less pointedly projecting
posteriorly, completely covering lobes of ovipositor. Abdominal
sternites of the male asymmetrical; ninth sternite much wider than
long, not quite as long as eighth sternite, longer than seventh ster-
nite. Male clasper simple, curved mesad, slightly swollen before
apex with short, narrow, fingerlike apex.

Location of type. Holotype male, Bougainville I., July 1 to Sept.
15, 1944, A. B. Gurney in the United States National Museum at
Washington, D. C.

Distributional data. As given for the type specimen.

Comparative notes. This species is near N. laticolUs (Guerin-
Meneville ) and N. mixta ( Montandon ) . Separated from the former
by the lateral margin of the pronotum, from the latter by the lesser
concavity of the embolium and from both by the shape of the male
clasper. The writer has examined a female specimen which he
believes to be this species, but since there is not a label on the pin,
it seems best not to make this specimen the allotype or even a
paratype. This specimen is in the Francis Huntington Snow En-
tomological Museum at the University of Kansas via the Torre-
Bueno Collection.

Nerthra mixta (Montandon)

(PI. XII, fig. 116)

1899. Mononi/x mixtus Montandon, Bulletin de la Societe des Sciences de
Bucarest-Roumanie, An. VIII, Nos. 4 and 5, p. 406.

1900. M. mixtus Breddin, Senckenbergischen naturforschenden Gesellschaft,
Band XXV, Heft 1, p. 175.

1901. M. mixtus Breddin, Abhandlungen der naturforschenden Gesellschaft, zu
Halle., XXIV, p. 24.

Size. Male: Length, 9.4 mm.; width of pronotum, 7.7 mm., width

The Family Gelastocoridae (Hemiptera) 421

of abdomen, 6.5 mm. Female, 9.4 to 9.6 mm.; width of pronotum,
7.7 to 7.9 mm.; width of abdomen, 6.5 mm.

Color. Uniformly dark reddish-brown except for front femora
and area about bases of the other legs which are paler.

Structural characteristics. Apex of head provided with five broad
toothlike tubercles, apical tubercle ventrocaudad of the others;
ocelli present. Pronotum extremely wide, much wider than abdo-
men; the median portion of the lateral margins more or less
rounded, anterior and posterior parts of the margin distinctly con-
verging medially; posterior margin concave before base of the
scutellum, convex level with the base of the embolium. Scutellum
moderately large, a tumescence on each side and at the apex. Hem-
elytra extending to or beyond the end of the abdomen; membrane
well developed; base of embolium narrow, apical two thirds broadly
and roundly expanded. Connexivum not at all prominent, scarcely
visible in the male. Bristles dark brown, clavate, forming rows and
clumps on the hemelytra, scutellum and pronotum. Abdominal
sternites of male asymmetrical; last three sternites small, distance
between caudolateral angles of sixth sternite less than half the width
of the posterior margin of the fourth sternite. Clasper of male
constricted at the middle and then again constricted half way be-
tween the middle and the apex, simple and sickle-shaped. Abdomi-
nal sternites of the female nearly symmetrical, projecting posteriorly
and nearly covering lobes of ovipositor, apex of posterior margin
is very slightly emarginate.

Location of type. Unknown. Montandon did not specify which
of the specimens before him was the type. The type of this species
is not in the British Museum with other Montandon types, at least
it was not seen by Doctor H. B. Hungerford in 1928. There are
two males in the Paris Museum, "Nouv. Bretagne, LIX — 1889" which
were before Montandon at the time of the description. These two
specimens may be considered as typical until such time as the type,
if existant, is located.

Distributional data. Montandon studied specimens from Aus-
tralia, New Guinea, New Britain and Amboina. This worker has
seen only three specimens, all of which are from New Guinea. They
are from the following localities: Mt. Hansemann, Astrolabe B.,
1901, Biro, 1 female; Lae, June, 1944, F. E. Skinner, 1 male and 1
female, (CAS).

Comparative notes. Most closely related to Nerthra omani n. sp.
but easily separated by the much wider pronotum and by the fact

422 The University Science Bulletin

that the clasper is twice constricted in this species while in IV. omani
n. sp. this is not the case.

Nerthra omoni n. sp.

(PL XI, fig. 99)

Size. Male: Length, 9.6 to 10.5 mm.; width of pronotum, 7.2
to 7.6 mm.; width of abdomen, 6.8 to 7.6 mm. Female: Length,
10.6 mm.; width of pronotum, 7.7 mm.; width of abdomen, 7.4 mm.

Color. Light brown to black above, the bristles a little darker;
ventrally brown, reddish-brown or black except for some yellow
spots on the mid and hind femora and on the pleurites about these
legs.

Structural characteristics. Apex of head with five large toothlike
tubercles, apical one ventrocaudad of the others; ocelli present.
Pronotum widest at transverse furrow, distinctly projecting beyond
the base of the embolium, usually slightly wider than abdomen;
the median portion of the lateral margins parallel or nearly so,
anterior and posterior parts of the margin distinctly converging
medially; posterior margins of the pronotum concave before the
scutellum then extending in a more or less straight line to the lateral
angle. Scutellum moderately large, a tumescence on each side and
at the apex. Hemelytra extending to or beyond the end of the
abdomen; membrane well developed; base of embolium narrow,
apical two thirds broadly and roundly expanded. Connexivum not
at all prominent, scarcely visible in the males. Bristles dark
brown, clavate, forming rows and clumps on the hemelytra, scu-
tellum and pronotum. Abdominal sternites of male asymmetrical;
last three sternites small, distance between caudolateral angles of
sixth sternite less than half the width of the posterior margin of
the fourth sternite. Clasper of male of nearly equal width for the
entire length, but slightly swollen at apical third and then tapering
to ovate apex which is recurved. Abdominal sternites of the female
nearly symmetrical, projecting posteriorly and nearly covering lobes
of ovipositor.

Location of type. Holotype male, allotype female and 2 male
paratypes, Guadalcanal, 1944, L. J. Lipovsky, in the Francis
Huntington Snow Entomological Collections at the University of
Kansas. Three male paratypes, Guadalcanal, June 15, 1942, Paul
Oman, in the United States National Museum at Washington, D. C.

Distributional data. Known only from Guadalcanal Island in the
Solomon Islands.

The Family Gelastocoridae (Hemiptera) 423

Comparative notes. Similar to Nerthra mixta (Montandon) but
differing in that the pronotum is not nearly so wide in proportion
to the width of the abdomen and in that the clasper of the male
is nearly of uniform width for the entire length while in the latter
species it is strongly constricted at the middle and again half way
to the apex.

Nerthra femoralis (Montandon)

(PI. XII, fig. 105)

1899. Mononyx femoralis Montandon, Bulletin de la Societe des Sciences
de Bucarest-Roumanie, an. VIII, nos. 4 and 5, p. 407.

1902. M. femoralis Horvatli, Temieszetrajzi Fiizetek, XXV, p. 612.

1925. M. femoralis Hale, Arkiv. for Zoologi utgivet av Kongliga Svenska
Vetenskapsakademien, Band 17A, No. 20, p. 16.

Size. Male: Length, 7.8 mm.; width of pronotum, 5.7 mm.; width
of abdomen, 5.7 mm. Female: Length, 8.8 to 10.0 mm.; width of
pronotum, 6.1 to 6.6 mm.; width of abdomen, 6.1 to 6.8 mm.

Color. Yellowish-brown to dark reddish-brown; mostly dark be-
low, trochanters and femora of legs yellow to yellowish-brown
suffused with red on the trochanters and basal halves of the femora,
femora dark at apices.

Structural characteristics. Front of head with five rather prom-
inent tubercles, four on frons and one at the apex, the latter not
visible from a dorsal view. Pronotum widest at middle, slightly
projecting beyond base of embolium, about as wide as abdomen;
lateral margins more or less rounded, anterior and median portions
nearly straight; posterior margin broadly and weakly sinuated
before scutellum; posterior third of pronotum crossed by seven
indistinct longitudinal carinations. Scutellum not very elevated, a
moderate tumescence at each side, mesad of each tumescence a
small oval depression. Hemelytra extending beyond end of ab-
domen in the male, not quite reaching end of abdomen in the
female, entirely covering the abdomen laterally in the male, some
of abdomen exposed laterally in the females; embolium slightly
dilated laterally, lateral margin convex for entire length; hemelytra
constricted at nodal suture. Groups and lines of black clavate
bristles present on pronotum, scutellum and hemelytra. Mesosternal
process apically blunt to moderately pointed, lateral margins nearly
straight. Abdominal sternites of female symmetrical; last visible
abdominal sternite dilated posteriorly, nearly covering lobes of
ovipositor, posterolateral angles slightly and roundly projecting
posteriorly. Abdominal sternites of male asymmetrical, last three
small; ninth sternite wider than long, slightly longer then eighth

424 Thi; University Sciexce Bulletin

sternite, about twice as long as seventh sternite. Male clasper sim-
ple, curved medially, slightly rounded and swollen apically.

Location of type. In the British Museum at London, England.

Distrihutional data. Montandon's specimens were from Champion
Bay, Sidney, Australia. The specimens examined by this writer
are all from Western Australia. They are from the following locali-
ties :

Bridgcton, Nov. 11, Darlington, 2 females, (MCZH); Pember-
ton, Nov. 10, W. M. Wliceler, 1 female, (MCZH); Rottnest Isl.,
Oct. 24, P. J. Darlington, 1 female, (MCZH); Wallcliffe, Margaret
P., Oct., Darlington, 1 female, (MCZHj; Wongong R., Perth, Jan.,
VV. S. Brooks, 1 male, (MCZHj.

Comparative notes. Most closely related to N. gurneyi n. sp. but
differing in that the clasper of the male is stouter and by the fact
that the lateral margin of embolium is not concave basally and that
there is a suffusion of red on the trochanters and basal halves
of the femora.

It should be pointed out that in the original description Mon-
tandon states that the front of the head has only four toothlike
tubercles. It is possible that he did not notice the apical tubercle
which is not visible from above or that he did not consider it to
be on the front of the head, or, this species may only have four
tubercles on the head. Hale, 1925, also suggests that N. femoralis
( Montandon ) has five tubercles on the head. A comparison with
type and with my drawing of the male clasper of one of the
above specimens has been made by Doctor R. L. Usinger. He
found them to be nearly identical; however, he indicated a slight
difference in the apex of the clasper which may possibly have
been due to a difference in the plane in which the clasper was
studied. Unfortunately I did not have him check the tubercles of
the head. If the type of N. femoralis (Montandon) does have but
four toothlike tubercles on the head there is the possibility that the
specimens herein described are not this species but are N. annulipes
(Ilorvath). It will be necessary to check also the genitalia of
the type of the latter species if it is a male. Horvath's
description would seem to indicate that he had only a single female
specimen before him. For the present I prefer to place the speci-
mens studied as N. femoralis. The original description of N. an-
nulipes (Ilorvath) will be presented further on in this paper.

The Family Gelastocoridae i^Hemutera) 425

Nerthra nudata n. sp.

I PI. XI, Bv:. 100^

Size. Miile: Length, 9.2 to 9.6 mm.; width of pronotiim, 6.0 to 6.1
mm.; width of abdomen, 6.2 to 6.5 mm.

Color. Light browni to bh\ck; bases of legs reddish, apices of
femora bhick, rest of legs yellowish-brown; abdominal segments
coppery or reddish-brown except last three segments which are
browii.

Structural characteristics. Front of head withont prominent tnber-
cles, a faint medial tubercle of frons present in one specimen. Pro-
notimi widest at a level of the transverse furrow, biureh- projecting
later;ill\- beyond base of emboliinn, not quite as wide as abdomen;
lateral margin with medial and anterior portions weakh- concave;
posterior miirgin slightly sinuated before scutellum; posterior third
of pronotum crossed b>- three broad indistinct longitudinal carina-
tions. Scutellmn ele\ated, tumescences at sides and apex, weakh-
depressed medially. Hemehtra extending be>ond end of abdo-
men, entirely covering abdomen; embolium elongate, lateral margin
smoothly convex; hemehtra hardly constricted at nodal suture.
Five patches of short cla\ate bristles on each hemelytron and a
patch on each basal timiescence of the scutellum. Mesosternal
process short, lateral margins rather broad and straight, apex very
blunt, broadly flattened. Abdominal steniites of male as>mmetrical;
last three sternites small; ninth sternite rather oval, longer tlian
seventh or eighth sternites, slightK" wider than long. Male clasper
simple, sickle-shaped; aedeagal furrow visible from a ventral view
for a short distance laterally at apex, also for a short distance me-
dialK- at basal fourth.

Location of type. Holot\-pe male and two male parat\-pes, Bris-
bane, Q., Australia, March 3, 1932, P. J. Darlington, in the Museum
of Comparative Zoolog>- at Harvard Universit>-. One male parat>pe,
McPherson Range, Nat. Park, Q.. Austr;ilia, Nhirch 16, 1932, P. J.
Darlington, in the Francis Huntington Snow Entomological Mu-
seum at the University of Kansas.

Distributional data. As given for the t^-pe series.
Comparative notes. Easily recognized by the absence of tubercles
on the front of the head and by the shape of the clasper.

426 The University Science Bulletin

Nerthra annulipes (Horvath)

1902. Mononyx annulipes Horvath, Termeszetrajzi Fuzetek, vol. XXV, pp.

611-612.
1902. M. annulipes Froggatt, Agricultural Gazette of N. S. Wales, Misc. Publ.

no. 538, p. 7, pi. II, fig. 25.
1943. M. annulipes McKeown, Australian Museum Magazine, vol. 8, no. 5,

pp. 176-180.

The writer has not seen this species, or at least has not been able
to recognize it.

Original description: "Beviter ovalis, supra sordide griseus; capite
antice quinque-tuberculato; pronoto longitudine sua duplo et di-
midio latiore, abdomini aequilato, disco rugis longitudinalibus ir-
regularibus praedito, marginibus lateralibus dilatatis parce subtili-
terque nigro-granulatis, subrectis, parallelis, tantum apice subito
introsum curvatis; scutello tuberculis tribus nigrogranulatis, duobus
subbasalibus majoribus, uno anteapicali minore, instructo; hemelytris
apicem abdominis subattingentibus et margines laterales abdominis
pone m e d i u m anguste detectos relinquentibus, margine costali
sensim arcuato, hand sinuato, corio venis elevatis nigro-granulatis
instructo et praesertim prope marginem costalem parce subtiliterque
nigro-granulato, membrana rudimentaria, sed distincte valvante,
intus fusco-limbato; corpore subtus fusco-nigro, luteo-variegato;
ventre minute granulato; tuberculo mesosterni in processum longi-
sculum, antrorsum curvatum producto; pedibus luteis, maculis fe-
morum anticorum, annulis duobus femorum posteriorum et tibiarum
intermediarum, tibiis posticis (basi excepta) tarsisque fusco-nigris.
" 5 Segmento ultimo ventrali apice rotundato-truncato, segmenta
genitalia tegente, marginibus lateralibus subrectis. Long. 7.75, Lat.
5.25 mill.

"Clarence River.

"By the rudimentary membrane allied to M. femoralis Montd.,
from which it may be separated by the smaller size, the head having
five tubercles in front, the mesosternal tubercle curved forward
and the dark annulated legs."

The possible relationship of this species to N. femoralis ( Montan-
don) has been discussed in the comparative notes of that species.

Nerthra ampUata (Montandon)

(PI. XII, fig. Ill)

1899. Mononyx ampliatus Montandon, Bulletin de la Societe des Sciences

de Bucarest-Roumanie, an. VIII, nos. 4 and 5, p. 404.
1906. M. ampliatus Distant, Fauna British India, vol. V, p. 313.

Size. Male: Length, 9.5 to 10.4 mm.; width of pronotum, 6.9
to 7.4 mm.; width of abdomen, 6.8 to 7.3 mm. Female: Length,

The Family Gelastocoridae (Hemiptera) 427

9.4 to 11.1 mm.; width of pronotum, 6.5 to 7.8 mm.; width of
abdomen, 6.6 to 7.8 mm.

Color. Brown above, ventrally mostly dark brown, a small
yellowish-brown area near apex of hind femora.

Structural characteristics. Front of head provided with five tu-
bercles, four on frons and one on apex, the latter not visible from
above. Pronotum widest at middle, distinctly projecting laterally
beyond the base of embolium; lateral margins more or less rounded;
posterior margin broadly but shallowly sinuated before scutellum;
posterior third of pronotum crossed by three slight and four very
faint longitudinal carinations. Scutellum elevated, tumescent at
middle of lateral margins and along median line. Hemelytra ex-
tending beyond end of abdomen and laterally almost completely
covering abdomen; lateral margin of embolium slightly concave
at basal half, posterior part dilated; hemelytra constricted at nodal
suture. Abdominal sternites of male asvmmetrical; ninth sternite
more or less oval, as long as eighth sternite, longer than seventh,
wider than long. Male clasper swollen at apex, indistinctly lobed
laterally, concavely depressed basad of the broadest part of the
dilation.

Location of type. In the British Museum at London, England.

Distributional data. This species is known only from New
Guinea. This worker has examined specimens from the following
localities:

Orazay, Dec, 1945, Heifer, 1 male; Slephansort, Astrolabe Bay,
1900, Biro, 1 female; Aligaba Crk., May 20, 1943, B. E. Rees, 1
female, (USNM); Laloki, Apr., 1909, F. Muir, 2 males and 7
females, (CAS); "N. Guinea," no date, B. P. Clark, 1 female,
(USNM); "N. Guinea," no date or collector, 1 female.

Comparative notes. This species is similar to Nerthra rohusta
n. sp. and Nerthra macrostyla n. sp., but differing in that it is a
little smaller and in the shape of the male genitalia. The clasper
of the male is much smaller than that of either of the species men-
tioned above. It also differs from N. rohusta n. sp. in that the
dilation of the apical half of the clasper is not angulate laterally
but is lobed. In N. macrostyla n. sp. this portion of the clasper is
also lobed, but more distinctly so, and the lobe is constricted at
the base while this is not true in the case of IV. ampliata
(Montandon).

428 The University Science Bulletin

Nerthra macrosttjia n. sp.

(PI. XI, eg. 103)

Size. Male: Length, 11.5 mm.; width of pronotum, 7.5 mm.;
width of abdomen, 7.5 mm. Female: Length, 11.3 to 11.7 mm.;
width of pronotum, 7.2 to 7.6 mm.; width of abdomen, 6.9 to 7.6
mm.

Color. Dark reddish-brown, embolium lighter with distinct, ir-
regular black spots; mostly dark brown below, yellowish-brown
area near the apex of the intermediate and hind femora and small
yellowish-brown spots at the posterior lateral angles of each ab-
dominal segment.

Structural characteristics. Front of head provided with five
rather large sharply pointed tubercles four on frons and one at
apex, the latter not visible from a dorsal view. Pronotum widest
at middle, distinctly projecting laterally beyond the base of the
embolium; lateral margins more or less rounded, median portion
nearly straight parallel to longitudinal axis of the body, anterior
and posterior portions converging obliquely to the eye and base
of embolium respectively; posterior margin sinuated before scutel-
lum; posterior third of pronotum crossed by three slight and six
very indistinct longitudinal carinations. Scutellum rather large,
elevated, tumescent laterally and along median line. Hemelytra
extending beyond end of abdomen, completely covering abdomen
laterally; embolium narrow and concave at base dilated behind
concavity; hemelytra slightly constricted at nodal suture. Groups
of blackish-brown clavate bristles on the pronotum, scutellum and
hemelytra. Abdominal sternites of male asymmetrical; ninth sternite
wider than long, not as long as eighth sternite but longer than
seventh sternite. Male clasper very large, swollen apically, a large
lobe on the lateral surface, concavely depressed basad of the lobe.

Location of type. Holotype male and allotype female, Bougain-
ville I., July 1 to Sept. 15, 1944, A. B. Gurney, and 3 female para-
types, Bougainville I., June 5, 1955, A. B. Gurney, in the United
States National Museum at Washington, D. C. One female para-
type. Treasury I., Solomon Is., Aug. 14, 1944, J. H. Faullus, in the
Francis Huntington Snow Entomological Collection at the Univer-
sity of Kansas. One female paratype. Naval Air Base, Bougainville,
Solomon Is., Apr. 1945, G. E. Bohart, in the Collection of the
California Academy of Science, at San Francisco, California.

Distributional data. As given for the type series. This species
appears to be restricted to the Solomon Islands, however, too few

The Family Gelastocoridae (Hemiptera) 429

specimens are known to definitely state that this species has such
a Hmited distribution.

Comparative notes. Very closely related to IV. ampliata (Mon-
tandon) and N. rohusta n. sp., but easily separated in the males
by the shape of the clasper. The females are difficult to distinguish
from the other species, but seem to be more slender. As the females
are so difficult to identify, one must rely upon distributional data,
size, and association with the males.

Nerthra rohusta n. sp.

(PI. XI, fig. 102; PI. XV, fig. 140)

Size. Male: Length, 11.2 to 12.5 mm.; width of pronotum, 7.6
to 8.1 mm.; width of abdomen, 7.6 to 8.0 mm. Female: Length,
11.7 to 12.5 mm.; width of pronotum, 8.3 to 8.4 mm.; width of
abdomen, 7.9 to 8.2 mm.

Color. Light brown above, scutellum darker, lateral margin of
the embolium with faint yellowish-brown spots; below mostly
dark brown, a light spot before the apex of femur of intermediate
and hind legs.

Structural characteristics. Front of head provided with five large,
rather pointed tubercles, four on frons and one at apex, the latter
not visible from a dorsal view. Pronotum widest at the middle, dis-
tinctly projecting beyond base of embolium; lateral margins more
or less rounded, medial portion sometimes straight, anterior and
posterior portions converging obliquely to the eye and base of
embolium respectively; posterior margin broadly sinuated before
scutellum; posterior third of pronotum crossed by seven indistinct
longitudinal carinations. Scutellum elevated, tumescent laterally
and along median line. Hemelytra extending to or beyond end of
abdomen, laterally in the male completely covering abdomen; em-
bolium narrow at base, lateral margin at base distinctly concave,
dilated laterally on apical half; hemelytra slightly constricted at
nodal suture. Groups of black clavate bristles on the pronotum,
scutellum and hemelytra. Mesosternal process apically moderately
pointed, lateral margins nearly straight, slightly rounded at apex
of process. Abdominal sternites of male asymmetrical; ninth
sternite wider than long, about same length as eighth sternite, longer
than seventh sternite. Male clasper greatly swollen apically, lateral
margin of dilation forming almost a right angle at widest point.

Location of type. Holotype male, allotype female, 3 male and 2
female paratypes, Nadzab, Markham R. val., New Guinea, June,

430 The University Science Bulletin

1944, K. V. Krombein in the United States National Museum at
Washington, D. C. One male paratype, same data as above and
one female paratype, same place, July, 1944, same collector, in the
Francis Huntington Snow Entomological Collections at the Uni-
versity of Kansas.

Distributional data. As given above for the type series.

Comparative notes. Near Nerthra ampliata (Montandon) and
Nerthra macrostyla n. sp., but differing from these in that the dila-
tion of the apical half of the clasper of the male is distinctly angulate
laterally while in the two species mentioned above the lateral
margin of the dilation is more or less lobed.

Nerthra luteovaria (Distant)

(PI. Xll, fig. 107; PL XV, fig. 136)

1904. Mononyx luteovarius Distant, Annals and Magazine of Natural History,

ser. 7, vol. XIV, p. 63.
1925. M. luteovarius Hale, Arkiv. for Zoologi utgivet av Kongliga Svenska

Vetenskaps-Akademien, Band 17a, No. 20, p. 15.

Size. Male: Length, 7.8 mm.; width of pronotum, 5.7 mm.;
width of abdomen, 5.5 mm. Female: Length, 7.8 to 8.5 mm.; width
of pronotum, 6.0 to 6.4 mm.; width of abdomen, 5.8 to 6.0 mm.

Color. Uniformly bright yellowish-brown above; below mostly
yellowish-brown, tarsal claws dark shiny brown, abdomen, basally
and medially, brown.

Structural characteristics. Five sharp-pointed tubercles on front
of head, four on frons and one on the apex of the head, the latter
not visible from above. Pronotum widest at the middle, distinctly
projecting beyond base of embolium, slightly wider than abdomen;
lateral margins more or less rounded, anterior portion very slightly
concave; posterior margin sinuated before scutellum; posterior
third of pronotum crossed by two moderate and three indistinct
longitudinal carinations. Scutellum elevated, a tumescence at each
side. Hemelytra extending posteriorly beyond end of abdomen,
nearly entirely covering abdomen in male, connexivum more ap-
parent in female; embolium shghtly dilated, lateral margin more
or less rounded, very slightly concave at basal third. Mesosternal
process with lateral margins more or less rounded, apex blunt.
Abdominal sternites of the female symmetrical, last visible abdomi-
nal sternite broadly expanded posteriorly, covering lobes of ovi-
positor. Abdominal sternites of the male asymmetrical, ninth
sternite about as long as eighth, wider than long. Male clasper
short and broad, apex distinctly narrowed and bent laterally.

The Family Gelastocoridae (Hemiptera) 431

Location of type. In the British Museum at London, England.

Distributional data. The specimens examined by the writer were
from Kinberley, Dist., N. V., Australia, Jan., Mjoberg, 1 male and
2 females.

Comparative notes. A small species, characterized by the shape
of the mesosternal process and the clasper of the male.

Nerthra tuberculata (Montandon)

(PI. XII, fig. 104; PL XV, fig. 137)

1899. Mononyx tuberculatus Montandon, Bulletin de la Societe des Sciences
de Bucarest-Roumanie, An. VIII, Nos. 4 and 5, p. 403.

Size. Male: Length, 8.5 to 8.9 mm.; width of pronotum, 6.1 to
6.4 mm.; width of abdomen, 6.0 to 6.3 mm. Female: Length, 9.2
to 9.9 mm.; width of pronotum, and abdomen, 6.5 to 7.0 mm.

Color. Black, covered with minute white granules which give
the insect a gray appearance; front of head with white tubercles
forming an inverted V; anterior edge of front trochantur and femur
whitish-yellow; apex of femur, tibia and tarsus of intermediate and
hind legs yellowish-brown; rest of ventral surface dark brown to
black.

Structural characteristics. Front of head with five large, sharply
pointed tubercles, four on the frons and one on the apex, the latter
not visible from a dorsal view. Pronotum projecting beyond base
of embolium, about as wide as abdomen; lateral margin more or
less rounded; transverse furrow crossed by two prominent longi-
tudinal carinations; posterior margin deeply sinuated before scu-
tellum. Scutellum elevated, timiescent laterally and at apex.
Hemelytra extending beyond end of abdomen, overlapping in such
a manner as to give the posterior end of the body a pointed effect;
embolium narrow at base, quite wide at nodal suture, lateral margin
nearly straight, rounded posteriorly; hemelytra distinctly con-
stricted at nodal suture. Abdomen almost entirely covered by the
hemelytra. Body covered with dark shiny reddish-black globoid
tubercles. Mesosternal process with lateral margins more or less
straight, apex moderately blunt. Abdominal sternites of male
asymmetrical; last three sternites narrow; ninth about same length
as eighth, longer than seventh, wider than long. Male clasper
swollen ventrally near apex; in ventral view, lateral and mesal
margins wedge-shaped apically.

Location of type. In the Museum of Brussels, Brussels, Belgium.

432 The University Science Bulletin

Distributional data. The type was from Port Denison, Australia.
The writer has seen specimens from the following localities:

Soutliwestern Australia: Margaret R., Oct. 31, P. J. Darlington, 2
males and 2 females, ( MCZH ) .

Western Australia: Pemberton, W. A., Nov. 10, P. J. Darlington,
1 male, (MCZH); Worgong R., Perth, W. Austr., Jan., W. S. Brooks,
1 female, (MCZH).

Comparative notes. The white tubercles on the front of the head,
shiny reddish-black globoid tubercles on the body, hemelytra
pointedly projecting posteriorly and the male clasper distinguish
this species from others.

Nerthra alaticollis (Stal)

(PI. XII, fig. 114; PI. Xlll, fig. 118; PL XIV, fig. 127)

1854. Mononyx alaticollis Stal, Ofversigt af Kongl. Vetenskaps-Akademiens
Forhandlingar, arg. 11, p. 239.

1859. M. alaticollis Dohrn, Catalogus Hemipterorum, Herausgegeben von dem
Entomologisehen Verein zu Stettin, p. 54.

1861. Matiniis alaticollis (Stal); Stal, Ofversigt af Kongl. Vetenskaps-Aka-
demiens Forhandlingar, arg. 18, p. 201.

1863. Af. alaticollis Stal, Berliner Entomologische Zeitschrift, VII, p. 407.

1876. M. alaticollis Stal, Kongl. Svenska Vetenskaps-Akademiens Handlingar,
Band 14, No. 4, p. 139.

1900. M. alaticollis Montandon, Bulletin de la Societe des Sciences de Buca-
rest-Roumanie, an. VIII, no. 6, pp. 4-5.

1906. M. alaticollis Kirkaldy, Trans. Am. Ent. Soc, vol. XXXII, p. 149.

Size. Male: Length, 6.7 to 7.3 mm., width of pronotum, 5.0 to
5.5 mm., width of abdomen, 5.0 to 5.6 mm. Female: Length, 7.4
to 8.5 mm., width of pronotum, 5.5 to 6.4 mm., width of abdomen,
5.2 to 6.3 mm.

Color. Quite variable, but in general, grey to light brown in
over-all appearance. Ground color runs from light yellow to black.
Disc of pronotum, scutellum, and head usually darker than rest of
body; in some these parts are black while the hemelytra and lateral
expansions of the pronotum are light yellow. Front of head may
have apical half orange or white or may be entirely black. Anterior
margin of front femora, bases and apices of middle and hind fe-
mora may be almost white. The ventral abdominal segments gen-
erally dark reddish-brown with the spiracles and area immediately
surrounding the spiracles lighter. The lateral margins of the em-
bolium and the pronotum with alternating dark brown and yellow
spots in most specimens.

