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O •" fw I

HARVARD UNIVERSITY

LIBRARY

OF THE

Museum of Comparative Zoology

UNIVERSITY OF KANSAS

SCIENCE BULLETIN

mi. CCMP. ZOQL
LSSRARY

SEP 2 4 1953

UNIVERSITY OF KANSAS PUBLICATIONS

University of Kansas Science Bulletin - Vol. XXXV - Part II

September 10, 1953
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 600 to 1,000 pages of reading
matter, with necessary illustrations. Exchanges with other institu-
tions and learned societies everywhere are solicited. All exchanges
should be addressed to

The University of Kansas Science Bulletin,
Library of the University of Kansas,

Lawrence, Kan.

Editor Edward H. Taylor

Editorial Board

Paul G. Roofe, Chairman
Frank E. Hoecker

H. B. HUNGERFORD

Arthur J. Mix

PUBLICATION DATES

The actual date of publication ( i. e., mailing date ) of many of the
volumes of the University of Kansas Science Bulletin differs so
markedly from the dates bourne on the covers of the publication or
on the covers of the separata that it seems wise to offer a corrected
list showing the mailing date. 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— October 1, 1932.
Vol. XXI— November 27, 1934.
Vol. XXII— November 15, 1935.
Vol. XXIII— August 15, 1936.
Vol. XXIV— February 16, 1938.
Vol. XXV— July 10, 1939.
Vol. XXVI— November 27, 1940.
Vol. XXVII, Pt. I— Dec. 30, 1941.
Vol. XXVIII, Pt. I— May 15, 1942;
Vol. XXVIII, Pt. II— Nov. 12, 1942.
Vol. XXIX, Pt. I— July 15, 1943;

Vol. XXIX, Pt. II
Vol. XXX, Pt. I-
Vol. XXX, Pt. II-
Vol. XXXI, Pt. I-
Vol. XXXI, Pt. II
Vol. XXXII— Nov.
Vol. XXXIII, Pt. I—
Vol. XXXIII, Pt. II
Vol. XXXIV, Pt. I-
Vol. XXXIV, Pt. II
Vol. XXXV, Pt. I-

— Oct. 15, 1943.
-June 12, 1944.
-June 15, 1945.
•May 1, 1946.
-Nov. 1, 1947.
25, 1948.
April 20, 1949.
-Mar. 20, 1950.
Oct. 1, 1951.
-Feb. 15, 1952.
July 1, 1952

UNIVERSITY OF KANSAS

SCIENCE BULLETIN

DEVOTED TO

THE PUBLICATION OF THE RESULTS OF

RESEARCH BY MEMBERS OF THE

UNIVERSITY OF KANSAS

Volume XXXV, Part II

University of Kansas Purlications

Lawrence, Septemrer 10, 1953

PRINTED BY

FERD VOILAND. JR.. STATE PRINTER

TOPEKA. KANSAS

1953

24-6490

S- 4 U

Contents of Volume XXXV, Part II

No. PAGE

6. The Effect of Radiation from Intraperitoneally Injected

Radium Chloride upon the Hemopoietic Activity of the
Bone Marrow in Albino Rats. Plates LXVIII to LXXII.

Paul G. Roofe and Hal G. Bingham, 945

7. Bees of the Genus Perdita in the Collection of the Uni-

versity of Kansas ( Hymenoptera, Apoidea).

P. H. Timherlake, 961

8. Comparative Morphological and Systematic Studies of Bee

Larvae with a Key to the Families of Hymenopterous
Larvae. Text figs. 1 to 287 Charles D. Michener, 987

9. A Revision of the North American Species of Typhlocyba

and its Allies ( Homoptera, Cicadellidae ) . Plates
LXXIII to XCII Paul G. Christian, 1103

10. The Ambrysus of Mexico (Hemiptera, Naucoridae).

Plates XCIII to C Ira La Rivers, 1279

11. A Revision of the Genus Buenoa (Hemiptera, Notonec-

tidae). Plates CI to CXVII Fred S. Truxal, 1351

12. A Review of the Lizards of Ceylon . Edward H. Taylor, 1525

13. Fourth Contribution to the Herpetology of San Luis

Potosi. Plates CXVIII-CXIX, map Edward H. Taylor, 1587

14. Report on a Collection of Ceylonese Serpents.

Edward H. Taylor, 1615

15. Frogs of the Family Centrolenidae from Brazil.

Edward H. Taylor and Doris M. Cochran, 1625

^sTcoMpTzoon f :

LIBRARY

\*W 2 4 1953

THE UNIVERSITY OP KANSAS

SCIENCE BULLETIN

Vol. XXXV, Pt. II] September 10, 1953 [No. 6

The Effect of Radiation from Intraperitoneally Injected

Radium Chloride Upon the Hemopoietic Activity

of the Bone Marrow in Albino Rats *

Paul G. Roofe and Hal G. Bingham f
Department of Anatomy, University of Kansas, Lawrence

INTRODUCTION

In the major portion of experiments in the past, radiation of the
bone marrow has been accomplished by an external source, X-rays
or gamma-rays in most cases. The literature relating to the effect
of roentgen rays on the blood-forming organs is extensive and con-
tradictory. The lack of unanimity among various reports may be
attributed in part to differences in the experimental conditions in
regard to radiation technique, the experimental animals used, and
the time interval between radiation and examination of tissues.

Previous investigations of the effect of ionizing radiation on bone
marrow, using radium as the internal source of radiation, were con-
ducted over varying periods of time from a minimum of a few
months to a maximum of several years duration. The results ob-
tained after a few months of low dosage internal radiation are gen-
eral hyperplasia of the erythrocytic tissue (Sabin, Doan, and Fork-
ner, '32). Continued exposure over a longer period of time
produces hyperplasia of granulocytic tissue and hypoplasia of the
erythrocytic tissue ( Rosenthal and Grace, '36 ) . After an even longer
time, however, radium causes the marrow to lose its ability to pro-
duce either erythrogenic or granulogenic tissue (Martland, '31).

The present experiment is an investigation of the effects of radium
chloride on the hemopoietic activity of the cells of the bone marrow
after exposure of short duration. In addition the effects of varying
dosage are reported.

° This investigation was carried out under contract with the Office of the Surgeon Gen-
eral, U. S. Army, Research Contract, No. W49-007-MD 466.

f Paul G. Roofe, Hal G. Bingham, Department of Anatomy, University of Kansas, Law-
rence, Kansas.

(945)

946 The University Science Bulletin

Following intraperitoneal injection of radium, elimination is rapid
during the first three days. The rate decreases and becomes es-
sentially zero after fifteen to twenty days. Since the alpha particle
penetrates about 25[x in tissue, the maximum damage occurs during
the first two or three days when the radium is still found in the
circulating blood. Approximately thirty percent of the injected
radium chloride is permanently retained in the bone matrix proper
(Hoecker and Roofe, '49). When radioactive material is fixed in
the bone, it is eliminated very slowly and the major portion remains
as a source of alpha particle radiation.

METHODS AND MATERIALS

Thirty-two Sprague-Dawley male albino rats were used. All
animals used were young adults and approximately the same weight.
Seven of these were untreated controls to establish the normal bone
marrow picture. Twenty-five animals were divided into five groups
of five. The first group received lOjxg of intraperitoneally injected
RaCl2 and each successive group received an increase of l(fy.g, so
that groups receiving 10, 20, 30, 40, and 50j;.g of RaCL were ob-
tained. An animal from each group was sacrificed on the third day
after injection; and one from each group was sacrificed at three-day
intervals thereafter.

After the rats were anesthetized with sodium nembutal the femurs
were quickly removed. The method of Mayer and Ruzicka ('45)
was found to yield well-preserved pencils of marrow without pre-
vious decalcification. The marrow was fixed in Zenker-formol, im-
bedded in paraffin and sectioned serially at 8 micra. Sections were
stained with hematoxylin-eosin-azuren •

The bone marrow cell count was made with the aid of a camera
lucida which projected a calibrated area onto especially printed
sheets for recording the various cell types. The results of these
counts were converted into number of cells per mm3 to make com-
parison possible with other hemopoietic organs and peripheral
blood. Twelve sections were chosen from each tissue so that the
entire bone marrow was adequately sampled. Six fields or foci of
cell concentration were counted on each of these bone marrow sec-
tions. An eosinophil field, a neutrophil field, an erythroid field,
and two myeloiderythroid fields 50 X 50 micra were counted under
oil immersion. A megakaryocyte field 500 X 500 micra was counted
under low power.

In most cases the Maximow and Bloom ('42) terminology and
method of cell identification were used. The various cell types in
the neutrophil and eosinophil series were readily identified by the

Effect of Radium on Bone Marrow 947

presence of their characteristic cytoplasmic granules. In addition
the myeloid and erythroid series were not difficult to distinguish.
However, the maturing cell forms within these series were difficult
to distinguish from each other. To avoid this problem the pro-
myelocyte and myelocyte forms of the myeloid series were grouped
together as immature cells and the metamyelocyte and adult cells
were counted as separate cell types. In the erythroid series two
cell forms were distinguished, the erythroblast and the normoblast.
The term erythroblast includes all the immature erythroid elements
up to the cell form containing hemoglobin in its cytoplasm. The
term normoblast includes all those cells with hemoglobin in their
cytoplasm and maturing cell forms up to the adult red blood cell.

The other bone marrow cells were also identified and named by
the Maximow and Bloom ('42) terminology. However, plasma
cells, lymphocytes and basophils were found with such low fre-
quency that they proved to be of little consequence in this study
and were not included in the results.

In order to determine what the effects of the radiation were on
the peripheral blood of the experimental animals, counts were made
on the peripheral blood by other workers in this laboratory. The
blood for the peripheral count was taken from the tail of the rats
just before sacrifice and standard methods of counting were carried
out.

RESULTS

Table I shows the mean and range of each hemopoietic cell type
in mm3 found in both the non-injected control animals and the 25
experimental. The 25 experimental animals are represented in the
table as five groups of five animals. The mean and the range of
each group pertain to those five animals sacrificed on the same day.
Each animal in the group of five received a different dose of RaCl2
ranging from 10[xg to 50>g.

Statistical analysis of the material in Table I is not included in
this paper. Superimposed on the variation due to the smallness of
the sample and due to the difficulty in identifying the cells, was the
great variability of the individual animals in response to the treat-
ment. As a consequence, the range of the cell counts is great in
many cases.

The data were plotted in the following manner: In each graph
the ordinate represents the number of cells and the absissa repre-
sents time. The points on the vertical lines represent counts for
individual animals with different doses killed on the same day. The
curve represents the mean cell count of each day.

948 The University Science Bulletin

A. Granulocyte Series

As can be seen in Table I, the counts of the immature neutrophils
for the experimental animals fall within the normal range. Because
of the extreme variation in both control and experimental animals,
it is not possible to tell if any change has occurred. On the other
hand, the metamyelocyte and adult cell forms exhibit an incidence
above normal with increase in time ( Figures 1 and 2 ) .

The trend of the count of the three cell types in the eosinophil
series is to increase above normal as is shown in figures 3, 4, and 5.
One of the 12 day animals gave exceptionally high counts for each
of the cell types. These three values are indicated on the graphs,
and were also included in the calculations of the means. All eosino-
phil cell types demonstrate a trend similar to that of the develop-
ing neutrophil cells in that they generally increase in number with
time. Furthermore the adult forms of both series appear to reach
a maximum at about the 12th day.

B. Erythroid Series

The cell counts for both erythroblasts and normoblasts drop be-
low normal initially, then rise back to normal by the 14th day ( Fig-
ures 6 and 7).

C. Other Bone Marrow Cells

Reticular cells. The cell count shows an initial rise, with a return
to normal by the 9th to 12th days (Fig. 10). On the 14th day the
count is again above normal.

Hemocytoblasts. It can be seen in figure 11, that the cell count
is generally above normal and tends to increase with longer time.

Megakaryocytes. The cell number decreases reaching minimum
values on the 9th and 12th days but returns to normal by the 14th
day (Fig. 12).

D. Peripheral Blood

Megakaryocytes are compared with peripheral platelets in figure
12. The platelets show the same trend as the megakaryocytes but
demonstrate the initial decrease and subsequent increase in count
later than the megakaryocytes. It appears then that the platelet
curve shows about a 3 day time lag when compared with the mega-
karyocyte curve. This observation has been published in a paper
by Roofe, Bingham, and Comer ('51) and is discussed more ex-
tensively there.

The peripheral polymorphonuclear leukocyte count is consider-
ably decreased from normal and remains that way for the duration
of the experiment.

Effect of Radium on Bone Marrow 949

The peripheral red blood cell count did not show much variation

from normal.

E. Dosage Effects

In general the effects with increasing dose were comparable to
the changes obtained with increase in time. However, it was evi-
dent that in most cases time was the important factor in determining
the response to the injected radium.

The one apparent exception to this was the case of the erythro-
blasts and normoblasts. As figures 8 and 9 show, the number of
erythroblasts was lowest with the smaller dose, while with increas-
ing dose the counts tended to be more in the normal range. How-
ever, the normoblast count decreases with increase in dose. (The
normoblast curve is included to show the more expected trend of
cell count with increased dose. )

DISCUSSION

In the neutrophil series an increase in the metamyelocyte and
adult cell forms was observed. This observation suggests that the
rate of maturation of the earlier forms to the more mature forms
may be accelerated. Lingley, Gall, and Hilcken ('40) also ob-
served a similar trend in their studies with roentgen radiation.
However, Martland ( '31 ) and Rosenthal and Grace ( '36 ) , in experi-
ments of longer duration than those discussed here, found that with
radiation from radium the immature cells constituted the majority
of the neutrophils. As has already been shown it is impossible to
determine what, if any, trend has been established in the immature
cells in this experiment.

In the case of the eosinophils, an accelerated maturation of im-
mature cells forms similar to that in the neutrophil series is sug-
gested by the results. Also an increased proliferation of the im-
mature cell forms probably occurred, since all cell forms of this
series increased to above normal incidence with increase in time
and in dosage. Isaacs ('32) also found an increase in the number
of eosinophils following roentgen radiation, as did Rosenthal and
Grace ('36) with radium. Martland ('31) found that the only cell
of the granulocytic series which proceeded to maturation was the
eosinophilic myelocytes. The observed increase in the number of
immature cells in the eosinophil series is in disagreement with the
observations of Osgood ('42) and Osgood, Albersold, Erf, and
Packham ( '42 ) , who found that roentgen radiation inhibited mitotic
division of all progranulocytes, which would be expected to de-
crease the production of all cell forms. However in this case larger
dosages and longer duration were involved.

950 The University Science Bulletin

The peripheral polymorphonuclear leucocyte count dropped con-
siderably below normal and remained there throughout the experi-
ment. This suggests that some mechanism associated with irradia-
tion causes the mature granulocytes to be retained in the bone
marrow. Dunlop ( '42 ) , who worked with roentgen rays, also found
an increased bone marrow neutrophil count with a concomittant
decreased peripheral polymorphonuclear leucocyte count. Mart-
land ('31) and Jacobson, Marks, and Lorenz ('49), using radium,
observed a reduction in leucocytes in the peripheral blood and
Martland ('31) also reported a decrease in the bone marrow neu-
trophil count.

Both the erythroblast and normoblast show an initial decrease
with a return to the normal range at a later time. Rosenthal and
Grace ( '36 ) found that continued action of radium inhibited matu-
ration of the erythroblast as evidenced by a slight increase in num-
ber of erythroblasts despite a decided decrease in the total marrow
erythrocytic elements. In the X-ray studies of Lingley, Gall, and
Hilcken ('40), the lowered number of normoblasts was explained
as due to an accelerated maturation of this cell into adult red blood
cells. The lack of appreciable variation of the peripheral red blood
cell count from normal is in agreement with the radon studies of
Nemenow and Gurewitsch ( '34 ) . On the other hand Dunlop ( '42 )
using roentgen rays and Jacobson, Marks and Lorenz ('49) using
RaCl2 found a peripheral anemic condition after a longer period of
irradiation.

In regard to dosage the erythroid series showed a different kind
of response than did the other bone marrow cells. Although the
normoblast cell count is lower in animals that received greater doses,
this was not the case with the erythroblasts. The greatest decrease
in the number of erythroblasts was found in the animals receiving
the lowest dose of radium. Interpretation of this phenomen is be-
yond the scope of the present experiment.

In the literature the myeloid-erythroid ratio is frequently used as
an index of change occurring in the cells of the bone marrow. The
value for normal animals found in this experiment was 0.739, which
falls in the normal range reported in the literature, 0.500 (Brown,
'51)-1.940 (Endicott and Ott, '45). The myeloid-erythroid ratio for
the experimental animals was increased to 0.910 on the third day,
and to 1.020 on the fourteenth day. Cellular processes of prolifera-
tion and maturation were increased more in the myeloid tissue, as
has already been shown, leading to a cell count above normal for
both the eosinophil and neutrophil series. In the erythroid tissue,

Effect of Radium on Bone Marrow 951

however, there was an initial decrease in both cell types with a
return to normal with longer time. This observation is not in agree-
ment either with Bauer ('40) or Chien-Liang and Ma ('40) who
both worked with roentgen radiation. They found a hyperplasia of
the erythroid cells and a hypoplasia of the myeloid cells. However,
in both of these cases a larger dose of radiation was employed.

Both the hemocytoblast and reticular cell counts were increased,
which is in agreement with the findings of Lingley, Gall and Hilcken
( '40 ) . They found that primitive cells in general proliferated more
rapidly in animals subjected to roentgen radiation. Warren, Mac-
Millian, and Dixon ('50) using P32 as the internal source of radia-
tion found that the stem cells were not injured at any time during
their experiment.

The number of megakaryocytes was found to be below normal
and remained that way throughout the duration of the experiment.
Dunlop ('42) working with X-rays and Warren, MacMillian and
Dixon ('50) using P32 also found that megakaryocytes were reduced
in number. The relationship between the megakaryocyte and peri-
pheral platelet curves is in agreement with the theory that the mega-
karyocytes fragment to become platelets. Three days pass before
the peripheral platelets show any change in number resulting from
the initial megakaryocyte decrease, indicating that they possibly
survive this length of time in the peripheral blood after entering it.
The newly produced megakaryocytes apparently do not fragment
into platelets for approximately three days after they are produced
because the peripheral platelet curve does not show a similar in-
crease in number until this period of time has passed. Jacob-
son, Marks, and Lorenz ('49) also noted a reduction in the peri-
pheral platelet count after injecting parenterally 0.94tj.g of radium
chloride, but did not correlate this with the megakaryocyte changes
in the bone marrow.

The sum total of the various cell responses to the initial irradia-
tion results in a decreased total cell count for the bone marrow and
a normal cell count for the peripheral blood. Some bone marrow
cells increase while others decrease in number and this is probably
due to a change in the rate of proliferation and/or maturation. With
increased time the bone marrow cells appear to proliferate and ma-
ture rapidly but not to pass into the peripheral circulation, which
tends to increase the marrow count and to slightly decrease the
peripheral blood count. In contrast to these results, Lingley, Gall,
and Hilcken ('40), using roentgen irradiation, observed a secondary
or regenerative return of the bone marrow to normal cellular con-

952 The University Science Bulletin

tent between the second to fourth weeks, while Warren, MacMillian
and Dixon ('50) found that maximum damage occurred at about
10 days with return of myelopoiesis 10 to 15 days after injection
of P32.

SUMMARY

Radium chloride emitting chiefly alpha particles was injected
intraperitoneally into the albino rats. The RaCL is eliminated rap-
idly from the animals during the first three days after injection so
it is mainly found in the circulatory system during that period.
Eventually, approximately 30 percent of the injected RaCL becomes
deposited in the bone tissue. Both the dosage and, more important,
the duration of exposure determined the kind of effect obtained. In
general, however, dosage effects and duration effects caused the
same type of cell responses. These effects are a change in the rate
of proliferation of the immature cells and a change in maturation
time of the more mature cells. Considering the effects on the dif-
ferent marrow cells separately:

1. Neutrophils and eosinophils. The more mature neutrophils
show an acceleration of maturation, but there is neither a decrease
nor an increase in the rate of proliferation of the immature forms.
The eosinophils follow this same tendency except that their im-
mature cells increase in the rate of proliferation.

2. Erythroid cells. Both the normoblast and erythroblast num-
ber decreases initially with a return to the normal range at a later
time. However, increase in dosage caused the normoblast to de-
crease, while the erythroblast decreased initially, it later returned
to normal.

3. Myeloid-erythroid ratio. This ratio is increased because the
myeloid cells increase above normal and the erythroid cells decrease
below normal.

4. Stem cells. The reticular cells and hemocytoblasts show an
increased proliferation.

5. Megakaryocytes and platelets. The two curves of these cells
are very similar in shape but the platelet curve shows a time lag of
about 3 days to the changes occurring in the megakaryocyte curve.
This supports the theory that platelets are formed from fragmenting
megakaryocytes .

6. Granulocytes and peripheral polymorphonuclear leucocytes,
also erythroid cells and peripheral red blood cells. The peripheral
blood picture does not indicate adequately the processes occurring
in the marrow. The decreased peripheral polymorphonuclear leuco-

Effect of Radium on Bone Marrow 953

cyte count gives no indication of the increased granulocyte produc-
tion in the marrow, nor does the apparently normal peripheral red
blood cell count reflect the initial decrease with later return to nor-
mal of the erythroid production in the bone marrow.

7. Total bone marrow and total peripheral blood count. Initially
the sum total of the various cell responses to the irradiation causes
a total cell decrease in the bone marrow but the total peripheral
blood count remains normal. With longer exposure to irradiation
the peripheral blood count tends to decrease but the bone marrow
count increases. This suggests a greater proliferation and matura-
tion of the marrow cells and a mechanism which prevents the pas-
sage of these marrow cells into the periphery.

954

The University Science Bulletin

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Effect of Radium on Bone Marrow 955

BIBLIOGRAPHY

Bauer, von Robert. 1940. Effects of variously administered roentgen rays

on marrow and its cell elements. Strahlentherapie, 67: 424-501.
Brown, J. Wm. 1951. The effect of protein deficiency on the hemopoietic

organs and the blood of the albino rat. Unpublished manuscript.
Chien-Liang, Hsu, and W. C. Ma. 1940. Direct and indirect effects of roentgen

radiation on the blood-forming organs of rats. Am. J. Cancer 39: 319-333.
Dunlop, C. E. 1942. Effects of radiation on the blood and the hemopoietic

tissues. Arch, of Path., 34: 592.
Endicott, K. M., and M. Ott. 1945. The normal myelogram in albino rats.

Anat. Rec, 92: 61-69.
Hoecker, F. E., and P. G. Roofe. 1949. Structural differences in bone matrix

associated with metabolized radium. Radiology, 52: 856-864.
Isaacs, R. 1932. Maturing effect of roentgen rays on blood-forming cells.

Arch. Int. Med., 50: 836-842.
Jacobson, L. O., E. K. Marks, and E. Lorenz. 1949. The hematological

effects of ionizing radiations. Radiology, 52: 371-395.
Lingley, Gall, and Hilcken. 1940. Comparative experimental studies of 200

kilovolt and 1000 kilovolt roentgen rays; biologic effects on bone marrow of

albino rats. Am. J. Rath., 16: 845-854.
Martland, H. S. 1931. The occurrence of malignancy in radioactive persons

Am. J. Cancer, 15: 2435.
Maximow, A. A., and W. Bloom. 1942. Testbook of Histology. W. B.

Saunders Co., Philadelphia and London, 4th Ed.
Mayer, E., and A. Z. Ruzicka. 1945. A method for studying numerical and

topographic problems in the whole femoral marrow of rats and guinea pigs,

with the use of uncalcified sections. Anat. Rec, 93: 213-231.
Nemenow, N., and Gurewitsch. 1934. Effects of intravenous injection of

radon on blood. Strahlentherapie, 50: 693-704.
Osgood, E. E. 1942. Investigation as to whether or not the action of roentgen

rays are direct or indirect on human marrow culture. Am. J. Roentgenol.,

35: 786-789.
Osgood, E., P. Albersold, L. A. Erf, and E. Packham. 1942. Effects of

million volt roentgen rays, 200 kilovolt roentgen rays, radioactive phos-
phorus, and neutron rays by marrow culture technique. Am. J. M. Sc,

204: 372-381.
Roofe, P. G, H. G. Bingham, and R. Comer. 1951. Effect of radiation

from intraperitoneally injected radium chloride upon megakaryocyte and

blood platelet production in albino rats. Kans. Acad, of Sci., 54: 391-394.
Rosenthal, M., and E. J. Grace. 1936. Bone marrow and lymph node

changes in rabbits produced by oral administration of radium sulfate. Am.

J. M. Sc, 191: 607.
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 injection of radium chloride and mesothorium in rabbits. J. Ex.

Med., 56: 267-289.
Warren, S., J. C. MacMdllian, and F. J. Dixon. 1950. Effects of internal

irradiation of mice with P32; spleen, lymph nodes, thymus, bone marrow.

Radiology, 55: 375-389.

956

The University Science Bulletin

PLATE LXVIII

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THE UNIVERSITY OF KANSAS

SGIENGE BULLETIN

Vol. XXXV, Pt. II] September 10, 1953 [No. 7

Bees of the* Genus Perdita in the Collection of the Uni-
versity of Kansas ( Hymenoptera, Apoidea)*

P. H. TlMBERLAKE,

University of California Citrus Experiment Station
Riverside, California

CONTENTS

PAGE

Subgenus Xerophasma Coekerell 962

Perdita bequaertiana Cockerel! 962

Subgenus Cockerellula Strand 963

Perdita bidenticauda sp. nov 965

Male 965

Female 966

Perdita laticauda sp. nov 967

Male 967

Female 967

Perdita lobata sp. nov 967

Male 967

Subgenus Macroteropsis Ashmead 968

Perdita latior Coekerell 968

Perdita arcuata dinognatha Coekerell 968

Unnamed group 969

Perdita trifasciata sp. nov 969

Male 969

Unnamed group 970

Perdita ainsliei Crawford 970

Perdita peculiaris sp. nov 970

Male 970

Subgenus Cockerellia Ashmead 971

Key to the species of Cockerellia, females 971

Perdita utahensis Coekerell 974

Male 974

Perdita baileyae Coekerell 975

° Paper No. 758, University of California Citrus Experiment Station, Riverside, Cali-
fornia.

(961)

962 The University Science Bulletin

PAGE

Perdita perpulchra flavidior subsp. nov 975

Female 975

Perdita bequaerti indianensis Cockerell 976

Perdita verbesinae Cockerell 976

Perdita fracticincta sp. nov 976

Female 976

Perdita lacteipennis lacteipennis Swenk and Cockerell 977

Perdita lacteipennis pallidipennis Graenicher 978

Perdita tricincta sp. nov 978

Female 978

Male * 979

Perdita albipennis Cresson 980

Perdita scopata sp. nov 980

Female 981

Male 981

Perdita lepachidis lepachidis Cockerell 982

Female 982

Perdita lepachidis levifrons subsp. nov 983

Female 983

Male 983

Perdita coreopsidis coreopsidis Cockerell 983

Perdita coreopsidis kansensis subsp. nov 984

Female 984

Male 984

Perdita coreopsidis obscurior subsp. nov 984

Female 985

Male 985

Literature cited 985

Abstract: This article is a revisionary study of the bees of the genus
Perdita Smith, subgenera Xerophasma, Cockerellula, Macroteropsis, Cock-
erellia, and two other un-named groups, in the collection of the University of
Kansas. Twenty-one species are treated; of these, eight species are new.
Four new subspecies are also described.

This is a first report on a large collection of bees of the genus
Perdita submitted to the author in the fall of 1950 by Charles Dun-
can Michener, and collected mostly in the states of Kansas and
Texas. Types of new forms will be found in the collection of the
University of Kansas, at Lawrence, Kansas.

Subgenus XEROPHASMA Cockerell
Perdita bequaertiana Cockerell

Xerophasma bequaerti Cockerell, Amer. Mus. Novitates vol. 66, 1923, p. 2, 2

(not Perdita bequaerti Viereck, 1917).
Perdita bequaertiana Cockerell, in Muesebeck, Krombein, and Townes, U. S.

Dept. Agri. Monog. No. 2, 1951, p. 1087.

One male, 11 females, Big Springs, Howard Co., Texas, June 20,
1947 (R. H. Beamer).

Timberlake: Bees of the Genus Perdita 963

No flower record accompanies these specimens, but several of the
females bear the coarse cobwebby pollen of Oenothera.

This is a large pallid bee ( very large for the genus Perdita ) , and
in the female the ocellar region, usually including the face above
level of antennae and the vertex behind ocelli more faintly, may
be slightly infuscated. On the abdcmen a pale brownish, or dilute
fuscous, preapical crossband, not quite reaching lateral margins, is
usually present in tergites 1 to 5. The structural characters have
been described rather fully by Cockerell, but one feature not men-
tioned by him is the fact that the antennae are inserted unusually
high on the face or above the middle of the eyes, in which respect
members of Xerophasma differ from all other Perdita. The mouth
parts are moderately elongate. Maxillary palpi are long and six-
jointed, the first five joints subequal, the sixth somewhat shorter.
Labial palpi are about equal to the maxillary pair, the first joint
very long, the last three joints each about equaling the apical joint
of the maxillary pair.

The male resembles the female except in the primary sexual
characters. The only specimen at hand lacks the abdominal band-
ing, and the dark cloud on the head is restricted to the area between
the ocelli. Aedeagus composed of a rather large quadrate capsule,
about one half longer than wide and somewhat wider than high.
Stipites produced ventrally on each side in a slender lobe, which is
abruptly bent inward and upward at the extreme tip. Sagittae
moderately stout, tapering toward apex, which is narrow and thinly
compressed dorsoventrally. Volsellae very large, the inferior pair
reaching middle of sagittae and provided with numerous minute
black tubercles along the dorsal margin; dorsal volsellae reaching
middle of inferior pair. Length of male, about 9 mm.; anterior
wing, 7 mm.

This species was described from a female collected about 10 o. m.,
July 9, 1917, at Fabens, El Paso Co., Texas.

Subgenus COCKERELLULA Strand

In 1922 (Amer. Mus. Novitates, vol. 47, pp. 1-4) Cockerell de-
scribed Perdita opuntiae and erected the subgenus Ltitziella for its
reception. Because Ltitziella Cockerell was preoccupied by Lutzi-
ella Enderlein, Strand, in 1932 (Folia Zool. Hydrobiol. Riga, vol. 4,
p. 196), proposed to rename it Cocker ellula.

The group Cockerelhda is similar to Macrotera Smith, but differs
in having the tongue short and the abdomen of the male modified in

964 The University Science Bulletin

various ways, either at the apex or on the ventral surface. It is quite
possible that discovery of additional species may reveal that these
characters are gradational and only of specific importance, since
members of both groups seem to be largely, although not com-
pletely, dependent on the pollen of various species of Opuntia.