Structural characteristics. Head with five rather small, sharp-
pointed tubercles, apical tubercle not visible from above; front rather
flat; ocelli present. Pronotum widest at the middle; lateral margin

The Family Gelastocoridae (Hemiptera) 433

more or less rounded, median half nearly straight in some speci-
mens; disc elevated and pitted with many small, irregular depres-
sions; posterior margin trisinuate. Scntellum slightly elevated;
small to moderate lateral tubercle at middle of each side, a small
pitlike depression postero-mesad of these; apex may or may not be
elevated. Hemelytra usually extending to or beyond end of abdo-
men; bluntly pointed apically; membrane absent, hemelytra entirely
coriaceous, not fused; embolium extremely expanded apically. Con-
nexivum variable in degree of expansion, broadly expanded in some,
not so much so in others. Hemelytra and scutellum nearly glabrous,
lacking black, clavate bristles. Abdominal sternites of female nearly
symmetrical; projecting posteriorly and almost completely covering
the lobes of ovipositor; posterior margin of last visible sternite pro-
jecting posteriorly laterad of apex which is slightly concave. Ab-
dominal sternites of male with last three segments small, less than
one-half the width of the posterior margin of the fourth sternite;
ninth sternite oval, slightly wider than long, longer than eighth
sternite; eighth longer than seventh. Clasper of male sickle-shaped,
slightly swollen basally, apex slender, slightly recurved.

Location of type. In the Museum of Stockholm, Stockholm,
Sweden.

Distributional data. This species is known only from Australia.
The writer has seen specimens from the following localities:

Brisbane, June 11, 1915, Bridwell, 1 male and 1 female, (USNM);
same place, no date, H. Hacker, 1 female; Caloundra, Oct. 10, 1912,
no collector, 1 female; Leura, N. S. Wales, no date, W. M. Mann, 1
female, (MCZH); Megalong Vy., Blue Mts., N. S. Wales, Jan. 20,
1932, Darlington, 1 female, (MCZH); Moreton Bay, Stradbrokel,
Queensland, Sept. 20, 1915, J. C. Bridwell, 1 female, (USNM); N. S.
Wales, no date, Hy. Edwards Collection, 1 female; same place, no
date, same collector, 1 female, (AMNH); Weathcote, N. S. Wales,
Dec. 7, 1914, C. T. Brues, 1 female, (MCZH); Wentworth Falls,
N. S. Wales, Dec. 29, 1931, W. M. Wheeler, 1 male, (MCZH).

Comparative notes. This species is most closely related to Nerthra
adspera (Stal) and N. stali (Montandon). It may be separated
from both by the shape of the lateral margin of the pronotum.

434 The University Science Bulletin

Nerthra adspersa (Stal)

(PI. Xlll, fig. 122)

1863. Matinus adsperstis Stal, Berliner Entomologische Zeitschrift, VII, p. 407.
1876. M. adspersus Stal, Kongl. Svenska Vetenskaps-Akademiens HandUngar,

Band 14, No. 4, p. 139.
1900. M. adspersus Montandon, Bulletin de la Societe des Sciences de

Bucarest-Roumanie, An. VIII, no. 6, p. 5.

Size. Length, 6.0 mm., width of pronotum, 5.2 mm., width of
abdomen, 5.2 mm. This specimen has had the posterior portion of
abdomen eaten by dermestids, and this makes the determination
of sex impossible, in view of present knowledge.

Color. Pale reddish-brown, almost pink. Scutellum and macula-
tions on margins of hemelytra and pronotum darker, the former
wi*"h a reddish cast, the latter brown. Lateral tubercles and apex
of head nearly white. Below, with abdominal segments dark brown
with large light yellow areas along the lateral margin of each
segment. Front femora dark reddish-brown, tibiae lighter.

Structural characteristics. Head lacking sharp-pointed tubercles,
a large blunt tubercle on the lateral margin immediately below
the inner margin of the eye; front quite flat; vertex nearly trans-
verse, scarcely convex; apex of head depressed, base of depression
wider than distance between ocelli. Pronotum greatly expanded
laterally, widest at a point just an*^erior of the humeral angle;
lateral margin with posterior two-thirds subparallel, gradually con-
verging anteriorly, anterior one third abruptly converging toward
the eye; posterior margin trisinuate. Scutellum weakly elevated,
lacking prominent elevations; a shallow depression present at the
middle on either side of the median line. Hemelytra very broad,
completely covering abdomen, membrane apparently lacking, but
as this portion is slightly damaged also it is not possible to say
definitely; embolium greatly expanded laterally.

Location of type. In the Museum of Stockholm, Stockholm,
Sweden.

Distributional data. Stal described this species from specimens
from Western Australia. The specimen studied by this worker is
also from Western Australia, Kirkaldy Collection, which is now
owned by the University of Kansas.

Comparative notes. This species is closely related to N. alaticollis
(Stal) and N. stali (Montandon). It can be separated from the
former by the shape of the pronotum and from the latter by the
large lateral tubercles of the head and by the fact that the base

The Family Gelastocoridae (Hemiptera) 435

of the depression of the apex of the head is wider than the distance
between ocelh. Montandon examined one of the types and mentions
that the tubercles of the head are quite visible, especially the
lateral tubercles which are located below each eye. The length
of the type was also sHghtly smaller than the specimen before the
writer, measuring 5.5 mm.

Nerthra stali (Montandon)

(PL XIII, fig. 125; PI. XIV, fig. 133)

1900. Matinus stali Montandon, Bulletin de la Societe des Sciences de
Bucarest-Roumanie, An. VIII, No. 6, pp. 5-6.

Size. Female: Length, 7.7 to 8.1 mm., width of pronotum, 6.8
to 7.0 mm., width of abdomen 6.9 to 7.3 mm.*

Color. Superficially appearing greyish-brown, ground color actu-
ally yellowish-brown and this is maculated with reddish-brown;
head, disc of pronotum and connexivum slightly darker than rest
of dorsum. Below; abdomen dark reddish-brown, except caudo-
lateral portions of each segment which are yellow or yellowish-
brown; legs brown with the trochanters, bases and apices of femora,
and bases of tibiae, yellow.

Structural characteristics. Head with five small tooth-like tuber-
cles, apical one is ventrad and caudad of the others which are
on anterior margins of head; front of head rather flat; ocelli present.
Pronotum broadly expanded laterally; disk elevated, pitted; lateral
margins of pronotum nearly straight for posterior two thirds, but
converging anteriorly, anterior third more abruptly converging to-
ward the eye; posterior angle projecting beyond the base of the
embolium, nearly forming a right angle; posterior margin of pro-
notum almost straight, very slightly concave before scutellum and
a small projection even with the base of the claval suture. Scutel-
lum slightly elevated, especially near the middle of the lateral
margins and at the apex. Hemelytra entirely coriaceous, not fused
together, tapering apically to a blunt point, not quite extending
to the end of the abdomen; embolium rather expanded laterad for
its entire length. Connexivum broadly expanded, widest part of
insect, but only slightly wider than hemelytra and pronotum. Ab-
dominal sternites of female symmetrical; last visible abdominal
sternite projecting posteriorly, nearly covering lobes of ovipositor,
not emarginate.

* A male has subsequently been examined by the writer. Since he was unable to in-
clude an illustration of the genetalia in this paper, the specimen will be discussed in a
separate paper.

436 The University Science Bulletin

Location of type. In the Museum of Stockholm, Stockhohn,
Sweden.

Distributional data. Montandon described this species from a
female specimen from occidental Australia. The worker has studied
the following specimens also from Western Australia:

Augusta, Febr., W. S. Brooks, 2 females, (MCZH); "Austral.
Occident.," no date or collector, 1 female, (Museum of Stockholm).

Comparative notes. This species is most closely related to N.
adspersus (Stal) and N. aJaticoUis (Stal). It may be separated
from the former by the number and type of tooth-like tubercles on
the head, by the pitting of the discs of the pronotum and by the
shape of the base of the depression of the apex of the head. From
the latter it may be separated by the shape of the lateral margin
of the pronotum.

Nerthra elongata ( Montandon )

1900. Matinus elongatus Montandon, Bulletin de la Societe des Sciences de
Bucarest-Roumanie, An. VIII, No. 6, p. 7.

The writer has not seen any specimens which agree with Montan-
don's description of this species. Original description:

"Forme allongee, teinte brunatre obscure assez uniforme sur tout
le corps. Tete ruguleuse, subarrondie en avant, sans tubercules
dentiformes apparents ni sur le bord externe, ni sur la surface.

Pronotum a cotes lateraux peu dilates, subparalleles sur les deux
tiers postcrieurs, assez largement arrondis antcrieurement. Tangle
posterieur etroitement arrondi, bord posterieur presque droit, tres
obtusement subsinue devant I'ecusson et tres faiblement oblique
lateralement de chaque cote. Rides et sillons du pronotum peu
accentues, tres obtus.

Ecusson a peine un peu plus long que large.

Elytres tres obtusement arrondies au bord externe, sans aucune
sinuosite sur la marge, recouvrant entierement le connexivum et
depassant faiblement I'extremite de I'abdomen; membrane a peu
pres nulle, presque indistincte, a peine valvante.

Dessous du corps assez uniformement brunatre, plus clair sur la
poitrine pres des hanches. Pattes brunatres, plus claires avec trois
grandes taches flaves mal limitees sur la partie inferieure des femurs
intermediaires et posterieurs, la partie superieure de ces femurs
regardant le corps au repos, presque entierement flave. Les femurs
anterieurs assez dilates, bruns en dessous, presque entierement
flaves a leur partie superieure.

The Family Gelastocoridae (Hemiptera) 437

Tubercule du mesosternum pas tres eleve, etroit des la base et
obtus au sommet.

Dernier segment de I'abdomen $ largement tronque a rextremite,
laissant apercevoir une tres faible portion des pieces genitales. Avant
dernier segment abdominal non ou imperceptiblement re treci au
milieu Australie (Fischer 1878) exemplaire unique. K. K. Hof
Museum Vienne."

Neiihra tasmaniensis n. sp.

(PI. XV, fig. 143)

Size. Male: Length, 6.3 mm.; width of pronotum, 4.4 mm.; widtli
of abdomen, 4.4 mm.

Color: Light brown above covered with many white granula-
tions. The white granules combine with white pigment to form
some fairly large patches of white on the hemelytra. Connexivum
with the basal halves of the segments dark brown and the apical
halves orangish-brown. Apical half of the head much lighter than
the basal portion, the extreme apex is a very pale yellow. On the
underside the body is dark brown except for a small yellow spot at
the posterocaudal angles of each abdominal segment. Basal seg-
ments of forelegs pale yellow to just before apex of femora, dorsal
surface of femora tinged with pink and the apex reddish-brown;
tibio-tarsi are reddish-brown and the tarsal claw is black. Inter-
mediate and hind legs with the basal half of the femora dull brown
followed by a pale yellow area and with a reddish apex; tibiae, tarsi
and tarsal claws reddish-brown.

Structural characteristics. Apex of head depressed, width of base
of depression less than distance between ocelli; head with five small
toothlike tubercles, apical one is ventrad and caudad of the others
which are on the anterior margins of head; front of head rather flat;
ocelli present, but very small. Pronotum broadly expanded laterally;
disk elevated, pitted; lateral margins of pronotum with anterior two
fifths straight, strongly converging toward the eyes, posterior two
fifths with the two sides subparallel, slightly convergent anteriorly,
slightly concave; humeral angle rounded; posterior margin nearly
straight, very slightly convex; pronotum widest level with the trans-
verse furrow, but only slightly exceeding in width the body at the
base of the embolia. Scutellum apparently not noticeably elevated,
but it is very badly damaged by the pin and accurate description is
not possible. Hemelytra extending to end of abdomen, entirely
coriaceous, membrane absent, not fused; embolia greatly expanded
basally, lateral margins nearly straight; edge of hemelytra con-

438 The University Science Bulletin

stricted at embolial suture. Connexivum barely visible, parts of but
two segments appearing from above. The last three ventral abdomi-
nal sternites of the male small, less than one-half the width of the
posterior margin of the fourth sternite; ninth sternite oval, slightly
wider than long, longer than eighth sternite; eighth longer than
seventh. Clasper (apical half) simple, tapering rapidly to rather
slender apex, very slightly sickle-shaped nearly straight.

Location of type. Unique, holotype male, L. St. Clair Res., Tas-
mania, October, 1939, D. M. in the British Museum, London, Eng-
land.

Distributional data. As given for the type.

Comparative notes. This species is very closely related to N. alati-
coUis ( Stal ) . The clasper of the male is very similar to the clasper
of the above-mentioned species. The pronotum is quite diflFerent
and the specimen is from a locality from which I have seen no
other material. It may be that further material from Tasmania will
show that the specimen before the writer is abberant, but until
such time as more specimens are available, the pronotal differences
seem sufficient for specific designation.

Nerthra suberosa (Erichson)
1842. Mononyx suberosus Erichson, Archiv fur Naturgeschichte, 8, p. 285.

It is possible that this is the species previously described as
Nerthra tasmaniensis n. sp. since both specimens were from Tas-
mania. The description does not agree with latter species in re-
spect to size (3| lin. is equal to 7.8 mm.) or in regard to the shape
of the pronotum. Since the writer has been unable to recognize
this species, the original description is given at this point.

"Mononyx suberosus: Fuscus, fronte fossulata, tliorace basi sub-
trisinuato, angulis posterioribus obtusiusculis. — Long. 3f lin."

"Fuscus, opacus. Caput scrobiculatum, griseo-squamulosum.
Thorax abdomine paulo latior, lateribus anterius rotimdatis, leniter
transversim bisulcatus, sulcis inter latera depressa et discum sub-
elevatum foveolatis, dorso postice obsolete tricarinato; fusco-
squamulosus. Scutellum basi utrinque carinatum, carinis nigro-
hirtis. Hemelytra nervis elevatis, fusco-squamulosa, utroque medio
lituris tribus minutis nigrohirtis. Corpus infra cum pedibus fuscum,
unicolor."

The Family Gelastocoridae (Hemiptera) 439

Nerthra walked n. sp.

(PI. XV, fig. 142)

Size. Male: Length, 9.3 mm.; width of pronotiim, 7.4 mm.;
width of abdomen, 7.3 mm. Female: Length, 9.4 to 9.7 mm.; width
of pronotum, 7.3 to 7.6 mm.; width of abdomen, 7.3 to 7.5 mm.

Color. Nearly concolorous, brown, rather dense, black bristles
making the specimens appear somewhat darker than they actually
are.

Structural characteristics. In general appearance strongly re-
sembling Nerthra macrothorax ( Montrouzier ) . Front of head
provided with five large, conical tooth-like tubercles, one apical,
two marginal and two superapical tubercles. Ocelli present. Width
of pronotum equal to or slightly greater than width of abdomen,
widest at a level with the transverse furrow; disc slightly elevated;
lateral margins subparallel for most part, abruptly converging
toward eyes anteriorly, posterior portions of lateral margins slightly
convergent from level with transverse furrow toward bases of em-
bolia; posterior margin of pronotum nearly straight, broadly but
very shallowly sinuate. Scutellum somewhat depressed basally;
clumps of bristles at middle of each lateral margin. Hemelytra ex-
tending beyond end of abdomen; entirely coriaceous and fused to-
gether; embolia greatly expanded laterally at the base. Connexivum
moderate. Body, especially scutellum and hemelytra covered with
rather dense clumps of short, clavate bristles. Abdominal sternites
of male, small, asymmetrical. Abdominal sternites of female nearly
symmetrical; last visible sternite projecting posteriorly and covering
the lobes of the ovipositor. Clasper of male stout, slightly recurved
and slightly swollen apically.

Location of type. Holotype male, allotype female and 2 female
paratypes, Adelaide R., N. W. Australia, J. J. Walker in the British
Museum, London, England. One female paratype, same data, in
the Francis Huntington Snow Entomological Collection at the
University of Kansas, Lawrence, Kansas.

Distributional data. As given for the type series.

Comparative notes. This species appears to be most closely re-
lated to N. grandis (Montandon) and N. femoralis (Montandon),
but diflFers from those two species in the shape of the pronotum,
fused hemelytra and in the slightly differently shaped clasper of the
male. The presence of ocelli and the toothlike tubercles on the
head will distinguish this species from N. macrothorax (Mont-
rouzier) which it resembles superficially.

440 The University Science Bulletin

Nertlira siniiosa n. sp.

(PI. XIV, figs. 131, 134)

Size. Female: Length, 7.5 to 7.8 mm., width of pronotum, 4.8 to
5.2 mm., width of abdomen, 5.0 to 5.4 mm.

Color. Light brown or yellowish-brown, darker on disc of
pronotum, base of scutellum and scattered areas on the ventral
abdominal segments.

Structural characteristics. Head with four fairly small, marginal
toothlike tubercles; ocelli present. Pronotum widest at a point
level with the transverse furrow, but only slightly wider than angle
at anterior one-fourth of lateral margin, lateral margin between
these two points strongly concave; antero- and posterolateral
margins slightly concave, converging mesad; posterior margin tri-
sinuate. Scutellum rather flat, a moderately large tumescence on
each side at middle of margin, apex slightly elevated. Hemelytra
rather pointed apically; membrane reduced to a narrow strip ex-
tending from apex along inner margin toward scutellum; extending
beyond the end of the abdomen; embolium not very expanded.
Connexivum completely covered by hemelytra. Abdominal ster-
nites of female more or less symmetrical, last visible sternite with
the apex produced, almost completely hiding lobes of the ovipositor,
rather broadly keeled, extreme apex very faintly emarginate.

Location of type. Holotype female, Dorrigo, N. S. Wales, Aus-
tralia, no date, W. Heron in the Museum of Comparative Zoology
at Harvard University. One female paratype, same data, in the
Francis Huntington Snow Entomological Collection at the Univer-
sity of Kansas.

Distributional data. As given for the type series.

Comparative notes. This species seems to be close to A/, laticollis
( Guerin-Meneville ) on the basis of the lateral margin of the pro-
notum, but it has the membrane of the hemelytra reduced and is
somewhat lighter in color than the latter species.

Nerthra grandis (Montandon)

(PI. Xll, fig. 106; PL Xlll, figs. 123, 126; PI. XIV, fig. 129)

1900. Matinus grandis Montandon, Bulletin de la Societe des Sciences de
Bucarest-Roumanie, an. VIII, no. 6, pp. 6-7.

Size. Male: Length, 9.1 mm., width of pronotum, 6.5 mm., width
of abdomen, 6.7 mm. Female: Length, 9.0 mm., width of pronotum
6.4 mm., width of abdomen, 6.7 mm.

Color. Almost uniformly dark reddish-brown, nearly black.

The Family Gelastocoridae (Hemiptera) 443

Structural characteristics. Front of head with five rather large
toothhke tubercles, apical one ventrad and caudad of the others
which are on the anterior margin of the head when viewed from
above; ocelli present, but very small. Pronotum slightly less in width
than abdomen; lateral margin with the posterior three fifths slightly
convex, nearly straight, anterior two fifths converging toward the
eye, straight; posterior margin trisinuate; transverse suture of disc
crossed by three distinct longitudinal carinae. Scutellum depressed
at the median basal portion, elevated at the apex and at the middle
of each lateral margin, the three elevations connecting to one an-
other. Hemelytra entirely coriaceous with the veins slightly elevated,
extending as far as end of abdomen; embolium expanded laterally
for its entire length. Connexivum visible in both sexes, rather ex-
panded laterad. Body covered black, clavate bristles which occur
in clumps on the hemelytra, the scutellum and the pronotum. Ab-
dominal sternites of the female symmetrical; last visible sternite
projecting posteriorly, covering the ovipositor lobes. Last three
abdominal sternites of male small, last sternite about two times
as wide as long, about equal in length to the eighth sternite and
one and one-half times as long as the seventh, ninth sternite only
about one-fifth as wide as fourth segment. Clasper of the male
simple, sickle-shaped, rather wide at base and tapering gradually
to apex which is recurved.

Location of type. In the Museum of Paris, Paris, France.

Distributional data. The specimens Montandon described were
from Australia. The writer has examined a pair of this species
labeled, Victoria, no date, P. R. Uhler Colin., (USNM).

Comparative notes. The above specimens were determined by
Montandon as this species in 1909. It is most closely related to
N. femoralis (Montandon), but may be distinguished from that
species by the entirely coriaceous hemelytra, by the absence of red
on the anterior femora and by the shape of the clasper of the male.
Montandon's description states that the membrane is reduced, but
the two specimens examined by this worker had the hemelytra
entirely coriaceous.

442 The University Science Bulletin

LITERATURE CITED

BlLLBERG, G. J.

1820. Enumeratio Insectorum in Museo Billberg. Holm. 138 pp.

Blatchley, W. S.

1925. Some Additional New Species of Heteroptera from Sonthem United
States, with Characterization of a New Genus. Entomological
News, vol. 36, pp. 49-52.

Champion, G. C.

1901. Biologia Centrali-Americana, Rhynchota Heteroptera, vol. 2, 416 pp.

Fabricius, J. C.

1798. Supplementum Entomologiae Systematicae. Hafniae. Proft et
Storch. 572 pp.
Guerin-Meneville, F. E.

1844. Note sur la Naucoris rugosa de J. Desjardins, formant un nouveau
genre d'Hemipteres, et description de plusiers especes des genres
Pelogonus et Mononyx. Revue Zoologique Travaux Inedits, pp.
112-114.

HUNGERFORD, H. B.

1922. The Life History of the Toad Bug. University of Kansas Science
Bulletin, vol. 14, pp. 145-171.

Kevan, D. K. McE.

1942. Some Observations on Mononyx nepaeformis ( Fabricius ) . Pro-
ceedings of the Royal Entomological Society of London (a) 17,
pts. 10-12, pp. 109-110.

KiRKALDY, G. W.

1897. Synonymic Notes on Aquatic Rhynchota. The Entomologist, vol.
30, p. 258.
Laporte, F. L. de

1833. Essai d'une Classification Systematique de L'order des Hemipteres.
Paris. 88 pp.
Martin, C. H.

1929. An Exploratory Survey of Characters of Specific Value in the
Genus Gelastocoris Kirkaldy and Some New Species. University of
Kansas Science Bulletin, vol. 18, no. 4, pp. 351-369.
Maxwell-Lefroy, H.

1909. Indian Insect Life. Calcutta and Simla. Thacker, Spink & Co.,
786 pp.
Melin, D.

1929. Hemiptera from South and Central America I. ( Revision of the
Genus Gelastocoris and the American species of Mononyx).
Zoologia Bidrag Fran Uppsala, Band 12, pp. 151-194.
MiLLSPAUGH, D. C.

1939. Bionomics of the Aquatic and Semi-Aquatic Hemiptera of Dallas
County, Texas. Field and Laboratory, vol. 7, no. 2, p. 82-83.

The Family Gelastocoridae (Hemiptera) 443

Nelson, H. G.

1949. A Method of Cleaning Insects for Study. The Coleopterist's Bul-
letin, vol. 3, no. 6, pp. 89-92.

Say, T.

1832. Descriptions of New Species of Heteropterus Hemiptera of North
America. New Harmony, Ind. 39 pp.

SxaL, C.

1861. Nova methodus famihas quasdam Hemiptera disponendi. Ofversigt

af Kongliga Vetenskaps Akademiens Fordhandlingar, Arg. 18, pp.

195-212.
1876. Enumeratio Hemipterorum. Kongliga Svenska Vetenskaps-

Akademiens. Handlingar, Band 14, no. 4, pp. 137-140.

ToRRE-BuENO, J. R. de la

1906. On Some Aquatic Hemiptera from Costa Rica, Central America.
Entomological New, vol. 17, p. 54.

444 The University Science Bulletin

INDEX

(Italics indicate synonyms.)

PAGE

adspera (Stal) (Nerthra) 434

alaticollis (Stal) (Nerthra) 432

amazonensis Melin ( Gelastocoris ) 325

americana ( Montandon ) ( Nerthra) 381

ampliata ( Montandon ) ( Nerthra ) 426

amplicollis ( Stal ) ( Nerthra ) 384

andinus Melin ( Gelastocoris ) .' 327

angulatus ( Melin ) ( Gelastocoris ) 339

annulipes ( Horvath ) ( Nerthra ) 426

apureensis Melin ( Gelastocoris ) 342

asiatica ( Horvath ) ( Nerthra ) 410

ater ( Melin) ( Nerthra) 389

hadius Herrich-Schiiffer (Mononyx) 393

barberi Torre-Bueno (Gelastocoris) 299

bergi DeCarlo ( Gelastocoris ) 332

bipunctatus Melin ( Mononyx ) 373

bipunctatus Stal ( Mononyx) 371

bolivianus DeCarlo ( Gelastocoris) 332

borealis ( Melin ) ( Nerthra ) 375

bracchialis n. sp. ( Nerthra ) 400

bridarollii DeCarlo ( Montandonius) 339

brunnea Blatchley ( Glossoaspis) 413

buenoi n. sp. ( Nerthra ) 365

bufo ( Herrich-SchaiFer ) ( Gelastocoris ) 322

californiensis Melin (Gelastocoris) 300

corudis n. sp. ( Nerthra ) 392

cucullatus Martin (Gelastocoris) 316

duplicatus Martin ( Gelastocoris ) 327

ecuadorensis ( Melin ) ( Nerthra ) 387

elongata ( Montandon ) ( Nerthra ) 436

femoralis ( Montandon ) ( Nerthra ) 423

flavtis (Gtierin-Meneville) (Gelastocoris) 331

f uscipes ( Guerin-Meneville ) ( Nerthra ) 393

fusco-conspersus Stal ( Monomjx) 366

fuscus Martin ( Gelastocoris ) 336

Galgidus Latreille 296

Gelastocorinae Champion 296

Gelastocoris Kirkaldy 296

Glossoaspis Blatchley 345

grandicollis (Germar) (Nerthra) 401

grandis ( Montandon ) ( Nerthra ) 440

grossus Montandon ( Mononyx ) 410

gumeyi n. sp. ( Nerthra ) 419

hirsuta n. sp. ( Nerthra ) 417

hungerfordi Melin (Gelastocoris) 316

hungerf ordi n. sp. ( Nerthra ) 398

The Family Gelastocoridae (Hemiptera) 445

PAGE

indica ( Atkinson ) ( Nerthra ) 405

lata ( Montandon ) ( Nerthra ) 389

laticoUis ( Guerin-Meneville ) (Nerthra) 418

limigcnus Stdl ( Monontjx) 401

lobata ( Montandon ) ( Nerthra ) 407

luteovaria ( Distant ) ( Nerthra ) 430

macrostyla n. sp. ( Nerthra ) 428

macrothorax ( Montrouzier ) (Nerthra) 414

major Montandon ( Gelastocoris ) 327

manni n. sp. ( Nerthra ) 396

mansosotoi DeCarlo ( Montandonius ) 339

martinezi DeCarlo ( Gelastocoris) 336

martini Todd ( Nerthra ) 358

Matinus Stdl 344

mexicana ( MeHn ) ( Nerthra ) 356

mixta ( Montandon ) ( Nerthra ) 420

Mononyx Laporte 343

montandoni ( Melin) ( Nerthra) 380

Montandonius Melin 339

nebulosus (Guerin-Meneville) (Gelastocoris) 331

nepaeformis ( Fabricius ) ( Nerthra ) 371

Nerthra Say 343

Nerthrinae Kirkaldy 343

nervosa ( Montandon ) ( Nerthra ) 416

niger Melin ( Mononyx) 368

nndata n. sp. ( Nerthra ) 425

obscurus Melin ( Mononyx) 376

obscurus Stdl ( Mononyx) 393

oculatus ( Fabricius ) ( Gelastocoris ) 298

omani n. sp. ( Nerthra ) 422

paraguayensis DeCarlo (Gelastocoris) 332

parvula ( Signoret ) ( Nerthra ) 362

Peltopterus Guerin-Meneville 344

peruensis Melin ( Gelastocoris ) 329

peruviana ( Montandon ) ( Nerthra ) 379

Phintius Stdl 344

planifrons ( Melin ) ( Nerthra) 388

projectus Distant ( Mononyx) 405

pulcher ( Stdl) ( Gelastocoris) 311

quadrimaculatus (Guerin-Meneville) (Gelastocoris) 331

quinquedentata ( Melin ) ( Nerthra ) 363

ranina ( Herrich-SchiiflFer ) ( Nerthra ) 368

raptoria ( Fabricius ) ( Nerthra ) 366

robusta n. sp. ( Nerthra ) 429

rotundatus Champion ( Gelastocoris) 314

rotundicollis Signoret ( Mononyx) 401

rudis ( Melin ) ( Nerthra ) 390

rugosa ( Desjardins) ( Nerthra) 412

Scylaecus Stdl 344

446 The University Science Bulletin

PAGE

serrata ( Montandon ) ( Nerthra ) 409

sinuosa n. sp. ( Nerthra ) 440

sordidus Herrich-Schciffer ( Mononyx) 401

spissa ( Distant ) ( Nerthra ) 412

stali Melin ( Phintius) 401

stall ( Montandon ) ( Nerthra ) 435

stali Torre-Bueno ( Gelastocoris) 331

stygica Say ( Nerthra ) 354

suberosa ( Erichson ) ( Nerthra ) 438

subsimilis Blatchleij (Gelastocoris) 300

tasmaniensis n. sp. ( Nerthra ) 437

tenebrosa n. n. ( Nerthra ) 376

terrestris ( Kevan ) ( Nerthra ) 373

tuberculata ( Montandon ) ( Nerthra ) 431

turgidula ( Distant) ( Nerthra) 406

unicornis ( Mehn ) ( Nerthra ) 378

usingeri Todd ( Nerthra ) 360

variegatus (Guerin-Meneville) (Gelastocoris) 311

vianai DeCarlo ( Gelastocoris ) 332

vicinus Champion (Gelastocoris) 319

viridis n. sp. ( Gelastocoris ) 338

walked n. sp. ( Nerthra ) 439

williamsi n. sp. ( Nerthra ) 383

willineri DeCarlo ( Montandonius ) 339

PLATE I

Fig. 1. G. oculatus (Fabricius). Dorsal view.
Fic. 2. G. oculatus (Fabricius). Ventral view.

The Family Gelastocoridae (Hemiptera)

447

PLATE I

OCELLUS

ANTERO-LATERAL
MARGIN.

POSTERO-LATERAL
MARGIN

SCUTELLUM _ / j

CONNEXIVUM.

MEMBRANE

OCELLOCULAR
SPACE

ANTERIOR ANGLE
DISC OF PRONOTUM
LATERAL ANGLE

-EMBOLIUM
CLAVUS

' CORIUM

ANTENNA

HEMELYTRON

448 The University Science Bulletin

PLATE 2

Fig. 3. N. ranina ( Herrich-Schaffer ) . Dorsal view.
Fig. 4. N. ranina ( Herrich-SchaflFer ) . Ventral view.