The several species belonging in Cockerelhda may be distin-
guished as follows:

1. Males 2

Females 5

2. Basal ventrite without a median fold 3

Basal ventrite strongly puckered out in middle of apical margin

to form a strong fold, the following ventral segments unmodi-
fied 4

3. Second ventrite with a broad thickening on middle of apical

margin, this thickening strongly dentate at outer ends; apical
tergite very broad, with a slender, almost fingerlike lobe at
the apical corners, and evenly truncate between the lobes;
head and thorax black or blackish brown, with pale-yellow
face marks; abdomen ferruginous; length, about 5.5 mm.;
Colorado (White Rocks near Boulder) and South Dakota

( Pine Ridge ) opuntiae Cockerell

Second ventrite only slightly thickened at middle of apex, the
third ventrite with a strong thickening which is unarmed at
outer ends and presents a transverse disk as viewed from be-
hind; apical tergite quadrate, with apical corners slightly
dentate, the disk a little concave and produced medially in
a rounded angle a little beyond and below the dentiform
lateral angles; head and thorax dark blue-green, the face
below antennae pale yellow; abdomen ferruginous; length
about 4-4.5 mm.; Texas (Big Bend Park) . . bidenticauda sp. nov.

4. Tergite 7 extremely broad, transversely quadrate, not at all nar-

rowed to apex, which is truncate and at most with a small
median notch; head and thorax dark blue-green, the face be-
low antennae yellowish white, abdomen ferruginous; length,
about 4 mm.; Texas (15 miles southeast of Dryden),

laticauda sp. nov.
Tergite 7 similar to that of laticauda but more convex, distinctly
grooved medially and emarginate at apex, thus forming a
broad, slightly oblique lobe on each side; head and prothorax
brown, the face below antennae pale yellow; thorax otherwise
dark blue-green, more or less washed with brown on pleura,
sternum, and apical truncation of propodeum; abdomen fer-
ruginous; length, 4-4.5 mm.; Texas ( 15 miles northwest of
Mission) lobata sp. nov.

5. Head and thorax dark green; smaller species 6

Head and thorax black, the abdomen ferruginous; mesoscutum

shining, delicately tessellate and with minute close punctures;
length, about 6 mm opuntiae Cockerell

Timberlake: Bees of the Genus Perdita 965

6. Mesoscutum tessellate but shining, finely punctured, the punc-
tures about two to three puncture widths apart; abdomen

ferruginous; length, about 4-4.5 mm bidenticauda sp. nov.

Similar to preceding and without good distinguishing characters
in the female sex laticauda sp. nov.

Perdita bidenticauda sp. nov.

Allied to P. opuntiae Cockerell but distinguished by its smaller
size and dark-green color, and by the very small lateroapical teeth
of the apical tergite of male.

Male. — Head and thorax dark green, slightly bluer on meta-
notum and propodeum and gradually changing to brown on under
surface of head and at anterior end of cheeks. Narrow inferior
orbits near base of mandibles, mandibles, except the rufous tips,
and entire face below level of antennae, pale yellow, with the
broad upper margin of lateral marks oblique and reaching upper
level of antennal sockets and intruding slightly between the fovea
and eye margin on the orbits. Tubercles and anterior margin of
pronotum brownish. Abdomen ferruginous red, with only the
lateral foveae of second tergite black. Legs black, more brownish
at base, the front tarsi and the front tibae, except a brownish line
behind, dull yellow. Antennae almost uniformly pale brown.
Tegulae pale testaceous yellow. Wings nearly clear hyaline, the
nervures brown, the subcosta more fuscous.

Head but little enlarged, broader than long, with the inner orbits
slightly diverging below. Mandibles nearly straight, robust, and
tapering to the blunt apex which has a small inner tooth. Labrum
twice as broad as long. Subantennal plates very small, the lateral
plates broad and becoming much broader at anterior ends. Clypeus
short and transverse. Frontal foveae about three times as long as
wide and well impressed. Cheeks simple and moderately wide.
Abdomen moderately broad and depressed. Apex of last tergite
rather broad and truncate, the apical corners a little dentiform, the
disk slightly concave and medially produced a little beyond and
well below level of the dentiform angles; as seen from behind and
beneath, the lower apical margin is carinate and forms a broad
ogival curve. First segment of venter normal, the second segment
slightly, and the third strongly, swollen in middle of apical margin;
this swelling elongate-oval as seen from behind, declivous, and with
outer ends acute but not at all prominent or dentiform. Marginal cell
about one fourth smaller than first submarginal cell, broad and ob-
lique at apex, with the apendiculation of radius distinct and elongate.

966 The University Science Bulletin

First recurrent nervure received near apex of first submarginal cell.
Stigma small and very narrow. Head and thorax minutely tessellate
and dullish, the disk of mesoscutum more shining than other parts,
the mesopleura duller, with the sculpture more lineolate and the
basal middle of propodeum comparatively coarsely tessellate.
Punctures of frons and mesoscutum minute and close, those of
vertex sparser, and those of disk of clypeus coarser and shallow.
Mesopleura impunctate except toward the sternum. Abdomen
moderately shining. Pubescence whitish, short and sparse, that of
abdomen extremely fine and subappressed, but a row of coarse, stiff
hairs present across disk of tergite 6, and similar hairs on outer side
of hind tibiae. Length, 3.45-4 mm.; anterior wing, 3 mm.

Female. — Dark green, or blue-green, the metanotum and pro-
podeum more bluish, the abdomen ferruginous red. Clypeus,
supraclypeal area extending in a triangle well above antennae, and
narrow orbital margins of face, black. Labrum brownish. Mandi-
bles testaceous yellow, with rufous tips. Antennae rather dark
brown, the flagellum dull yellowish beneath. Legs dark brown,
the tarsi paler, the front knees with a small yellowish spot, the spurs
reddish. Tegulae brown at base and pale testaceous on outer
margin. Wings as in male.

Head as wide as long, the clypeus much larger and more pro-
duced than in male. Inner orbits of eyes parallel. Mandibles
simple at apex. Frontal foveae about five times longer than wide
and deeply impressed. Pygidium of abdomen acute at apex. Hind
knee plates distinct. Spurs of middle and hind tibiae slightly
hooked at apex and minutely serrate beneath. Claws with a small
inner tooth just beyond the middle. Sculpture as in male except
as follows: labrum, clypeus, and supraclypeal area with coarse,
shallow punctures; frons and vertex with well-separated punctures
that are much sparser than those of mesoscutum; abdomen duller,
more strongly lineolate than in male, and impunctate. Pubescence as
in male, the apical fimbria of abdomen tinged with fuscous. Scopal
hairs of hind tibiae sparse and moderately long; those of dorsal
margin with minute, close-set, short branches. In comparison with
opuntiae, the scopal hairs are definitely shorter and sparser. Length
about 3.45-4.5 mm.; anterior wing, 3-3.2 mm.

Holotype male, allotype, and 6 males, 2 females (paratypes) at
flowers of Opuntia, Cooper's store, Big Bend Park, Texas, April 11,
1947 (Michener and Beamer).

Two of the females bear the extremely coarse pollen from
Opuntia, and the third a much finer pollen from some other flower.

Timberlake: Bees of the Genus Perdita 967

Perdita laticauda sp. nov.

Allied to P. bidenticauda, but the male has distinctive differences
in the ventral segments and apical tergite of the abdomen. The
females of the two species are apparently indistinguishable.

Male. — Remarkably similar to the male of bidenticauda except
in regard to the special characters of abdomen. Color the same,
with the yellow mark on cheeks near base of mandibles either large
or evanescent. First ventrite of abdomen strongly folded in middle
of the apical margin, thus producing a large projecting pucker.
Following ventrites little modified, although the third has a trans-
verse preapical band of short dense hair, occupying the middle third
of the width. Apical tergite extremely large and broad, more than
twice as broad as long, and broadly truncate at apex, the trunca-
tion with a small median notch. Length, 3.5-4 mm.; anterior wing,
2.8-3 mm.

Female. — Like the female of bidenticauda. First tergite some-
times infuscated on basal half. Length, 3.45-4.5 mm.; anterior wing,
2.8-3 mm.

Holotype male and allotype, 15 miles southwest of Dryden, Ter-
rell County, Texas, at flowers of Gilia acerosa, April 13, 1949 ( Mich-
ener and Beamer). Also the following paratypes taken at the same
time and place: 10 males, 8 females on Gilia, and 2 males, 6 females
on flowers of Chamaesaracha conioides.

The females bear a moderately fine-grained yellow pollen, which
probably was derived from the Gilia flowers, as it seems to be
slightly coarser and less pale yellow than the pollen borne by other
species of Perdita that have been taken at flowers of Chamaesaracha.

Perdita lobata sp. nov.

This species is similar to P. laticauda but has the apical tergite
even broader, more convex from side to side, rather strongly grooved
medially, and broadly and shallowly emarginate in middle to form
two bread, truncate apical lobes.

Male. — Head enlarged, nearly as bulky as thorax, the temples
and cheeks broad. Inner orbits distinctly diverging anteriorly.
Mandibles with a weak inner tooth at apex. Frontal foveae well
impressed, about five times as long as wide. Abdomen depressed
and much broader than thorax. First ventrite with a strong fold in
middle of apical margin, the fold producing a puckerlike process.
A band of dense short hair on the third ventrite. Apical tergite as
described above. Structural characters, sculpture and pubescence
otherwise about as in laticauda and bidenticauda.

968 The University Science Bulletin

Thorax dark bluish green, a little suffused with brown, especially
below the wings and on the sides and apex of propodeum. Entire
prothorax and the head reddish brown, the frons with a slight green-
ish tinge. Mandibles, except piceous tips, labrum, face below level
of antennae, and mark at anterior end of cheeks, pale yellow. The
yellow of face gradually merges into the brown and extends slightly
above the level of antennae on each side. Abdomen ferruginous
red, nearlv the same color as the head. Foveae on lateral margins
of tergite 2 very short and black. Legs brown, the front tibiae on
anterior side and front tarsi dull yellow. Antennae nearly uniformly
pale brown. Tegulae pale testaceous. Wings hyaline, the nervures
brown, the subcosta darker brown. Length, 4-4.5 mm.; anterior
wing, 3 mm.

Two males (holotype and paratype), 15 miles northwest of Mis-
sion, Hidalgo County, Texas, on Opuntia, March 30, 1946 (C. D.
Michener ) .

Subgenus MACROTEROPSIS Ashmead
Perdita lotior Cockerell

Sixteen males, 5 females from the following localities in Arizona:
6 miles east of Douglas, Aug. 11, 1940; Portal, Aug. 12, 1940; Cave
Creek, Chiricahua Mts., Aug. 12, 1940 (Michener); and Rustler's
Park, Chiricahua Mts., July 5^ 1940 (D. E. Hardy).

This is a common species in New Mexico and Arizona at flowers
of Sphaeralcea.

One specimen of a new species of this group was also taken by
Michener near Portal, Arizona, at flowers of Sphaeralcea. This will
be described elsewhere.

Perdita arcuata dinognatha Cockerell

Many of both sexes from Mazourka Canyon, 6,000 feet, Inyo Mts.,
California, on Sphaeralcea ambigua, May 23, 1937 (Michener).
Also taken at Mountain Springs Canyon, Argus Mts., 5,000 feet, Inyo
County, May 22, 1937; Andreas Canyon near Palm Springs, April 6,
1939; Ribbonwood, San Jacinto Mts., May 21, 1940 (Michener);
Pifion Flat, San Jacinto Mts., May 18, 1939 (E. G. Linsley); San
Bernardino County, May 1. 1920 (W. Benedict); and Westgard
Pass Plateau, Inyo County, May 27, 1937.

This form differs from P. arcuata arcuata Fox from Baja Cali-
fornia merely in having the very short transverse clypeus of the
male yellow.

Timberlake: Bees of the Genus Perdita 969

Perdita trifasciata sp. nov.

This little species belongs to a new group which will be accorded
subgeneric standing in a subsequent paper. In this group belong
five species from the Colorado Desert of California, which are
usually found at flowers of Coldenia. The New Mexico species
differs from all except one of the California species in having the
abdomen dark, and from the exception in having the face marks
reduced and the pale abdominal bands very narrow and only three
in number.

Male. — Head and thorax dark olive-green, with yellowish-white
markings as follows: basal half mandibles (these gradually chang-
ing to red at apex), labrum, clypeus, except usual dots and two
broad brown stripes ( somewhat farther apart than their own width )
on disk, transversely linear lateral marks not reaching above level
of clypeus, and a small spot on tubercles and at posterior corners
of pronotum. Middle of supraclypeal area brownish around a very
small whitish spot. Antennae nearly uniformly brown above and
testaceous yellow beneath. Legs blackish, the front and middle
knees, tibiae and tarsi, base of hind tibiae, and base of both hind
tibiae and tarsi, pale yellow; front and middle tibiae blotched with
brown behind. Abdomen black with a very narrow subapical
yellowish-white band on tergites 1 to 3, the band on tergites 1 and 2
emarginate behind on each side, and that on tergite 3 broadly inter-
rupted on each side. Tegulae testaceous brown. Wings somewhat
dusky, the nervures and margins of stigma brown.

Head quadrate, as broad as long, the face below antennae promi-
nent, the temples moderately wide. Disk of clypeus about one half
as long as wide and shorter than the transverse length of the lateral
extensions. Stigma narrowly lanceolate. Substigmatal and post-
stigmatal portions of marginal cell equal. Claws bifid. Sculpture
of head and thorax microscopically tessellate, imparting a soft, dull,
satiny appearance. Abdomen considerably smoother and more
shining than thorax. Pubescence white, rather short, sparse, and
coarse; that on mesoscutum subappressed. Length, about 2.5-3
mm.; anterior wing, 2.1-2.5 mm.

Two males (holotype and paratype), White Sands, New Mexico,
June 27, 1940 (R. H. Beamer).

In Cockerell's table ( 1896 ) this species does not run easily, but
probably goes best to P. austini Cockerell, which is a somewhat
larger, shining species, with the lateral face marks reaching above
level of the antennae.

970 The University Science Bulletin

Perdita ainsliei Crawford

One male, Sioux City, Iowa, July 26, 1924 (C. N. Ainslie).

The ainsliei group, which contains the following and some eight
other undescribed species from the Southwest, apparently deserves
subgeneric recognition.

Perdita peculiaris sp. nov.

A dark blue-green species, differing from P. ainsliei Crawford in
the black abdomen and in having the light markings restricted to
the mandibles and a narrow anterior band on disk of clypeus. ( In
ainsliei the abdomen is ferruginous and the clypeus and quadrate
lateral marks are yellowish white.) In Cockerell's table (1896)
this new species runs to a female variety of P. phymatae Cockerell
which is only superficially similar.

Male. — Head and thorax dark blue-green, the abdomen black.
Mandibles, except rufous tips, and a band on anterior margin of
disk of clypeus, yellowish white. Remainder of clypeus black.
Labrum brown. Antennae blackish, the flagellum more brownish
and becoming broadly rather light brown beneath. Legs black,
the small joints of tarsi brown. Tegulae testaceous brown. Wings
somewhat dusky hyaline, the nervures and margins of stigma sepia
brown.

Head about as wide as long, rather thick fronto-occipitally, the
face below antennae strongly convex. Clypeus prominent, its
anterior margin rounded out and well elevated. Cheeks much
narrowed anteriorly and simple. Subantennal plates about as broad
as high. Inner anterior corners of lateral plates of face rounded
and somewhat tumescent next to disk of clypeus. Abdomen broadly
ovate, convex above, a little broader than thorax. Stigma of fore
wing narrowly lanceolate. Marginal cell broadly and obliquely
truncate at apex, with the portions beneath and beyond the stigma
equal. Outer nervure of second discoidal cell obsolescent. Claws
bifid. Mandibles moderately long, curved, and simple. Maxillary
palpi rather long and six-jointed. Head and thorax minutely tes-
sellate and moderately shining, the metanotum and propodeum
strongly tessellate and more dullish than other parts. Puncturation
of mesonotum very fine and close, and that of frons moderately
sparse. Abdomen also dullish from a minute dense puncturation
(in which it differs from ainsliei as well as almost all other species
of Perdita) except on the depressed apical margin of the tergites.
Pubescence fine, sparse, and inconspicuous. Length, about 3-4 mm.
(the abdomen much recurved at apex); anterior wing, 2.5-3 mm.

Timberlake: Bees of the Genus Perdita 971

Aedeagus of the type of P. ainsliei; the long slender apical lobes
of stipites abruptly thickened and bent downward at apex (acumi-
nate and very sharp at apex in ainsliei). Sagittae forming a fusi-
form body, tapering to the acute apex, as seen from side, but the
tip of each rod expanding acutely outward, as seen from above, and
armed with a slendar upward-directed spine at the inner apical
corner (in ainsliei rather similar, but without the apical spine and
clothed beneath with a loose fascicle of hairs close to the apex).
Volsellae reduced to one pair, which are rather long and acuminate,
as seen from above ( two pairs of volsellae in ainsliei ) .

Two males (holotype and paratype), 15 miles southeast of Dry-
den, Terrell Co., Texas, on Chamaesaracha conioides, April 13, 1949
( Michener and Beamer ) .

Subgenus COCKERELLIA Ashmead

The type of Cocker ellia is P. hyalina Cresson, which has been
usually considered the male of P. albipennis Cresson, although the
synonomy is doubtful. Excepting Xerophasma, the species of
Cockerellia are the largest in the genus Perdita and are found
almost exclusively at flowers of Compositae, mostly of the genera
Helianthus, Rudbeckia, Ratibida, Raileya, Verbesina, Coreopsis,
Geraea, and Gaillardia, but with records also from Erigeron, Hetero-
theca, Pedis, and Prionopsis.

Key to the Species of Cockerellia Females

1. Head and thorax mainly green 2

Mainly bright yellow, the lower half of pleura black; clypeal

dots, facial foveae and flagellum above, also blackish; mesono-
tum with short, dense, erect yellow hair heata Cockerell

2. Clypeus yellow or whitish, the lateral marks triangular and reach-

ing level of antennae or beyond 3

Face marks restricted to spots on clypeus and transverse lateral
marks, the latter not extending above level of clypeus 6

3. Mesonotum minutely, densely punctate, the punctures not much

more than their own diameter apart; supraclypeal area en-
tirely dark 4

Mesonotum polished, finely punctured, the punctures although
very numerous, several times their own diameter apart;
clypeus, triangular lateral marks, and a vestigial supraclypeal
mark, yellowish white; prothorax, legs in large part and ab-
domen yellow, the abdomen with four black bands,

utahensis Cockerell

4. Head and thorax brassy green, the mesonotum a little tessellate

and dullish between the punctures 5

972 The University Science Bulletin

Head and thorax dark blue-green; mesonotum shining, the dense
punctures very minute; base of mandibles, clypeus, lateral
marks, prothorax and legs in large part, and abdomen, yel-
lowish white, the abdomen with five black bands; pubescence
white baileyae Cockerell

5. Clypeus and lateral marks white; prothorax, legs in large part,

and abdomen, pale yellow or yellowish white, the abdomen
with five rather broad black bands, and also black on middle

of basal declivity of tergite 1 perpulchra perpulchra Cockerell

Similar, but light parts of thorax, legs and abdomen bright yellow,
the first tergite entirely yellow except for a narrow apical black
band perpulchra flavidior subsp. nov.

6. Frons dull or opaque, obscurely punctate or impunctate 7

Frons at most dullish, distinctly although minutely punctate 12

7. Wings slightly dusky hyaline, the nervures and stigma testaceous

or pale brown; hair of hind tibiae more or less fuscous 8

Wings whitish or milky hyaline, the nervures pale yellow or
whitish, the stigma frequently yellow or pale orange; hair of
hind tibiae usually but not always light 9

8. Wings rather distinctly dusky, the nervures and stigma pale

brown; clypeus, except two dark stripes which are abbreviated
anteriorly and more or less widened behind, transverse lateral
marks, and a broad band on tergites 2 to 5, narrowly inter-
rupted medially on 2 and 3, yellow ... bequaerti bequaerti Viereck
Wings more faintly dusky, the nervures more yellowish; dark
stripes on clypeus often enclosing a median yellow stripe,
which expands anteriorly bequaerti indianensis Cockerell

9. Frons dull but not opaque, its punctures obscure 10

Frons opaque and virtually impunctate 11

10. Clypeus strongly produced, the width of the anterior truncation

a little less than length of the oblique sides to base of mandi-
bles, the face below antennae prominent in profile; immacu-
lately dark green, with the abdomen blackish, or spot or streak
on middle of clypeus, slender transverse lateral marks, two
spots on pronotum, and interrupted band on tergites 2 to 4,

yellow verbesinae Cockerell

Clypeus less produced, the anterior truncation broader than
length of oblique sides, the face below antennae little prom-
inent in profile; more brassy green than verbesinae; median
streak on clypeus, slender, transversely curved lateral marks,
scape of antennae beneath, cuneate marks on pronotum,
tubercles, and a broad, interrupted band on tergites 2 to
4, yellow fracticincta sp. nov.

11. Large, robust species, about 9 mm. long; face entirely dark, or,

typically, with median stripe on clypeus and transverse lateral
marks, yellow; tubercles at apex, two spots on pronotum,
and broad band, usually reaching lateral margins on tergites
2 to 5, also yellow; hair on hind tibiae whitish to pale
brownish lacteipennis lacteipennis Swenk and Cockerell

TlMBERLAKE: BEES OF THE GENUS PERDITA 973

Slightly smaller; abdominal bands rather narrow, not quite reach-
ing lateral margins and narrowly interrupted on tergites
2 and 3; marks on pronotum and tubercles large; face marks
large, the clypeus yellow except for two triangular dark areas
on disk, with the yellow extending triangularly upward be-
tween them; hair of hind tibiae sometimes pale fuscous; length
about 7-8 mm lacteipennis pallidipennis Graenicher

12. Clypeus more prominent, and more produced, the disk fully as

long or a little longer than wide 13

Clypeus less produced and prominent, the disk somewhat broader
than high; head therefore appears to be slightly broader than
long, with face below antennae much less prominent in profile, 15

13. Frons minutely tessellate, shining, and minutely punctate; yellow

bands of abdomen usually broad, and a yellow mark present

on each side of disk of tergite 1 14

Frons rather strongly tessellate, more dullish and much more
strongly punctate than in albipennis; a rather narrow yel-
low band, not reaching lateral margins on tergites 2 to 4;
tergites 1 and 5 immaculate, or 5 with two yellow spots which
are usually small or faint tricincta sp. nov.

14. Head and thorax dark olive-green; hair of legs light, the scopa

of hind tibiae pale yellowish, yellow abdominal bands usually
reaching lateral margins; face marks pale yellow, the upper
part of clypeus disk with two dark blotches which usually

outline a yellow "T" mark albipennis Cresson

Head and thorax brassy green; hair of legs fuscous to blackish,
especially on middle and hind legs; abdominal bands broad
and even but failing to reach lateral margins; clypeus usually
black, with a medium pale-yellow line scopata sp. nov.

15. Abdomen always dark, the yellow bands nearly or quite reaching

lateral margins on tergites 2 to 5, and tergite 1 dark with a
yellow mark on each side of disk; median yellow mark on
upper part of clypeus not much narrowed above and often

T-shaped 16

Abdomen sometimes yellow, with dark subapical and basal spots
on each side of segments; or sometimes dark, with enclosed
yellow band on tergites 2 to 5, broadest at middle and nar-
rowed toward sides (in which case tergite 1 lacks yellow
lateral marks on disk, but may be more or less yellow at base ) ;
dark blotches on upper part of clypeus large, the yellow intru-
sion between them pointed above 17

16. Frons more or less distinctly tessellate and dullish, the punctures

minute; yellow bands on tergites 2 to 5 rather even and entire,
narrow on tergite 2 and gradually wider on successive seg-
ments, that on tergite 5 subabbreviated,

lepachidis lepachidis Cockerell
Frons almost polished, the punctures consequently very dis-
tinct; markings as in lepachidis, but bands on tergites 2 and 3
generally more or less notched medially behind, and that on 2
sometimes distinctly interrupted. . lepachidis levifrons subsp. nov.

974 The University Science Bulletin

17. Abdomen and legs not almost entirely yellow; the abdomen with

dark markings, at least on tergites 1 to 4 18

Abdomen and legs almost entirely yellow; four small fuscous
spots on disk of tergite 1, and two small subapical spots on
tergite 2; femora of front and middle legs with a pale-brown
blotch on inner side coreopsidis kansensis subsp. nov.

18. Abdomen yellow with a dark band at junction of tergites 1-2 to

4-5, more or less reduced to a basal spot on lateral margins
and a subapical spot on each side; tergite 1 also with a dark
blotch on each side at summit of basal declivity,

coreopsidis coreopsidis Cockerell
Abdomen dark above with an enclosed yellow band on tergites
2 to 5, broadest in middle and narrowed at outer ends; ter-
gite 1 also more or less yellow at base,

coreopsidis obscurior subsp. nov.

Perdita utahensis Cockerell

Five males, 2 females, 10 miles south of Tucson, Arizona, Aug. 7,
1940 (C. D. Michener); 5 males, 1 female collected by the writer
at the same place and time were found at flowers of Verbesina
encelioides. Cockerell described only the female of this species
from a specimen collected in southwestern Utah.

Male. — Head and thorax dark blue-green. Spot at anterior end
of cheeks, mandibles, except rufous tips, labrum, clypeus, large tri-
angular lateral marks reaching level of antennae, first three joints of
antennae broadly beneath, collar and posterior band on pronotum,
narrowed mesad, and the tubercles, bright yellow. Sometimes
upper margin of subantennal plates yellow, and this even confluent
with lateral marks. Legs blackish, the trochanters in part, the
femora broadly at apex and an anterior stripe on front and middle
pair, the tibiae except behind, and the tarsi, yellow. Flagellum
orange ferruginous, becoming dusky above; the pedical and apex
of scape fuscous above. Abdomen shining blackish, the apical
tergite testaceous yellow, and the apical depression of tergites 1
to 6 whitish subhyaline. Preceding the depression is a yellowish-
white band on tergites 1 to 6, this band obliterated except on lateral
margins on tergite 1, dilated on each side and covering reflexed
sides on tergites 2 to 6, and with remaining part of band broadly
interrupted, or almost so, sublaterally, and notched or subinter-
rupted medially on tergites 3 to 5, but the same part on tergite 2
generally obliterated. Tegulae yellow at base and almost hyaline
on outer margin. Wings whitish, with a slight dusky-yellowish
opacity. Nervures and stigma yellowish, the margins of stigma
generally slightly brownish.

TlMBERLAKE: BEES OF THE GENUS PERDITA 975

Head quadrate, the cheeks broad. Face below antennae polished,
impunctate. Vertex and frons minutely granular-tessellate, the lat-
ter moderately shiny and minutely punctate. Mesonotum nearly
polished and set with minute, moderately close punctures. Pubes-
cence white and moderately dense. Length, about 6-7 mm.; an-
terior wing, 4.3-4.9 mm.

Perdita baileyae Cockerell

One male, Rice Co., Kansas, July 3, 1923 (C. H. Martin).

Cockerell described the male of this species from Mesilla, New
Mexico. I have seen a female from San Xavier Mission, Arizona
(E. P. Van Duzee).

Perdita perpulchra flavidior subsp. nov.

Cockerell recorded perpulchra from Las Cruces, New Mexico,
and described the type as having the abdomen white with black
bands, the black on the basal tergite extending triangularly to the
base, with the white on each side enclosing a black subapical spot.
A female taken 15 miles west of Holbrook, Arizona, at flowers of
Erigeron, Sept. 3, 1930 (Timberlake) agrees well with Cockerell's
description, except that the abdomen is more yellowish. In flavidior
the light parts, except those of face, are bright yellow, and the basal
tergite is black only at apex.

Female. — Head and thorax dark olive-green. Basal half of man-
dibles, clypeus, and triangular lateral marks reaching level of an-
tennae, pale yellow. Scape of antennae, except spot at apex above,
under side of pedicel and of first joint of flagellum, prothorax and
legs in large part, and abdomen, bright yellow. Coxae at base,
under side of front and middle trochanters and femora, spot at apex
of hind femora above, posterior surface of front and middle tibiae,
hind tibiae entirely and all tarsi, except basal joint of front pair, dark
brown or blackish. A black band at juncture of tergites 1-2 to 4-5,
with the broad intervening yellow bands distinctly notched on each
side behind by sublateral swellings of the black, especially on ter-
gites 2 and 3. Tergite 5 with a narrow, preapical blackish band.
Venter pale brownish yellow at base; segments 2 and 3 each with
two fuscous patches, and the apical segments extensively fuscous.
Tegulae pale testaceous, with a yellow spot at base. Wings some-
what whitish hyaline, slightly tinged with dusky yellowish. Ner-
vures and stigma testaceous yellow.

Head slightly broader than long, the clypeus little produced and
prominent. Frons slightly duller, with somewhat closer and fainter

32—6490

976 The University Science Bulletin

punctures than in albipennis. Mesonotum minutely and very
closely punctate and covered with fine very short, erect hair. This
rather dense pubescence of mesonotum and that of vertex distinctly
ochreous. Wings, in comparison with albipennis, small and slightly
dusky. Length, 8 mm.; anterior wing, 4.8 mm.

One female (holotype), Great Bend, Barton Co., Kansas, on
Heterotheca subaxillaris, Sept. 9, 1949 (Michener and Beamer).

Perdita bequaerti indianensis Cockerell

In preparing the preceding key to the species of Cockerellia,
I noticed that a specimen from Gary, Indiana, that had been de-
termined by Cockerell as P. pallidipennis Graenicher, had some-
what dusky wings, a character which would relegate it to P. be-
quaerti Viereck. I conclude that indianensis that was based on a
male from Lafayette, Indiana, must be the same as the Gary speci-
men, since the margins of stigma and end of marginal cell could
not be described as fuscous unless the wings were slightly dusky,
rather than milky hyaline as in pallidipennis.

This is a rather weakly developed race of bequaerti, and was
described as a race of pallidipennis. It and P. wickhami Cockerell
were published in the same paper and must be the sexes of prac-
tically the same thing (new synonymy).

Perdita verbesinae Cockerell

Five males, 9 females, Portal, Arizona, on Verbesina encelioides,
Aug. 12, 1940 (C. D. Michener).

Specimens of this species, even those of a series taken at the same
spot, vary greatly in the extent of the yellow markings, and some-
times lack the markings altogether. Such immaculate specimens
were named var. nigior by Cockerell in his original description.

Perdita fracticincta sp. nov.