The Family Gelastocoridae (Hemiptera)

449

PLATE II

OCELLUS
AMTtRlOR ANSLE-

DISC OF PRONOTUM

LATERAL ANCLE-

SCUTELLUM

«ONNEXIVUM

MEMBRANE

OCELLOCULAR SPACE

ANTERO-UATERAL

MARGIN

POSTERO-LATERAL
MARGIN

CLAVUS

-J-.-EMBOLIUM

CORIUM

ANTENNA
ROSTRUM

25t EYE

15—6730

450 The University Science Bulletin

PLATE III

Fig. 5. G. oculatus (Fabricius). Antenna.
Fig. 6. A/, ranina ( Herrich-Scha£Fer ) . Antenna.
Fig. 7. N. raptoria (Fabricius). Portion of foreleg.
Fig. 8. G. oculatus (Fabricius). Foreleg.
Fig. 9. IV. ranina (Herrich-Schaffer). Foreleg.
Fig. 10. N. buenoi n. sp. Portion of foreleg.

The Family Gelastocoridae (Hemiptera)

451

PLATE III

452 The University Science Bulletin

PLATE IV

Fig. 11. G. oculattis (Fabricius). Lateral view of head.

Fig. 12. N. ranma (Herrich-Schaffer). Lateral view of head.

Fig. 13. G. viridis n. sp. Abdominal sternites of male.

Fig. 14. G. angtilatus ( Melin ) . Abdominal sternites of male.

Fig. 15. G. bufo (Herrich-Schaffer). Pronotum.

Fig. 16. G. oculattis variegatus (Guerin-Meneville). Pronotum.

Fig. 17. G. vicinus Champion. Pronotum.

Fig. 18. G. rotundatus Champion. Pronotum.

The Family Gelastocoridae (Hemiptera)

453

PLATE IV

13.

14

15.

16.

17.

18.

454 The University Science Bulletin

PLATE V

Fig. 19. G. htingerfordi Melin. Pronotum,

Fig. 20. G. fuscus Martin. Pronotum.

Fig. 21. G. major Montandon. Pronotum.

Fig. 22. G. amazonensis Melin. Pronotum.

Fig. 23. G. viridis n. sp. Pronotum.

Fig. 24. G. nebulosus ( Guerin-Meneville ) . Pronotum.

Fig. 25. G. angulatus (Melin). Pronotum.

Fig. 26. G. peruensis Melin. Pronotum.

Fig. 27. G. oculatus (Fabricius). Male genitalia.

Fig. 28. G. oculatus ( Fabricius ) . Left clasper of male genitalia.

Fig. 29. G. angulatus (Melin). Male genitalia.

The Family Gelastocoridae (Hemiptera)

455

PLATE V

■■mi^ KEEL HOOO

.FRINGE

'\W":^~~v- PAN

TUMESCENCE OF
RIGHT CLASPER
»IOHT CLASPER HOOK
LEFT CLASPER

-RIGHT CLASPER

.BULB

-KEEL

456 The University Science Bulletin

PLATE VI

Keel and Right Clasper of Male Genitalia.

Fig. 30. G. rotundatus Champion.

Fig. 31 G. oculatus variegatus (Guerin-Meneville).

Fig. 32. G. vicinus Champion.

Fig. 33. G. hungerfordi Melin.

Fig. 34. G. bufo (Herrich-Schaffer).

Fig. 35. G. nehulosus (Guerin-Meneville).

Fig. 36. G. amazonensis Melin.

Fig. 37. G. viridis n. sp.

Fig. 38. G. peruensis Melin.

Fig. 39. G. major Montandon.

Fig. 40. G. fusctis Martin.

The Family Gelastocoridae (Hemiptera) 457

PLATE VI

458 The University Science Bulletin

PLATE VII

Right Clasper of Male Genitalia.

Fig. 41. IV. ranina ( Herrich-Schaffer ) .

Fig. 42. N. martini Todd.

Fig. 43. N. mexicana (Melin).

Fig. 44. N. parvula (Signoret).

Fig. 45. IV. bracchialis n. sp.

Fig. 46. N. raptoria (Fabricius).

Fig. 47. N. peruviana (Montandon).

Fig. 48. N. terrestris (Kevan).

Fig. 49. N. nepaeformis (Fabricius).

Fig. 50. IV. tenebrosa n. n.

Fig. 51. N. hungerfordi n. sp.

Fig. 52. N. fuscipes (Guerin-Meneville).

Fig. 53. IV. amplicollis (Stal).

Fig. 54. N. manni n. sp.

The Family Gelastocoridae (Hemiptera)

459

PLATE VII

460 The University Science Bulletin

PLATE VIII

Fig. 55. N. montandoni ( Melin ) . Abdominal stemites and portion of right
clasper of male genitalia.

Fig. 55a. N. montandoni (Melin). Enlarged portion or right clasper of
male genitalia.

Fig. 56. N. ater ( Melin ) . Abdominal stemites and portion of right clasper
of male genitalia.

Fig. 56a. N. ater (MeHn). Enlarged portion of right clasper of male geni-
talia.

Fig. 57. N. unicornis ( Melin ) . Abdominal stemites and portion of right
clasper of male genitalia.

Fig. 57a. N. unicornis ( Melin ) . Enlarged portion of right clasper of male
genitalia.

Fig. 58. N. ecuadorensis (Melin). Portion of right clasper of male genitalia.

Fig. 59. N. borealis (Melin). Portion of right clasper of male genitalia.

Fig. 60. N. hungerfordi n. sp. Abdominal stemites of male.

Fig. 61. N. tenebrosa n. n. Abdominal stemites of male.

Fig. 62. N. terrestris ( Kevan ) . Abdominal stemites of male.

Fig. 63. N. fuscipes (Guerin-Meneville). Abdominal stemites of male.

Fig. 64. N. raptoria ( Fabricius ) . Abdominal stemites of male.

Fig. 65. N. peruviana ( Montandon ) . Abdominal sternites of male.

Fig. 66. N. manni n. sp. Abdominal stemites of male.

Fig. 67. N. parvula (Signoret). Abdominal stemites of male.

Fig. 68. N. amplicollis (Stal). Abdominal sternites of male.

The Family Gelastocoridae (Hemiptera) 461

PLATE VII!

462 The University Science Bulletin

PLATE IX

Fig. 69. N. mexicana ( Melin ) . Abdominal stemites of male.

Fig. 70. N. martini Todd. Abdominal stemites of male.

Fig. 71. N. bracchialis n. sp. Abdominal stemites of male.

Fig. 72. N. nepaeformis (Fabricius). Abdominal stemites of male.

Fig. 73. N. ecuadorensis (Melin). Abdominal stemites of male.

Fig. 74. N. nepaeformis (Fabricius). Front margin of head.

Fig. 75. ZV. martini Todd. Front margin of head.

Fig. 76. N. fuscipes ( Guerin-Meneville ) . Front margin of head.

Fig. 77. N. unicornis (Melin). Front margin of head.

Fig. 78. N. hungerfordi n. sp. Abdominal stemites of female.

Fig. 79. N. bracchialis n. sp. Abdominal stemites of female.

Fig. 80. N. manni n. sp. Abdominal stemites of female.

The Family Gelastocoridae (Hemiptera) 463

PLATE IX

SUPERAPICAL TUBERCLE^ 73
LATERAL TUBERCLE

75.

MEDIAN TUBERCLE

APICAL TUBERCLE.

77

78.

80

464 The University Science Bulletin

PLATE X

Abdominal Sternites of Female.

Fig. 81. N. ranina ( Herrich-Schaffer ) .

Fig. 82. N. buenoi n. sp.

Fig. 83. N. peruviana ( Montandon ) .

Fig. 84. N. quinquedentata (Melin).

Fig. 85. N. montandoni (Melin).

Fig. 86. N. ecuadorensis ( Melin )

Fig. 87. N. terrestris (Kevan).

Fig. 88. N. raptoria (Fabricius).

Fig. 89. N. fuscipes (Guerin-Meneville).

Fig. 90. N. rudis (Melin).

Fig. 91. N. martini Todd.

Fig. 92. N. tenehrosa n. n.

Fig. 93. N. nepaeformis (Fabricius).

Fig. 94. N. mexicana ( Melin )

Fig. 95. N. atnplicollis (Stal).

The Family Gelastocoridae (Hemiptera) 465

PLATE X

466 The University Science Bulletin

PLATE XI

Right Clasper of Male Genitalia

Fig. 96. N. grandicollis ( Germar ) . This does not represent a true ventral

view, but rather a ventrolateral view.

N. lobata (Montandon).

N. indica (Atkinson).

N. nudata n. sp.

N. omani n. sp.

N. spissa (Distant)

N. robusta n. sp.

N. macrostijla n. sp.

Fig.

97.

Fig.

98.

Fig.

99.

Fig.

100.

Fig.

101.

Fig.

102.

Fig.

103.

The Family Gelastocoridae (Hemiptera)

467

PLATE XI

468 The University Science Bulletin

PLATE XII

Right Clasper of Male Genitalia

Fig. 104. N. tuberculata (Montandon).

Fig. 105. N. femoralis (Montandon).

Fig. 106. N. grandis (Montandon).

Fig. 107. N. luteovaria ( Distant ) .

Fig. 108. N. laticollis (Guerin-Meneville).

Fig. 109. N. gurneiji n. sp.

Fig. 110. N. stygica Say. Apical portion.

Fig. 111. N. ampUata (Montandon).

Fig. 112. N. williamsi n. sp.

Fig. 113. N. macrothorax ( Montrouzier ) . Apical portion.

Fig. 114. N. alaticollis (Stal).

Fig. 115. A/, americana (Montandon).

Fig. 116. N. mixta (Montandon). Apical Dcrtion.

The Family Gelastocoridae (Hemiptera)

469

PLATE XII

470 The University Science Bulletin

PLATE XIII

Fig. 117. N. stygica Say. Abdominal stemites of female.

Fig. 118. N. alaticollis (Stal). Abdominal stemites of male.

Fig. 119. N. macrothorax ( Montrouzier). Abdominal sternites of male.

Fig. 120. N. americana (Montandon). Dorsal view of male.

Fig. 121. N. rugosa (Desjardins). Dorsal view of female.

Fig. 122. N. adspersa (Stal). Dorsal view.

Fig. 123. A/, grandis (Montandon). Abdominal stemites of male.

Fig. 124. N. nervosa (Montandon). Portion of pronotum and left heme-

lytron of female.

Fig. 125. N. stali (Montandon). Dorsal view of female.

Fig. 126. JV. grandis (Montandon). Dorsal view of female.

The Family Gelastocoridae (Hemiptera) 471

PLATE XIII

472 Till-: Univehsity Science Bulletin

PLATE XIV

Fig. 127. N. alaticollis (Stal). Abdominal sternites of female.

Fig. 128. N. asiatica (Horvath). Abdominal sternites of female.

Fig. 129. N. grandis (Montandon). Abdominal sternites of female.

Fig. 130. N. sttjgica Say. Dorsal view of female.

Fig. 131. N. sinuosa n. sp. Abdominal sternites of female.

Fig. 132. N. hirsuta n. sp. Abdominal sternites of female.

Fig. 133. N. stall (Montandon). Abdominal sternites of female.

Fig. 134. IV. sinuosa n. sp. Dorsal view of female.

Fig. 135. N. macrotJwrax (Montrouzier). Dorsal view of female.

The Family Gelastoc;oridae (Hemiptera)

473

PLATE XIV

474 The University Science Bulletin

PLATE XV

Fig. 136. N. luteovaria (Distant). Abdominal stemites of female.

Fig. 137. N. tuherculata (Montandon). Abdominal stemites of female.

Fig. 138. N. lohata ( Montandon ) . Abdominal stemites of female.

Fig. 139. N. laticoUis (Guerin-Meneville), Abdominal stemites of male.

Fig. 140. N. robusta n. sp. Abdominal stemites of male.

Fig. 141. N. usingeri Todd. Right clasper of male genitalia.

Fig. 142. N. tvalkeri n. sp. Right clasper of male genitalia.

Fig. 143. N. tasmaniensis n. sp. Right clasper of male genitalia.

Fig. 144. N. corudis n. sp. Abdominal stemites of male and apical portion
of right clasper.

Fig. 145. N. corudis, n. sp. Enlarged apical portion of right clasper of
male genitalia.

The Family Gelastocoridae (Hemiptera)

475

PLATE XV

145.

THE UNIYEHSITY OF KANSAS

SCIENCE BULLETIN

Vol. XXXVII, Pt. I] October 15, 1955 [No. 12

Studies of the Agar-Plate Precipitin Test ^

BY

Charles A. Leone, A. Byron Leonard
and Carlon W. Pryor ~

Department of Zoology, University of Kansas
Lawrence, Kansas

Abstract: Modifications of Ouchterlony's Petri -plate, agar-diffusion method
for performing precipitin tests have been described. Minor changes in the
preparation of the agar, the use of plastic molds to form the v^'ells which
receive the serological reagents, and the addition of dyes to color the agar are
discussed. A simple technique for making permanent photographic records of
the zones of precipitation is reported. Homologous and cross reactions are
illustrated. A manner in which independent serological systems may approxi-
mate true cross reactions in their appearances is illustrated. Photographic
records of the development of zones of precipitation using the a- and R- pre-
cipitin tests are presented.

INTRODUCTION

Ouchterlony ( 1948 ) described an excellent method for qualita-
tively analyzing soluble antigens and antibodies on a Petri plate
containing agar. Antigens and antibodies diffuse through the agar
from wells so arranged that the planes of diffusion are at right
angles to each other. A visible zone of precipitation forms where
the reagents combine in proper proportions. Bjorklund ( 1952a,
1952b ) employed the method in a preliminary analysis of serological
components in hemopoietic tissues of rabbits. Oudin (1952), by
means of his "simplifying hypotheses," discussed the theoretical
basis for antigen-antibody reactions in gels. He pointed out (p.
371 ) that the relative concentrations of the antigens and antibodies
influenced both the location of the zone of precipitation and the time
of its appearance. In order to detect all zones, he suggested that

1. These studies were aided by a contract between the Office of Naval Research, De-
partment of the Navy, and the University of Kansas, NR 163-012.

2. Public Health Service Predoctorate Research Fellow of the National Microbiological
Institute.

(477)

478 The University Science Bulletin

the plates be examined at intervals over a period of several days.
In multiple systems, where several antigens and antibodies are
interacting, the successive appearance of the zones, and the pro-
gressive manner in which each zone develops require that a series
of permanent records be made of the zones of precipitation in the
plates in order to facilitate accurate analyses. Various types of
tracings, diagrams and photographs of the zones have been used
to illustrate the end results of experiments. No reports have been
published in which the daily changes in the agar-plates are accu-
rately recorded in simple fashion. Oudin's suggestion (p. 365) to
place the plates on photographic paper and to make contact prints
seemed to us to be the most promising.

We are applying the Ouchterlony method to the study of the
precipitin reaction itself, and in comparative serological analysis of
the hemocyanins of several arthropods. We modified the method
in several ways that improved the quality of our results. The pur-
poses of this report are to describe the Petri-plate, agar-diflFusion
method as it has been employed in our laboratory, to present a
simple technique for making permanent photographic records of
the zones of precipitation as they occur in the agar, to illustrate the
development of the zones of precipitation during a 7-day difiFusion
test, to picture homologous and cross reactions, to show how inde-
pendent serological systems may approximate true cross reactions
in their appearances, and to illustrate the influence of dilution of the
antigen and the antiserum upon the characteristics of the zones of
precipitation.

MATERIALS AND METHODS

Agar. Agar-preparations must be as transparent as possible in
order not to mask small, diffuse zones of precipitation. Various
brands and qualities of agar were used as were various chemical
and mechanical means of clarifying the preparations. Best results
were obtained with the following procedures:

Place 66 grams of granulated, Difco Bacto-agar into a four-liter Ehrlenmeyer
flask with two liters of distilled water and heat in a steam sterilizer at 18 pounds
of pressure for 15 minutes. Remove from autoclave, add, gradually, 300 ml. of
0.5 percent calcium chloride to the hot agar, and manually rotate flask gently
until the flocculation of calcium sulfate occurs. Pour the hot agar into two,
1000 ml. beakers and permit it to solidify. Cut the solidified agar into cubes
approximately one cm. on a side, place cubes into a large container and wash
in running tap water for three days. Drain off excess water and remelt cubes.
Dilute liquid agar with an equal volume of 1.6 percent solution of sodium
chloride buffered to pH7 with M/75 dibasic sodium phosphate and M/75 mono-

The Agar-Plate Precipitin Test 479

basic potassium phosphate. While the agar is still hot, add 120 mg. of trypan
blue, and 52 ml. of a 1.0 percent solution of "Merthiolate." Reheat the agar
in the steam sterilizer for 15 minutes at 15 pounds of pressure. Filter the
preparation in a Buckner funnel through No. 5 Whatman paper into a warm
four-liter, side-ann flask, applying gentle suction to speed the filtration. Pour
the filtered agar into large, clean beakers and permit it to solidify. Cut agar
into cubes. Store at room temperatures in plastic covered containers. Portions
of the stock agar-preparation can be remelted and poured into Petri i^latcs as
desired.

During storage of the agar, water is gradually expressed from
the cubes but seems not to alter the usefulness of the agar. We
have stored preparations for periods up to 30 days without apparent
changes in the results obtained from serological tests in which the
agar was used. Aluminum foil is not satisfactory as a dust cover
for the agar. During prolonged storage, vapor from the agar
causes the aluminum to deteriorate.

Petri plates containing agar should be used within a few hours,
and never more than 24 hours, after plates are poured. After longer
periods, sufficient dehydration occurs in unused plates, even when
covered, to cause aberrant results. Moreover, when "old" plates
are used in serological tests, cracks may appear in the agar, usually
at the corners of the wells, after four or five days and interfere
with the tests. Storing the stock of the agar-preparation, or the
poured-plates, in the refrigerator is not recommended because the
agar becomes opalescent and does not completely clarify when
brought back to room temperatures or higher.

Mold assembly. For making the reservoirs into which the solu-
tions of antigens and antibodies are placed, Ouchterlony employs
two or three metal molds 10 mm. on a side, mounted together in a
fixed position. Each mold is pierced by a central hole to avoid
sucking up agar when the assembly is withdrawn. We have found
that an assembly of five plastic molds ( 10 mm. x 10 mm. x 15 mm. )
made of transparent cellulose acetate is satisfactory. "Plexiglass"
from the Rohm and Haas Company or "Lucite" from the Tennessee
Eastman Corporation have been used to make the assembly. Four
molds are arranged in a hollow square so that adjacent surfaces are
20 mm. apart. The fifth mold is centered among them so that its
corners are in apposition to the corners of the outer molds (Pi. I).
Mounting the molds in fixed symmetrical positions provides a con-
stant geometry which is useful when critical comparisons are to be
made among different preparations of proteins or among several
antisera.

480 The University Science Bulletin

Pouring the agar. Following Ouchterlony's method, the bottoms
of the Petri plates are first covered with a thin layer of agar which
is allowed to cool; the mold-assembly is placed on the surface of the
agar, and another layer of agar approximately three mm. in depth
is poured. Capillary migration of the liquid agar up the sides of the
molds elevates the rims of the wells above the plane surface of the
gel. The elevated rims are objectionable in that they interfere with
the photographic recording of zones of precipitation situated close
to the wells. In an attempt to avoid the elevated rims we coated the
molds with materials such as paraffin, detergents, and silicone to
change the amount of wetting by the agar. General Electric's Sili-
cone SF-81 (403) -13308 N-*8, gives the best results, although some
capillary movement still occurs. Permitting the agar-solution to cool
to 50-56° C before pouring it is an additional aid. The mold-assem-
bly is removed with a quick, vertical pull as soon as the agar is
completely gelled.

"Merthiolate" was added as a bacteriostatic agent to the solutions
of antigens and antisera, as well as to the agar. We found a dilution
of 1:10,000 (0.01 percent) to be most satisfactory. The covers of
the Petri plates can be removed frequently for the addition of re-
agents, for the observation of the zones of precipitation, or for pho-
tographic purposes, but excessive drying should be avoided.

Normal practice in our laboratory is to fill the wells with the
solutions of serological reagents, and then refill the wells as neces-
sary. Twelve hours is the most common interval between fillings.
Five to six ml. of solution is sufficient to run a test for seven to ten
days. Most satisfactory for transferring solutions to the wells is a
dropping pipette with a tube eight inches long, and one-fourth inch
in diameter manufactured by Glasco Products, Chicago, Illinois.
All tests have been run at room temperature (24-26° C). Wet in-
serts of filter paper or paper toweling fitted to the inside of the lids
of the Petri plates reduce evaporation of the serological reagents
and prevent the formation of droplets of liquid which could fall on
the plate and influence the results of the test. The paper is soaked
with water, as necessary, to make it adhere continuously to the
glass.

In comparisons of antigens, the antiserum generally is placed in
the central well, while in the investigation of properties of different
antisera it is convenient to place the antigen in the central well.
Appropriate control solutions may be employed as required.

Photographic record. Daily photograms of the zones of precipitate
are readily made by placing the plates over photographic paper,

The Agar-Plate Precipitin Test 481

making a suitable exposure, and treating the exposed paper as an
ordinary photographic print.

Dyes may be used to color the agar and improve the photographic
contrast between the zones and the surrounding medium. Bjorklund
(1952a) took ordinary photographs of the precipitates in his Petri-
plates and used methyl-orange and blue sensitive film to obtain
pictures of high contrast. We have employed methyl-orange and
red-insensitive Kodabromide No. 5 photographic paper to make
satisfactory photograms. Hemoglobin in the solutions results in
the appearance of white "halos" around the wells on the photograms
of the plates (PI. II, figs. 5 and 6) colored with methyl-orange.
Since the zones of precipitate are recorded as white also, the white-
ness of the "halo" tends to mask them. Solutions of proteins which
are essentially colorless are not troublesome in this respect.

We experimented with a selection of dyes and photographic

exposures in order to improve the photograms. The agar was

colored with the following kinds and amounts of dyes:

Methyl orange 33 mgs/100 cc of agar

Methyl green 0.1 cc of a 1% aqueous solution/50 cc of agar

Methyl violet 0.1 cc of a 1% aqueous solution/50 cc of agar

Crystal violet 30 mgs/1000 cc of agar

Trypan blue 30 mgs/1000 cc of agar

Photograms were made by placing a Petri plate, with its cover
removed, over a piece of Kodabromide paper No. 5 of sufficient size
to record the entire plate. An ordinary photographic enlarger, with
the light at an arbitrarily determined, but uniform, distance from
the plate was used as a light source. While it is convenient, an
enlarger is not necessary, and any concentrated light source em-
ployed in an otherwise darkened room, would serve as well. The
photograms of the plates colored with trypan blue gave greatest
contrast between the zones of precipitation and the agar. The
'Tialo" effect obtained in methyl orange plates was absent, and
the boundaries of the basins were clearly delimited. Photograms of
the blue-plates were sensitive enough to record the lint which
adheres to the surface of the agar, finger prints on the glass and
agar, defects in the glass, and other miscellaneous marks and
smudges. All of these artifacts are easily identified in the photogram
and hence, make possible the most accurate reading of the serologi-
cal reaction which has occurred in the agar. None of the dyes
interfered with the formation of the zones of precipitation. A blue,
glass-marking (or wax) pencil was used to record pertinent infor-
mation about each test on the underside of the dish. If the pencil-

16—6730

482 The University Science Bulletin

markings are printed or written in mirrored writing, they are
recorded properly on the photogram, and together with zones of
precipitate form a complete record of the test on the photographic
paper. By appropriately altering the concentration and the color
of the dye, the time of exposure, and the kind of photographic
paper used, useful photograms can be obtained under a variety
of conditions.

Antigen- antibody system. The hemocyanins from two species of
arthropods, the king crab Linmlus pohjphemus and the blue crab
Callincetes sapidiis, were injected intravenously and concurrently
into a rabbit. Three series of %, /2, 1, and 2 ml. doses of each
antigen were given (Leone 1952). The resulting antiserum was
tested with each antigen and with a pooled sample composed of
both antigens (PL I).^ The concentration of the antigens was 3.0
grams percent in the wells containing the hemocyanins of either
species. Protein levels in the wells containing the pooled sample
were adjusted so that the effective concentration of each antigen
was also 3.0 grams percent. The solution of hemocyanin of one
species was used as diluent for the other.

A hyperimmune antiserum was produced in a rabbit against the
hemocyanins of the edible crab Cancer magister. Five series of in-
jections, of /4, /2, 1, and 2 ml. doses, were given (Leone, 1952).
The antiserum was used to analyze the influence of the relative
concentrations of antigens and antibodies on the zones of precipita-
tion which formed in the agar (Pis. Ill and IV). In the tests in-
volving varying concentrations of antigen and a constant concentra-
tion of antibodies, a doubling-dilution sequence of the hemocyanins
was prepared, beginning with a solution of 6.39 grams percent
protein, a dilution of 1:15.625 (Well No. 1, PI. Ill), and ending
with a dilution of 1:256,000 (Well No. 16, PI. III). The doubling-
dilutions were placed in the wells in the regular sequence (Pi. Ill,
Cell Nos. 1 through 16). Undiluted antiserum was placed in the
central well (xm). In the tests involving varying concentrations of
antibodies and a constant concentration of antigen (Pi. IV), a
doubling-dilution sequence of the antiserum was prepared, the
greatest dilution being 1:128 (PI. IV, Cell Nos. 1 through 8). A
solution of antigen containing 6.39 grams percent of hemocyanin
was placed in the central well (m) and the dilutions of antisera
were placed in sequence in the outer wells.

Descriptive terms. We have found it necessary to adopt a series
of terms to describe uniformly the serological reactions which

The Agar-Plate Precipitin Test 483

occur in the agar. Oudin (1952) has provided a number of useful
terms such as: zone of precipitation, coalescence of zones, partial
coalescence of zones, and intersection of zones. Zones are quahta-
tively differentiated in our laboratory as to appearance by designat-
ing them as broad or narrow, and dense when they were opaque,
or diffuse when they were translucent. In some tests, more precise
comparisons among zones are possible by measuring their widths
on the photograms. The characteristics of a zone of precipitation
changes as a reaction progresses. For example, a zone after 1 day
may appear as short, narrow and dense; after 7 days it may be
greatly elongated, broad and dense in its medial region, and be
narrowing and diffuse in its lateral regions. The inedial region of
a zone is that portion which develops between the adjacent corners
of the well containing antigen and the well containing antiserum.
As the zone continues to grow in length its lateral portions may
coalesce with, or intersect, other zones. In multiple precipitating
systems several zones of precipitation often appear between the
wells containing the reactants. The position of a zone is described
as being more or less central, according to its relative nearness to
the central well. Zones usually are referred to as being opposite a
given reference-well when they occur between the wells containing
the reactants, and between two wells when the wells in question
contain only one kind of serological reagent, that is either antigens
or antisera,

EXPERIMENTAL RESULTS

Plate I illustrates the daily growth and development of zones of
precipitation, some of which coalesce whereas others intersect.
The zones grow in length and thickness. Ry three days of diffusion-
time two zones of precipitation appeared between the wells con-
taining antiserum (xcl) and the hemocyanin of Callinectes (c),
and three zones appeared between the walls containing the anti-
serum and the hemocyanin of Limulus (l). The wells marked cl
contained a mixture of the hemocyanins of the two species. Only
two zones of precipitation appear opposite the CL-wells at four
days. Five zones could be expected in view of the fact that all of
the zones between the c- and L-wells intersect. The rabbit produced
no antibodies which were reactive with the hemocyanins of both
Callinectes and Limulus. It is obvious that the zones of precipita-
tion opposite the CL-wells are superimposed upon one another.
As would be expected, between the CL-wells, all of the zones

484 The University Science Bulletin

coalesce, and between the CL-well and the c-well or. L-well some of
the zones coalesce while others do not.

Plate II, Figure 1 pictures a duplicate of the test shown in Plate I,
with the exception that in the second test the wells were not refilled
with the serological reagents. A comparison of results obtained by
repeatedly filling the wells with those obtained by a single filling is
possible. The 0.3 ml. of reagent required to fill each well, developed
zones in seven days; the zones were approximately equivalent to the
zones shown for four days in Plate I. The zones of precipitation are,
however, more sharply defined in Plate II, Figure 1, than in Plate I,
and where coalescence and overlapping of the zones have occurred
the reactions involving the proteins of either Callinectes or Limulus
are more easily identified. The less central zone between the c- and
XCL- wells (PI. I) did not appear in this test (PL II), whereas the
most central zone between the l- and xcL-wells was barely \'isible.
There probably was insufficient reagent available to permit the dif-
ferentiation of these less distinct zones, or to develop the coalescence
and intersection of the zones to the degree obtained in Plate I,
Figure 6.

Plate II, Figure 2 illustrates the appearance of homologous and
cross reactions between the antiserum (xm) produced against the
hemocyanins of Cancer magister and the sera of four closely related
species of crabs: C. magister (m), C. pagurus (p), C. productus
(d) and Carcinus maeniis (e). The "partial-identity reactions"
(Ouchterlony 1948) visible between m- and P-wells and m- and
E-wells are similar in appearances to the overlapping and independ-
ent reactions, pictured in Plate I, Figure 6.

The homologous reaction in Plate II, Figure 2 developed three
zones of precipitation, each distinctly different from the other. The
most nearly central zone was broad and dense in its medial region,
and dense and narrowing in its lateral regions. The middle zone was
uniformly narrow and dense. The outer, or least central, zone was
broad and diffuse. The variable characteristics of the zones of pre-
cipitations for the cross reactions indicate that there are qualitative
differences among the hemocyanins of these species. Between
p-well (C. pagurus) and the antiserum only a single dense zone of
uniform width developed. Between D-well (C. productus) and the
antiserum tliree zones developed the least central of which was
barely visible. C. productus more nearly approaches C. magister
in reactivity than does C. pagurus. The coalescing zones between
p- and D-wells obviously are equivalent whereas those between d-
and E-wells are not. The characteristics of the zone produced by the

The Agar-Plate Precipitin Test 485

cross reaction of hemocyanins from Carcinus (e-wcII) with the
antiserum are distinctly different, that is to say, they are more dif-
fuse and tapering, from the characteristics of the cross reaction
involving the proteins of C pagurus (p-well). The partial coales-
cence of the Carcinus zone ( E-well ) with the C. magister zone ( m-
well ) obviously is different from the partial coalescence of the latter
and the zone for C. pagurus (p-well). The photogram is the only
form of recording known to the authors which could detect such
qualitative differences among precipitin reactions in gels.

Plate III contains the record of the zones of precipitation obtained
from a standard a-precipitin test. Photograms of the Petri plates
are so arranged that the development of the zones of precipitation
for any particular dilution of antigen may be compared in the hori-
zontal rows of figures, and zones of precipitation for all of the dilu-
tions of antigen, at any given time, may be compared in the columns
of figures. A study of Plate III reveals that the concentration of
antigen of a homologous precipitin reaction conducted in agar, under
the conditions stipulated above, influences (1) the characteristics
of a zone, that is to say, whether it is dense or diffuse, broad or nar-
row, of uniform width or tapering; ( 2 ) the number of zones of pre-
cipitation which develop and the number of zones which are
detectable; (3) the relative position of the zones of precipitation
between the wells containing the reagents; (4) the time of appear-
ance of a zone of precipitation; (5) the time required for identical
zones to coalesce. All five of these factors are variables which must
be kept in mind when this agar-diffusion technique is used to analyze
protein systems.

The antigen-dilutions of higher concentration were responsible
for the development of the largest number of zones. At seven days,
opposite Well No. 1 (PI. Ill, Fig. IF), containing 6.39 grams percent
protein, there were eight distinct zones, plus another "zone" which
appeared to hug the walls of the well. Well No. 2 had seven zones;
Well Nos. 3 and 4 had six zones, etc. As the concentration of anti-
gen decreased, the number of visible zones decreased and the posi-
tions for the zones moved closer to the antigen-wells. The outer,
or less central, zones which are so apparent in Figure IF, and that
are visible to a lesser extent opposite Well Nos. 5 and 6 (Fig. 2F),
have disappeared from the agar in the region opposite Well No. 9
(Fig. 3F), and only the zone corresponding to the broad, most
central zone in Figure IF is present. This still-present zone is dis-
placed away from the antiserum-well, as compared to the position it
occupied in Figure IF. The zones of precipitation which formed

486 The University Science Bulletin

opposite Well Nos. 5 and 6 (Fig. 2C) illustrate a situation where
the region of coalescence reveals two distinct zones at three days,
but by seven days (Fig. 2F) the distinction is obliterated.

No zones were visible opposite Well Nos. 11 and 12 after one
day of diffusion (Pi. Ill, Fig. 3A), but by two days a diffuse zone
had appeared. The reactions around Well No. 13 are of special
interest. The antigen in the well had a dilution of 1:64,000 (1.81
mgs. percent). No zone of precipitation developed in the agar
in the seven days of diffusion. A visible precipitate was deposited,
however, around the edges of the Well No. 13 which were closest
to the antiserum-well ( Figure 4F ) .

Plate IV illustrates the occurrence of zones of precipitation in a
two-dimensional agar-diffusion precipitin test in which /? technique
(constant antigen-varying antiserum) was used. This hemocyanin-
antihemocyanin test is the reciprocal of the a-test illustrated in
Plate III. At all of the dilutions of antiserum for which precipita-
tions were visible, the zones were broad medially and narrowing
laterally. The densities of the zones were a function of the concen-
tration of the antibodies. Two principal zones of precipitation oc-
curred; the more nearly central zone was narrow and the other
zone was broad. As the broad zone developed, it became obvious
at the region of coalescence that the precipitate was not deposited
homogeneously (PI. IV, Fig. IF). Seven zones of precipitation
were visible between Well Nos. 1 and 2 after seven days of diffu-
sion. To test whether or not the development of the several zones
was an artifact which was a consequence of repeated fillings of the
wells, or due to minor fluctuations in temperature during the ex-
periment, the test was repeated employing but a single filling of
the wells and a carefully controlled temperature (26° + 0-5° ^)-
The results of the test are shown in Plate II, Figures 3 and 4. Except
for the fact that the zones of precipitation had not developed, after
seven days, to the same extent as the zones in Plate IV, Figures IF
and 2F, no differences were apparent which could not be ascribed
to the limited amounts of reagents available in the control-test.
Four zones were visible in the region of coalescence (Pi. II, Fig. 3)
and this approximately corresponded to the five-day stage of the
original test (PL IV, Fig. IE).

Only in the "undiluted" and "1 + 1" wells (PI. IV, Fig. IF, Well
Nos. 1 and 2) were the antibodies sufficient to cause the distinct
representation of the several zones in the region of coalescence.
The zones of precipitation between other wells were too difiFuse to

The Agar-Plate Precipitin Test 487

make possible a sure subdivision of the broad zone. The highest
dikition of antiserum at which a zone of precipitation was visible
was 1:64. This dilution and the next higher dilution (1:128) are,
in our experience, typical agar-titers for powerful precipitating
antisera produced against hemocyanins.

The principal effect of varying concentration of antibodies upon
the zones of precipitation is on their density. To a lesser extent
the proximity of a zone to the source of antibodies is also influenced.

DISCUSSION

We have found the fixe-mold assembly as easy to handle as
Ouchterlony's three-mold unit. With the former, simultaneous com-
parisons among a larger number of serological reagents, and the
use of various types of control solutions on the same plate as test
solutions, are possible. Grabar and Williams ( 1954 ) , have em-
ployed as many as six wells around the central well.

Diffusion of the solutions of antigens and antibodies from wells
of \'arious depths were tried. If the wells were less than 3 mm. deep
diffusion and evaporation occurred so rapidly that refilling was
needed at frequent intervals. Deeper wells, while making possible
the demonstration of the ribbonlike character of the zones when
they are viewed at an angle oblique to the vertical, have the dis-
advantage of introducing the problem of a discontinuous diffusion
of the solutions through the agar as a result of the lower portions
of the basins always being wet, and the upper portions occasionally
being dry.

The photograms proved to be more sensitive than the unaided eye
in detecting the presence and extent of the zones of precipitate.
For making quantitative measurements of the position of the zones,
the photograms have a distinct advantage over Ouchterlony's sug-
gestion of employing a graduated mechanical stage and a cross-hair
eye-piece. The positions of the zones may be precisely determined
from the photograms by superimposing on the prints a transparent
sheet of cellulose acetate paper on which a millimeter grid has been
ruled. A standard bacterial-colony counter with a modified grid
may also be used for this purpose. Almost unlimited magnification
of the plates can be obtained by using a transparency projector
such as a Vu-Graph (Charles Beseler Co., Newark, N. J.). This
machine will project an image of the plate to any desired size, and
at the same time, permit a critical focus of the zones. Large-scale,
accurate tracings of the zones can be made readily.

488 The University Science Bulletin

In our experience the practice of refilling the wells with serological
reagents resulted in patterns of precipitation which yielded more
information concerning the number of reactions going on in the
system, than was obtained in experiments where the wells were
filled just once. Variations in the results obtained between the
two types of tests were only those of degree of development of the
zones, and were not any of the effects which could be ascribed to a
discontinuous diffusion of reagent through the agar-medium. In
addition, we have found that, when solutions of antigen or anti-
serum were spilled on the agar, the entire plate can be liberally
flushed in cold tap water, drained, the wells blotted dry and refilled
immediately with the reagents without causing any detectable
changes in the patterns of precipitation. If the spilled reagents are
not quickly flushed away, precipitates will form on the surface of
the agar and these precipitates will mask the underlying zones of
precipitation and spoil the photographic qualities of the Petri plate;
the spilled reagent will distort also the shapes of the underlying
zones, by means of actual immunochemical interference.

An electrophoretic and ultracentrifugal analysis of the serum of
Cancer magister used in these experiments revealed that only two
component-hemocyanins were present. Hemocyanins are known
to be the only precipitin-inducing substance in the sera of Crustacea
(Tyler and Scheer, 1945). Oudin (1952) has concluded that the
appearance of a zone of precipitation in an agar-medium is an indi-
cation of a unique reaction between an antigen and its antibody.
The fact that more zones of precipitation occur (Pi. Ill, Fig. IF)
than can be accounted for in the serum on the basis of physico-
chemically detectable components requires some explanation.
Hooker and Boyd ( 1934) have shown that native proteins of a given
species have more than one kind of antigenic radical. Since the
most zones of precipitation in the agar-diffusion tests are obtained
when both of the reagents are in high concentration (PL III, Fig.
IF, PI. IV, Fig. IF), it is reasonable to assume that the several
zones are serological realities and not artifacts or phenomena analo-
gous to Liesegange rings.

In applying the Ouchterlony method to the analysis of complex
mixtures of antigens such as sera or extracts of organs care must
be taken that cross reactions are realities and not artifacts which
could result from the suprapositioning of unrelated serological re-
actions. Plate I, figure 4 (or PI. II, fig. 1) shows zones of precipita-
tion which overlap each other but which are the results of inde-

The Agar-Plate Precipitin Test 489

pendent serological reactions as evidenced by the intersection of
all the zones in the region between the c-well and the L-well.

From theoretical considerations (Oudin, 1952; Ouchterlony, 1953)
and the empirical knowledge of Ouchterlony (1948) and from
the data described above, the location and time of appearance of
a zone of precipitation in the agar in the Petri plate are dependent
upon the concentrations of the antigens and antibodies involved.
Dependent also upon concentration are the number of zones of
precipitation which appear, whether or not coalescence or inter-
section of zones can be realized, and other characteristics of the
zones such as their density and width. In tests involving compari-
sons of reactions of several related antigens with a single antiserum,
the most useful data would be obtained when the components
of the antigens were in equal and relatively high concentrations
unless the concentrations of the components are themselves a vari-
able under investigation.

SUMMARY

Modifications of Ouchterlony's Petri-plate, agar-diffusion method
for performing precipitin tests have been described. Minor changes
in the preparation of the agar, the use of plastic molds to form the
wells which receive the serological reagents, and the addition of
dyes to color the agar are discussed. A simple technique for making
permanent photographic records of the zones of precipitation is re-
ported. Homologous and cross reactions are illustrated. A manner
in which independent serological systems may approximate true
cross reactions in their appearances is illustrated. Photographic
records of the development of zones of precipitation using the a-
and /3-precipitin tests are presented.

LITERATURE CITED

BjORKLUND, B.

1952a. Specific inhibition of precipitation as an aid in antigen analysis

with gel diffusion method. Proc. Soc. Exp. Biol, and Med., vol. 79,

pp. 319-324.
1952b. Serological analysis of components in hemopoietic tissue. Proc. Soc.

Exp. Biol, and Med., vol. 79, pp. 324-328.
Grabar, p., and C. Williams

1954. A report before the 10th Annual Conference on Protein Metabolism,

Rutgers University, New Brunswick, New Jersey. Jan. 29-30.
Hooker, S. B., and W. C. Boyd

1934. The existence of antigenic determinants of diverse specificity in a

single protein. II In two natural proteins; crystalline duck egg

490 The University Science Bulletin

albumin and crystalline hen egg albumin. J. Immunol., vol. 26, pp.
469-479.
Leone, C. A.

1952. Effect of multiple injections of antigen upon the specificity of
antisera. J. Immunol., vol. 69, pp. 285-295.

OUCHTEKLONY, O.

1948. Antigen-antibody reactions in gels. Arkiv. Kemi. Min. och Geol.,
vol. 26B, pp. 1-9.

1953. Antigen-antibody reactions in gels. IV Types of reactions in coor-
dinated systems of diffusion. Acta Path, et Microbiol. Scandinav.,
vol. 32, pp. 231-240.

OUDIN, J.

1952. Specific precipitation in gels. Methods in Medical Research., vol. 5,

pp. 335-378.
Tyler, A. and B. T. Scheer

1945. Natural heteroagglutinins in the serum of the spiny lobster Panulirus

interruptus. II Chemical and antigenic relation to blood proteins.

Biol. Bull., vol. 89, pp. 193-200.

PLATE I

A two-dimensional agar-diffusion test illustrating the development of coalesc-
ing and intersectiong zones of precipitation during 7 days of diffusion-time.
The peripheral wells contained the hemocyanins of Callinectes (c), or Limulus
(l), or a mixture of both of them (cl). The central well (xcl) contained
antiserum which was produced against the mixture of antigens. The inter-
section of all the zones of precipitation between well c and l indicate that
no antibodies were present which could combine with both antigens. Magni-
fication of illustrations: X 1.

The Agar-Plate Precipitin Test

491

PLATE I

492 The University Science Bulletin

PLATE II

Fig. 1. A duplicate of the reaction pictured in Plate I with the exception
that these wells were not refilled with reagents. In the regions where they
overlap each other, the zones are more sharply defined than in Plate I. The
zones were not developed as much in the regions of coalescence and intersec-
tion.

Fig. 2. The appearance of homologous and cross reactions between tlie anti-
serum (xm) produced against the hemocyanins of Cancer magister and the
sera of four closely related species of crabs: Cancer magister (m), C. pagurus
(p), C. prodtictus (d) and Carcinus maenus (e).

Figs. 3 and 4. A duplicate of the ^-precipitin test which is illustrated in
Plate IV with the exception that the wells were not refilled with the reagents.
The doubling-dilutions of antiserum were placed in sequence in the peripheral
weWs and the antigen was placed in the central well of each plate.

Fig. 5. An illustration of the 'lialo" effect which is obtained in photograms
when an antiserum containing hemoglobin ( central well ) difi^uses into agar
which has been colored with methyl orange.

Fig. 6. A photogram with "halos" obtained from four antisera (peripheral
wells) containing hemoglobin. Agar was colored with methyl orange. The
numbers in the wells are prints of the mirrored writing on the undersurface of
the Petri plate.

Magnification of illustrations: X 1-

The Agar-Plate Precipitin Test

493

PLATE II

494 The University Science Bulletin

PLATE III

A record of the zones of precipitation obtained in the agar from a standard
a-precipitin test with a hemocyanin-antihemocyanin system. Photograms of
the Petri plates are so arranged that the development of the zones for any par-
ticular dilution of antigen may be compared in the horizontal rows of figures,
and the development of the zones for all of tlie dilutions of antigen, at any
given time, may be compared in the columns of figures. The doubling-dilution
series of antigen begins in Well No. 1 with one part protein per 15.625 parts
saline, and ends in Well No. 16 with a dilution of 1:256,000. Antiserum is
located in the central well (xm) of each plate. The serological system is the
hemocyanin in the serum of the crab Cancer magister and its homologous
antiserum. Magnification of illustrations: X '2.

The Agar-Plate Precipitin Test

495

PLATE III

496 The University Science Bulletin

PLATE IV

A record of the zones of precipitation obtained in agar from a standard
^-precipitin test with a hemocyanin-antihemocyanin system. Photograms of the
Petri plates are so arranged that the development of zones for any particular
dilution of antiserum may be compared in the horizontal rows of figures, and
development of the zones for all of the dilutions of antiserum, at any given time,
may be compared in the columns of figures. The doubhng-dilution series of
homologous antiserum begins with undiluted antiserum in Well No. 1 and ends
in Well No. 8 with a dilution of 1:128. A 6.0 percent solution of the hemo-
cyanins of Cancer magister is situated in the central well ( M ) . Magnification of
illustrations: X 2/3.

The Agar-Plate Precipitin Test

497

I DAY

PLATE IV

2 DAYS

3 DAYS

THE UNIVERSITY OF KANSAS

SCIENCE BULLETIN

Vol. XXXVII, Pt. I] October 15, 1955 [No. 13

Additions to the Known Herpetological Fauna of Costa
Rica with Comments on Other Species, No. II

By Edward H. Taylor

Abstract: This paper treats of six caecilians, seven Salientia, ten lizards,
and five snakes from Costa Rica. Of this number the following are described
as new: Dermophis costaricense, Dermophis occidentalh, Dermophis glandii-
losus, Eleutherodactyhis n^innis, Eleutherodactylus vocator, Agahjchnis saltator,
Anadia metallica arborea, Anadia metaUica attenitata, Leposoma southi orient-
alis, Lepidophijma reticulatum, Lepidophyrna anomalum, Lepidophyma ophi-
ophthaJmum.

Atelopus varius loomisi is elevated to species rank and the following are
added to the list of Costa Rican species: Dermophis parviceps, Eumeces mana-
guae, Leptotyphlops ater.

A fourth journey to Costa Rica was made by me in the summer
of 1954. Much of my effort in collecting was made along the
Panamanian border of Costa Rica and along the Nicaraguan border
in the north. Political difficulties between the two latter countries
prevented me from carrying out plans for more extensive collect-
ing in the Caribbean drainage area of the northern border.

Accompanied by Prof. Marco Tulio Pacheco I visited the town
of Tilaran and collected chiefly in an area to the northeast at Finca
Bosco and Lake Arenal. One trip was taken along the Pan-American
Highway to the northern border. One journey was made to Cocos
Island, another to the upper reaches of the Saripiqui river.

For the most part my efforts yielded gratif>dng results. However,
an unknown Teiid lizard, seen at an elevation of about 2,425
meters on the west slopes of Cerro de la Muerte, escaped from
under my hand, leaving only a wiggling tail as evidence that
such a creature occurs. Numerous toad voices of a species not
identified, but believed to be a species of Microhyla, were heard

(499)

500 The University Science Bulletin

at Liberia in late August, but none could be found in the lush
vegetation growing in the pools from whence the calls issued.

Despite the collections I have made on four journeys much work
must be done before the entire herpetological fauna is collected;
and before the details of the distribution of the various species
within the country can be authoritatively known.

I wish to acknowledge my grateful thanks to Professor Marco
Tulio Pacheco for very numerous specimens and for the assistance
rendered by him and members of his family; and to Dr. and Mrs.
Leslie Holdridge who were my hosts for a part of the summer.

The following genera and species are treated in this paper:

PAGE

Dermophis Peters 501

Dermoi3his parviceps Dunn 502

Dermophis occidentalis sp. nov 503

Dermophis costaricense sp. nov .506

Dermophis glandulosus sp. nov • 509

Gymnopis Peters 510

Gymnopis multiplicata multipHcata Peters 511

Gymnopis multiplicata proxima (Cope) 511

Atelopus varius varius ( Lichtenstein and von Martens) 513

Atelopus senex Taylor 515

Atelopus loomisi Taylor 517

Eleutherodactylus mimus sp. nov 517

Eleutherodactylus vocator sp. nov 522

Eleutherodactylus diastema ( Cope ) 525

Agalychnis saltator sp. nov 527

Eumeces Wiegmann 531

Eumeces managuae Dunn 531

Anadia Gray 534

Anadia metallica metallica ( Cope ) 535

Anadia metallica attenuata subsp. nov 537

Anadia metallica arborea subsp. nov 542

Leposoma southi orientalis subsp. nov 546

Lepidophyma A. Dumeril 547

Lepidophyma flavimaculatum flavimaculatum A. Dumeril 549

Lepidopliyma flavimaculatum obscurum Barbour ( not Costa Rican ) 550

Lepidophyma reticulatum sp. nov 551

Lepidophyma anomalum sp. nov 552

Lepidophyma ophiophthalmum sp. nov 558

Anomalepis dentatus Taylor 562

Leptotyphlops ater Taylor 562

Conophis lineatus nevermanni Dunn 563

Leptodeira sp 565

Coniophanes piceivittis piceivittis Cope 566

Neopareas bicolor Giinther 568

Additions to Fauna of Costa Rica 501

CAECILIIDAE

Two genera of this order, Dcrmophis and Gyninopis, are rec-
ognized in Costa Rica. These genera may be separated by the fol-
lowing key:

Eye present, not covered by bone, usually visible through the skin. One
mandibular tooth-series ( inner lacking ) Dermophis

No visible eye, the eye wholly or partially covered by bone in the fully
grown specimen; two mandibular tooth-series, the inner consisting of
one or more teeth ( splenial ) on each side Gymnopis

Genus Dermophis Peters

Dcnnnphis Peters, Monatsh. Akad. Wiss. Berlin. 1879, p. 937 (no type men-
tioned); Boulenger, Catalogue of the Batrachia Gradientia s. Caudata and
Batrachia Apoda in the collection of the British Museum, 1882, p. 97 (re-
peats in English Peters' definition. Includes six species ( one of which is
African), some of which have the inner mandibular series, others do not);
Noble, Bull. Amer. Mus. Nat. Hist., vol. 49, no. 11, 1924, p. 305.

Gymnopis Dunn (part.) Bull. Mus. Comp. Zool. Harvard Coll., vol. 91, no. 6,
1942, p. 461 (combines Dermophis, Gymnopis and Cryptosophis).

The original definition of Dermophis Peters reads as follows:
"Aiigen sichtbar. Tentakelgrube kreisformig vor und unter dem
Auge, Tentakel kugelformig; Korper mit dachziegelformigen Schup-
pen. UnterkieferzJihne in zwei Reihen." "America und Afrika."
Peters listed under the genus, Dermophis mexicanus, proximus,
simus, hrasiUensis, brevirostris, and thomensis. No type was des-
ignated but Dr. G. K. Noble, in 1924, named mexicanus the type,
perhaps without being aware that mexicanus did not agree fully
with the generic description. Dunn, 1942, likewise seemingly was
unaware of this.

Peters stated that the genus had two mandibular tooth-rows but
included mexicanus that has only a single row. Since the latter has
becoiue the type of the genus by designation, it is necessary to cor-
rect the generic definition or by a suspension of rules designate a
type that has two rows — for example, proximus. Certainly less
confusion exists by redefining the genus as having no inner mandib-
ular tooth-series, thus excluding those species having an inner man-
dil)ular tooth-series. Were it possible to designate a new t\'pe of
the genus and proxima were designated, then Dermophis and
Gymnopis would become synonyms since muhipUcata and proxima
are congeneric, and a new generic designation would be necessary
for the group of species having an eye ( usually visible ) not covered
by bone and the single mandibular tooth-series. Other workers
have not solved the problem of throwing the two genera together.
If they have found forms that differ in the structures of the skulls

502 The University Science Bulletin

with relation to the eye, they have probably overlooked or disre-
garded the possibihty that still another genus was involved.

It may be that Gymnopis midtiplicata oaxacae with a visible eye,
should be regarded a Dermophis. Dunn is somewhat ambiguous
regarding the presence of the inner mandibular row of teeth. In
his general discussion he says of the genus Gymnopis (combined
Dermophis and Gymnopis) "the inner mandibular tooth row is
poorly developed or absent (p. 441)". "In Gymnopis and in
Siphonops the teeth on the lower jaw are uniform but larger than
those on the upper. The inner mandibular row is reduced to one
tooth on a side (in oligozona and in multiplicata) or is entirely
absent." In the discussion of individual species he states: "Forms
with visible eyes have on the whole, no teeth in the inner mandib-
ular row." Under Gymnopis multiplicata oaxacae Dunn gives
"inner mandibular teeth" as a character. One of the specimens
listed by Dunn (EHT 16869) has no trace of the inner mandibular
teeth. Mertens does not mention the character. Should it prove
that others of tlie series referred to oaxacae have a visible eye not
roofed by bone (as in EHT 16869) and lack the second series of
mandibular teeth they should be referred to the genus Dermophis
as a distinct species.

Dermophis parviceps Dunn

Siphonops parviceps Dunn, Occ. Papers. Boston See. Nat. Hist., vol. 5, July
9, 1924, pp. 93-94 (type locality. La Loma, Bocas del Toro Province, Pan-
ama, elevation 1,200 ft. [not 2,000 ft. as originally stated]); Dunn, Proc.
New England Zool. Club., vol. 10, Oct. 11, 1928, pp. 74-75.

Gymnopis parviceps Dunn, Bull. Mus. Comp. Zool. Harvard Col., vol. 91, no.
6, Dec. 1942, pp. 462, 478-479.

Diagnosis: Eye visible; lacking the inner mandibular teeth; pri-
maries reduced, 96-97; secondaries 13-15; 5-10 complete; tentacle
close to lip and nearer eye than nostril; ratio of diameter of body to
total length, 21-24; probably under 200 mm. in length.

Description: (From KUMNH No. 36276, Moravia de Chirripo,
Limon Province, Costa Rica). Head oval, slender, the width of
head (5 mm.) at mouth angle, 'distinctly less than width of neck
(7 mm.) or body (8mm.); head length to first transverse nuchal
groove (dorsally) 7.3 mm.; head to angle of mouth 4.5 mm.; width
of body in total length (191 mm.), 21.3 times; tentacle close to lip,
its distance from eye ( 1.25 mm. ) less than its distance from nostril
(2mm.); distance between eye and nostril 3 mm.; snout extending
beyond mouth 1 mm.; distance between eyes, 3.8 mm.; first nuchal
groove passes across head then runs down and somewhat forward,

Additions to Fauna of Costa Rica 503

crossing the chin 6.2 mm. back from tip of snout; second groove
crosses throat 3.3 mm. back of first; third groove separated on throat
from second by 4 mm.; each of the annular folds, formed by the
three grooves, with secondary grooves on dorsal surface only; on
each side of the neck, beginning on back part of head, a longitudinal
lateral fold limited above by a slight longitudinal groove, and ex-
tending back to near end of body, the ventral coloration covering
the anterior part of fold. Palatal region rather short; maxillary-pre-
maxillary tooth-series total 30; vomero-palatine series 28; mandib-
ular teeth total 24; no inner mandibular (or splenial) teeth; outer
mandibular teeth larger than maxillary or other teeth above; pri-
mary grooves 97; secondaries 14, v^dth five of the latter complete;
vent with a primary groove entering its sides, its anterior edge with
two large lobules; a pair of narrow lateral lobules and five lobules
bordering posterior part of vent.

Color in life: Above deep plumbeous lavender with some scat-
tered irregular whitish spots or flecks, becoming lighter at the be-
ginning of the lateral ridge; lighter grayish-lavender low on sides
and venter; head pinkish flesh. In preservative head has become
brownish gray, a slightly darker tone than the venter; ventral colora-
tion limited by this lateral fold or ridge except for the first three
annuli on the fold.

Measurements in mm.: Total length, 192 (180); snout to vent,
191; head width, 5(5); width of neck, 7; width of body, 8(8); head
to first annular groove, dorsally, 7.3; head to angle of mouth, 4.5;
width of body in total length, 21 times; tentacle from eye, 1.25;
tentacle to nostril, 2; between eye and nostril, 3; length of snout be-
yond mouth, 1.1. (Measurements in parenthesis those given for the
type.)

Dermophis occidentalis * sp. nov.

(Fig. 1.)

Type: KUMNH No. 36298, collected on the Dominical Road ap-
proximately 15 km. WSW of San Isidro del General in open meadow
under stones, Aug. 3, 1954; Edward H. Taylor, coll.

Paratypes: KUMNH No. 36296, 36297, topotypes, taken July 19,
1952, same collector.

Diagnosis: A presumed diminutive species with secondaries 29-
37; the primaries 112-113; secondaries completely surrounding body
10-12; length 186-204; body width in length 30-33.

* Latin, occidentalis =: western.

504

The University Science Bulletin

Fig. 1. Dermophis occidentalis sp. nov. Type. KUMNH No. 36298, 15
km. WSW San Isidro del General, San Jose Province, Costa Rica. Total length,
186 mm.

Additions to Fauna of Costa Rica 505

Description of the type: Body slender, its width (5.9 mm.) con-
tained in body length ( 186 mm. ) 31.5 times; head narrow, oval, its
width at mouth angle, 5 mm.; tentacle separated from mouth by
.5 mm.; eye from mouth .92 mm.; nostril to eye 2 mm.; eye to tenta-
cle .75 mm.; nostril from tentacle 1.5 mm.; snout projecting beyond
mouth 1.1 mm. Snout to first annular groove 5.6 mm.; snout to
third annular groove 10.5 mm., dorsal measurement (ventral meas-
urement 10 mm.); primary annular grooves, 112 to vent, followed
behind vent by an annular fold partly divided; secondary annuli,
28; secondary annuli completely surrounding body, 9; maxillary-
premaxillary teeth (total) 26; vomero-palatine teeth 22 (total);
outer mandibular series (total) 21; inner mandibular series absent.
The vent is transverse, bordered anteriorly and posteriorly by several
fleshy denticulations.

Scales are first found on the posterior part of the body along the
primary groove following the beginning of the secondaries. At
first the scales are small, forming short transverse rows wider than
long ( .4 X .9 mm. ) ; they form a strongly overlapping series, usually
only one row discernible at first; farther back the scales are much
larger, arranged in at least three overlapping transverse rows in
each annulus (posterior to the secondary groove and posterior to
the primary groove a series of three or four similar transverse rows ) .
These follow the grooves completely around the body. Seen in
lateral view when one cuts through the skin, the scales of the back
part of the preceding series overlap those of the following series,
although they are separated by a series of recumbent glands which
open near the groove.

Scales on the posterior part of the body are varied in size, but
usually are considerably larger than those occurring anteriorly,
some measuring 1.5 mm. x 1.3 mm. Some are cycloid or subcy-
cloid but many of these are more or less truncate anteriorly.

Color in life: Head, on top and sides, light grayish tan; body
plumbeous lavender; entire underside a lighter shade growing
still lighter under neck; chin and throat lighter than top of head;
area about vent dull cream.

Measurements in mm. and data on type and paratypes: Nos.
36298, 36296, 36297, respectively; total length, 186, 194, 204; width
of body, 5.9, 6, 6.8; snout to first annular groove (dorsal), 5.6, 5.8,
5.7; snout to third annular groove (dorsal), 10.5, 10.6, 10.4; primary
annular grooves, 112, 112, 112; secondary annular grooves, 28, 37,
33; complete secondary grooves, 9, 10, 12; maxillary-premaxillary

506 The University Science Bulletin

teeth (total), 26, 26, 26; vomero-palatine teeth (total), 22, 26, 26;
outer mandibular teeth (total), 21, 21, 21; inner mandibular, 0, 0, 0.

Variation: The two paratypes agree well with the type. The
eye is dim in No. 36296, but can be discerned at least on one side.
The specimens were taken from under rocks or logs in a meadow.
Two other species of caecilians occur in this same general area.

Dunn summarily states that "there are never more than two rows
of scales or rings to a segment (in Rhinatrema, Gijmnopis and
Caecilia) and that there is no overlapping of the scales of one
ring by those of another."

Nieden has mentioned that several rows may occur in an annulus
and that overlapping may occur, both of which statements are
true, at least in certain forms.

Dermophis costaricense * sp. nov.

(Fig. 2)
Type: KUMNH No. 36343. Collected, Cinchona, Heredia Prov-
ince, Costa Rica, July 14, 1954, by Edward H. Taylor.

Paratypes: KUMNH Nos. 36337-36340, 36342, 36344-36347, all
topotypes; July and August, 1954; Edward H. Taylor, coll; Nos.
36447-36448, Moravia de Chirripo, Limon Prov. Costa Rica.