This species is allied to P. lepachidis Cockerell and P. coreopsidis
Cockerell in the shape of the head, but in the strongly tessellate and
dull frons, the greatly reduced face marks, and the interrupted ab-
dominal bands it strongly resembles verbesinae.

Female. — Brassy green, the labrum and clypeus black, the latter
green, however, on upper margin of disk. Small longitudinal
median spot on clypeus, a small transverse spot on each lateral
extension of clypeus, and slender to transversely arcuate lateral
marks, yellow. The lateral marks bordering anterior margin of
lateral plates of face are slightly interrupted on one side and have

TlMBERLAKE: BEES OF THE GENUS PERDITA 977

a short extension upward on orbits. Interrupted band on hind
margin of pronotum, and the tubercles, yellow. Abdomen blackish,
a small spot on lateral margins of tergite 1 (mostly on the reflexed
ventral portions) and interrupted band on tergites 2 to 4, yellow;
band on tergite 2 reaching the lateral foveae, and that on 3 having
a faint, slender posterior extension to lateral margins. Mandibles
and pygidium rufo-testaceous, the former more rufous at apex.
Antennae blackish above, the scape yellow, the flagellum brown
beneath. Legs blackish, a small spot on front knees and the front
tibiae on anterior side, yellow. Tegulae pale testaceous. Wings
milky hyaline, nervures nearly colorless, the stigma pale yellow.

Head hardly longer than wide, the clypeus little produced and
prominent. Frons and vertex minutely granular-tessellate and
rather dull, the minute shallow punctures of frons only moderately
distinct. Face below antennae polished, with sparse minute punc-
tures. Mesoscutum polished on disk, although minutely tessellate
anteriorly, and provided with rather close minute punctures. In
some ways, including shape of head, pubescence, size, etc., agreeing
closely with coreopsidis. Length, 7 mm.; anterior wing, 4.9 mm.

One female (holotype) 29 miles south of Sarita, Willacy Co.,
Texas, on Coreopsis, Apr. 14, 1950 ( Michener, Beamer, Stephen and
Bozen ) .

Perdita lacteipennis lacteipennis Swenk and Cockerell

Unquestionably a common species in western Kansas at flowers
of Helianthus, as there is a series of 127 females and 145 males in
the University of Kansas collection from various localities in the
Counties of Chase, Beno, Sheridan, Ottawa, Hamilton, Greeley,
Meade, Ford, Finney, Pawnee, Osborne, Barton, Pratt, Russell,
Rawlins, Saline, Ellsworth, Comanche, Stafford, Clark, Decatur,
Rice, and Rooks. Nearly all the specimens that are labeled with the
flower record were taken on Helianthus petiolaris, but 6 males from
Chase and Hutchinson were on Prionopsis ciliata. The dates of
capture range from June 14 to Sept. 6.

Also 79 specimens from La Junta, Colorado, two of them labeled
as from Helianthus annuus; 2 males, Lamar, Colorado (Snow); 1
male, Artesia, Moffat Co., Colorado, on Helianthus petiolaris, July
22, 1950 (Michener); and 1 male each from Leonard, North Dakota,
July 25, 1937, and Lusk, Wyoming, July 14, 1937 (C. L. Johnston).

Specimens from western Kansas are mostly typical lacteipennis,
although about 10 per cent of the females have the face marks very
small or absent and thus agree with the form canadensis Crawford.

978 The University Science Bulletin

Of 28 females from La Junta, Colorado, 6 verge toward canadensis
(face marks obsolescent), 6 verge toward pallidipennis (clypeus
with a yellow spot on each side besides the median mark), and 16
are typical lacteipennis.

Perdita lacteipennis pallidipennis Graenicher

Thirty-two females, 15 males, Douglas Co., Kansas, on Hclianthus
annuus, Sept. 9, 1949 (Michener and Beamer); 3 females, Topeka,
Kansas, Sept. 17, 1910 (E. G. Titus).

These specimens are nearly typical pallidipennis, the female of
which has the clypeus yellow, with a dark blotch above on each
side of disk, or the dark color frequently reduced to a short strip
on each side. Males have the basal half of mandibles, labrum, and
anterior border of face yellow.

Specimens from Clay Co., Kansas (J. C. Bridwell), average
somewhat closer to pallidipennis than to typical lacteipennis but
probably should be classed as intermediates.

In the original description of lacteipennis it is stated on the au-
thority of Prof. Myron Swenk that the species does not occur in
eastern Nebraska. However, the species does occur there, as I
have a female of pallidipennis from Nebraska City on Helianthus,
Sept. 14, 1901 (M. A. Carriker). The form pallidipennis has the
face markings much as in albipennis and hence must have been
confused with albipennis by Swenk. The dull frons of lacteipennis
and the shining punctate frons of albipennis easily distinguish the

two species.

Perdita tricincta sp. no v.

Nearest P. albipennis Cresson, but distinguished by the duller,
more tessellate, and more coarsely punctate frons, and by the three
narrow bands on abdomen (the yellow on the fifth tergite restricted
to two small spots or absent). The male has a dark, shining abdo-
men and transverse yellow face marks.

Female. — Head and thorax dark green, the mesonotum some-
what brassy. Labrum and clypeus blackish, the latter with a broad,
pale-yellow, median stripe and a suffused yellowish mark on each
side. Lateral marks pale yellow, rounded at inner ends and gen-
erally somewhat less than twice as broad as high. Mandibles
testaceous brown, the tips dark rufous. Collar of pronotum ob-
scurely yellowish, the hind margin and tubercles bright yellow.
Abdomen blackish, with a rather narrow yellow band on tergites
2 to 4, that on 2 ending at the lateral fovea, the others not quite

TlMBERLAKE: BEES OF THE GENUS PeRDITA 979

reaching lateral margins, all slightly notched medially, those on
tergites 2 and 3 behind and that on tergite 4 in front. Tergites 1
and 5 entirely black, or the latter frequently with two submedian
yellow spots. Pygidium ferruginous brown. Legs blackish, the
front tibiae yellowish on anterior side. Antennae blackish, the
scape narrowly clear yellow and the flagellum brown beneath.
Tegulae pale testaceous. Wings milky hyaline, the nervures nearly
colorless, the stigma pale yellow.

Head about as broad as long, with the clypeus produced and
prominent. Facial foveae about one third as long as eyes. Sculp-
ture as in albipennis, except the frons duller, with a stronger tessella-
tion and coarser punctures. Pubescence of vertex and mesonotum
more ochreous than in albipennis. Scopa of hind tibiae a pale
yellowish brown. Length, 7.5-9 mm.; anterior wing, 5-5.2 mm.

Male. — Head and thorax dark green. Base of mandibles, an-
terior band on clypeus (more testaceous far laterad), sometimes a
median streak on clypeus to summit, and lateral face marks, about
twice as wide as high, yellow. Mandibles testaceous in middle and
rufous at tips. Labrum testaceous. Antennae orange-brown, a
little dusky on apical half of flagellum above, becoming darker
toward base of flagellum, and blackish on pedicel and on the scape
except at base, with underside of scape and of first joint of flagellum
yellow. Collar of pronotum yellowish, especially toward the mid-
dle, a small cuneate spot on posterior corners of pronotum clear
yellow, but tubercles dark. Abdomen shining blackish, the apical
tergite testaceous, the apical depression of preceding segments sub-
hyaline. In some specimens, including the allotype, there is a trace
of yellow just in front of the apical depression on tergites 5 and 6
and on the reflexed sides of the preceding segments. Legs blackish;
all tarsi, front tibiae broadly and middle tibiae narrowly on anterior
side, and front and middle knees, yellow. Tegulae and wings as in
female. Head quadrate, the cheeks broad. Sculpture as in female,
the pubescence white. Length, 5.3-8 mm.; anterior wing, 4-5 mm.

Holotype female and allotype, Southmost, Cameron Co., Texas,
on Helianthus annum, Apr. 13, 1950 (Beamer, Michener, Stephen,
and Rozen). Paratypes as follows: 30 females, 40 males taken
with the types; 1 male, 5 miles east of Brownsville, Apr. 13, 1950;
3 females, 4 males, Progresso, Hidalgo Co., on Helianthus annuus,
Apr. 12, 1950; 1 male, 6 miles east of Rio Grande, Starr Co., Texas,
Apr. 12, 1950 ( all, Michener, Beamer, Stephen, and Rozen ) .

980 The University Science Bulletin

Perdita albipennis Cresson

The material recorded below belongs almost entirely, in respect
to the females, to the variety lingualis Cockerell, which is the best-
known and the most common form of albipennis in collections. This
more highly colored form is, acording to Cockerell, possibly a valid
race. The male variety helianthi Cockerell, with yellow markings
on abdomen and a more or less blackish flagellum, flies in about
equal numbers with males having the flagellum orange brown or
ferruginous and the abdominal markings obsolete, and both fly
with lingualis. The P. hyalina Cresson, without face or abdominal
markings indicated in the original description, seems distinct from
albipennis. The variety pasonis Cockerell, with a dull frons, is
more likely allied to P. lacteipennis than to albipennis. Altogether,
therefore, it is necessary to have a much fuller collection of albi-
pennis in order to understand its variation and possible subspecia-
tion.

The following material of albipennis has been examined:

Kansas: 14 females, 17 males, Clark Co., 1,950 feet, Aug. 23,
1911; 2 females, 1 male, Rawlins Co., 2,850 feet; 1 female, Norton
Co., 2,270 feet, Aug. 12, 1912; 2 females, Smith Co., 1,800 feet, Sept.
4, 1912; 2 females, 1 male, Stevens Co., 2,700 feet, Aug. 10, 1911;
1 male, Sherman Co., 3,690 feet; 1 male, Ellis Co., 2,000 feet, July
18, 1912; 1 male, Decatur Co., 2,560 feet (all F. X. Williams); 1
male, Ness Co.; 10 females, 4 males, Sharon Springs, Wallace Co.,
on Helianthus annuus, Aug. 23, 1931 ( Timberlake ) ; 1 male, Lakin,
Kearny Co., on Helianthus petiolaris, July 23, 1950 (C. D. Mich-
ener ) .

Colorado: 4 males, La Junta, July 3, 1949 (R. H. Beamer and
J. R. White).

Wyoming: 1 male, Lingle, Goshen Co., July 14, 1937 (C. L.
Johnston ) .

New Mexico: 1 female (probably belonging here, but it cannot
be identified positively as the head is missing), Cuervo, Guadalupe
Co., June 23, 1940 (E. E. Kenaga).

Arizona: 1 male, Fort Apache, Navajo Co., Aug. 26, 1897 (Snow,
No. 2113).

Perdita scopata sp. no v.

This species is much like P. albipennis, but the hair of middle and
hind legs and that of front tarsi more or less blackish, a character
more faintly indicated in some of the other species of Cockerellia,
such as P. bequaerti and P. lacteipennis pallidipennis, which, how-

Timberlake: Bees of the Genus Perdita 981

ever, have the frons dull and impunctate. The male of scopata has
the face markings greatly reduced or absent and the abdomen im-
maculate, opaque black.

Female. — Head and thorax dark green, the frons, vertex, and
mesonotum with a strong brassy luster, the labrum and clypeus
black. Line on under side of scapes, median line on clypeus, not
reaching anterior margin, and slightly transverse lateral marks,
whitish. The lateral marks, rounded at inner ends, extend about
half way to level of antennae. Sometimes a transverse whitish spot
on lateral margins of disk of clypeus. Mandibles black at base and
rufous on apical third or half. Antennae blackish, the flagellum
slightly rufescent beneath. Tubercles and hind margin of pronotum
pale yellow, the anterior margin narrowly yellowish. Abdomen
black, the pygidial plate dark rufous. Tergites 2 to 5 each with a
broad yellow band, these bands more even than in albipennis and
falling distinctly short of the lateral margins. A slender, oblique,
more or less sinuate and broken yellow line on each side of the base
of the disk of tergite 1. Legs blackish, the front knees and a line
on anterior side of front tibiae suffused with yellow. Tegulae tes-
taceous. Wings whitish hyaline, the nervures and stigma pale tes-
taceous yellow (in comparison with albipennis the wings have a
barely perceptible dusky tinge and the nervures are less pallid).
Structure, sculpture, and pubescence as in albipennis, with the fol-
lowing differences: cheeks less broadened and clypeus less prom-
inent; punctures of mesonotum slightly closer; abdomen dull, with
an excessively minute tessellation; pygidium very broad and broadly
subtruncate at apex; basal middle of propodeum minutely rugose;
pubescence flavo-ochraceous on top of head and on mesonotum,
more or less dark fuscous on middle and hind legs and front basi-
tarsi, and dilute fuscous at apex of abdomen. Length, 7-9 mm.;
anterior wing, 5-5.5 mm.

Male. — Dark green, the mesonotum with a brassy luster, the base
of mandibles, labrum and clypeus shining black. Abdomen opaque
black, the apical depression of tergites more shining but not palles-
cent; tergite 7 becoming brownish testaceous on apical half. No
light facial thoracic or abdominal markings, except rarely a small
pale-yellow spot on each side of disk of clypeus and another opposite
anterior corners of eyes. Mandibles rufous at apex. Antennae
blackish, the scape pale testaceous at extreme base, the flagellum
brown beneath. Legs blackish, a small spot on front and middle
knees pale yellowish, the front and middle tarsi testaceous yellow,
and the hind tarsi brownish. Tegulae testaceous. Wings milky

982 The University Science Bulletin

hyaline, with pale-yellow nervures and stigma (wings distinctly
whiter than in female). Structure, sculpture, and pubescence simi-
lar to condition in male of albipennis, except the frons slightly
duller, with less distinct punctures, mesonotum with somewhat
closer punctures, base of propodeum with a small, triangular, finely
rugose area, and abdomen dull from an excessively minute thimble-
like tessellation. Length, 6-8 mm., anterior wing, 5-5.5 mm.

Holotype female and allotype, 29 miles south of Sarita, Willacy
Co., Texas, on Coreopsis, Apr. 14, 1950 (Beamer, Michener, Stephen,
and Rozen). Paratypes as follows: 25 females, 61 males, taken
with the types; 10 females, 36 males, same locality and date, at
flowers of Helianthus annuus; 7 females, 4 males, Southmost,
Cameron Co., on Helianthus annuus, Apr. 13, 1950; 1 male, 5 miles
east of Brownsville, Apr. 13, 1950 (Beamer, Michener, Stephen,
and Rozen ) ; 1 female, 20 miles south of Sarita, on yellow composite,
April 2, 1946 (C. D. Michener); 1 female, Hidalgo Co., Texas,
May 5, 1929 (H. B. Parks); 1 female, 1 male, Brownsville, June
(F. H. Snow, No. 2428); and 1 female, 1 male, Galveston, May
(Snow, No. 1824).

The Brownsville specimens taken in June ( Snow ) differ in having
considerably larger face marks than usual.

Perdita lepachidis lepachidis Cockerell

This species was described in 1896 from Santa Fe and Socorro,
New Mexico. Only the male was known to Cockerell, who col-
lected it at flowers of Ratibida tagetes (the plant was then called
Lepachys tagetes ) .

Female. — Brassy green, much as in coreopsidis, but the median
pale-yellow area of clypeus briefly crossed at the top like a "T" and
expanding below, or sometimes isolated and Indian club-shaped
(in coreopsidis triangular and pointed above). Anterior and pos-
terior margin of pronotum and tubercles bright yellow. Abdomen
blackish, with a spot on each side of tergite 1 and a crossband on
tergites 2 to 5, bright yellow; these bands reaching lateral margins
or nearly so, and that on tergite 2 rather narrow, but those on
tergites 4 and 5 broad and emarginate behind on each side. Legs
dark, the anterior side of front and middle tibiae and a small spot
on knees, yellow. Scape of antennae yellow except at apex above,
the flagellum brown above and yellowish brown beneath but be-
coming yellow beneath on the basal joint and on the pedicel.
Tegulae pale testaceous. Wings milky hyaline, the nervures and
stigma pale yellow. Length, about 7 mm.; anterior wing, 5-5.3 mm.

Timberlake: Bees of the Genus Perdita 983

Four females, Lakin, Kearny Co., Kansas, on Ratibida columnaris,
July 23, 1950 ( C. D. Michener ) ; and 1 female, Sheridan Co., Kansas,
2,650 feet (F. X. Williams).

The specimen from Sheridan Co. differs in having the clypeus
yellow with two dark stripes which are irregular, that on the left
side forming a figure 7 and that on the other side confluent with
the usual lateral dot.

I have examined, also, a pair from Alpine, Texas, July 8, 1942
(H. A. Scullen), one female from Sterling, Logan Co., Colorado,
May, 1923 (Grace Sandhouse), and two males from Santa Fe and
Socorro, New Mexico (Cockerell).

Perdita lepachidis levifrons subsp. nov.

This race differs from typical lepachidis in the smoother, more
shining frons and in the more restricted facial and abdominal
markings.

Female. — Brassy green. Lateral face marks and clypeus pale
yellow, the disk of latter with a blackish blotch (becoming green
on outer margin above) on each side, isolating a rather slender
median yellow streak, which is briefly crossed at summit to form a
"T." Yellow abdominal bands not quite reaching lateral margins,
that on tergite 5 generally much abbreviated, those on tergites 2 and
3 notched medially behind, and that on tergite 2 frequently narrowly
interrupted in middle. Flagellum more ferruginous and the dark
part of legs, especially on tibiae, more brownish than in lepachidis.
Frons so delicately tessellate as to be almost polished, its punctures
minute but distinct. Length, 6.5-7.5 mm.; anterior wing, 4.8-5.1 mm.

Male. — Not appreciably differing from male of lepachidis from
New Mexico, except that frons is nearly polished and the wings
are smaller. Length, 6-7 mm.; anterior wing, 4.3-4.6 mm. (length
of wing in New Mexico male, 5 mm. ) .

Nine females, 4 males (holotype female, allotype, and paratype),
Brownsville, Texas, June (F. H. Snow, Nos. 2416 and 2417). Three
of the females lack the abdomen.

Perdita coreopsidis coreopsidis Cockerell

This species was described from specimens collected at Cotulla,
La Salle Co., Texas, at flowers of Coreopsis, but it apparently prefers
Gaillardia to any other flower.

Texas: 2 females, 1 male, Catarina, Dimmit Co., on Monarda
punctata coryi, Apr. 11, 1950 (Beamer, Michener, Stephen, and
Rozen ) .

984 The University Science Bulletin

Kansas: 9 males, Dodge City, Ford Co., on Gaillardia, June 15,
1949 (Michener and Beamer); 1 female, Satanta, Haskell Co., on
Gaillardia, June 16, 1949 (Michener and Beamer); 1 female, 5 miles
north of Quinter, Sheridan Co., on Gaillardia, June 17, 1950 (Mich-
ener); 1 male, Ashland, Clark Co., on Gaillardia, June 12, 1949
(Michener and Beamer); 4 females, 4 males, Lamed, Pawnee Co.,
on Gaillardia, June 14, 1949 (Michener and Beamer); 2 females,
Clark Co., 1,962 feet, May and June (F. H. Snow, Nos. 1160 and
1161); 2 females, Gove Co., 2,813 feet (F. X. Williams).

Perdita coreopsidis kansensis subsp. nov.

This form flies with the typical subspecies on the northwestern
border of the known range of coreopsidis, but farther south and
west it apparently attains the status of a race. It differs from
typical coreopsidis in having the legs and abdomen of the female
nearly all yellow, and the abdomen of male yellow with dark bands.

Female. — Like typical coreopsidis except in color of legs and
abdomen. Legs yellow, the anterior and posterior surface of front
femora beneath except at apex, and anterior side of middle femora
beneath except apex, brown or fuscous, and the hind margin of
middle and hind tibiae and basitarsi slightly darkened. Abdomen
yellow, with four fuscous spots on tergite 1 and sometimes two
spots on tergite 2, these spots being situated as in the spotted form
of coreopsidis, but small.

Male. — Yellow face markings similar to those of female ( obsolete
or nearly so in typical male). Apex of femora, the tibiae and tarsi
nearly all yellow (the yellow mainly restricted to anterior side of
front tibiae in typical male). Tergite 1, except a narrow apical
light crossband, and a crossband at base of tergites 2 to 5, black,
the rest of abdomen yellow, except for a small transverse preapical
dark spot on each side of tergites 2 and 3. Apical depression of
tergites whitish subhyaline as in the typical male.

Two females, 1 male (holotype female, allotype, and paratype),
Gove Co., Kansas, at 813 feet (F. X. Williams); 1 male (paratype),
Dodge City, Ford Co., Kansas, on Gaillardia, June 15, 1949 ( Mich-
ener and Beamer); and 1 male (paratype), Van Horn, Culberson
Co., Texas, May 23, 1932 (E. G. Linsley).

Perdita coreopsidis obscurior subsp. nov.

This race of coreopsidis appears to be restricted to the coast of
Texas, from the mouth of the Rio Grande to Galveston. It differs

Timberlake: Bees of the Genus Perdita 985

from the typical form in the less extent of yellow markings, and in
the slightly sparser puncturation of the mesoscutum.
Female. — Like coreopsidis except in color of abdomen. Tergite

1 nearly all dark except at base, and the following tergites fuscous
or blackish around the margins to enclose a broad yellow band that
is more or less narrowed and acute at outer ends. On tergites 4 and
5 the dark margin sometimes broadly interrupted in middle apically
and more rarely also at the sides, thus more or less isolating two
dark spots before the apical depression. Length, 6-7.5 mm.; anterior
wing, 4.1-4.6 mm.

Male. — Mandibles except tips and lateral extensions of clypeus,
yellow; disk of clypeus black, with the yellow sometimes intruding
slightly on each side anteriorly. Abdomen with the yellow mark-
ings evanescent or absent and never forming more than a thin line
interpolated between the dark base and the subhyaline apical de-
pression. Punctures of mesoscutum moderately close. Length, 4.8-
6.5 mm.; anterior wing, 3.9-4.4 mm.

Sixteen females, 59 males (holotype female, allotype, and para-
types), Boca Chica, Cameron Co., Texas, on Gaillardia, Apr. 13,
1950 ( Beamer, Michener, Stephen, and Rozen ) . Also the following
paratypes: 1 female, Brownsville, July 3, 1938 (R. H. Beamer);

2 females, 12 males, Brownsville, Apr. 13, 1950 (Beamer); 6 fe-
males, 2 males, 18 miles north of Harlingen, Cameron Co., on
Gaillardia, Apr. 2, 1938 (C. D. Michener); 1 female, 1 male, Gal-
veston, May (F. H. Snow, Nos. 1825, 1826); and 3 females, 1 male,
Corpus Christi, June 28, 1942 (E. S. Ross).

LITERATURE CITED

COCKERELL, T. D. A.

1896. The bees of the genus Perdita F. Smith. Proc. Acad. Nat. Sci.
Philadelphia, vol. 48, 1896, pp. 25-107.

THE UNIVERSITY OF KANSAS

SCIENCE BULLETIN

Vol. XXXV, Pt. II] September 10, 1953 [No. 8

Comparative Morphological and Systematic Studies of

Bee Larvae With a Key to the Families of

Hymenopterous Larvae lp 2

By Charles D. Michener
CONTENTS

PAGE

Introduction 988

Acknowledgments 988

A brief account of hymenopterous larvae 989

A key to some groups of hymenopterous larvae 993

Evolutionary trends 995

Relative specialization of principal types of bee larvae 997

Reversibility in evolution 999

An account of evolution among bee larvae 1000

Some general problems of larval evolution 1005

Techniques 1006

External morphology of Anthophora stanfordiana 1007

Descriptive accounts 1015

Colletidae 1015

Halictidae 1023

Andrenidae 1033

Megachilidae 1040

Xylocopinae 1060

Anthophorinae 1066

Apinae 1084

Abstract: This paper is a study of known bee larvae from all parts of the
world. It contains a key to the larvae of the major groups of Hymenoptera
and an account of evolutionary trends among bee larvae. A detailed study of
the external morphology of Anthophora is presented, ( 1 ) as a contribution to
insect morphology, ( 2 ) as a basis for the establishment of a suitable terminology
for the structures, and (3), as a basis for comparison of other genera. De-
scriptive comments on known bee larvae are presented, together with fuller
descriptions and figures of forms from the Western Hemisphere.

1. Prepared with the aid of grants from the University of Kansas General Research
Fund and the Kansas Industrial Fellowship Fund.

2. Contribution number 802 from the Department of Entomology of the University of
Kansas.

(987)

988 The University Science Bulletin

INTRODUCTION

The purposes of this paper are to shed light on the evolutionary
trends and phylogeny of the bees as exemplified by larval character-
istics, to interpret insofar as possible problems of larval evolution
generally, and to make possible identification of certain bee larvae
when not accompanied by associated adults.

The scope of the work is necessarily limited by the relatively few
species which have been available for study. It is a deplorable
fact that while many fine papers on bee biologies have been written,
particularly in Europe, and while larvae of many bees have been
seen, often sketched or photographed, most of these larvae have
never found their way to museums nor have they been described or
illustrated in detail. The magnificently illustrated works of Guido
Grandi provide the most important exception; his descriptions and
illustrations are virtually as useful as specimens themselves.

The scope of the study is also limited almost exclusively to mature
larvae. The earlier stages are even less available in collections than
are mature larvae. Except as otherwise indicated, all descriptions
and figures in this paper are based upon mature larvae.

ACKNOWLEDGMENTS

Special thanks are due to Dr. J. W. MacSwain of the University
of California who assembled so many of the larvae used in this
study that the work would have been impossible except for his
generous loan of specimens. Others who have provided specimens
of interest from their own collections or from collections of which
they have charge include Drs. E. G. Linsley and P. D. Hurd of the
University of California; Dr. B. D. Burks of the Division of Insect
Identification, U. S. Department of Agriculture; Dr. G. E. Bohart of
the Bureau of Entomology and Plant Quarantine, Logan, Utah;
Dr. P. O. Ritcher of Oregon State College; and Dr. Warwick E. Kerr
of the University of Sao Paulo.

Mention should be made of several associates at the University
of Kansas who have made observations on bee habits and obtained
some larvae: Jimmie R. White, Alvaro Wille, Wallace E. LaBerge,
and Roland L. Fischer.

Special thanks are due to those who have helped with the prep-
aration of the illustrations, Barbara L. Rozen, Julie Sokal, and
Barbara Trotter.

Michener: Comparative Studies of Bee Larvae 989

A BRIEF ACCOUNT OF HYMENOPTEROUS LARVAE

In order to establish which of the characteristics of the various
bee larvae are primitive and which are specialized, a general survey
of hymenopterous larvae seems appropriate. As Peterson (1948)
says, probably less is known about the larvae of Hymenoptera than
about those of any of the other great orders. This applies par-
ticularly to the Clistogastra, many students having been impressed
by the fact that larvae of this group all look more or less alike as
a result of reduction and loss of sclerotization and structures. No
doubt this reduction is correlated with their habitats; none of the
larvae of Clistogastra are free-living, all are inhabitants of plant
or animal tissues or live in protected places (nests) constructed by
their parents or their hosts. A recent author remarks on the sim-
plified soft bodies of these insects and even goes so far as to say
that sclerotized mouthparts are usually wanting. This statement
no doubt results from an examination of the larva of Apis, whose
mouthparts, even the mandibles, are relatively unsclerotized. A
casual examination of figures of the mandibles of other bees will
show its inaccuracy.

No attempt has been made to give an exhaustive treatment of
hymenopterous groups or to make a full survey of the literature
other than that on bees. The following remarks, pertaining to
mature larvae, are based on the principal literature and on the
specimens at hand, special emphasis being placed on characters
that vary among bee larvae because of the light which this infor-
mation sheds on the relative specialization of such characters among
bees.

The larvae of the suborder Chalastogastra usually have thoracic
legs and often abdominal prolegs as well; the body usually bears
setae, and there are often sclerotic projections at the end of the
abdomen. Free-living forms have eyes above the antennae. The
antennae and maxillary and labial palpi are usually several seg-
mented. The cardo is distinct from the stipes, while a lacinia and
galea are recognizable. The salivary opening is single, not guarded
by lips or sclerotic projections. The mandibles are robust, with
the inner surfaces concave and with several stout apical teeth.
Some works describing and illustrating these larvae are Peterson
(1948), Yuasa (1922), MacGillivray (1914), Short (1952) and
Tokunaga, Tsujita, and Yamazaki ( 1951 ) .

The larvae of the suborder Clistogastra differ from those of the
Chalastogastra in lacking legs, in being unsclerotized and pale

990 The University Science Bulletin

in color except for the spiracles and portions of the head capsule
and mouthparts, and in lacking eyes. The antennae and maxillary
and labial palpi have at most one projecting segment. The laciniae
are absent or not recognizable.

The wasps have larvae more like those of Chalastogastra than do
any other Clistogastra. In the Vespidae the body often bears a
few setae and the antennae are reduced to mere convexities each
of which bears a slightly more convex central area which is pro-
vided with a few sensillae. The maxillae and labium are distinct
projecting structures but the maxillary and labial palpi are one-
segmented, papillalike. The former are similar to the galeae, so
that each maxilla appears to bear two papillae. The cardo is dis-
tinct from the stipes, while the lacinia is unrecognizable. The
salivary opening is a transverse slit guarded by thin, slightly scle-
rotized lips. The mandibles bear several large teeth at their apices,
or the apices may be attenuate, bidentate or simple. Vespid larvae
are well described and illustrated by several authors ( Grandi, 1928a,
1930, 1934, 1934a, 1935, 1937; Maneval, 1936, 1939; Micheli, 1930,
1934; Parker, 1943; Reid, 1942; Short, 1952; Soika, 1933, 1934).
Reid has given an extensive treatment with a key to the subfamilies.

The larvae of Scoliidae (Grandi, 1940), Formicidae (Athias-
Henriot, 1947; Menozzi, 1936; Starke, 1949; W. M. Wheeler, 1918;
G. C. Wheeler, 1928, 1935, 1943, 1948, 1950; G. C. and J. Wheeler,
1951, 1952), Pompilidae (Grandi, 1926, 1939; Maneval, 1936, 1939),
Chrysididae (Maneval, 1936; Soika, 1934), and Sphecidae (Grandi,
1926, 1928, 1928a, 1929, 1930, 1931, 1934a; Maneval, 1936, 1939;
Micheli, 1929, 1930, 1933, 1937; Soika, 1932, 1934) are essentially
similar to those of the Vespidae. The body setae are present in some
( e. g. Sphecidae ) . The antennae are frequently a little better de-
veloped than in the Vespidae with a distinct projecting papilla
bearing the sensillae. The salivary opening is similar to that of the
vespids in all of the above mentioned wasps groups except that in
many Sphecidae it is divided into two openings, each of which may
project as a long process. All intergrades exist within the Sphecidae
from the single slitlike opening found in the Sphecinae (Ammo-
phila, Sceliphron) and Astatinae, to the two small separate open-
ings on projections characteristic of most other sphecids (see
Michener, 1952). The maxillary galea is reduced in size in some
sphecids (Cerceris, Grandi, 1926, 1928a; Philanthus, Grandi, 1931;
and Miscophus, Maneval, 1939) and is in fact entirely absent in
one minute sphecid, Ammoplaniis (Maneval, 1939). While absence
of this papilla (galea) is ordinarily a character of the Apoidea, the

Michener: Comparative Studies of Bee Larvae 991

Ammoplanus larva does not show other apoid characteristics, for it
has the several apical mandibular teeth and the paired salivary
openings characteristic of related sphecids.