Diagnosis: A Dermophis related to, but differing from mexicanus
in having a higher average number of primaries 112-119 instead of
97 to 100; secondaries 80 to 93 instead of 51-80; total primaries
and secondaries, 192-203 instead of 152-186.

Description of the type: Body not especially slender, its width
(11 mm.) in total length (330 mm.) 30 times; head oval, its length
from first annular groove 12 mm.; to third annular groove 22 mm.
(dorsal measurement); when measured on ventral surface the
measurements are 10 mm. and 20 mm. respectively; eye visible,
not covered by bone, forming a slightly elevated mound, its distance
from tip of snout 5.4 mm.; distance between eye and nostril 4 mm.;
distance between edge of eye and tentacle 1.9 mm.; between
tentacle and nostril 2.8 mm.; tentacle surrounded by a horseshoe-
shaped groove, fastened behind to surface skin; angle of mouth
to snout tip, 20.5 mm.; primary annular grooves (counting the
three grooves of the first two modified annular folds) 113; second-
aries (beginning on the 31st annular fold) 83; two partial annuli
posterior to vent; first two modified folds each with a transverse
groove which is to be interpreted as incomplete secondary folds,
but which are not counted.

* Of Costa Rica.

Additions to Fauna of Costa Rica

507

Fig. 2. Dermophis costaricense sp. nov. Type. KUMNH No. 36343,
Cinchona, Heredia Province, Costa Rica. Actual total length 330 mm.

508 The University Science Bulletin

Scales begin as far forward as the 18th primary in the dorsolateral
part of the annulus, and continue to the end of body, the trans-
verse series growing longer posteriorly; in each annular fold there
are four, sometimes five, rows of scales of varying size, contiguous
or overlapping laterally, but each strongly overlapping the more
posterior series. The scales usually present curved or rounding pos-
terior edges, while the anterior edge is usually truncate.

The ventral surface of the body is more or less flattened, the
posterior ventral region especially so; the vent shows two larger
fleshy "denticulations" with a smaller median one; anterior to
vent, laterally and posteriorly, there are 6 "denticulations," the sur-
faces of the "denticulations" showing a few indistinct papillae.

Maxillary-premaxillary teeth total 38, the maxillaries much larger
than the premaxillaries; vomero-palatine series 38, much smaller
than the premaxillaries; outer mandibular series 28, the teeth un-
equal in size, the largest double the size of the largest maxillaries;
no inner mandibular tooth series.

Color in life: Above dark plumbeous, the head showing some
lighter variegations; tentacular area and area about nostril lighter;
ventral region somewhat lighter than dorsum; region of vent,
cream; an indefinite lighter region under neck.

Measurements in mm.: Total length, 330; width of body, 11;
width of head at mouth angle, 8.9; width of head at first annular
fold, 9.1.

Variation: A series of paratypes from within a few hundred
yards of the exact type locality shows some variation in the propor-
tion of body width to body length, as well as certain variation in
the number of primaries, secondaries and the number of complete
secondaries.

The table on page 509 indicates much of this variation.

Remarks: This is not regarded as a subspecies of mexicanus since
Dermophis mexicanus occurs from north central Veracruz to Pan-
ama (fide Dunn) and gracilior a subspecies likewise in Panama
presumably shows closer relationship to mexicanus than does this
species. Since mexicanus also occurs on the Caribbean drainage
north of the Cordillera Central and on the eastern slopes of the
Talamanca Mountains, it is probable that the two ranges overlap
considerably. A specimen of M. mexicanus has been taken on the
northeastern slope of Volcan Poas, within the probable range of
this species.

Additions to Fauna of Costa Rica
Measurements and data on Dermophis costaricense sp. nov.

509

1

IS

J

E

•i

us

•B
C

o

IB

SO

^^

el

O

Maxillary,
premaxillary
teeth

, c

>

c

OS

Inner

mandibular

a

T3 a>

36447

111

113
114
115
111
117
114
117
113
113
115
113
113

90
83
79
87
89
75
86
95
59
93
88
81
83

13

12

11

10

10

9

12

14

8

11

9

9

8

0
0
0
0
0
0
0
0
0
0
0
0
0

387
330
306
290
305
305
282
275
265
260
212
195
170

15
11

10.5
10.8
10.6
10.5
9
8

8.2
8.8
9.8
7.6
6.7

26-1-

36343

36340

38
42
41
35
38
31
38
33
35
32
32
30

38
46
42
42
40
39
40
36
37
38
37
32

28
30
28
26
28
28
28
24
24
26
26
24

30

29 +

36337

27 —

36448

28 +

36344

36342

36347

36341

29 +
31—
34 +
32

36338

36346

36345

30—

27 +
26 —

36339

27—

Nos. 36337-36347 Cinchona.

Nos. 36447-36448 Moravia de Chirripo.

Dermophis costaricense ranges in altitude from about 1,900 feet
at Moravia to approximately 5,000 feet at Cinchona.

Dermophis glandulosus * sp. nov.

Dermophis mexicanus gracilior Taylor, Univ. Kansas Sci. Bull., vol. 34, pt. 2,
Feb. 15, 1952, pp. 781-784, pi. 88.

Type: KUMNH No. 29979, San Isidro del General, San Jose
Province, Virgil Cave collector.

Diagnosis: Eyes distinct, the tentacle closer to eye than nostril;
93 primaries (not counting nuchal grooves); 49 secondaries; no
inner mandibular tooth-series; grooves throughout most of body
showing a series of enlarged glands. Dorsum purplish lavender,
somewhat darker on outer edges; lateral fold lighter lavender; ven-
tral surfaces dull cream dimly clouded with lavender; 30 teeth in
the maxillary-premaxillary series, the premaxillary teeth larger than
others; prevomerine-palatine series total 40, the teeth small; outer
mandibular row, 26, the teeth unequal in size. 22 of the 46 body
secondaries encircle the body. Total length of type 159 mm.

This specimen was referred by Taylor to Dermophis mexicanus
gracilior with some hesitation in the above cited work. Further
study of this genus suggests that it is not mexicanus and is here
given a different specific designation. Since the type was described
at length in the above publication, it is not repeated here. The
accompanying photograph shows something of the curious color
pattern and the size and character of the enlarged annular series
of glands.

* Latin, glandulosus — glandulose.

510

The University Science Bulletin

Fig. 3. Dermophis glandulosus sp. nov. Type, KUMNH No. 29979, San
Isidro del General. San Jose Province. Costa Rica. Actual length, 159 mm.

type of genus

Genus Gymnopis Peters

Gymnopis Peters, Monatsb. Akad. Wiss. Berlin, 1874, p. 616
Gijmnopis multiplicata Peters ) .

The genus Gymnopis was proposed by Peters in 1874, with a
very brief description and a single species muHipUcata (which
becomes the type) was described. The genus was described in
1879 as follows: "Augen unter den Schadelknochen gelegen Ten-
takelgrube etwas hoher, sonst wie bei SipJwnops, weit hinter dem
Nasenloch gelegen, ringformig das kugelformige Tentaculum
einschliessend. Korperhaut mit dachziegelformig gelagerten Schup-

Additions to Fauna of Costa Rica 511

pen. Unterkieferziihne in zwei Reihen. Ini Schadelbau Sipho-
nops naher stehen."

Gymnopis multiplicata multiplicata Peters

Gijmnopis multiplicata Peters, Monatsb. Akad. Wiss. Berlin, 1874, p. 616, pi. 1,
fig. 1 (type loeality, Vcragua, Panama).

The follo^^'ing specimens are in the collection, previously unre-
ported: KUMNH No. 36667, 5 km. NE Tilaran, Guanacaste Prov-
ince; No. 36668, San Isidro del General; No. 36670, 15 km. WSW
San Isidro del General, Puntarenas Province?; Nos. 33669, 36671,
5 km. E (on highway) of San Isidro del General, San Jose Province.

In all the specimens the eyes are concealed under the bony sur-
face of the skull, and the inner mandibular tooth row is present.
Specimens from higher elevations have, on the average, fewer
primaries (7-10) and a concomitant reduction in secondaries.
Older, larger specimens (usually) have more teeth than young,
smaller specimens, thus approaching the counts for multiplicata
proxima — in fact, there seems to be an overlap in the count in
certain specimens. The range of primaries in these specimens is
121-132, of secondaries 98-114, while in the specimens of proxima
the range is 112-121, 87-102 respectively (see accompanying table
of data and measurements).

Gymnopis multiplicata proxima (Cope)

Siphonops proximus Cope, Proc. Amer. Philos. Soc., vol. 17, 1877, p. 90 (type
locality, eastern Costa Rica.) [^Limon, fide Dunn.]

The following specimens are in the collection: KUMNH No. 36675
Cariblanco, Limon Province; Nos. 36672, 36673, 36674, 36676, 36678
Turrialba, Cartago Province; No. 36677 Los Diamantes, Limon
Province. The accompanying table shows much of the variation
occurring in this series.

The scales in both subspecies usually appear six to eight folds
in advance of the beginning of the secondaries. There are several
transverse rows (three or four) of scales in each segment. These
are contiguous transversely (or somewhat overlapping) and each
row imbricates broadly with the following rows. The scales of
one segment somewhat overlap those of the following segment
in the posterior part of the body, but are separated by a row of
elongated recumbent glands.

At birth, the maxillary-premaxillary tooth-series has appeared
and 14 teeth are in evidence on each side. The vomerine and
palatine teeth are differentiated. There are five or sLx palatine
teeth protruding through the gums. Except for a large, median.

512

The University Science Bulletin

hooked tooth in the mid-Hne of the palate the vomerine teeth have
not appeared above the gums. Subsequently this tooth is probably
shed since a differentiated median tooth is not evident in older
specimens. In the lower jaw the dentaries have four longer or
shorter rows of teeth, irregularly disposed, perhaps 20-25 on each
side; an inner mandibular series is not developed at this time.
The dentary teeth of these juveniles do not resemble those of the
adults. They are somewhat spatulate, each bearing several, fine,
clawlike denticulations. A few of them have not yet risen above
the surface of the gums but most of them are well above the
surface. The two specimens showing this condition are RCT Nos.
331-332. One, 331, was taken in a small pile of rocks with a large
adult specimen of Gymnopis multiplicata proximo; the other. No.
332, was in the uterus of the same adult specimen. These measure
129 and 125 millimeters respectively. The arrangement of the
mandibular teeth is reminiscent of the condition obtaining in certain
adult Ambystoma — Amhystoma schmidti and young Anibystoma
texanurn.

I find it difficult to account for the development of these special
embryonic teeth. Two explanations are suggested: one that they
represent a recapitulation of an ancestral condition, the other that
they subserve some intra-uterine function.

The median hooked tooth on the vomer is not present in the
intra-uterine specimen.

Gymnopis multiplicata multiplicata Peters

t

u

u

>>

CO

>1 CO

ete

ndary

ves

S =3

2.1

i-'-S

bO

c

^

i

s

"5

O

is

a 0

O t.

ompl
seco
groo

eeth,
max
pren

eeth,
vom
pala

eeth,
oute
man

eeth,
inne
man

C3

Z

^i

Oh

a:

O

H

H

H

H

J

rt

36667

Tilaran

122

101

13

43

47

36

2

384

16

24

36668

San Isidro. .

132

114

16

40

42

36

2

348

13

26.7

36669

5 km. E

San Isidro. .

121

98

8

38

44

33

2

323

9 8

32

36670

15 km. wsw

S»,n Isidro. .

129

110

14

39

40

27

2

244

5.8

42

36671

5 km. E

San Isidro. .

122

102

10

30

30

24

2

148

4.8

31

Gymnopis multiplicata

proxima (Cope)

36672

Turrialba. . .

118

96

12

40

48

36

2

394

16.8

23.5

36673

Turrialba. . .

118

95

9

50

54

38

2

391

17

23

36674

Turrialba. . .

112

87

9

42

48

34

2

360

18

20

36675

Cariblanco. .

116

90

13

45

52

36

(?)

344

(19?)

29

36676

Turrialba. . .

119

93

10

39

38

30

2

279

13

21

36677

Los

Diamantes. .

121

102

11

34

34

26

2

216

7

31

36678

Turrialba. . .

116

97

10

33

32

22

2

179

7.8

22

Additions to Fauna of Costa Rica 513

SALIENTIA

Ateloptis varius varius ( Lichtenstein and von Martens)

(Fig. 4)

Phrijnidium varium Lichtenstein and von Martens, Nomenclator Reptilium et
amphihiorum Musei Zoologici Berolinensis, 1856, pp. 40, 41 (type locality,
Veragoa = Veragua, Panama).

This brilliantly colored Ateloptis, black with markings of red and
yellow above, was taken at several localities as follows (the first
three localities are in the Caribbean, the last four in Pacific drainage
areas ) : No. 32894, Moravia de Chirripo, Limon Province; No.
30299, a problematical specimen from Ujarras or Navarro (M.T.
Pacheco, coll.), No. 32885 Guabata, Mt. Paloma. Nos. 32863-32879,
15 to 20 kms. WSW of San Isidro del General; No. 32894, 4.5 km. E
San Isidro del General, along highway; Nos. 32880-32883 near
Sarchi, San Jose Province; Nos. 32886-32888 Palmar, Puntarenas
Province. Two specimens, one, No. 32880, a large female from near
Sarchi, and a male, No. 32866 from WSW of San Isidro del General
are figured. All of the specimens from the Pacific drainage area, ex-
cept one, agree in the general structural and color characteristics.
Most of them are males and smaller than the females.

The dorsal colors are red or red-orange (rarely pinkish), with
one or more mesial marks of bright yellow (rarely absent), and
other reddish marks present on the limbs; the tips of the digits
are yellow, the venter and chin light yellow-flesh, and an area on
soles and palms somewhat orange-yellow. A few specimens have
one or more small black ventral spots. Both lips are immaculate
yellow and the area below the vent is colored like the venter.

The specimen from 4.5 kilometers east of San Isidro del General
differs in having the dorsal markings very narrow, more numerous,
and the venter has numerous, heavy, rounded or vermiculate marks
on a yellow or pinkish-yellow venter. Both lips are blackish. The
webbing on the feet is a little more extensive.

The specimens from the Caribbean drainage area do not agree
among themselves and two may represent subspecific variants.

No. 32885 agrees in considerable detail with the Pacific group
save that the lips are black and there are six rounded black spots on
the venter.

No. 32894 has the venter unspotted save for three small spots
on the breast and one on the chin. The lower lip is bordered with
black and the upper has some dim black marks. The general color
pattern shows the dorsal reddish marks narrowed and the middle

17—6730

514

The University Science Bulletin

Fig. 4. Atelopus varitis varius ( Lichtenstein and von Martens ) . Upper fig.
KUMNH No. 32866 i ; snout to vent, 30 mm.; 15 km. SW San Isidro del Gen-
eral, San Jose Province. Lower figure, KUMNH No. 32880 9; length, 43 mm.;
near Sarchi, San Jose Province.

Additions to Fauna of Costa Rica 515

of the back has numerous yellow dots. There is a large black area
below the vent on the thighs.

No. 30299 may well represent a different form since the snout is
more angulate and pointed, more flattened and projecting in pro-
file. Much of the area under the thighs is black and four large
rounded black spots are present.

This specimen from an uncertain locality (Ujarras or Navarro)
was collected by, and presented to me by Prof. Pacheco. No color
notes were taken and I do not know what the color may have been
in life. It is probably from an elevation higher than those I have
collected. I suspect its relationship may be with loomisi and may
represent it in the region where variiis occurs.

Atelopus scncx Taylor

(Fig. 5)

Atelopus senex Taylor, Univ. Kansas Sci. Bull., vol. 35, pt. 1, July 1, 1952, pp.
630-631 (type locality near the divide, between Volcan Poas and Volcan
Barba, on western slope of Volcan Poas, Pacific drainage, elevation about
6,800 ft., Costa Rica).

Atelopus senex was described from a single specimen in the Rich-
ard Clark Taylor collection. In 1952 I acquired three specimens,
KUMNH Nos. 32314-32316 from Cuabata, Mt. Palomo, Costa
Rica, collected by Prof. Marco Tulio Pacheco; and in 1954 another
series of live specimens, KUMNH Nos. 35914-35922, from Navarro,
Cartago Province, Costa Rica, presented to me by Prof. Pacheco.

The structural characters of these specimens agree well with those
given for the type; however the color patterns differ much among
themselves as well as from the color characters of the type.

The specimens have in common the following characteristics from
which they differ from other known Costa Rican forms.

The skin on the head and most of the dorsal surface is smooth
(lacking minute spicules, visible under magnification); the skin on
snout (sometimes behind snout) shows one or more fine longi-
tudinal grooves, none of which are in straight lines; the skin above
the "supratympanic" ridge bears a paratoid thickening, cream white
in color; the type and all of the specimens from Navarro have the
venter dull gray with some indefinite lighter areas. However the
three specimens from Cuabata, Mt. Palomo, have the venter lighter
with discrete black spots or flecks. The webbing on the feet reaches
the terminal discs on all save the fourth toe and is not or but very
slightly incised between. The three Cuabata specimens are illus-
trated by a photograph showing the dorsal coloration.

516

The University Science Bulletin

Fig. 5. Atelopu.s senex Taylor. Upper figure, KUMNH No. 32314 $ , snout
to vent length, 44 mm. Lower left figure, No. 32315 S , length, 32 mm. Lower
right figure. No. 32316 $ , length, 33 mm. All from Guabata, Mt. Palomo,
Costa Rica.

The Navarro specimens are for the most part variegated, dark gray
above, often with dim light Hnes following the sides (seen from
above), and one along the anterior part of the median line; some
have guanine flecks or spots, and one specimen ( No. 35916 ) has the
top and sides of the head, and almost all of the dorsum and sides

Additions to Fauna of Costa Rica 517

of body covered with large cream blotches; a few even appear on
limbs. Another shows similar scattered spots and flecks only on
head and shoulders. The guanine spots may be entirely lacking.
The thickened (glandular?) area on the "supratympanic" ridge is
invariably present and cream in color.

Atelopus loomisi Taylor

(Fig. 6)

Atelopus varitis loomisi Taylor, Univ. Kansas Sci. Bull., vol 35, pt. 1, July 1,
1952, pp. 625-627, fig." 7 (type locality, Isia Bonita [Cinchona], eastern
slope of Volcan Poas, Heredia Province, Costa Rica, approximately 5,000
ft. elevation ) .

This species has been traced north and down to an elevation
of approximately 1,800 feet at Cariblanco, where two specimens
were taken along the Saripiqui River. The lighter dorsal areas
have a little more pigment than specimens taken at higher eleva-
tion. The venters, in life, had the same bright orange-red colora-
tion and large black spots as did the type. Since more Atelopus
from Costa Rica have been available for study I now regard this
form a species distinct from varitis. A problematical specimen
occurring south of the Meseta Central of Costa Rica may represent
a subspecies of this form. See comments under Atelopus varius
varius.

The figure shows two of the typical series: KUMNH No. 24742,
paratype, 41 mm.; type, No. 24744, 44 mm.

Eleutherodactylus mimus * sp. nov.
(Fig. 7)
Type: KUMNH No. 37128, 5 km. NNE Tilaran, Guanacaste
Province, Costa Rica, Aug. 13, 1954; Edward H. Taylor, coll.

Paratopes: KUMNH Nos. 37126-37127; 37129-37132, topotypes,
Aug, 13-J5; same collector.

Diagnosis: In the goUmeri group (including noblei) but differ-
ing from goUmeri in having a slenderer head and body, and larger
discs on fingers and toes. The first finger longer than second; a
strongly defined, free-edged, inner tarsal fold about half tlie length
of tarsus; toes at least two-fifths webbed (or somewhat more), the
webs continuing as a strong fringe to the terminal discs; web reduced
between the fourth and outer toe (less than one-fourth webbed);
tympanum of male circular, a little more than three fourths of the
diameter of eye; strong supratympanic fold continuing back in a
slightly diagonal line, a branch from which goes behind tympanum.

* From Gr. niiinikos= mimic.

518

The University Science Bulletin

Fig. 6. Atelopus loomisi Taylor. Upper figure, KUMNH No. 24742 9-
Snout to vent length, 41 mm.; paratype. Lower figure, KUMNH No. 24744 9,
paratype, snout to vent length, 44 mm. Both from Isla Bonita, Heredia Prov-
ince, Costa Rica.

Additions to Fauna of Costa Rica

519

Fig. 7. Eleutherodactijlus mimus sp. nov. KUMNH No. 37128, Type.
Five km. WE Tilaran, Las Canas, Guanacaste, Costa Rica. Snout-vent
length, 33.2 mm.

Description of species: Head slender, its length ( 18 mm. ) much
greater than its width ( 12.4 mm. ) ; snout rather poin'^ed, the tip
rounded, the canthus distinct but slightly rounded; loreal region
sloping precipitously,* not concave; a slight depression indicated
between nostrils; head flat, except for a small ridge across the in-
terorbital region; length of eye (4 mm.) equal to its distance from
nostril; length of snout (median) 6 mm.; diameter of tympanum
(3.15 mm.) at least equal to three fourths of eye leng h; distance

* The figure is somewhat misleading, since the dark sides of the upper jaws are not
distinguishable from the background.

520 The University Science Bulletin

between eye and tympanum 1.5 mm.; snout projecting beyond
mouth 1.2 mm.; tongue (7.5 mm. x 6.5 mm.) subcordiform, some-
what notched behind; choanae smaller than raised area of vomerine
teeth, largely lateral; vomerine teeth behind, but within inner
level of choanae, separated from choanae and each other by
about the width of one series; no vocal slits or vocal sac; Eustachian
tube openings much larger than choanae.

Arm brought forward, the wrist reaches tip of snout; first finger
distinctly longer than second; subarticular tubercles large; three
large distinct palmar tubercles, the median and outer may partially
fuse; supernumerary tubercles on palm prominent; a lateral ridge
on fingers scarely discernible; terminal pads grooved transversely,
outer pads a little wider than those on two inner digits, those
on third and fourth a half wider than the adjoining part of finger.
Leg long, the tibiotarsal articulation extending beyond tip of
snout for a distance equal to one third of the tibial length; when
legs are folded at right angles to the body they overlap 9.5 mm.;
a sharp-edged, free, inner tarsal fold, half as long as tarsus; web
reaching to middle of first finger, two thirds the length of the
second and third, about one fourth the length of outer finger;
discs on toes larger than those on fingers; a broad fringe on both
sides of toes to disc which is wider than digit with fringe; no
tubercles on sole; an elongate inner metatarsal tubercle and a much
smaller rounded outer one.

Skin on much of dorsal surface smooth; two pairs of moderately
distinct tubercles in shoulder region, the outer a little the larger,
inner a little farther forward than outer; a strong supratympanic
fold runs diagonally back from eye and continues as a free fold
of skin along the sides; a pair of low tubercles behind lower edge
of tympanum; a strong posttympanic fold branches from the supra-
tympanic fold and runs down closely behind the tympanum; sides
with a few minute granules; venter and chin smooth; a ventral
disc limited by distinct folds laterally and posteriorly; posterior
lumbar region finely granular, with larger granulations in region of
vent and below it; granules present on inner part of the under side
of thighs.

Color in life: Above generally dark lavender-brown; the snout
to middle of interorbital region clay-white, bordered behind by a
straight, narrow black line; a hair-fine median cream line, not
reaching tip of snout; a dim, darker, symmetrical, dorsal pattern
consisting of a diamond-shaped figure with radiating lines and a
lighter center, greatly narrowed in the middle of back, then sud-

Additions to Fauna of Costa Rica 521

denly widened, sending two branches back diagonally; some trans-
verse marking on back of lumbar region; side of head and tympanic
area dark lavender, the color continuing on side to near venter;
edge of lip a little lighter; venter and breast flesh-white; throat
and chin with a scattered powdering of blackish pigment; an area
under thighs and tibia pinkish. Thighs above with at least four
irregular brown bars with lighter edges, separated by grayish
lavender which may have narrow secondary bars; tibia with
four or five, more or less chevron-shaped bands with lighter edges;
tarsus and foot with indications of bars; an irregular brownish-
black area at knee continued as a blackish stripe on front of
tibia; sole purplish; top of foot and digits flecked with darker;
palm much lighter than sole; black spots on front of arm; a dark
arch about region of vent; back of thigh uniform light brown.

Measurements in mm.: Snout to vent, 33.2; width of head, 12.4;
length of head, 18; arm, 17; leg, 65; tibia, 13; tarsus and foot, 27.

Variation: The general color pattern can be discerned in the
paratypes if they are submerged in clear liquid but the colors
on the whole are less intense in the younger specimens. One speci-
men has three light rounded spots on the middle of the back
arranged in triangular pattern. Some of the specimens have a
pair of small tubercles on the dorsal surface of the snout and the
dorsal surface of the body is slightly more shagreened. The sur-
face of the thigh and tibia also may show very fine granulations.
The transverse interorbital fold, the character of the webbing of
the digits, the size of the digital discs, are constant. The median
light line rarely reaches the top of the head. The lateral skinfold
may not always be in evidence. The females have smaller tympani
that are higher than wide.

Remarks: At this type locality a specimen of Eleutherodactylus
nohlei was taken, but the related Eleutherodactylus gollmeri was
not found. I have examined the types of Lithodytes lanciformis
Cope. The head is much wider (9.8 mm. x 11 mm.). It is not im-
possible that lanciformis is identical with nohlei but the present
condition of the types makes it impossible to be certain. The pinkish
coloration mentioned by Cope is characteristic of young specimens
of nohlei.

In the U.S. National Museum I have examined a specimen of
this new species ( number and locality not recorded by me ) having
a snout-vent length of 43.5 mm., arm, 22; leg, 88; tibia, 31.6; foot
and tarsus, 37; heel extending beyond snout 13.5 mm. It is identi-
fied as lanciformis.

522 The University Science Bulletin

Eleutherodactylus vocator * sp. nov.
(Fig. 8)

Type: KUMNH No. 37001, Agua Buena, Puntarenas Province,
Costa Rica, July 25, 1954, Edward H. Taylor, collector.

Paratypes: KUMNH Nos. 36999, 37000, topotypes, same collec-
tor, July 24, 1954.

Diagnosis: A diminutive frog, the head much narrower than
the body; maximum known size, $ 17.6 millimeters; first finger
distinctly shorter than second, lacking webs on both hands and feet;
digital discs small on outer fingers, that on first not wider than
digit and those on inner toes are not perceptibly wider than digits,
while the third and fourth have discs slightly wider than adjoining
part of digits and pointed rather than rounded terminally; heel
reaches to eye; interorbital distance a half wider than an eyelid; a

Fig. 8. Eleutherodactylus vocator sp. nov. Type and paratype. Figure on
left, KUMNH No. 37000 i , paratype, Agua Buena, Puntarenas Province;
figure on right, KUMNH No. 37001 S , type, Agua Buena, Puntarenas Province,
Costa Rica, snout-vent length, 16.2 mm.

strong notch at median point of upper lip. Male with a very ample
vocal pouch, the vocal slits extending back beyond mouth angle;
vomerine teeth absent; the choanae lateral, concealed when seen
directly from below; venter smooth, and pigmented.

* From Latin, vocator =: one who calls.

Additions to Fauna of Costa Rica 523

Description of type: Head relatively small, the snout pointed
(seen from above) and flattened, since the line of the mouth tends
to curve up anteriorly; canthus rostralis moderately distinct,
rounded; the loreal region sloping very abruptly (nearly vertical)
and not concave; nostril well back from the snout tip, the distance
from eye (1.2 mm) a little greater than its distance from median
point on upper lip (1.16 mm.); length of eye (2.1 mm.) shorter
than length of snout ( median measurement 2 mm. ) but less than
its distance from the median point on tip measured laterally (2.45
mm. ) ; tympanum covered with pigmented skin, the anterior out-
line visible, its greatest diameter .55 mm.; an indistinct fold from
eye runs diagonally back covering a portion of tympanum; interor-
bital distance (2.35 mm.) much wider than eyelid (1.5 mm.); a
strongly distensible, subgular, vocal sac which forms two strong
plies when not distended; tongue much narrowed in its anterior
half, then widening, its greatest length more than double its great-
est width; vocal slits elongate, running back behind angle of
mouth; choanae lateral, hidden when palate is seen from below;
vomerine teeth absent; openings of Eustachian tubes no larger
than choanae; a distinct notch in median point of upper lip; snout
extending beyond line of mouth .6 mm.

Arm brought forward the wrist reaches to tip of snout; first finger
much shorter than second without a widened disc, the length 1
mm.; disc on third finger pointed anteriorly; on other fingers the
discs are not or but minutely wider than adjoining part of digit;
subarticular tubercles flat; two palmar tubercles somewhat in-
distinct, the outer (median) large, subcircular.

Leg short, when folded at right angles to body heels fail to
meet by one millimeter; leg brought forward, the tibiotarsal artic-
ulation reaches to near middle of eye; no trace of a tarsal fold;
discs on digits not or a trifle wider than the digits, that on the
fourth toe pointed and somewhat asymmetrical; subarticular tuber-
cles flat, indistinct; some evidence of supernumerary tubercles on
palm and sole; a low, rather large, inner, metatarsal tubercle, its
outline not strongly defined, the outer distinct, somewhat more
elevated.

Skin on dorsal part of body and head finely shagreened (seen
under a lens); sides of abdomen with some granulation extending
to venter; posterior part of venter, with some very flat granules;
posterior and part of ventral surface of thighs with relatively large,
strong granules; skin of throat smooth, but distended; a pair of

524 The University Science Bulletin

tubercles below and behind tympanum and one preceding it. No
ventral abdominal disc.

Color: Gray above with a narrow interorbital darker line and
an elongate indefinite darker median area; a white spot on heel;
on each side of the chin a lighter area. Entire ventral surface and
concealed parts of the limbs pigmented, the pigment forming a
reticulation leaving very numerous small rounded light areas; this
likewise true of the sides of body; lips with several indefinite light
flecks; a dark line follows the supratympanic fold; a light spot at
(dorsal) base of each digital disc. Underside of hands and feet
pigmented.

Measurements of type and paratijpes 37001, 37000, 36999, respec-
tively: Snout to vent, 16.2, 16.1, 17.6; width of head at jaw angle,
6, 6, 6.1; length of head, 7, 6.9, 7; arm, 8.6, 8.5, 9; leg, 23, 22, 23;
tibia, 7.6, 7.2, 7.3; hand, 3.3, 3.1, 3.25; tarsus and foot, 10, 10, 9.9.

Variation: The male paratype agrees very closely in size as
well as in other characteristics with the type, save that the dorsal
pigment, and some of the lateral, tends to form dark flecks. A some-
what diagonal, lighter line follows below the diagonal dark line
marking the supratympanic fold. The female paratype is dark,
appearing nearly black on head and shoulders. However lighter
flecking occurs on the sides, rump, and to some extent on limbs.
Lateral light areas are present on sides of the chin and the white
flecks on the darker venter are more conspicuous, as is the heel-
spot. The female, as well as the males, is adult and the abdomen
is distended with four, large, yellowish eggs.

Remarks: A related species Eleittherodactyliis diastema was
described from a specimen taken by John M. Bransford on the
Panamanian (not Nicaraguan) Canal Survey of 1874. The exact
locality is given as "Camp Mary Caretta" but a manuscript label
is said actually to read Camp Margharetta, and the suggestion
has been made that Margarita, near Colon, is intended. In the
specimens I have identified as belonging to Cope's Lithodytes
diastema, the adult males and females are considerably larger than
males or females of this species. The female of diastema may lay
as many as ten eggs measuring four millimeters in diameter. The
yolk of the egg is cream completely lacking in dark pigment. At
Cervantes, Cartago Province, this species has been taken chiefly
in bromelias at about 4,800 ft. "elevation. Here the species is com-
mon, some plants yielding half a dozen specimens, all showing
considerable variation in color and marking. The eggs of that

Additions to Fauna of Costa Rica 525

species are frequently found in the bromelias. These are usually
laid so they are not submerged, the young having a direct trans-
formation typical of most, if not all, Costa Rican Eleutherodactylus.
Several females may lay their eggs together in a single plant, and
at different times, since sets, differing in degree of development,
may be found side by side.

Noble ( 1918 ) also described a form from Nicaragua under the
name of Hijla chica. He considered that the clawlike terminal
phalanges placed the frog in the family Hylidae. However the life
history involving the absence of a free swimming stage (as well as
other considerations) seem certainly to place his form in Eleuther-
odactylus. I have not examined the type but consider it likely,
that, if not a definitive form, it belongs with diastema rather than
with vocator.

The most outstanding differences between vocator and diastema
are that the former has smaller, differently shaped, terminal digital
discs, and the head is more pointed, and presumably the species
fails to attain as large a size. The species probably occupies the
Pacific slopes of the Talamanca Range, and the Brunquefia Moun-
tains. Many were heard at Agua Buena and presumably also in
the mountains above San Isidro del General but no specimens were
found in the latter locality. In northern Costa Rica along the
Pacific slopes of the Cordillera Volcanica they were neither heard
nor seen.

Eleutherodactylus diastema (Cope)

(Fig. 9)

Lithodytes diastema Cope, Joum. Acad. Nat. Sci. Philadelphia, ser. 2, vol. 8,
1876 (1875), pp. 155-156 (type locality "Camp Mary Caretta," Panama).

This species was not figured in my paper, "A Review of the Frogs
and Toads of Costa Rica" (Univ. Kansas Sci. Bull., vol. 35, pt. 1,
1952, pp. 701-703 ) hence I am presenting here a photograph of four
specimens to show color-pattern variation. This may serve as a
basis of comparison with the figure of Eleutherodactylus vocator
herein described.

Genus Agalychnis Cope

Agahjchnis Cope, Proc. Acad. Nat. Sci. Philadelphia, 1864 (Aug. 30),
p. 181 (type of genus Hijla callidryas Cope)*; idem, p. 182; Nat. Hist.
Rev., 1865, pp. 110 (embraces H. moreletii, H. holochlora and H. callidryas).

On the night of Aug. 14, 1943, a large pond was visited at a point
some 4 kilometers NNE of Tilaran. Large numbers of amphibians

* Taylor, Univ. of Kansas Sci. Bull., vol. 35, pt. 1, 1952, p. 801, incorrectly states that
Hyla moreletii is the type of genus.

526

The University Science Bulletin

Fig. 9. Eletitherodachjhis diastema Cope. Upper left, KUMNH No. 31643
$, snout to vent length, 19 mm.; upper right. No. 31633 9, length, 22.2 mm.;
lower left. No. 31672 9, length, 24 mm.; lower right. No. 31629 9, length, 24
mm. All from Cervantes, Cartago Province, Costa Rica.

Additions to Fauna of Costa Rica 527

composed of several species were calling from the shores or from
weeds partly submerged and together their voices blended into a
great chorus. Several were found breeding. In the weeds partly
covered with water and in trees and shrubs along the edge of the
pond very large numbers of breeding Agalychnis calUdryas were
found — the number of males seemingly exceeding the number of
females. Near one end of the pond (approximately 200 meters
in length), a second but smaller species of Agalychnis was found,
these seemingly occurring in smaller numbers and it was presumed
that these were not breeding. Ten adult males and one female
were collected and several young that were partially or completely
transformed. Certain young and one adult male were found on
plants and shrubs three hundred meters from the pond and per-
haps a hundred meters higher than the pond level. In contrast
more than fift\' adult calUdryas were collected at the pond and a
much larger number could have been acquired with ease. Many
of these were clasping and eggs were found on leaves above the
water. No young specimens or larvae of this species were taken.

The smaller species belongs in the section of the genus that in-
cludes dacnicolor, alcorni, and moreletii, the latter of which also
occurs in Costa Rica. However the species is distinct, and I believe,
unnamed. It is described below.

Agalychnis saltator * sp. nov.
(Fig. 10)
Type: KUMNH No. 35615, 4 km. NNE of Tilaran, Guanacaste,
Costa Rica, Edward H. Taylor, collector.

Paratypes: 35609-14; 35616-26. Topotypes. Prof. Marco Tulio
Pacheco, Srta. Georgiana Pacheco and Edward H. Taylor, collec-
tors.

Diagnosis: A small species, the largest male 44 mm., the single
female 52 mm.; eye reddish; body lacking lateral black and cream
markings; width of interorbital region equal to, or minutely greater
than, width of an eyelid; canthus rostralis indicated but rounded;
no paratoid; digits less than half webbed on feet, less than one
third webbed on hand; skin-fold above tympanun obsolete but very
strong behind tympanum, continuing above arm; heel to front of
eye; tympanum very distinct, its diameter slightly less than half
length of eye; green above with narrow, dim, wavy, darker, trans-
verse lines indicated on body; a cream spot at posterior corner
of eye.

* Latin, saltator =: leaper.

528

The University Science Bulletin

Fig. 10. Agalychnis saltator sp. nov. Type. KUMNH No. 35615, four
km. NNE Tilaran, Guanacaste, Costa Rica. Snout vent length, 44 mm. ( trans-
verse dark lines, visible to the eye, are scarcely shown in the photograph ) .

Description of type: Head triangular, wider than the body; no
paratoid present; canthus indicated, curving forward, rounded;
areas about nostrils swollen with a slight depression between;
in profile, snout sloping down to edge of lip; distance between
eye and nostril (4 mm.), greater than distance between nostril
and median notch on upper lip (3.2 mm.); loreal region sloping

Additions to Fauna of Costa Rica 529

broadly to lip, not or but slightly concave; narrowest interorbital
distance (4.3 mm.) greater than an eyelid (4.1 mm. flattened);
tympanum circular, its diameter (2.4 mm.) less than half length
of eye-opening (5.2 mm.); eye to tip of snout 5.3 mm. (median
measurement); upper eyelid folds down somewhat over eyeball;
lower lid with cream venation; large vocal slits in floor of mouth
extending but little forward of the mouth angle; tongue longer than
wide, notched behind, free for nearly half its length and free on
sides; choanae large, the vomerine teeth in two tiny fascicles, be-
tween the anterior borders of choanae, separated from each other
by a distance equal to one fascicle and from edge of the laterally
placed choanae by a somewhat greater distance; openings of the
palatal glands form an indefinite line nearer tip of palate than
anterior level of choanae; upper arm slender, pigmented above,
forearm thickened with a prominent "elbow"; a cream skinfold
beginning behind elbow runs along outer ventral surface of arm
and is more or less evident to base of outer finger; first finger much
shorter than second, the disc widened, a little smaller than tym-
panum; discs on two outer fingers a little larger than second, and
double width of first; web between first two fingers short, failing to
reach to level of tlie subarticular tubercle of first and not half way
to tubercle on second finger; between second and third to upper
level of tubercle on second, to base of tubercle on third; between
two outer fingers web reaches half way to tubercles — all webs con-
tinuing to discs as a thick fringe; tubercles on two outer fingers bifid;
palmar tubercles not strongly defined, that at base of first finger
flat, oval; two small tubercles on palm; well-defined supernumerary
tubercles; first finger on inner dorsal surface with a patch of small
brownish nuptial spines; leg relatively short, the heel reaching
front edge of eye or slightly farther; when legs are placed at right
angles to body, heels overlap three millimeters; tarsal fold very in-
distinct save for an irregular row of minute granules; outer fold
barely indicated by a white line; first toe shorter than second; two
inner toes less than a third webbed; between second and third,
web reaches to middle of distal tubercle of second toe; between
third and fourth, web reaches to base of distal tubercle of third
and between two outer toes to about the same distance; webs con-
tinued to discs as a fringe; discs on toes smaller than those on
second finger; inner metatarsal tubercle narrow, elongate, outer
farther forward, distinct, small; subarticular tubercles and super-
numerary tubercles present.

530 The University Science Bulletin

Skin on dorsum, head, and above thighs with small granules or
corrugations, not equally distributed, but very distinct especially
on sides of head; tympanum somewhat elevated, thinly covered with
skin; back edge of tibia rather sharp; region of vent sticking out
behind level of thighs, the skin Hap covering the vent does not
reach down quite to level of vent; on chin, skin wrinkled but
without distinct granulation; venter strongly granular; inner part
of thigh with small granules intermixed with larger granules, the
latter forming an indistinct row on posterior edge; the skinfold
from eye is obsolete above tympanum or dimly indicated, promi-
nent behind tympanum.

Color in life: Above dark leaf-green with dim indications of
transverse, darker, wavy lines or spots on dorsum; ventral and
concealed surfaces of limbs, yellow orange. The green color covers
only outer finger and part of two outer toes; front and back of
thigh and part of upper arm lavender, the green color forming a
narrow line on top of thigh. Area about vent somewhat whitish;
a cream-white spot at back corner of eye; eye reddish.

Measurements in mm. of type and No. 35622 $ : Snout to vent,
44, 52; width of head, 15, 17; length of head, 15.2, 18; arm, 29, 34;
hand, 13, 14; leg, 68, 78; tibia, 22, 25.4; foot and tarsus, 28, 33.

Variation: When first taken all the adults, were of practically
the same shade of green and orange. In preservation part of the
specimens have become brownish or brownish lavender with a
frosting of bluish white over the dorsal surfaces. The darker trans-
verse markings can be seen below this color, and in No. 35625 the
markings are rather clearly defined. The orange coloration is lost
leaving these pigmentless surfaces white.

Relationship: The relationship, based on the absence of lateral
cream and purple markings, is seemingly with Agahjchnis moreletii.
The latter species is much larger ( nearly 25 percent in linear meas-
urement and more than double the volume). The skin is smoother,
and the supratympanic fold well developed above tympanum; the
tympanum is less well defined and the webbing on fingers and toes
is greater. The vent opens downward at level of venter.

Remarks: Besides the series of Agahjchnis saltator, the following
species of amphibians were taken at the pond on the same evening:
SmiUsca hatidini, Hijla imderivoodi, Hijla loquax, Hyla ebraccata,
Bufo coccifer, Hyla phaeota, Agahjchnis callidryas, Leptodactyliis
melanonotus.

Additions to Fauna of Costa Rica 531

REPTILIA

Genus Eumeces Weigmann

Eumcces Wicgmann, Herpetologia Mexicana, 1834, p. 36 (type of genus
Scincus pavimcntutus = schneideri) .

This cosmopolitan genus occurs in the Western Hemisphere from
Canada throughout most of the United States and Mexico, and into
Central America but the number of species dwindles until only three
are known south of the Isthmus of Tehuantepec. Two of these
extend into northern Central America, Eumeces schwartzi having
been taken in northern Guatemala and E. sumichrasti in northern
Honduras. The third species, originally discovered in Managua,
Nicaragua, is here traced south into Costa Rica and presumably it
is the only known species of the genus occurring there. Four speci-
mens were taken.

Eumeces managuae Dunn

(Fig. 11)

Eumeces taeniolatus ( non Blyth ) Boulenger, Catalogue of the lizards in the
British Museum, vol. 3, 1887, p. 383 (type locality [in errore] "Punjab,
India").

Eumeces managuae Dunn, Proc. Biol. Soc. Washington, vol. 46, 1933, p. 67,
68 (tvpe locahtv, aviation field Managua. Nicaragua); Tavlor, Univ. Kan-
sas Sci. Bull., vol. 23, pp. 104-110, pi. 3, figs. 6, 7, 8.

Diagnosis: A large species of the genus characterized by a greatly
expanded series of median dorsal scutes; four pairs of expanded
nuchals, one postmental, a postnasal; large auricular lobules; sub-
caudals transversely widened; adpressed limbs widely separated;
gray or gray-brown above, the dark color tending to form elongate
lines.

Description of species. ( From KUMNH No. 34185, Tenorio,
Guanacaste, Costa Rica). Rostral large, much wider than high,
the part visible above forming a very obtuse angle, its length
less than that of the supranasals; latter moderate, broader than
long, the length of the median suture distinctly less than the great-
est length of the scale, which is less than its width; the nasal is
definitely divided; frontonasal much larger than the prefrontals,
wider than long, forming a suture with the frontal and at same
time separating the prefrontals; latter generally pentagonal, touch-
ing two loreals laterally, and touching the frontonasal, frontal, first
supraocular, and first superciliary; frontal angular anteriorly reach-
ing its greatest width between the first supraoculars, and then
narrowing slightly posteriorly, bounded laterally by two supra-
oculars and behind by the paired frontoparietals, and not making
a suture with the interparietal (as in the type). Interparietals a

532

The University Science Bulletin

Fig. 11. Eumeces managiiae Dunn. KUMNH No. 34185, Tenorio, Las
Canas, Guanacaste, Costa Rica. Total length, 282 mm.

Additions to Fauna of Costa Rica

533

little smaller than the prefrontals (much smaller in the type) theii*
mutual suture about one third of their length, and touching three
supraoculars laterally; parietals not separated by the interparietal,
their posterior inner edges forming a suture; four pairs of widened
nuchal scales; a postnasal present behind the posterior nasal; two
loreals, the anterior higher and much widened at top, and generally
slenderer than posterior; four supraoculars, the second largest;
three well-defined presuboculars, eight superciliaries, three or four
small preocular scales, one touching the loreal; four postsuboculars;
primary temporal small, quadrangular; secondary temporals large,
the upper elongate, the lower triangular; tertiary temporal present.
Seven supralabials followed by two superimposed pairs of post-
labials. About 27 irregular scales about ear-opening, three of the
anterior scales form lobules; five supralabials preceding the subocu-
lar supralabial; mental bordering mouth a somewhat longer dis-
tance than rostral, nearly twice as wide as deep; seven infralabials;
a single postmental followed by three pairs of chinshields, only
the first pair forming a median suture; scales on dorsum from
parietal to above vent, 67, of which four are nuchals, followed by
seven large paired scales, and these in turn followed by 56 widened
body scales; scales about anterior part of neck, 31; about narrow
part of neck, 23; about middle of body, 17; about axillary region,
26; about base of tail, 13. A total of 83 subcaudals, all about five
times as wide as long except two pairs following the vent (the
distal part of the tail is regenerated); a tiny patch of granular
scales in axilla and a similar elongate patch behind leg insertion.
Third and fourth fingers equal in length, each with twelve scales
on under surface; palm with three light padlike scales larger than
others; fourth toe distinctly longer than third with sixteen keeled
scales below; three large padlike scales on heel.

Color: Above rather yellowish brown, the head somewhat lighter,
rather more grayish brown on sides; a light lemon-yellow wash

Measurements of Eiimeces managuae Dunn

Number

Sex

Snout to vent . .

Tail

Snout to eye , . .
Snout to foreleg
Axilla to groin. .

Arm

Leg

Width of body.
Width of head . .

34185

125

157

8

36.5

74

22

30

19

16

34186

104.5

145.5

7.2

32

60

22

27

16.5

14

34187

yg

62

30 reg.

4.8
20
36.5
13
16
11

9

34188

yg

57

41 broken

4.4
20.2
33
12
15

9

8.5

534 The University Science Bulletin

on venter; chin whitish; the dorsal scales with one or two darker
areas that form discontinuous darker lines, two median, and one
each on other scale rows; chinshields and six ventral rows immacu-
late flesh with dark flecks appearing at base of tail and growing more
numerous toward tip; each labial with a dark spot; each head
scale with a darker mark, some with more than one.

Variation: Nos. 34186 and 34187 have the frontoparietals separated
instead of forming a suture, while 34189 has them touching at a
single point.

In the two young specimens the outer ventral scale rows may
have some dark flecks. The general color above is wood-brown
with the dark lineation strong and continuous. In the younger speci-
men the yellow of the venter is more pronounced. It is remarkable
that these specimens were collected on an airfield as was the type.

They were found burrowing under the edges of small boulders
and two were taken from a small pile of stones. Certain others
seen escaped into masses of growing cacti about rocks. This is
the southernmost record of the genus Eumeces in the Western
Hemisphere.

Genus Anadia Gray

Anadia Gray, Catalogue of the specimens of lizards in the collection of the
British Museum, 1845, p. 74 (type of the genus, Anadia ocellata Gray).

Diagnosis: Slender, elongate lizards, the head narrowed and
rather pointed. Head with regular shields; frontonasal present
separating nasals; nostril in a single nasal, or in a suture between
two nasals; eyelids developed, with a divided transparent disk on
lower lid; scales smooth, juxtaposed, or slightly imbricate, form-
ing transverse rows; ventrals equal to or a little larger than dorsals,
quadrangular, forming transverse and longitudinal series. Tail
cylindrical; lateral teeth compressed, tricuspid; tongue bifid, some-
what arrow-shaped; auricular opening present; limbs rather weak,
pentadactyl. At least some forms are arboreal.

This genus comprises some nine or ten species distributed chiefly
in northern South America. Only a single species is known in
Central America. This was described in 1875 by E. D. Cope as
Chalcidolepis metalliciis from a single specimen collected by Wil-
liam Gabb in the Aguacate Mountains of Costa Rica. Later Cope
( 1885 ) placed his Chalcidolepis metalliciis in the genus Leposoma.

The genus was reviewed by Arthur Loveridge in 1929, and a new
species, Anadia nicefori, was described from Colombia. In the
same paper he reports another specimen of Cope's species from
Costa Rica.

Additions to Fauna of Costa Rica 535

Burt and Burt in 1933 again list the South American species of the
genus. Dunn, in 1944, reviewed the Colombian species, describing
a new form, and listing, in all, six species for Colombia. He places
Anadia nicefori in the genus Ftxjchoglossus.

To the best of my knowledge the genus is still unreported in
Panama. The Costa Rican material of the genus has been scant.
Mr. Loveridge has loaned me two Costa Rican specimens from
the Harvard collection. There are three in my Costa Rican collec-
tions.

Key to forms of Anadia in Costa Rica

1. Femoral pores, 3-?; 23 scales about body; 56 transverse dorsal rows

between parietal and back level of thigh; brown above with a row

of blackish scales on each side metallica metallica ( Cope )

Femoral pores, 5 or" more; more than 25 scales around body 2

2. Femoral pores, 5-5; 28 scales about body; 63 transverse scale rows

between parietals and back level of thighs; back heavily spotted

with black metallica arhorea subsp. nov.

Femoral pores, 9-10; 28-31 scales about body; 52-56 scales between
parietal and back level of thighs; black spotting on back reduced or
absent metallica attenuata subsp. nov.

The material is inadequate to determine positively whether we
are dealing with species or subspecies. One may presume that
metallica occupies the Pacific drainage of the Aguacate mountains
and perhaps also of the adjoining Talamanca Range; arhorea the
Pacific slopes of the Cordillera Volcanica; and attenuata the Carib-
bean drainage of the Meseta Central and the eastern slopes of the
Talamanca Range. It may be doubted that the barriers are such
as to bring about complete isolation. An accumulation of series
from a given locality will permit one to determine the extent of
variation and a clearer interpretation of the relationships as to
whether they are specific or subspecific.

One other fact suggesting the closer relationship is that the light
color on the side of neck and upper lip (including lower eyelid)
seemingly is nearly identical in the three forms, and at least in two
there is a row of widely spaced light dots from above arm insertion
to the groin, some of which may have dark borders. However, this
condition obtains also in certain species occurring in South America,

Anadia metallica metallica (Cope) (new combination)

Chalcidnlcpis metallicus Cope, Journ. Acad. Nat. Sci. Philadelphia, ser. 2, vol.

8, 1876 (1875), pp. 116-117, pi. 24, fig. 5; pi. 28, fig. 2 (type locality,

"Aguacate Mountains," Costa Rica, William Gabb, coll.).
Ecpleopus (Chalcidolepis) metallicus Bocourt, Mission Scientifique au Mexique

et dans rAnit'-riquc Centrale; Etude sur les Reptiles et les Batraciens, livr.

6, 1879, pp. 371-372.

536 The University Science Bulletin

Leposoma metoUicum Cope, Proc. Amcr. Philos. Soc, vol. 23, no. 121, Jan.

1886, pp. 97-98; U. S. Nat. Mus. Bull. no. 32, 1887, p. 45.
Anadia metallica Giinther, Biologia Centrali-Americana; Rcptilia and Batrachia,

Aug. 1885, p. 30; Boulenger, Catalogue of the Lizards in the British

Museum, 2nd ed., vol. 2, p. 400; Loveridge, Proc. Biol. Soc. Washington,

vol. 42, Mar. 25, 1929, pp. 99-102 (part.).

Diagnosis: A slender lizard with a tail nearly twice length of
head and body; snout-to-vent length, 58 mm.; head narrowed,
pointed; limbs with five digits, all clawed; 23 scales in a transverse
row around body, subeqvial in size; frontonasal single, separating
nasals; 55 scales from parietals to level of posterior face of femur;
two large preanal scales preceded by a somewhat smaller pair;
occipitals enlarged. Teeth compressed with a central cusp Hanked
by a denticle on each side.

Description of species: (Data from Cope's type description). A
slender lizard with a very long tail and feeble limbs; head nar-
rowed and acute in front, with a produced rostral shield; frontonasal
as wide as long; frontal elongate, two prefrontals, two fronto-
parietals, two parietals separated by an interparietal; four supra-
oculars; interparietal elongate having sutures parallel to the larger
parietals; latter bounded externally by a large temporal, forming
with them a diagonal suture; these all are bounded posteriorly by
a series of four shields across the occiput, and these in turn by
a transverse series of seven scales larger than those on the nape
following them; nasal followed by a large loreal and this by a
smaller preocular; seven supralabials, 4,3,1,2,6,7,5, being their
diminishing order of size; two pairs of chinshields in contact
in median gular region, the posterior pair being twice as long as the
first. Nostril in a single nasal plate, tympanum distinct.

Dorsal scales all smooth in uninterrupted transverse annuli round
the body, the size subequal on the various regions including the
nuchal and gular; twenty scales in a cross-row between the angles
of the lower jaw; scales in pectoral region irregular, the last row
of the neck having the appearance of a collar; a series of 23 scales in
an annulus of the body; twelve transverse rows between large
postoccipital row and line of axillae; between this line and posterior
line of thighs, 43 scales. There are two large, longitudinal, anal
scales, which embrace a scale between them on the anal border;
these preceded by another pair, but of reduced size.

Toes 5-5, all clawed. Tail nearly twice head-body length; hind
limb one fourth head-body length. Limbs surrounded by large
scales except on concealed faces of humerus and femur, where

Additions to Fauna of Costa Rica 537

the scales are small and flat; lower eyelid dried so its characters are
obscured.

Color: Light gray with red and green metallic reflections; sides
brown, the middle of baek darker than a line above the brown
of the side; near the light dorsolateral lines, a few scales are black-
ish, forming a row on each side; venter dusted with brown; head
with deep brown sides and a white upper lip; sides of tail brown
with a zigzag upper margin.

Measurements in mm.: Total length, 165; head-body length, 58;
tip of snout to axilla, 17; length of head to tympanum, 11; length
of head to orbit, 4.5; length of arm, 13; length of leg, 14.5.

Remarks: Cope's figure 5, plate 24, is perhaps a bit diagram-
matic. There is no suggestion of the lateral series of separated light
spots. Figures 2a and 2b, plate 28 are presented on a small scale
but the conformation of the chinshields do not conform to the
description since the second pair is shown to be very small. A third
pair which is not mentioned are large, and separated from each
other.

Dr. Doris Cochran has kindly re-examined the type. She writes:
— "The type of Chalcidolepis metaUiciis, USNM 30568, is not in the
best shape, being pretty dry, and having lost its left leg. Its right
leg shows 3 femoral pores, the 3rd quite faint. Perhaps there are
only 21 or 22 scales around the body, — it is too brittle to say exactly.
The nasal and first labial are broadly in contact. The first pair of
chinshields are as long as the mental; the second pair is a little
longer than the first; the third pair is shorter than either pair 1 or
pair 2, and are separated by a small scale, while the first two are
entirely in contact."

It would appear that the specimen was dried when received by
Cope (since he at least mentions the fact that the eyelid is dried)
and the femoral pores were overlooked or an incorrect statement
set down concerning them. He states — "no femoral pores."

Anndia metallica attenuata* subsp. nov.
(Fig. 12)

Tijpe: KUMNH No. 34223 ^ from Pacuare, Rio Pacuare, on
road between Turrialba and Moravia de Chirripo, Cartago Prov-
ince, Costa Rica; Aug. 17, 1951, Edward H. Taylor, collector.

Paratype: KUMNH No. 34224, Moravia de Chirripo.

Diagnosis: Frontonasals separated from the first labial or not;

* Latin, attenuatus = drawn out.

538

The University Science Bulletin

Fig. 12. Anadia metuUica attenuata subsp. nov. Type and paratype.
Left figure KUMNH No. 34224, paratype, Moravia de Chirripo; figure on
right, type, KUMNH No. 34223, Pacuare, Rio Pacuare, on road between Tur-
rialba and Moravia de Chirripo; actual length, 177.5 mm.

Additions to Fauna of Costa Rica 539

rows of black dots on dorsal surface reduced or absent; dorsolateral
light line extending to tail, then breaking up into two series of small
clay-white spots; a series of small white or cream dots on sides
from axilla to groin, those near axilla with dark borders; 9-10
femoral pores; 28-31 scale rows around middle of body; 14 rows of
gular scales between chinshields and nuchal collar; scales from
parietals to back level of thigh, 53-56.

Description of type: Body slender, the basal region of the tail
plump and nearly as wide as body for some distance; rostral large,
the part visible above about half length of frontonasal; latter longer
than wide, separated from the first labial by the nasal; prefrontals
rather large, each nearly half the area of the frontonasal, touching
nasal, loreal, and first supraocular laterally; frontal widened anteri-
orly, narrowed posteriorly, about as long as frontonasal; four supra-
oculars, first small, three touching frontal; a pair of small fronto-
parietals, each touching frontal, two supraoculars, a parietal and
an interparietal; interparietal shaped somewhat like frontal, tlie
widened part posterior, the sides definitely not parallel; parietals
large, their posterior part forming less than a right angle, each
touching last supraocular, upper postocular, and three other large
scales; the parietal series bordered behind by a row of specialized
postparietals; some of the scales of the temporal area enlarged,
one bordering the auricular opening; nasal single, nostril above
middle of first labial; a large loreal, nearly as high as long; five
supraciliary scales; four postoculars; supralabials 7-7, the fourth
largest, the fifth smallest; ear-opening less than half length of
eye-opening; 6-6 infralabials, the last small; mental about size of
rostral; a large azygos postmental; three pairs of chinshields, the
last pair largest, forming a short median suture anteriorly, separated
posteriorly by two scales; first transverse gular row consisting of
eight scales, the outer very large, separated from sixth infralabial
by one narrow scale; 15 rows of gular scales including "collar"
series; lateral scales narrower than dorsals or ventrals; small scales
present about limb insertions; all scales smooth, the dorsals quad-
rangular, somewhat imbricating; very large scales on limbs save
on underside of upper arm, and ventral and posterior part of thigh;
soles, palms and basal joints of digits with padlike lamellae, only
one or sometimes two divided on each toe; on two terminal joints
of each digit, lamellae compressed; a strong terminal claw.

Scales in a row about middle of body, 31; dorsal scales from
parietals to back level of thigh, 52; scales from posterior chin-

540 The University Science Bulletin

shields to front level of thigh, 50; siibcaiidals (tip of tail regener-
ated), 95; six preanal scales, the four median longest, second row
with four scales.

Color in life: Dorsum somewhat mottled olive; head lighter with
a very indefinite pattern; a pair of dirty-white or clay dorsolateral
lines extending from anterior supraoculars to tail where they tend
to break up on each side into two more or less regular series of
cream-colored spots, but becoming more irregular towards tip of
tail; a pair of short light lines in nuchal region; a brown stripe
with darker edges begins behind eye, continues on neck and side of
body to tail, growing lighter and more indefinite posteriorly; a few
ocelli in axillary region continuing as a series of indefinite light
dots to groin; limbs variegated cream-white and light brown; chin,
throat, breast, and underside of arms almost pure white; venter
dirty white with a very fine sprinkling of gray pigment, usually con-
centrated into indefinite areas on each scale; underside of tail with
a Hght salmon wash and with some fine, scattered, brownish pig-
ment.

Measurements in mm: Snout to vent, 58.5; snout to arm insertion,
18.7; axilla to groin, 33.5; tail, 119 (tip regenerated); total length,
177.5; arm, 12; leg, 15.5.

Remarks: The type locality is only a few kilometers from Moravia
de Chirripo, where another specimen was obtained. The type
specimen was captured in a field recently cleared of forest trees.
The rather heavy claws suggest that the species is arboreal rather
than terrestrial.

Variation: A second specimen referred to this subspecies is
KUMNH No. 34224 from Moravia de Chirripo; and while not agree-
ing in all details it does agree in essential characters. The specimen
is a female taken from a bromelia in a forest tree at some distance
from the ground.