Larvae of the Evaniidae (Genieys, 1924) and Gasteruptionidae,
subfamily Gasteruptioninae (Short, 1952) differ from the wasps in
lacking maxillary galeae. In this respect they resemble the Apoidea,
from which they differ in having broader, tridentate mandibles.
Larvae of the Gasteruptionidae, subfamily Aulacinae ( Short, 1952 ) ,
have retained the galeae as distinct papillae and have paired salivary
openings as in most subfamilies of Sphecidae.

The Apoidea or bees, like the Evaniidae and the Gasteruptioninae,
differ from the wasps principally in lacking the maxillary galeae so
that there is only one papilla (the palpus) on each maxilla. Occa-
sionally, as in Centris ( ■= Eulaema ) there is a projection in the
position which would be occupied by the galea, but it is obviously
a mere projection and not a distinct papilla, as is the galea in wasps.
The bees also differ from wasps, ants, and the like in lacking the
maxillary cardines as distinct sclerites, although in some groups,
especially the megachilids and some halictids, they are represented
by swellings. Moreover the mandibles are simple or bidentate at
the apices, frequently with small teeth or serrations on the margins
and inner surfaces. This is in contrast to wasps which usually have
several large apical teeth and rarely have small teeth or serrations.
The salivary opening is a slit, with sclerotic lips, or the lips may be
absent; more rarely it is a round or oval opening, sometimes very
inconspicuous. It is never bifid as in many sphecids. Setae are
present on the body principally in the Megachilidae.

The Oryssidae, which on the basis of adult characters are merely
parasitic Chalastogastra, have larvae with mouthparts more re-
duced than in the bees. The mandibles are tridentate with the
median tooth longest, the whole mandible remarkably nearly bi-
laterally symmetrical, as shown by Rohwer and Cushman (1917).
The labium is separated from the maxillae by grooves which do
not reach far back on the sides of the head. The maxillary struc-
ture is difficult to interpret in the light of Rohwer and Cushman's
illustration but there appears to be a basal piece (possibly cardo)
at the level of the mandibular bases, not back toward the rear
margin of the head. If this is the case, there is a great difference
between the Oryssus structure and that of any other hymenopteron.
Thanks to Dr. B. D. Burks and the authorities of the United States
National Museum, I have been able to study some of the material
used by Rohwer and Cushman, but it is now mounted on slides and

33—6490

992 The University Science Bulletin

difficult to interpret. However, it does appear that the maxillae
were more or less correctly illustrated by Rohwer and Cushman.

The Chalcidoidea and Cynipoidea are characterized by the re-
duction of all mouthpart structures except the mandibles; the latter
are commonly acute and edentate although they are sometimes
toothed apically, particularly in phytophagous cynipoids. The
labium and maxillae are usually (in Chalcidoidea) indistinguishably
fused with the head and scarcely recognizable; rarely in chalcidoids,
more commonly in cynipoids, they are separated by grooves; at
their apices the palpi are absent or reduced to minute papillae or
the maxillary palpi are distinct tubercles in cynipoids; the galeae
are absent, and the cardines unrecognizable. The salivary opening,
when recognizable, is small, single, not guarded by lips, sometimes
a transverse slit. The body is often provided with setae.

Larvae of many species of chalcidoids are described and illus-
trated by Buscalioni and Grandi (1938), Parker ( 1924 ) , Salt (1931),
and Short (1952), while those of cynipids are described and illus-
trated by Short (1952), and Yasumatsu (1943).

The Ichneumonidae, Braconidae and Agriotypidae have remark-
able mouthparts in which the rather small, usually acute and
edentate, mandibles are the principal movable structures. The
other appendages and sclerites of the front of the head are reduced
to flat or slightly convex areas. The conspicuous structures in this
region (except in Collyria) are not these appendages and sclerites,
but the lines between them, which are strongly sclerotized. Sug-
gestions of such sclerotization may be seen in bees and wasps,
where they may indeed be conspicuous; however, in these forms
the maxillae, labium, etc., are always projecting lobes. In the
ichneumonoids, on the other hand, there are no projecting lobes or
but feeble ones. The sclerotized lines are as follows: (1) one
following the course of the epistomal suture, (2) one on each side
extending between the anterior and posterior mandibular articula-
tions ( pleurostomal thickening), (3) one extending from each pos-
terior mandibular articulation to the posterior tentorial pit (hypo-
stomal thickening), (4) one on each side separating the maxillae
from the labium ( labio-maxillary thickening or maxillary sclerome
of Vance and Smith, 1933 ) , ( 5 ) one dividing each maxilla between
cardo and stipes (stipital sclerome of Vance and Smith), and (6)
one separating the prementum from the postmentum ( labiostipital
sclerome of Vance and Smith). In these insects there is only one
maxillary papilla (presumably the palpus) and the salivary open-
ing is slitlike, without distinct lips, or is apparently absent.

Michener: Comparative Studies of Bee Larvae 993

Some excellent descriptive and illustrative work on larvae of
ichneumonoids is provided by Beirne (1941), Salt (1931), Short
( 1952 ) , Thorpe ( 1930 ) , and Vance and Smith ( 1933 ) . Beirne and
Short give keys for the separation of families and subfamilies.

A Key to Some Groups of Hymenopterous Larvae 1

( Based on mature larvae )

The following key to some groups of Hymenopterous larvae summarizes
some of the information contained in the preceding paragraphs. In preparing
it, I have borrowed freely from the keys of Peterson ( 1948) and Soika ( 1934).
It should be remembered that in the Clistogastra only an exceedingly small
percentage of the species have been studied. Therefore many exceptions will
probably be found to the statements of the key; it is merely an outline to be
improved as investigators obtain more information.

1. Antennae and maxillary and labial palpi distinctly several-seg-

mented (if one-segmented, then apex of abdomen bears scle-
rotic processes); lacinia distinct; eye spot usually present;
thoracic legs usually present and segmented; abdominal pro-
legs usually present ( suborder Chalastogastra ) 2

Antennae and maxillary and labial palpi one-segmented to ab-
sent (apex of abdomen not sclerotized ) ; lacinia indistinct or
absent; eye-spot absent; legs absent. ( Suborders Clistogastra
and Idiogastra) 13

2. Thoracic legs with claws 3

Thoracic legs without claws 10

3. No segmented subanal appendages on the tenth abdominal seg-

ment, if setiferous subanal knobs are present see couplet 8
(Xyelidae); thoracic legs usually stout, short and irregular in

shape; prolegs usually present 4

Paired, segmented, subanal appendages present on the sternum
of the tenth abdominal segment; thoracic legs slender, elon-
gated and straight; prolegs absent Pamphiliidae

4. Claws on thoracic legs usually conspicuous and with a distinct

pad or divergent lobe arising from the base of each tarsal

(claw) segment 5

Claws on thoracic legs usually small and without a pad arising
from the base of the tarsal (claw) segment. (If pads occur
they arise from the tibia ) 6

5. Suckerlike lateral protuberances on abdominal segments 2 to 4

or 5 and 8 Acorduleceridae

No suckerlike lateral protuberances on abdominal segments Argidae

6. Antennae with one (or two) segment(s) 7

Antennae with more than two segments 8

1. Since this paper was in proof the larva of an additional hymenopterous family, the
Rhopalosomatidae, has been described (Gurney, Ashley B., 1953, Proc. U. S. Nat. Mus., vol.
103, pp. 19-34). The mature larva would run to couplet 14 in this key, agreeing with the
Vespidae, etc., in having two papillae ( galea and palpus ) on each maxilla. It differs from
these families in the small rather than traverse and slit-shaped salivary opening and in the
presence of a long spine on either side of the salivary opening. The cardo isi not described
and may not be recognizable. The structures which Gurney considered as eye-spots seem
certainly to be something else; possibly they are pigmented spots at the anterior tentorial pits.
The preceding larval stage has wholly different and much reduced mouthparts.

994 The University Science Bulletin

7. Mid-abdominal segments with seven annulets; spiracles definitely

winged Cimbicidae

Mid-abdominal segments with one to four annulets; spiracles not
definitely winged (Fenusinae) Tenthredinidae

8. Antennae never with more than five segments 9

Antennae with six or seven segments. ( Prolegs on all abdominal

segments; setiferous knobs or protuberances on the subanal
areas of the caudal segment) Xyelidae

9. Antennae with three segments, the third peglike and the first and

second incomplete, crescentic, flattened areas; prolegs present
on abdominal segments 2 to 8 and 10 . . ( Diprioninae ) Diprionidae
Antennae with four or five segments, if only three, all are com-
plete or the third is short and not peglike; prolegs present on
abdominal segments 2 to 6, 2 to 7, or 2 to 8 and 10,

Tenthredinidae

10. Thoracic legs mammalike or absent; caudal segment with median

sclerotized process 11

Thoracic legs indistinctly four segmented but without claws;
caudal segment without median sclerotized process. (Prolegs
vestigial on segments 2 to 8 and 10, caudal pair united, form-
ing a single protuberance) ( Phyllotominae ) Tenthredinidae

11. No subanal appendages present; eye-spot not pigmented 12

Small and short subanal appendages present; eye-spot pig-
mented. ( Antennae with four or five segments ) Cephidae

12. Antennae one-segmented; labial palpi one- or two-segmented;

metathoracic spiracles conspicuous Siricidae

Antennae with three or four segments; labial palpi three-seg-
mented; metathoracic spiracles vestigial Xiphydriidae

13. Salivary opening paired, each opening on a projecting process.

Gasteruptionidae ( subfamily Aulacinae ) and Sphecidae ( part )
Salivary opening single (slit-shaped, oval, round) or absent 14

14. Maxillae each with two papillae (galea and palpus). (Cardo

separated from stipes by sclerotic line or at least by a fold ex-
cept in some ants) . . Vespidae, Scoliidae, Pompilidae, Formicidae,1

Chrysididae, and Sphecidae (part)
Maxillae with a single papilla (palpus) or without papillae 15

15. Most conspicuous structures of head, other than the mandibles,

the heavy framework of sclerotic rods which lie on the lines
between the primary structures of mouthparts and front of
head; labium and maxillae not produced; cardo and stipes

separated by sclerotic rod, the former very large 21

Lines between mouthparts and parts of head capsule not or feebly
sclerotic; cardo and stipes fused or nearly so, without a con-
spicuous sclerotic rod between them 16

16. Mandibles and head structures not sclerotized, only visible with

staining Ichneumonidae ( subfamily Collyriinae )

Mandibles and usually other parts of head and mouthparts rec-
ognizably sclerotic 17

1. In ants the galea and maxillary palpus are often represented by mere clusters of
sensillae, or the palpus is represented by a single sensilla in Atta. The peculiar larvae of
the ant Leptanilla (see Wheeler, 1928) does not run properly in this key.

Michener: Comparative Studies of Bee Larvae 995

17. Maxillae and labium projecting as distinct separate lobes, usually

distinguishable to their bases; salivary opening usually a slit. . 20
Maxillae and labium not projecting as separate lobes, either com-
pletely fused and indistinguishable or partly so; salivary open-
ing sometimes a slit, more often oval, round, or absent 18

18. Mandible symmetrical about longitudinal axis, apex tridentate,

middle tooth longest Oryssidae

Mandible asymmetrical, lower or outer tooth usually longer than
others 19

19. Maxillae and labium reduced and usually indistinguishably fused;

maxillary and labial palpi usually unrecognizable. . . . Chalcidoidea
Maxillae and labium distinct apically; maxillary and labial palpi
usually minute Cynipoidea

20. Mandibles with apices simple or bidentate Apoidea

Mandibles with apices tridentate,

Evaniidae, Gasteruptionidae (subfamily Gasteruptioninae )

21. Last abdominal segment ending in two slender hooks; mandibles

large Agriotypidae

Last abdominal segment without hooks; mandibles small 21

22. Accessory longitudinal tracheal commissure present in thorax,

Ichneumonidae ( most )
Accessory longitudinal tracheal commissure of thorax absent,

Braconidae

EVOLUTIONARY TRENDS

With the aid of the data assembled in the preceding sections, it is
possible to determine with some certainty whether some of the
larval characters of bees are primitive or specialized. Thus, since
setae are present on the bodies of the larva of primitive Hymen-
optera, an obvious conclusion would be that setae on the bodies
of bee larvae are primitive. Long and conspicuous antennal papil-
lae and maxillary and labial palpi would be considered primitive
compared to short ones or compared to the absence of these papillae
and palpi, since these structures are several-segmented in the
Chalastogastra and are represented by long and conspicuous papil-
lae in most wasps. Among bees the antennae range from scarcely
detectable convexities on the head capsule to distinct convex areas
each bearing a cylindrical papilla, which may be as much as three
times as long as broad. The palpi may be similarly long, or may
be entirely absent.

The salivary opening, in most wasps, is a transverse slit at the
apex of the labial lobe guarded by a pair of thin, sclerotized lips.
Such a structure is found in vespids, pompilids, chrysidids, and in
some sphecids. In other sphecids the salivary opening is divided,
so that there are actually two such openings, each at the apex of a

996 The University Science Bulletin

projection. This is a specialization of certain sphecids, a condition
not repeated elsewhere among the Hymenoptera except in the
Aulacinae and having no counterpart among the bees. Among bees,
therefore, the slitlike opening guarded by distinct lips would seem
to be the primitive condition. Reduction of the lips, and reduc-
tion of the slit to a straight or crescentic mark or to a small round
hole must be specializations.

The mandibles provide some of the most distinctive characteris-
tics of the various groups of bee larvae. The mandibles of most
sphecoid and vespoid wasps, chrysidids, evaniids, gasteruptionids,
and for that matter those of the Chalastogastra as well, have sev-
eral large sharp apical teeth. The greatest number of such teeth
found in mandibles of bee larvae is two and it seems likely that
acute, rounded, or truncate mandibular apices are derived from the
bidentate type. The bees are peculiar in having, in many cases,
numerous small teeth along the margins, those of the upper margin
forming a band which curves onto the inner surface forming a
"cusp". For bees this condition is considered primitive since it
seems readily derivable from that found in certain sphecoid wasps
in which there is a tooth on the inner surface of the mandible. In
bees the inner surface of the mandible beyond the cusp is often
hollowed out and the hollow margined by a more or less distinct
ridge. This hollow can be seen with the cusp still present in
Nomia, Augochlora, and Melissodes, although the cusp is much re-
duced in the last. In most forms in which this concavity is well
formed the cusp is gone, as are also most or all of the small teeth
of the mandible. Although the concavity is a specialized character,
it is obviously lost, a further specialization, in Trigona and Apis
so that here its absence represents a greater specialization.

In most bees the apices of the mandibles are much narrower than
the bases, often acutely pointed, yet the mandibles are not drawn
out into long attenuate apices. In some, however, the apices are
much attenuate. This may occur among relatives of bees with a
cusp (e.g., Colletes) or as an obvious derivation from forms with
a concavity and no cusp (e. g., Trigona).