Direct comparison with the specimen just described shows but
little difference in head squamation. The large frontoparietal
separates the nasal from the rostral, and the interparietal has
parallel sides. The supralabials are 7-6, the fourth and fifth being
fused on the right side; however, there are 6-6 infralabials due to
a segmenting of the large first infralabials. The scale counts from
occiput to posterior level of thigh is 54; there are 16 gular and 36
transverse ventral scale-rows and 28 scales in a row around body.

The color differences are due largely to a row of small dark dots
rarely covering a single scale completely. There is a small part

Additions to Fauna of Costa Rica 541

segmented from the third supraocular on the right side, two seg-
ments on the left side. The preanal series consists of five elongate
scales preceded by a series of four smaller scales. The female has
2-1 minute femoral pores, and these are smaller than those in the
male. There are two oviductal eggs present.

A specimen collected by Mr. C. H. Lancaster at Cachi, Cartago
Province, "arborea sobre el terra, Nov. 1910. (3,300 ft.?) on the
south bank of the Rio Reventazon," has been forwarded to me by
Mr. Arthur Loveridge. This specimen is the one mentioned by
Mr. Loveridge (Proc. Biol. Soc. Wash., vol. 42, 1929, pp. 99-102).

The specimen is somewhat faded and the dorsolateral light lines
are discernible when submerged in alcohol, and the inner edges
are darker than the outer. No black dorsal spots can be discerned.
The olive brown lateral stripe from snout passes through the eye
and along the side of the head and is lost on the side. The venter
is pigmented somewhat, the chin, lips, and throat cream. The
tail has been broken 26 millimeters behind the vent and the broken
portion of the tail is almost uniformly cream white with little or
no evidence of pigment. The basal portion of the tail shows con-
siderable pigmentation. There are 28 scale-rows around the middle
of body, and 52 scales in a longitudinal row from parietals to back
level of thigh; femoral pores, 8-9; caudal whorls, 113; third pair of
chinshields separated; six preanal scales, preceded by two scales;
snout-vent length 57 mm., tail 144 mm. On one side the nasal
touches the rostral; on the other it fails to touch, while the first
labial touches the frontonasal. There is no scale segmented from
the third supraocular.

Another Costa Rican specimen from "Santa Teresa" Costa Rica
(a locality I do not find on my available maps) collected by
Dr. Dodge "in jungle" was also loaned by Mr. Loveridge. This
specimen is presumably a recent hatchling and presumably is a
female. The minute femoral pores are 3-4; the caudal whorls are
109; the scales in a dorsal row 56; and there are 28 scales in a
row surrounding the middle of the body. The adpressed limbs
touch in this specimen while in all others here reported the ad-
pressed limbs are separated by from 10 to 14 annuli. I am uncer-
tain whether this is purely a juvenile character or not, but presume
that it is.

The color pattern is strongly defined. The area between the
dorsolateral lines shows a vermiculating reticulation of brown while
anteriorly on the head the color is dark olive, more or less uniform;

542 The University Science Bulletin

dorsolateral light lines are well defined; a lateral brown stripe be-
gins at the nasal, passes through eye, includes the upper half of
the auricular opening and continues to the groin; a row of about
13 unequally spaced cream spots surrounded by black color begin
above arm insertion and continue to above leg insertion, the poste-
rior ones less distinct and lacking dark borders.

In this specimen the frontonasal makes a considerable suture with
the labial anterior to the nasal and the interparietal is proportion-
ally more elongate than on the other specimen. A small scale is
segmented from the outer part of the third supralabial (as is true
of KUMNH Nos. 34223, 34224, and 34225, but this is not true
of MCZ No. 15386 from Cachi. Until the adult characters of a
male specimen is known the placing of this female form will re-
main in doubt.

This specimen measures 70 mm. in total length; snout to vent,
26 mm.; the greatest width of head, 4 mm.; arm, 6 mm.; leg, 8.2
mm.

Anadia metallica arhorea * subsp. nov.
(Fig. 13)

Tj/pe;— KUMNH No. 34225, from Las Flores, Tenorio, Las Caiias,
Guanacaste Province, Costa Rica, Aug. 22, 1952; John Baker and
Edward H. Taylor collectors.

Diagnosis: A slender, elongate species, the male with 5-5 femoral
pores; 63 transverse dorsal scale series between parietal and back
level of thigh; back heavily spotted with black, the spots covering
from two to four scales; 28 scales in a row about middle of body.

Description of type: Body elongate, slender, the tail twice head-
body length; head slender, the snout narrowed, rounded at tip;
rostral large, in contact posteriorly with two labials and the
frontonasal, the suture with the latter curving, the length of part
visible from above about half length of frontonasal; latter single,
seven-sided, forming a considerable suture with first labial anterior
to the nasal; prefrontals rather large, nearly half the area of fronto-
nasal, laterally touching the nasal, loreal, and the diminutive first
supraocular; frontal relatively small, larger than the prefrontals,
smaller but equally as long as the frontonasal; frontoparietals small
touching frontal, two supraoculars, interparietal, and parietals; latter
scales very large, terminating posteriorly in an acute angle, sepa-
rated by a parallel-sided interparietal; the parietals touch last supra-
ocular, upper postocular, and two temporals, the upper of which is

* Latin, arboreus — of the trees.

Additions to Fauna of Costa Rica

543

Fig. 13. Anadia metallica arborea subsp. nov. KUMNH No. 34225, Type.
Las Flores, Tenorio, Las Canas, Guanacaste, Costa Rica; actual length, 181 mm.

544 The University Science Bulletin

much enlarged; interparietal touches three enlarged postparietal
scales; transverse series of scales following the postparietals like-
wise modified, larger than the nuchal series which follow behind.

Nostril in a single nasal, separated from rostral by a distance
more than a third of its length, its size less than that of loreal which
borders it behind; a small preocular; nostril above latter half of the
first labial; four supraoculars, first very small, the series bordered
laterally by five supraciHaries, a smaller second loreal (frenocular)
and three small suboculars; four larger postoculars; about four series
of temporals, showing much variation in size; an enlarged scale
borders the auricular opening; lower eyelid well developed, with
a longitudinal series of transparent scales; ear-opening nearly circu-
lar, much smaller than eye-opening.

Seven supralabials, 1,4,3,6,2,7,5 being the diminishing order
of size; five infralabials, the last nearly concealed under the supra-
labial but bearing a brown spot; the following scale is small but
likewise might be counted as the sixth infralabial, lying as it does
below or under the seventh supralabial; labial border of mental
equal that of the rostral; azygos postmental large; three pairs of
chinshields, the first two broadly in contact, the third pair narrowly
separated, narrowed almost to a point on their inner border; the
second pair longer than first; three small scales lie between the
posterior parts of the third pair; these followed by the first gular
row consisting of nine scales, the two outer greatly enlarged and
touching fourth infralabial, separated by one scale from the fifth;
seventeen and two incomplete rows of gulars between third chin-
shields and (including) scales forming the nuchal collar; three
enlarged modified pectoral scales following the nuchal collar; 37
transverse scale rows from nuchal collar to the last transverse row
on abdomen; these scales square or rectangular save on breast and
in last row; the preanal series consists of a small, triangular, median
scale bordered laterally by two large scales that make a common
suture anterior to median scale, then are separated anteriorly by
another scale; these bordered laterally by a narrow elongate lateral
scale and these in turn by a very small narrow scale; anterior to
these is a second series of four scales.

Femoral pores 5-5, the innermost situated so that it might be
regarded a preanal pore, but it is practically continuous with the
femoral series; following vent there are two or three, irregular,
granular series of scales; first two annuli of caudals small, the total
caudal series being 108; from interparietal to back level of thigh,
62 transverse rows; the scales on side of neck, about ear and postaxil-

Additions to Fauna of Costa Rica 545

lary region, and about leg insertion, smaller and somewhat irregular;
scales in a row about middle of body, 28. There is a tendency for
the smaller axillary scales to be continued posteriorly on side for
some distance. Occasionally the scale annuli are broken on back
and scales tend to alternate.

Limbs slender, pentadactyl, all clawed; lamellae under palms
and soles, as well as on basal portion of digits, flattened, padlike;
two terminal joints of digits compressed, the lamella forming a
median ridge or angle; digits more or less angular, all with strong
claws; arm with large scales except on underside of upper arm; leg
with large scales save on posterior part of thigh and on part of
ventral surface of thigh.

Color in life: Generally light brownish olive above, the head
mottled with olive; a clay-colored dorsolateral line beginning on the
head passes over eyes, along neck and side of body; dorsum brown-
ish olive above with two irregular series of black spots from occiput
to tail, even continued some distance on tail but growing more
indefinite on latter half; some scattered dots between the rows; a
dark-edged brown stripe beginning on snout passes along side of
head, becoming less distinct on side of body and tail; on this are a
few small white flecks with darker borders, especially in axillary
region, and an indefinite row of dark spots; a cream-white line from
tip of snout covers the labials and extends back, becoming continu-
ous with the white or cream color of the chin and venter; on belly
the scales have a faint scattering of brownish pigment; chin and
throat pure white. Some black spots continued on to tail; on the
tail the clay-colored dorsolateral stripe interdigitates with the lateral
brown stripe on basal part of tail but it gradually merges in the
more uniform brownish color; a row of somewhat quadrangular light
spots along ventrolateral part of tail; limbs tan with numerous, more
or less distinct, small brownish spots.

Measurements in mm. and scale data of KUMNH No. 34225: Sex,
^ , snout to vent, 63; snout to arm insertion, 20; axilla to groin, 39;
tail, 118; total length, 181; arm, 12; leg, 15; femoral pores, 5-5; scales
around body, 28; scales from parietals to back level of thigh, 63;
scale-rows from chinshields to front subcaudals, 108; scales from
nuchal collar to front level of thigh, 37.

Remarks: This specimen was discovered on a tree bole some eight
feet from the ground by John Baker. Despite its short, rather weak
limbs it ran with dexterit}^ but I finally managed to capture the speci-
men by driving it down with a long pole until it was wdtliin reach.

18—6730

546 The University Science Bulletin

Leposoma soiithi orientalis * subsp. nov.

Leposoma souHii Ruthven and Gaige (part.). Papers Mus. Zool., Univ. Michi-
gan, no. 147, 1924, p. 1-3 (specimen from Museum of Comparative Zoology,
Harvard College, from Suretka on the Sixaola River, Costa Rica, that "cor-
responds fairly well with L. sauthi with the exception of the chin shields").

Leposoma dispar Burt and Burt (pari.). Bull. Amer. Mus. Nat. Hist., vol. 61,
art. 7, 1931, pp. 347-349 (two specimens, MCZ Nos. 18916-17).

Leposoma soiithi Ruibal (part.), Bull. Mus. Comp. Zool. Harvard Col., vol.
106, no. 11, June 1952, pp. 484-485 (at least Suretka specimens, MCZ
Nos. 18916-17).

Type: KUMNH No. 36124 ^ , from Volio, Limon Province,
Costa Rica; Edward H. Taylor, collector, June 24, 1954.

Parafype: No. 36125 5 , taken with the type, same place and
date.

Diagnosis: A small species, maximum known snout-to-vent
length, 36.5 mm.; preanal pores, 2-2; femoral pores, 4-4. Differs
from Leposoma soiiihi southi in having gular scales larger and in
fewer (two to four) rows; two pairs of chinshields, both in contact;
third chinshield replaced in this form by two scales of equal or un-
equal size and the two followed by three somewhat enlarged scales;
body more slender with five, rather than seven, dorsal scale rows.
Scales around body, 19 to 21; transverse rows on venter from breast
to preanal scales, 18-20; transverse dorsal scale rows from occiput
to back level of thighs 29-30; eye smaller and ear longer; eye disc
divided into four or five scales rather than two; dorsolateral stripe
very indistinct if present.

Color: The head is a dark brownish-black save that each of
the labials has a dark spot except the sixth which is uniformly
dark. The dorsal surface is brownish tan with some scattered pig-
ment except on the anterior part of a dorsolateral scale-row
which may lack pigment entirely. The sides are more heavily
pigmented forming a dark stripe from neck to tail. On the ventral
surface the chin is cream white, the venter and base of the tail being
somewhat pinkish.

Measurements and scale data on type and parafype: Nos. 36124
^ , 36125 2 '• snout to vent, 36, 36.5; snout to arm insertion, 13.8,
13; tail, 41,t 61; total length, 77, 97.5; arm, 11.8, 9.5; leg 13, 13.2;
axilla to groin, 18, 18.2; supralabials, 6-6, 6-6; infralabials 5-5, 5-5;
scales around body, 19, 20; occiput to back level of thigh, 31,
30; subcaudals, — , 66; scales on eye disc 4-4, 3-4; femoral pores 4-4,
0-0; preanal pores, 2-2, 1-1.

* Latin, orientalis = of the East,
f Regenerated.

Additions to Fauna of Costa Rica 547

Genus Lepidophyma A. Dumeril

Lepidophyma A. Diinic'ril, in Dumeril and Dumeril, Catalogue methodique
de la collection des reptiles (Paris Museum), 1851, pp. 137-138 (type of
genus, Lcpidopliiima flavimaculafus A. Dumeril); Cope, Proc. Acad. Nat.
Sci. Philadelpliia, 1883, p. 30.

Poriodo^a.ster A. Smith, in Gray, Proc. Zool. Soc. London, 1863, p. 154 (type
of genus Foriodogastcr grayii Smith ) .

Akieistops Miiller, Verh. naturf. Ges. Basel, vol. 6, 1878, p. 390 (type of genus,
Akleistops guatcmalcnsis Miiller ) .

Description of genus: Body and tail rather rounded; head wid-
ened posteriorly; eye without distinct eyelids; no teeth on palate;
scales of head thin, smooth except for minute tubercular rugosi-
ties; nostril surrounded by three scales. Frontal very large, divided
mesially; no supraoculars present; a row of small superciliaries;
lower labials and chinshields completely fused, the labials having
the general appearance of chinshields; anterior nasals forming a
median suture; frontonasal, frontoparietals, interparietal (bearing
an eyespot), and parietals present; three enlarged temporals. Body
covered on sides and back with granules intermixed with trans-
verse (vertical) rows of larger conical or trihedral tubercles; a pair
of longitudinal tubercular rows dorsally; tail segments covered
with transverse rows of complete or partial annuli; venter with
quadrangular scales in transverse and longitudinal rows; throat
covered with equal granules; a nuchal fold; usually a longitudinal
fold on side of neck from behind ear, and a pair of folds on neck.

The genus has entered Costa Rica from the north along both
coasts and has extended itself at least to the Canal Zone in Panama.
The literature references of the genus in lower Central America are
few. In some cases the southern specimens have been referred to
Lepidophyma flavimaculatiim A. Dumeril. The following have
been reported: one from Honduras (Werner 1896); one from
Nicaragua (Rio Mico, Caribbean drainage, Gaige, Hartweg and
Stuart 1937 ) ; none reported from Costa Rica; one from Panama ( re-
ferred to Poriodogaster grayii Peters 1874); one from Rio Chilibrillo,
Panama and one from Barro Colorado I., Canal Zone (Barbour
1924, described as L. /. ohscurum) .

On the whole, the genus is a conservative one and for the most
part general body form and general characteristics remain the same
throughout. However, certain characters are affected by evolu-
tion, producing differences worthy of nomenclatorial recognition.
At the same time other, perhaps major characters may be variable
within specialized populations.

One of the difficulties in studying the genus in Central America

548 The University Science Bulletin

is that only in a few cases are adequate museum series available
from any locality.

The specimens from Costa Rica in my collection number 58, of
which 19 are from six localities in the Caribbean drainage; while
39 are from five localities in the Pacific drainage area.

The Costa Rican specimens are regarded as belonging to three
forms, two presumably confined to the western. Pacific drainage,
and one to the eastern, Caribbean drainage. They are lowland
forms, none having been found above an elevation of 3,000 feet.
It is doubtful that any of the forms are completely isolated geo-
graphically. Two of them occur together at Tilaran, Guanacaste.

Whether these forms are species may be questioned. I here re-
gard them as species, aware that future study may bring evidence
to the contrary.

Mertens ( 1952 ) has recently reported Lepidophyma stnithi
smithi Bocourt from El Salvador. While he does not describe the
conformation of the squamation in detail he does give the follow-
ing significant information:

1. A median prefrontal present (one case a double scale).

2. Three scales between the whorls of larger caudal scales ( four
to a segment).

3. Femoral pores (1 specimen) 5-7.

4. Four dark longitudinal bands on brown ground color on
which are rows of yellow flecks.

5. Chin region of all with darker markings.

Mertens had four examples, the largest having a snout-vent
length of 66 millimeters, the tail 101 mm.; two smaller specimens
were 39-42 mm.; and 59-61 mm. respectively.

It will be apparent tliat the species found in El Salvador differs
markedly from the two found in the Pacific drainage of Costa
Rica.

Key to the Costa Rican Species of Lepidophyma

1. Throat region behind chinshields strongly marked with dark brown
or black reticulations enclosing rounded cream spots. Second pair
of chinshields separated by small scales; postocular scales wider.
First loreal higher than second nasal; no trace of lateral stripe;
14-17 femoral pores; Pacific drainage,

Lepidophyma reticulatum sp. nov.
Throat region behind chinshields lacking definite reticulations,
the area usually light with nearly equal distribution of pigment,
or a few hght brown dots; second pair of labials forming a median
suture; anterior loreal lower or higher than posterior nasal; post-
ocular scales narrower 2

Additions to Fauna of Costa Rica 549

2. Femoral pores 13-15; a pair of postparietals or partial sutures marking
the same; first loreal higher than the posterior nasal. Caribbean

drainage Lepidophyma anomahim sp. nov.

Femoral pores 16-18; no trace of postparietals or partial sutures
marking same. Five transverse complete or partial caudal annuli
dorsally on each segment of tail; posterior nasal higher than first
loreal, which is separated from frontonasal. Pacific drainage,

Lepidophyma ophiophthalmum sp. nov.

The literature on the genus, especially as it applies to Central
America, is scattered. I include the following series of biblio-
graphic references for the recognized forms, flavitnaculatum flavi-
maculatiim, f. ohscurum, smithii smithii, and the doubtful grayii.
I cannot vouch for the correctness of the synonymy since it is possi-
ble that some of the reports have been confused.

Lepidophyma flavimoculatum A. Dumeril

Lepidophyma -flavimaculatus A. Dumeril, in Dumeril and Dumeril Catalogue
methodique de la collection des reptiles (Paris Museum), 1851, pp. 138-139
(type locality: restricted locality, Rio de la Pasion); A. Dumeril, Rev. Mag.
Zool., 1852, p. 409, pi. 17; Sumichrast, Arch. Sci. Phys. Math., Mar. 1873,
p. 251.

Lepidophyma spec. Miiller, Verh. naturf, Ges, Basel, vol. 6, 1878, p. 627
(Verapaz, Guatemala).

Lepidophyma jlavomactdatum Bocourt, Mission Scientifique au Mexique et
dans I'Amerique Centrale; fitudes sur les Reptiles, livr. 5, 1878, pp. 306-309,
pi. 20 F, figs. 2, a-g (also spelled jlavimactdatum on page facing plate);
Boulenger {part.). Catalogue of the lizards in the British Museum (Natural
History), 2nd Ed. vol. 2, 1885, pp. 326-327 ( Tehuantepec, Mex.); Werner,
Verh. zool-bot. Ges. Wien, Jahr 1896, p. 4 (Honduras) p. 9 (Guatemala);
Abh. Wiss., Band 22, Abt. 2, p. 345; Barbour, Proc. New England Zool. Club,
vol. 9, Jan. 24, 1924, p. 9 (southeastern part of Mexico, Yucatan and the
Caribbean coastal plain of Guatemala, British Honduras and Nicaragua
[M.C.Z.]); Stuart, Copeia 1937, no. 1, Apr. 10, p. 69 (along Rio de la
Pasion near Peten-Alta Verapaz border [exact locality unknown] ) ; Misc.
Publ. Mus. Zool. Univ. Michigan, No. 69, June 12, 1948, p. 55 (El Peten,
3 spec); Cont. Lab. Vert. Biol. Univ. Michigan, no. 45, May 1950, pp. 23,
56 (below 900 m.).

Lepidophyma smithi smithi Bocourt

Lepidophyma s^mithii Bocourt, Journ. Zool., Paris, 1876, vol. 5, p. 403 (type
locality: restricted locality, Mazatenango, Guatemala); Mission Scientifique
au Mexique et dans I'Amerique Centrale, fitude sur les reptiles, livr. 6,
1878, pp. 309-312, pi. 20 F, fig. 3, a-b; pi. G. figs, 2, a-b; Miiller, Verh.
naturf. Ges. Basel, vol. 6, 1878, p. 627 (Mazatenango, Guatemala); Barbour,
Proc. New England Zool. Club, vol. 9, Jan. 24, 1924, pp. 9-10 (Pacific
coastal plain of southwestern Mexico and Guatemala).

Akleistops guatemaJensis Miiller, Nat. Ges. Basel, vol. 6, 1878, pp. 390-398,
pi. 1; pi. 2, figs. 1-5 (type locality: "Mazatenango, coste grande de Guate-
mala" ) .

Lepidophyma maculatnm ( part. ) Boulenger, Catalogue of the Lizards in the
British Museum, 2nd Ed., vol. 2, 1885, pp. 326-327.

Lepidophyma smithi smithi Smith, Proc. U. S. Nat. Mus., vol. 92, no. 3153,
1942, p. 380; Smith and Taylor, U. S. Nat. Mus. Bull. 199, 1950, p. 152;
Mertens, Abh. Senckb. Naturf. Ges., No. 487, 1952, p. 53, pi. 11, fig. 61
(San Salvador).

550 The University Science Bulletin

Lepidophyma graijii (A. Smith)

Puriodogaster grayii A. Smith, in Gray, Proc. Zool. Soc. London, 1863, p. 154, pi.
21 (unknown locality); Peters, Proc. Zool. Soc. London, 1874, p. 307.
( Specimen from Panama thought to be identical with this species, suggesting
Panamd as the type locality. Bocourt states that this specimen has 17
femoral pores on each side ) .

Lepidophyma flavimaculatitm obscurum (Barbour)

Lepidophyma flavonwculaturn obscurum Barbour, Proc. New England Zool.
Club, vol. 9, Jan. 4, 1924, pp. 9-10 (type locality, Rio Chilibrillo, Panama;
known also from Barro Colorado Island, Canal Zone).

Barbour has based his form on two specimens from Panama ( Rio
Chihbrillo, and Barro Colorado Island, Canal Zone). His descrip-
tion is extremely brief consisting only of the following: "In squama-
tion very similar to L. f. favotiiacidotmn. It differs in having rela-
tively a slightly smaller and narrower head; in having the longi-
tudinal rows of enlarged dorsal scales more straight, more even, and
more closely and regularly beset with longer spine-like scales. The
color is blackish brown, and the yellow rosette-like spots are only
about one half of the size, each, of those in the northern subspecies,
of which apparently it is an offshoot, typically colored for the humid
forest, as the other is for a more xerophilous life." Whether this is
identical with Peters' specimen ( see above ) can only be determined
when the two specimens are compared and these in turn compared
with the type of Poriodogaster grayii.

I have examined one Panamanian specimen identified as L. jlavi-
maculatiini obscurum ( USNM 65809 ) . In this there were no lateral
light spots on the back and sides of body but some at base of tail.
There are five labials preceding the subocular labial that have small
cream spots. The venter is uniform dark brown. The tail is broken
and a single segment remains, so that the character of the caudal
squamation cannot be ascertained. The femoral pores are 13-14
(female). The preanals consist of a posterior pair bordered laterally
by several tiny scales. The pair preceding these is smaller and
longer than wide and preceded by two small regular pairs. The
medial prefrontal is large, about half the area of the outer prefrontal.
There is no trace of postparietals.

This specimen measures 103 mm. snout to vent; snout to arm, 39.5
mm.; axilla to groin, 48 mm.; arm, 30 mm.; leg, 41 mm.; width of
head, 17 mm.; length of head, 25 mm.

Additions to Fauna of Costa Rica 551

Lepidophyma reticulatum * sp. nov.

(Fig. 14)

Type: KUMNH No. 36245 $ , Agiia Buena, Puntarenas Province,
Costa Rica, July 25, 1954; Edward H. Taylor collector,

Parafypes: Nos. 36238-36244, 36246-36249, topotypes, same date
and collector; Nos. 34189-34193, 3 km. E San Isidro del General,
San Jose Province; Nos. 34194-34202, 34204, 15-17 km. WSW San
Isidro del General, Puntarenas Province; Nos. 36230-36237, 4-5 km.
NE Tilaran, Guanacaste.

Diagnosis: Large sf)ecies with approximately 38 caudal segments,
each, except first, with five rows of partial or complete scale annuli
crossing dorsal surface of tail (rarely six), the terminal segments
not clearly marked; a median prefrontal absent (or present in some
specimens in the San Isidro del General region); no trace of post-
parietals; femoral pores in both sexes, 14-17 (average 15.4); the
first loreal higher than posterior nasal and touching frontoparietal;
only first pair of chinshields in contact behind mental. Throat
strongly reticulated with dark blackish-brown enclosing cream
spots; scales with small glandular pores.

Description of type: Head nearly triangular seen from above;
snout rather pointed, the rostral wider than high, narrowly visible
above; anterior nasals broader than high, forming a median suture;
frontonasal eight-sided, broader than long; two prefrontals, their
outer edges bent down over canthus rostralis, smaller than the di-
vided frontals; outer edges of frontal separated from orbit by a small
row of supraciliaries; no supraoculars; interparietal slightly wider
anteriorly than posteriorly, pushing in between posterior part of
frontal scales, separating broad frontoparietals completely, and
touching both parietals; latter longer than wide bordered laterally
by two temporals the anterior of which is more than triple size of
posterior; nostril surrounded by three scales: first labial, anterior
and posterior nasals; first loreal higher than nasals, its length about
one third the length of the second loreal which is nearly as high; two
presuboculars and a very narrow linelike preocular; five supracili-
aries; five postoculars; two minute suboculars separate fifth supra-
labial from the orbit; first upper temporal twice as wide as postocu-
lars, lower small; between sixth and seventh labials and second
upper temporal two rows of small temporal scales; seven supra-
labials (eighth scale very small); the seventh and eighth scales

* Latin, reticulatus = reticulated, netlike.

552

The University Science Bulletin

Fig. 14. Inner figure, Lepidophyma reticulatum sp. nov. Paratype. KUMNH
No. 34194 5. 10 mi. SW, San Isidro del General. Outer figure, Lepidophyma
anomalum sp. nov. KUMNH No. 34207 ? . Type. Actual length 233 mm.

separated from mouth by granules; a row of somewhat larger tuber-
cles border edge of ear opening, and these preceded by two or
three still larger moundlike scales; infralabials (fused labials and
chinshields) four, only the first pair in contact; second pair largest,
narrowly separated mesially; fourth about one-fifth times as large as

Additions to Fauna of Costa Rica 553

third; posterior part of lower lip bordered by several granules;
mental large, its labial border three times that of rostral border; on
throat and sides of neck, conical granules; ear-opening large, its
greatest diameter approximately 4 mm.; vertical diameter of orbit,
3 mm.; length of eye to tip of snout, 7.5 mm.; distance from orbit to
ear-opening 8 mm.; a pair of ventral longitudinal folds on neck;
a pair of short lateral folds running back from ear; a transverse
nuchal fold.

Body covered with small conical or moundlike granules inter-
mixed with larger conical, trihedral or irregularly shaped tubercles
that tend to form vertical rows on sides of body; a pair of longitudi-
nal rows on dorsum separated from each other by four or five rows of
granules; tail segments marked by series of annular rings of scales
increasing in size posteriorly; the first two definite segments with
large tubercular whorls, and these separated by two small granular
rows; first typical segment with five partial or complete annular
scale rings, the scales growing larger towards the ventral part so
that the median ventral part of each segment has only three scales;
tip of tail regenerated in type. Scales on limbs varied in size,
the hind leg especially with numerous larger tubercles; palm and
sole with somewhat conical granules; digits with varied ventral
scales, those on proximal regions conical or trihedral; in medial
parts digital lamellae divided unequally, distally they are complete
(or an occasional one divided).

Scales on breast flat, subimbricate, becoming elevated laterally;
venter with flat juxtaposed scales arranged in eight longitudinal
rows with an addition of a partial lateral row of more elevated scales
on each side; about thirty transverse rows from axiliary level to the
inguinal level.

Scales on under side of thigh somewhat subimbricate anteriorly;
femoral pores 17-15; two pairs of larger (slightly irregular) preanal
scales preceded by a very small pair.

Color in life: Head brown with some indefinite dark marks tend-
ing to form a somewhat symmetrical (somewhat radiating) pattern.
Supralabials with cream marks on sutures; infralabials each with a
large black spot or diagonal bar separated by cream; throat strongly
reticulated with brownish black enclosing cream spots. On side of
head and neck rounded cream spots surrounded by black; bodv
black on back and sides with six longitudinal series of irregular
cream spots; arms and legs with a few conspicuous spots. A pair
of larger cream spots on tail above vent; segments of tail with a
series of vertical cream lines chiefly on the largest tubercular scale

554 The University Science Bulletin

series; median part of dorsal surface black; ventral surface with
cream predominating anteriorly and black posteriorly; femoral
pores and adjoining scales cream; venter dirty cream with some
dark flecks along ventrolateral region; fingers and toes ringed with
cream; anal scales bluish white.

Measurements in mm.: Snout to vent, 92; tail, 120; length of head,
20; width of head, 16.2; snout to arm, 33; axilla to groin, 45; arm, 30;
leg, 41.

Variation: The topotypic specimens agree closely with the type
in most characters. There is no trace of the median prefrontal; the
first loreal has the same relation to the posterior loreal and the
posterior part of the nasal, and the femoral pores average approxi-
mately 16 on each side. Two have the preanal scales irregular.
The scale rows on the dorsal part of the tail segments are five, but
in two specimens six may appear on one or two segments. The
total number of segments is approximately 38, however, on the
posterior part of the tail the segments are not well differentiated.

The specimens that occur at San Isidro del General differed most
in that a median prefrontal was present in those from near the town,
while in those from a somewhat lower elevation, at a locality 15-17
kilometers west-southwest of the town, five have the prefrontal
present, four have the scale partially segmented. Two of these
specimens have six annular rings on several segments of the tail.

Those taken still farther north at Tilaran lack all trace of the
median prefrontal. The head markings are very indefinite. One
specimen has a bifid tail, where it has been regenerated. There are
other smaller, less significant characters that can be used to separate
these various groups of specimens. In all, the throat maintains the
heavy reticulation, the postocular series is enlarged, and the anterior
temporal is proportionally small.

Lepidophyma anomalum * sp. nov.
(Fig. 14)

Type: KUMNH No. 34207 5, Los Diamantes, Guapiles, Limon
Province, Costa Rica; Sept. 3, 1947; Edward H. Taylor and Richard
C. Taylor, collectors.

Paratypes: KUMNH Nos. 34205-34206, 34213, topotypes, same
date and collectors; No. 34210, Bataan, Limon Province; Nos. 34211-
34212, 34214-34216 La Lola, Limon Province, Nos. 36256-36257,
Puerto Viejo, Limon Province; No. 34208, Morehouse Finca, Turri-

Greek, anomalos = abnormal.

Additions to Fauna of Costa Rica 555

alba, Cartago Pro\ ince; Nos. 36251-36255, Suretka, Liiuou Pro\ iiice;
No. 34209, Costa Rica (uncertain proNcnance).

Diagnosis: A large species, with approximately 35 caudal seg-
ments, the basal seven to twelve with four complete or partial scale
annuli, dorsallv; the following segments with five annuli; terminal
three or four segments not clearly differentiated; a median prefrontal
normally present; first t\vo pairs of infralabials in contact behind
mental; femoral f)ores in males and females 13-15 (average about
14); throat cream with a few tiny, scattered, brown dots the size
of the scales between neck folds, but no reticulation of black or
brown and no cream spots; usually a pair of postparietals completely
or partially segmented from parietals; body and caudal scales each
with a small glandular pore. Two dorsal rows of spots, light tan;
lateral spots cream.

Description of type: Head, seen from above, subtilangular,
pointed anteriorly, the border of the head plates tending to be
slightly angular posteriorly on the occiput, and with a slight median
longitudinal depression in the parietal region; rostral wider than
high, angular medially; anterior nasals form a median suture about
two thirds the greatest width of the scale; frontonasal eight-sided,
and distinctly wider than long, touching first loreal; three pre-
frontals, the median little more than one-third times the area
of the outer scales; frontal divided mesially, separated from
orbit by superciliaries, angular posteriorly; interparietal bearing an
eye-spot, subhexagonal, only slightly wider anteriorly, somewhat
narrowed near the middle, separating completely the frontoparietals;
latter scales with their long axis diagonal, touching one supraciliary,
one postocular and two temporals; parietals distinctly longer than
wide; postparietals only partially indicated by a pair of grooves
directed diagonally backward from the sutures with the interparie-
tal; second nasal much higher than first, much higher than long
(wide), not as high as second loreal, not separating it from fronto-
nasal; first loreal as high as second but only half its length ( width ) ;
a slender linelike preocular four times as high as wide; one pre-
subocular; four narrow postoculars and five small supraciliaries;
first temporal much wider (four times) than adjoining postoculars;
second temporal large, about five times area of first temporal, about
four times area of second; eight supralabials, first high forming with
the nasals the border of nostril, fifth below eye and separated from
the orbit by a linelike scale, sixth largest, touching two or three
postoculars, but separated from the first and second temporals by

556 The University Science Bulletin

one or two rows of granular temporals, last labial very small; three
anterior infralabials ( fused labials and chinshields ) large, both form-
ing median sutures, the second pair partially separated on the me-
dian line; last two small with two differentiated scales bordering
them; mental large, angular posteriorly; a small anomalous triangular
scale (true infralabial? ) lying between the mental and first large
infralabial on each side; auricular opening large, with a row of
about seven larger tubercles along its outer border, its height four
millimeters; orbit of eye circular; no evidence of eyelids; orbital
diameter three fourths the height of ear-opening; distance between
orbit and tip of snout (7 mm.) less than distance between orbit and
ear; a pair of longitudinal folds along outer ventral surface of neck;
lateral fold from ear to shoulder, and a transverse nuchal fold;
body and sides covered with small, conical or moundlike granules,
intermixed with larger, conical, trihedral, or pyramidal tubercles,
the latter forming vertical, transverse rows laterally; a pair of longi-
tudinal rows of tubercles on back, separated by five rows of sub-
equal granules; chin (partly) and throat covered with subequal
granules; breast and venter covered with flat, juxtaposed, quad-
rangular scales, forming ten longitudinal rows, and between axilla
to groin levels, about 27 transverse rows; scales on breast smaller,
somewhat irregular; tail with each segment bearing posteriorly a
whorl of larger tubercles; on first basal segment the large whorls
are separated by two, on second by four smaller whorls, while the
following six segments have them separated by three complete or
partial whorls; from here on they are separated by four whorls.
Thus each segment has either four or five complete or partial whorls
dorsally; but ventrally, where the scales are larger, there are two or
three only.

Arms and legs covered with granular scales, with some intermixed
tubercular scales, especially on upper parts of hind leg; those on
arm semispinose; some much enlarged scales on outer ventral sur-
face of tibia; subdigital scales may be granular near base, the lamel-
lae tending to be unequally divided except on the distal half. Female
with fifteen pore-scales, most of which are pierced, and the three
or four proximal and four or five distal ones show larger, perhaps
functional pores. First preanal scales large, longer than wide, with
a small scale on each side, and preceded by a second pair; these
in turn preceded by a row of four scales.

Color in life: Dorsal part of head brown, with evidence of some
darker marking, the area above eye distinctly darker; body black

Additions to Fauna of Costa Rica

557

dorsally and laterally, with six rows of spots, those on the dorsal
surface somewhat indistinct, tan in color; lateral rows of spots
brownish cream; tail blackish brown with very narrow vertical
cream lines marking tubercular whorls on sides of tail; a vertical
cream line in front of eye; cream dots on supralabials between black
spots; infralabials with black spots or bars separated by brownish
cream (fawn); chin dirty white, with a few fine brown flecks; cream
spots present on cheeks and on sides of neck; the periphery of the
ventral scales with much brownish pigment leaving the central part
of scales lighter; preanal scales bluish white; subcaudal scales brown,
mixed with cream; femoral pore-scales cream; arm and leg blackish
brown with some cream spots.

Variation: The femoral pores are smaller in this species and the
number is somewhat reduced varying between 12-15 ( one specimen
has 16 on one side). Four of the specimens (the type, and Nos.
34208, 34215, 34216) have small intercalary "labials" following the
mental on the border of mouth. In the last three specimens and
No. 36257 there is a small scale segmented from the outer anterior
part of the parietals on one or both sides; only two specimens have
the median interparietal completely missing; two specimens have
five labials anterior to the subocular but this is a case of the second
or the fourth labial being segmented. Two specimens have the large
temporal segmented into two unequal scales, the segmented section
being the outermost part. The most striking variable character
is the development of a pair of scales ( postparietals ) segmented
from the inner parts of the parietals behind the interparietal. Eight
of the specimens (6 of them from Bataan and La Lola) have the
pair complete or practically so; six others show partial suturing of

Table of Measurements of Lepidophyma anomalum sp. nov.

Number

Snout-
vent

Tail

Axilla

to
groin

Arm

Leg

36257

34207

36254

104
96
84
81
71
67
62
44
42.5
42
41
41
38
38
35

132*

137

117

*

*

*

86

62

*

*

53

53

50

47

*

31

29

29

27

22

22

21

16

14.5

15

13.2

13

12.2

12

12

42

40

39.5

38

30

30

26

19

18

21

18

18

17

17

16

49
47
39

34211

36255

36253

36
31
30

34208

34205

28
19.5

34206

18.2

36256

23

36252

18

36251

18

34213

16.2

34215

16

34212..

16

Tail regenerated or partly missing.

558 The University Science Bulletin

the scales while five from Suretka, Los Diamantes, and Puerto
Viejo have no trace of this character. It was noted that in three
tiny specimens from La Lola the loreal was not higher than the pos-
terior nasal and in one case the posterior nasal excluded the first
loreal from the frontonasal. Two specimens showed some variation
in the character of the anal scales by having a partial row of small
scales separating first and second pairs. In the younger specimens
the basal subcaudals seem to be slightly keeled. In one, there were
two presuboculars. The pores in the body and caudal scales are
small; those on the venter can be distinguished only with difficulty.

Lepidophyma ophiophihalmum* sp. nov.
(Fig. 15)
Type: KUMNH No. 36250; 5 km. NNE Tilaran, Guanacaste,
Costa Rica, Aug. 16, 1954; Edward H. Taylor, coll.

Pawtypes: KUMNH Nos. 34217-34219, Las Flores, Tenorio, Las
Canas, Guanacaste (near base of Volcan Tenorio), Aug. 22, Edward
H. Taylor and John Baker, colls.

Diagnosis: Superficially closely resembling Lepidophyma reticti-
lattim in color and marking. Median prefrontal present or absent;
chin lacking reticulation, light colored save for brown dots the size
of granules; posterior nasal much higher than anterior, and higher
than first or second loreal; first loreal separated from frontonasal;
second pair of labials forming a common suture; vertical diameter
of ear-opening a half greater than diameter of orbit; frontonasal
hexagonal; 35 caudal segments all with five complete or partial
annular scale rings except terminal three or four where segments
are ill defined and may be reduced to four annuli; parietals propor-
tionally longer and narrower than in reticulatum; frontoparietal
narrowed laterally where it touches the last supraciliary and upper
postocular; first pair of preanal scales bordered laterally by a small
preanal; a partial or complete row of small scales between first
(larger) and second preanal pair; femoral pores 16-18; anterior
temporal distinctly larger than in L. reticulatum.

Description of type: Head outhne generally triangular seen from
above; snout pointed; rostral wider than high, rising to a sharp
angle mesially, distinctly visible seen from above; anterior nasals
wider than high, their inner median height at suture little more than
half of outer height at nostril; frontonasal hexagonal, slightly broader
than long; two large prefrontals, their outer edges bent down over

* Serpent eye. Greek. From ophi snake and ophthalm eye.

Additions to Fauna of Costa Rica

559

Fig. 15. Lepidophyma ophiophthalmum sp. nov. Type. KUMNH No.
36250, five km. NNE Tilaran, Las Canas, Guanacaste Province, Costa Rica.
Total length, 183 mm.

560 The University Science Bulletin

canthal edge, laterally touching posterior nasal, both loreals, first
supraciliary and preocular; frontal large, its front borders straight,
transverse, angular posteriorly, separated from orbit by a row of
three or four somewhat projecting supraciliaries; frontoparietals
six-sided, narrowed anteriorly; interparietal wider anteriorly, sub-
hexagonal, separating completely the frontoparietals, with an "eye-
spot" that is white with a black spot showing in its middle; parietals
(6 mm. X 3/2 mm.) six-sided; first temporal relatively large (2.5 mm.
X 2 mm ) ; second temporal very large, third, less than one-third times
size of second; scales in preauricular region small, variable in size;
auricular edge bordered by a row of seven heavier scales nearly
hiding another smaller inner row; a pair of longitudinal skin folds
on outer ventral surface of neck; a less distinct fold from behind ear
to nuchal fold; a transverse nuchal fold. Eye circular without evi-
dent lids, the diameter of orbit ( 2.8 mm. ) less than height of auricu-
lar opening (3.6 mm.); distance between tip of snout and orbit
(6.5 mm.) less than distance between orbit and ear. Body covered
with small conical or moundlike granules, interspersed with large
conical or trihedral, spinelike tubercles forming vertical lateral rows;
a pair of longitudinal rows on dorsum, separated from each other by
four or five irregular rows of subequal small granules; the larger
tubercular scales form continuous series around tail, one large series
on posterior part of each tail segment. Nostril surrounded by three
scales; posterior nasal higher than loreals or anterior nasal, widened
at top, separating the first loreal from frontoparietal, touching pre-
frontal; anterior loreal twice as high as long, one third length
(width) of second but nearly as high; a long, narrow preocular;
one presubocular; three postoculars, lowest largest; fifth labial be-
low orbit separated from it by two granular scales; sixth labial
separated from first two enlarged temporals by a small lower anterior
temporal and two granular rows of temporals; first (upper) tem-
poral more than three times width of upper postoculars. Seven
supralabials, first reaching up to nostril, sixth largest, nearly twice
as long as high; last small, separated from edge of lip; four lower
labials very large ( fused labials and chinshields ) ; two anterior pairs
making mesial sutures, the second pair much larger than others;
mental with a labial border double that of rostral, angular posteri-
orly; posterior part of the mouth opening bordered by granules.
Tail segments approximately 36, each covered with five complete
or partial rows of subconical scales, the posterior row composed
of much larger tubercles; ventrally each segment with three, rarely

Additions to Fauna of Costa Rica 561

four scales, larger than the lateral or dorsal scales; arms and legs
covered with small granules intermixed with larger ones, some of
those on under surface of thigh subimbricate; femoral pores 16-16
in small single scales; subdigital lamellae variable, sometimes di-
vided or broken up into small tubercular scales at base of third and
fourth toes; distal lamellae usually complete; scales on venter flat,
quadrangular, subequal except those on breast which are smaller,
arranged in 10 longitudinal rows and 26 transverse rows from level
of axilla to level of groin; on part of the transverse rows one counts
12 scales, but the outer row on each side is irregular, variable, the
scales not flat. Most if not all scales on body and tail with pores or
glands visible under magnification. Preanal scales consist of a
posterior row of two much enlarged scales and two smaller lateral
scales; median row broken leaving the third pair on left side touch-
ing a large posterior scale; third row has two scales, larger than
those in median row.

Color in life: Head brownish with some indefinite darker mark-
ing; black spots on loreals and supralabials separated by small cream
spots; infralabial and mental with dark bars, separated by white;
chin dirty white with a few brown granules; sides of neck with large
cream spots on black background; body black with six irregular
rows of lighter spots; venter scales heavily dusted with brown leav-
ing a whitish central area; outer rows of ventral scales with a nearly
continuous, irregular white line; femoral pore-scales white. Arms
and legs spotted white; tail with irregular white spots on sides, one
to each segment, variable in size and position; under side of tail
blackish with some white scales; preanals bluish white.

Measurements in mm.: Snout to vent, 77; tail, 106; head width,
13.8; head length, 18; snout to foreleg, 31; snout to ear, 18.5; axilla
to groin 36; arm, 23.5; leg, 35.

Variation: The three paratypes are from a locality 20 kilometers
northwest of the type locality, at Las Flores on the large cattle ranch
of Tenorio, a place situated near the base of Volcan Tenorio but at
about the same elevation as the type locality. The two smaller
specimens have a median prefrontal which is lacking in the adult.
The median row of preanals is complete in the young, broken in the
adult. There are 16 to 18 femoral pores present in all.

562 The University Science Bulletin

Anomalcpls dentatus Taylor

Anomalepis dentatus Taylor, Proc. New England Zool. Club, vol. 17, June 26,

1939, pp. 90-91, pi. 5, figs. 1-3 (type locality, Barro Colorado Island, Canal
Zone); Taylor, Univ. Kansas Sci. Bull., vol. 34, pt. 1, Oct. 1, 1951, pp. 24-25.

A small snake of the genus Anomalepis taken at a point about
four kilometers southeast of Los Angeles de Tilaran, Province of
Guanacaste, proves to belong to this species. The specimen is 156
mm. in total length, of which 3.1 mm. represents the length of the
tail.

The scale counts are as follows: Scales from rostral to tip of tail,
271. On the ventral surface from mental to vent, there are 264
scales; from vent to end of tail seven subcaudals. The longitudinal
scale-rows about body are 28, 24, 22, 20. When scale-rows drop out
they do so by a fusing of two scale-rows, thus forming some en-
larged scales. The subsequent scales become gradually smaller.
Two mandibular teeth are present, separated by a mesial diastema.
The conformation of the head scales is very much the same as de-
picted by Taylor, loc. cit. save that the scales, lateral to the frontal,
are slightly larger than they appear in the dorsal view of figure 3
since the scales are drawn in perspective. The preanal scales are
in contact posteriorly.

No evidence whatever has been brought to bear that dentatus is
a synonym of Anomalepis mexicanus. The differences as portrayed
in the characteristics of the large head scales, especially their pro-
portions, is strong evidence to the contrary.

Leptotyphlops ater * Taylor

Leptotyphlops ater Taylor, Univ. Kansas Sci. Bull., vol. 26, no. 15, Nov. 15,

1940, pp. 536-538, fig. 4 (type locality, Managua, Nicaragua).

A Costa Rican specimen of this species was taken at Bagaces at
a point where one of the streets enters the Pan-American Highway.
It was in loose earth below a flat rock. It is presumably a juvenile
specimen, its total length being 76 millimeters, of which five are in
the tail. The tail turns down and ends in a rather sharp spine.

The dorsal and lateral surfaces are dark blackish but when sub-
merged in a clear liquid it shows a very faint suggestion of darker
medial areas on the scalerows. This probably is not an adult con-
dition since the type displays no trace of such markings. This speci-
men differs but little from the type in the characters of the squama-

* Taylor I. c. p. 540, couplet 22 states: "only spine on tail cream"; and "tail in tail
length, 14"; This should read, cream spot on tail including about four transverse scale rows;
(as stated on p. 533) and, tail in tail length 16. Dunn and Saxe 1950 point out this error.

Additions to Fauna of Costa Rica 563

tion. The conformation of the head scutes is practically identical
with that shown in the figure of the type — the nasal is completely
divided, a supraocular bears the same relation to the rostral and
anterior (superior) nasal, the labials, parietals, frontal and inter-
parietals likewise, are practically identical proportionately. The
eye is scarcely discernible in the ocular. The last labial and the
ocular have their lower parts whitish and the infralabials are light.
The anterior part of the snout is dark gray. The anterior part of the
venter is grayish white, growing darker posteriorly.

There are 14 scale-rows about body and 10 about tail and approx-
imately 245 transverse scale-rows from the rostral to the tip of the
tail. There are 19 subcaudals. The anal is a large single scale pre-
ceded by a narrow, transversely widened scute, at least as wide as
the anal.

The width of the head is 1.6 mm. and of the body 1.7 mm.*

The ratio of tail length to total length is 15; the ratio of the width
of the body to the total length is 47.5. In the type these two ratios
are 18.9 and 60 respectively. The scales from lip to vent number
225. It will be necessary to verify the number of scales in the type
as the statement "259 scales from snout to vent" is probably intended
for "snout to tip of tail." Even if this were true the present specimen
has fewer transverse scale rows on the body than the type.

This species is not a member of the olbifrons group.

Conophis lineatus nevermanni Dunn

(Fig. 16)

Conophis nevermanni Dunn, Copeia 1937, no. 4, pp. 214-215 (type locality,
Rio Poas de Aserri [near San Jose] Costa Rica); Taylor, Univ.' Kansas Sci.
Bull., vol. 34, pt. 1, Oct. 1, 1951, p. 145; Savage, Trans. Kansas Acad. Sci.,
vol. 52, no. 4, 1949, pp. 484.

A specimen of this species, KUMNH No. 35630, taken from under
a stone in an open meadow, two miles south of Hotel Yomale, some
32 kilometers north of Barranca, Puntarenas Province, but in, or near,
the boundary of Guanacaste Province, is referred to this species
with some hesitancy. The markings and scale characters of the
specimen follow:

The venter is ivory to yellow ivory with a row of black dots on
the lateral part of ventials throughout the length of the body. A
black line running through the middle of the first (outer) scale-row

* The width of the body in the t>pe is given as 305 miUimeters, an obvious typographical
error for 3.03.

564

The University Science Bulletin

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Fig. 16. Conophis lineatus nevermanni Dunn. KUMNH No. 35630; about
32 km. N Barranca, near the boundary of Guanacaste and Puntarenas Provinces,
Costa Rica.

is continued a little past the base of the tail. Above this, an ivory
line covering one whole scale-row and the edges of adjoining scale-
rows runs from the tip of the snout along the body and onto the tail
bearing a series of black dots along the latter half of the body and

Additions to Fauna of Costa Rica 565

posteriorly is lost in the ventrolateral color of tail. The broad black
band, beginning on the tip of the snout runs along the neck where
it covers about four scales, then narrows, occupying all except the
outermost edges of the third and fourth rows. A lighter streak
is present in the middle of this stripe, evident only for a part of its
length. Above this is an olive-gray stripe on the fifth and sixth rows,
becoming lighter cream or ivory posteriorly, and throughout it is
lighter on the lower part of the fifth row. This is bordered above
by a narrow black line not more than one scale wide, on the seventh
row anteriorly, on the sixth posteriorly. The five rows (four pos-
teriorly) lying between the seventh row and its fellow of the
opposite side are blackish on the back of the head and neck, but
become olive-gray more posteriorly. The color is not uniform. The
median dorsal row has a line of dashlike black marks, and a row of
darker flecks occur on the edges of the eighth and ninth rows of
each side. The upper part of the supralabials have a cream or
ivory line with a ragged lower edge while the lower part of the
labials show pigmented areas of black, the color not solid. The
infralabials, chin, and part of throat pigmented with brown, enclos-
ing two short rows of rounded ivory spots.

The following scale characteristics obtain: ventrals 169, sub-
caudals 70, anals 2, supralabials 8, the fourth and fifth enter orbit;
infralabials 9, preocular 1, postoculars 2, anterior temporals 2, scale
rows, 21-19-19-17. ,

This form definitely is not Conophis lineatus dunni Smith from
Nicaragua and it is possible that it is not identical with Dunn's
nevermanni. The later is very briefly described and the type is said
to be in Dunn's private collection.

Leptodeira sp.

Hypsiglena torquata torqttota Tavlor, Univ. Kansas Sci. Bull., vol. 36, pt. 2,
no. 11, July 15, 1954, p. 712-713, fig. 9.

In the above listed publication I reported and figured a juvenile
snake as a species of Hypsiglena. The specimen had been badly
injured, the head having been subjected to extensive injury in the
palatal and dental regions. After the publication I was struck by
the large number of subcaudals, a character which should have
caused me to question the association with Hypsiglena from the
first. After careful re-examination of the specimen, a fragment of
the posterior mandible was found showing the base of a broken, en-
larged, posterior, grooved tooth. This fact, together with the scale
characters and the high subcaudal count, places the specimen in the
genus Leptodeira, as now understood.

566 The University Science Bulletin

I think it unwise to attempt to allocate the form specifically owing
to its mutilated condition and the fading of the colors and markings.
New material should be at hand before this is undertaken.

Coniophanes piceivittis piceivittis Cope

(Fig. 17)

Coniophanes piceivittis Cope, Proc. Amer. Philos. Soc, vol. 11, July 1869, pp.
149-150 (tvpe locality, Chihuitan, Oaxaca, Mexico); Bailev, Papers Michigan
Acad. Sci. Arts, Lett, vol. 24, pt. II, 1938 (1939), pp. 29-31, pi. 2, fig. 2
(Bebedero, Costa Rica); Taylor, Univ. Kansas Sci. Bull., vol. 34, pt. 1, no. 1,
Oct. 1, 1941, p. 142.

Coniophanes piceivittis piceivittis Mertens, Abh. senckenb. naturf. Gesel.,
no. 487, 1952, p. 61 (San Salvador).

In my paper on Costa Rican serpents I suggested that Bailey's
record for this species in Costa Rica might be questioned. The
finding of two specimens in western Costa Rica, four miles north of
Hda. Santa Rosa (on Pan-American Highway), Guanacaste Prov-
ince, in the summer of 1954, verifies its presence in the Covmtry.
These were taken by me in the vicinity of a small stream, in a pile
of chips about the base of a recently felled tree. The arrangements
of the markings on the body and head are as follows: first to third
scale-rows with edge of the adjoining row, lavender cream, pep-
pered lightly with lavender brown; anteriorly and posteriorly only
two whole scale-rows are involved; above this on the fourth to
eighth scale-rows a broad straight-edged black stripe which begins
on the rostral and runs back, covering upper part of labials, and on
neck, covers four whole scale-rows and the adjoining halves of two
others; it gradually narrows to three rows and parts of the adjoining
rows on the posterior parts of the body; a narrow, irregularly edged,
cream line begins on rostral, passes above eye along edge of parietal
to the neck where the edges become straight, and continues on to
tail, covering the ninth whole row and parts of the adjoining scale-
rows. The median black stripe covers the top of the head then con-
tinues to tail, covering five median scale-rows and the adjoining
edges of two others, but reduces to three and two half rows on pos-
terior part of body. The supralabials are cream, dark-edged above
with a splashed black spot on the lower part of each scale. The in-
fralabials, chinshields and scales at beginning of throat flecked with
black. The ventrals are uniformly cream except for a few darker
flecks on their upturned lateral edges. The second specimen agrees
in rather accurate detail with the first.

Scale characters of these specimens, KUxVINH No. 23553 and
23554 follows. Ventrals 165, 163; subcaudals 62 +, 83; scale rows
23, 25, 25, 19; 23, 25, 25, 19; supralabials, 8-8, 8-8; infralabials 10-10,

Additions to Fauna of Costa Rica

567

Fig. 17. Coniophanes piceivittis piceivittis Cope. KUMNH No. 23553, 4
mi. N Hda. Santa Rosa (on Pan-American Highway), Guanacaste, Costa Rica.
About natural size.

568 The University Science Bulletin

10-10; preoculars 2-2; postoculars 2-2; temporals 1 + 2 + 4, 1 + 2
+ 3; the scales on sides at least three inches in front and behind
level of vent strongly keeled or tubercled; the anal scale is divided.

Neopareas bicolor Giinther

(Figs. 18, 19)

Neopareas bicolor Giinther, Biologia Centrali-Americana; Reptilia and Batrachia,
July 1895, pp. 178-179, pi. 56, fig. C (type locality, Chontales Mines, Nica-
ragua); Taylor, Univ. Kansas Sci. Bull., vol, 34, 1951, p. 66 (incorrectly
spelled Neoparias ) .

Leptognathus bicolor Boulenger, Catalogue of the Snakes in the British Museum,
vol. 3, 1896, p. 460.

A specimen of this genus was obtained at Turrialba at the Inter-
American Institute of Agriculture in 1952. The animal had gained
admittance to the building of the Institute and was captured in one
of the dormitories. The characters of this specimen are as follows:

Description of KUMNH No. 31920: Body strongly compressed;
head short, the width of the neck (2.6 mm.) in width of head
(7 mm.) 2.7 times; rostral wider than high, narrowly visible above
as a broad triangle, the lower part of scale with an arched depres-
sion; internasals small, about one third area of the prefrontals, much
wider than long, their common suture half that between the pre-
frontals; latter scales broader than long, touching nasal, loreal, and
preocular laterally; both prefrontals and internasals with minute
pustular rugosities; frontal pentagonal, shield-shaped, the anterior
edge straight, transverse, its length (3.2 mm.) greater than its
width ( 2.8 mm. ) ; supraoculars narrowed anteriorly, widened pos-
teriorly, not reaching back as far as frontal, but extending somewhat
farther forward; parietals large, longer than frontal, their length
equal to their distance from the internasals. Nasal single, irregular
in shape, the back edge of the scale elevated and somewhat exca-
vated behind nostril; loreal large, higher than long, bordering orbit
below the smaller preocular; three postoculars, the lower smallest;
temporals, 2 + 2 + 3, ^ + L±A ; supralabials 10-9, the fourth to
seventh ( third to sixth ) entering orbit; mental small, much narrower
than the rostral; infralabials, 11-12, first five touching the large
azygos scale on chin, two (three) labials touching second azygos;
third and fourth azygos each in contact with two labials; first pair
of labials fused behind the mental; labials and azygos scales on chin
with minute rugosities ( male character ) . Eye very large, its diame-
ter (2.5 mm. ) greater than length of snout (2.1 mm. medial measure-
ment).

Scales smooth, lacking pits; scale formula: 19 (back of head),

Additions to Fauna of Costa Rica

569

Fig. 18. Neopareas bicolor Giinther. A. Head, lateral view. B. Head, dorsal
view. C. Head, ventral view. All much enlarged.

570

The University Science Bulletin

15, 15, 15; the median dorsal row not or but scarcely larger than
adjoining rows; two broadened, dorsal scales above anal region;
scales behind the fourth azygos not differentiated from typical ven-
trals and the ventral count begins at that point: ventrals, 186; anal
single; subcaudals, 118; total ventral-subcaudal count 304.

Color in preservative: Above black, banded with cream; the first

Fig. 19. Neopareas bicolor Giinther. KUMNH No. 31920, Turrialba, Costa

Rica; actual total length, 334 mm.

Additioxs to Fauna of Costa Rica 571

band on back of head about four scale-rows wide, dorsally widening
laterally to include the temporals, posterior supra- and infralabials,
the posterior azygos shield on chin and the following five ventrals;
a discrete black spot on a somewhat enlarged median postparietal
scale and a few very small dots of black on upper temporals; cream
bands on head and body 16, dorsally covering approximately four
transverse scale-rows, widening on the sides and covering five to
six ventrals; the intervening black color covers ten scale-rows ante-
riorly then eight or nine in middle region and reducing to seven
posteriorly on body; all becoming narrower on sides and venter;
nine cream bands on tail, all, except first cream band, have series of
black dorsal flecks.

Measurements in mm.: Total length, 334; snout to vent, 227; tail,
107; width of head, 7; length of head, 11.

Dentition: The dentition of this species is remarkable. The high
anterior, dentary teeth, surrounded by gums, form two high longi-
tudinal ridges in the anterior part of the floor of the mouth that fit
into deep depressions in the anterior part of the palate; the lower
jaws lie in the middle of the floor of the mouth separated by a dis-
tance of only 1.5 millimeters for much of their length. The sides of
the mouth-floor are muscular. The maxillaries lie horizontal, the an-
terior teeth somewhat erect, the posterior curving, smaller than
the teeth on dentaries. Maxillary teeth, 21, decreasing in size pos-
teriorly; dentary teeth, 22, decreasing in size posteriorly; palatine
and pterygoid teeth present.

Remarks: Giinther, who described the genus and species had only
a single specimen. The general characteristics of the teeth were not
mentioned except to state that they were small and ungrooved.
From data given by Giinther and Boulenger the type specimen
differs from my specimen as follows: twenty more scales in the
total ventral-subcaudal count (324-304); two more encircling white
rings (27-25); internasals larger, two thirds length of prefrontals
(half length); frontal as long as broad (longer than broad); third
azygos scale followed by a pair (fourth azygos scale undivided);
cream bands immaculate (bands bearing numerous dorsal black
flecks ) .

Since I did not see the specimen before preservation, it is possible
that the light bands were originally pink or red. The collector "be-
lieved that such was the case, but was uncertain."

The amount of variation to be expected is unknown. A series
may show constancy in the variations mentioned above.

572 The University Science Bulletin

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574 The University Science Bulletin

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D

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