Table 1 shows a list of eleven trends with the supposed primitive
alternatives on the right, specialized alternatives on the left. Inter-
mediate conditions occur in all cases, indicating intermediate de-
grees of specialization. All of these trends involve characters which,
from our present meager knowledge, appear to be of generic or
greater importance at least somewhere among the bees.

Michener: Comparative Studies of Bee Larvae

997

Table 1

Some trends involved in evolution of bee larvae, with primitive alterna-
tives at the left, specialized, at the right. Characters are judged as to primi-
tiveness on the basis of the Hymenoptera as a whole.

Primitive

1. Setae present on body

2. Antennal papilla long

3. Maxillary and labial palpi long

4. Maxillae and labium distinct

5. Salivary opening a large slit,

with lips

6. Prementum and postmentum

clearly separate

7. Mandibles with large apical

teeth

8. Mandibles with small teeth

forming a cusp

9. Mandible without defined inner

concavity

10. Apex of mandible rather broad

11. Peritreme present

1.
2.
3.
4.
5.

8.
9.

Specialized

Setae absent on body
Antennal papilla absent
Maxillary and labial palpi absent
Maxillae and labium fused
Salivary opening small, round,

without lips
Prementum and postmentum

indistinguishably fused
Mandible acute, rounded or

truncate
Mandible without cusp

Mandible with clearly defined
inner apical concavity

10. Apex of mandible attenuate

11. Peritreme absent

RELATIVE SPECIALIZATION OF PRINCIPAL TYPES

OF BEE LARVAE

Considering the small number of bee larvae which are adequately
known, it seems futile to attempt to establish a detailed scheme
of relationships or phylogenetic tree based upon their characters.
However, some general conclusions as to the light which the larval
characters shed on relationships can be reached.

No known bee larva agrees with all of the primitive characteristics
listed in table 1, nor is there any which agrees with all of the spe-
cialized features.

In general bee larvae may be divided into two large and inter-
grading groups, as shown below:

Group II

antennal papilla distinct

mandible without cusp, but with apical

inner concavity
salivary opening a long transverse slit

guarded by lips

Group I

antennal papilla reduced or absent
mandible with a multidentate area

or cusp on inner surface
salivary opening reduced, without

lips

In the first group fall the Colletidae, Halictidae, Andrenidae,
Melittidae, and the genera Neopasites, Nomada, Epeolus, and
Triepeohis of the Apidae. From adult characters it seems likely
that Exomalopsis and its relatives would fall here too, were their
larvae known. (Colletes, Hylaeus, Euryglossa, Epeolus, Triepeohis
and one species of Nomadopsis lack the mandibular cusps but the
mandibles are attenuate, showing no resemblance to those of group
II. Other larval characters show the obvious relationship of Epeolus

998 The University Science Bulletin

and Triepeolus to Nomada, which has cusps, and of the Nomadopsis
to another species of the genus which has them.)

In the second group fall the Megachilidae and the Apidae (sense
of Michener, 1944) except for the genera Neopasites, Nomada,
Epeolus, and Triepeolus. (Antennal tubercles are reduced or ab-
sent in Anthophora, although not in the similar larva of Melecta, and
in Melipona, Trigona and Apis, although not in the similar Bombus.
The mandibular concavity is absent in Stelis, although not in the
similar larvae of other megachilids, in Ceratina, although not in the
similar Xylocopa, and in Trigona and Apis although not in the simi-
lar Melipona. The salivary opening is reduced in Ceratina and
Xylocopa.

If table 1 is correct, as it appears to be from a survey of the
characters of the order as a whole, it is obvious that neither the
group I larvae nor the group II larvae can be established as primi-
tive, since group I combines primitive mandibles with specialized
antennae and salivary opening while group II combines specialized
mandibles, with primitive antennae and salivary opening. Making
use of other characters, one finds that labio-maxillary fusion, an
obvious specialization, occurs principally in group I (Colletes,
Panurginae, Nomada, Triepeolus, to a lesser extent in Hesperapis
and the Halictidae ) . Setae on the body surface, presumably primi-
tive, occur only in group II (Megachilidae, AUodape, a few in
Bombus). Large apical mandibular teeth, another primitive char-
acter, are found in both groups but principally in group II.

It is evident that the larvae of Megachilidae exhibit more sup-
posedly primitive characters than those of any other bees, agreeing
with the primitive alternatives of all trends listed in table 1 except
for numbers 8 and 9, which might almost be considered a single
trend. If this conclusion is accepted, then the other bees having
group II larvae are more specialized in larval characters than the
Megachilidae, while the group I larvae have even more specialized
features, some of them agreeing with the specialized alternatives
of most of the trends except numbers 8 and 9 in table 1.

It is scarcely necessary to mention that the combining of primi-
tive and specialized traits is very common, for probably no living
animal is primitive in every character. One may speak of a certain
structure process, or function as primitive in relation to another
structure, process, or function with some degree of safety, but when
one speaks of a primitive organism it must be with the understand-
ing that it has some specialized characters.

Michener: Comparative Studies of Bee Larvae 999

REVERSIBILITY IN EVOLUTION

A comparison of the foregoing with what is known of bee phy-
togeny, based on adult characters (see Michener, 1944) reveals the
curious fact that the group II larvae, i. e. those having a number of
primitive characters, are associated not with the adults which have
numerous primitive characters but with long-tongued adults of
specialized groups. The phytogeny of bees, in its general outlines
( e. g. evolution of the proboscis from a short one like that of wasps
to a much elongated one ) , is so clear and is supported by so many
adult characters that it can scarcely be doubted. Yet the summa-
tion of larval characters suggests an opposite evolutionary direction.

A further study of the larval characters themselves, however,
shows that some of them do fit into the general picture of bee
phytogeny. Those that do so are characters 8 and 9 of table 1.
The way in which these characters have probably evolved will be
shown below.

The remaining characters are all ones in which specialization in-
volves reduction or loss; all structures involved are relatively simple,
for example, papillae and setae. The specialized condition of these
characters in primitive bees could be explained as a series of parallel
developments. Thus, body setae may have been present in all of
the ancestors along the phyletic line leading to the Megachilidae,
and such setae lost independently in each of the stocks which
branched from this phyletic line. The same might have been true
of other structures, such as the antennal papillae.

For at least some of these characters, a different explanation
seems probable, namely that the structures were lost (or reduced)
in very early bee evolution and later regained. The explanation
seems more probable because it avoids the hypothesis of very
numerous parallelisms. Although the "law of irreversibility" in
evolution or Dollo's "law" is well known, there are many excep-
tions to it. Muller (1939) pointed out that there is no genetic
basis for it. Most structures are controlled by many genes so that
exact reversion is most unlikely but it is obvious that where only
one or a few genes are concerned, as might be the case where mere
length of a papilla or palpus is concerned, exact reversion might
occur as a result of reverse mutations. Another possible explana-
tion for the apparent reversions in larval characters of bees may be
that the structures are lost morphologically as a result of inhibiting
factors or complexes but the genetic determinants for the structures
retained. Later, if the inhibition were removed, the structures

1000 The University Science Bulletin

would reappear unless the loci which govern them were altered in
function during the time when the structures were not being pro-
duced.

Thus it seems that while the relatively complicated mandibular
structures do offer characters of easily understood phylogenetic
value, the other structures considered, probably because of rever-
sions, show a reversal of expected phylogenetic significance. If the
hypotheses presented are correct, the primitive and specialized
alternatives for these characters shown in table I should be reversed
for the bees.

AN ACCOUNT OF EVOLUTION AMONG BEE LABVAE

As indicated previously, the larvae of sphecid wasps usually have
mandibles with several large apical teeth. Presumably in asso-
ciation with the change of food from insects or spiders to pollen
and honey, the mandibles of bee larvae became smaller and no
known form retains more than two large apical teeth. The mandi-
bles acquired many small teeth, some of them grouped to form a
cusp on the inner surface some distance from the apex, so that there
is a concave surface between the cusp and the apex. Larvae having
such mandibles are found among the short-tongued families of bees
and as already indicated, in certain genera of Apidae. Presumably
this mandibular structure is in some way correlated with handling
the pollen food for, with some exceptions, all bee larvae have such
a concave space, although it is quite different in nature in some.

The most primitive of the bees would undoubtedly be placed in
the Colletidae. In this family the mother bee applies to the inner
surfaces of the cells a thin transparent or translucent substance.
This seems to take the place of the cocoon spun by the larva in
wasps, for colletids, unlike their presumed ancestors (Sphecidae)
spin no cocoons. It seems likely that the cell lining of the colletids
and the cocoon of the wasps serve for water conservation.

It is noteworthy that among bee larvae as a whole, those which
do spin cocoons have larger antennal papillae and palpi than those
that do not. Of course they also have a large salivary opening
which takes the form of a transverse slit guarded by lips, while
those which do not spin usually have the salivary opening reduced
to a curved slit, an oval, or a circle, and while it is sometimes pro-
duced, it does not have lips. It seems reasonable to suppose that
the antennae and palpi bear the sensillae that enable the larva to
apply its silk properly in cocoon construction. Among forms which
do not spin the palpi are often not so completely reduced as the

Michener: Comparative Studies of Bee Larvae 1001

antennal papillae. Therefore it seems not unlikely that the palpi
also bear sensillae utilized in some other activity, perhaps eating.

Considering the above it is not surprising that the larvae of the
Colletidae have reduced antennal tubercles and palpi and a re-
duced salivary opening ( i. e. they are group I larvae ) , for they
construct no cocoons. In the absence of the need for a cocoon,
there would be mutation pressures against the organs making
cocoon spinning possible with no corresponding selection pres-
sures maintaining these organs. Indeed, because of competition
among the organs for the nutrient materials available, the indi-
viduals without the organs necessary for spinning should be at an
advantage in a mixed population.

Among the Colletidae three of the genera studied (Hylaeus,
Euryglossa and Colletes, apparently also Lonchopria) have the
apices of the mandibles reduced and attenuate, the cusp absent.
Similar attenuation can be found among some of the other bees;
the reason is unknown but a suggestion is that it occurs in groups
whose provisions are usually liquid. Perhaps with such provisions
mandibles are not very effective. That there is no universal cor-
relation between mandibular structure and food consistency is
shown, however, by the fact that a cuckoo bee larva often has quite
different mandibles from the host whose provisions it eats (com-
pare, for example, Triepeolus and Melissodes) .

The Halictidae, Andrenidae and Melittidae, like the Colletidae,
have group I larvae with mandibular cusps and with reduced
antennal papillae, palpi, and salivary openings. The cells are lined
with a very thin layer of wax, so that cocoons are presumably un-
necessary. Systropha, however, is reported to spin a cocoon. It
may be that, like the higher bees, it has reacquired (or retained?)
the necessary organs.

The genus Neopasites has larvae which, in head characters, are
amazingly like certain halictids. Nomada likewise has larvae with
mandibular cusps, reduced salivary opening, etc. Epeolus and
Triepeolus are similar to No?nada except that their mandibles are
attenuate, the cusps gone. All of these genera are parasitic, but
it may be that their pollen-collecting ancestors among the Exo-
malopsini have similar group I larvae. Unfortunately larvae of
Exomalopsini are virtually unknown. Claude-Joseph (1926) briefly
describes and figures the larva of Exomalopsis. Details of mandib-
ular structure are not shown, but the small salivary opening sug-
gests a group I larva. The antennal papillae and palpi seem elon-
gate, however, as in group II larvae. At least some of the more

1002

The University Science Bulletin

ANTHOPHORA

Fig. 1. Inner views of mandibles of various bee larvae showing the develop-
ment of mandibular structure. The vertical series illustrates reduction and loss
of the multidentate cusp and development of the carina surrounding the inner
apical concavity of the mandible. The offshoots to the right illustrate attenua-
tion and reduction of the mandibular apices.

Michener: Comparative Studies of Bee Larvae 1003

primitive Apidae ( sense of Michener, 1944 ) have larvae much more
like those of the short-tongued families than those of the other
Apidae and the Megachilidae. This means that, unless the man-
dibular structure of the type II larva arose twice, the Megachilidae
arose from apid ancestors rather than from preapid ancestors as
indicated by me in 1944. It suggests, furthermore, that Neopasites,
Nomada, Epeolus, etc., together with the Exomalopsini might con-
stitute a subfamily (or family) division co-ordinate with the Xylo-
copinae, Anthophorinae, Apinae, and Fideliinae. This division
would be called the Nomadinae. Its limits are unknown at present.
Further study may indicate adult characters by which it can be
recognized. An alternative explanation of the occurrence of group I
larvae among the parasitic Apidae (Neopasites, Nomada, Epeolus,
Triepeolus) is that these larvae have mandibles, antennae, and palpi
adapted to the type of nest and provisions supplied by their hosts.
This suggestion could explain the characteristics of the first three
genera listed for they are parasites of bees having group I larvae.
Triepeolus, however, as already stated, is a parasite of Melissodes
which has fairly typical group II larvae although remnants of man-
dibular cusps persist.

The larva of Nomada is so similar to those of Epeolus and Trie-
peolus as to lend weight to a conclusion reached from a study of
adults of South American forms, namely that the Nomadini, Osirini,
and Epeolini had a common parasitic ancestor. Details of this will
be published elsewhere.

In certain group I larvae ( e. g. Augochlora, in the Halictidae ) ,
the space between the cusp and the apex of the mandible is strongly
concave and the concavity is bordered by a sharp line or ridge
cutting obliquely across the mandible. In group II larvae (Mega-
chilidae and Apidae except Neopasites, Nomada, Epeolus, and
their relatives) the cusp, and usually the small teeth of the man-
dibular margins, are gone but a distinct inner apical concavity
exists, margined basally by a transverse or oblique line or ridge.
In Melissodes a small remnant of a cusp is retained and the line
margining the concavity is oblique; in others (e.g. Emphor), the
cusp is entirely gone, the line oblique; in most group II larvae,
however, the line is transverse. As among the group I larvae, there
are group II forms in which the mandible becomes attenuate and
loses its characteristics group features. This is true of Ceratina,
Trigona, Apis, and some species of Stelis. In Apis the mandible is
so reduced as to not even be attenuate, as there is no longer a pro-
jecting apex.

1004 The University Science Bulletin

More or less concurrently in phyletic history with the develop-
ment of the type of mandible usual for type II larvae came the
return to cocoon spinning habits. This may be associated with the
fact that many of the bees of this lineage fail to line their cells with
wax or other waterproof material. The Megachilidae, for example,
do not do so. In any event the cocoon spinners among the bees
have, as already indicated, larger antennae and palpi than others
and large salivary openings with lips. It is interesting that, although
obviously a group II larva, cocoon spinning has been abandoned
in Anthophora. Its antennal papillae are reduced, as are its palpi.
Most interestingly, its salivary opening, while of the shape of that
of a typical group II larva, is much reduced in size, moved dorsally
and no longer at the apex of the labium, and is provided with small
nonsclerotic lips.

The larvae of the Xylocopinae are an exception to the rule that
forms with reduced salivary openings also have reduced antennae,
for the antennal papillae are of moderate size in spite of extreme
reduction of salivary openings.

With regard to the setae on the bodies of larvae, it is remarkable
that they are present only on a few of the most specialized of the
groups of bees. Thus, scattered setae occur in the social genus
Bombus, while setae are more numerous on the bodies of at least
some groups of subsocial AUodope, and on megachilids which are
among the most specialized of the solitary bees.

Some of the best specific characters found among bee larvae
occur among the spiracles. Thus Ritcher (1933) was able to dis-
tinguish every species of Bombus larva available to him by means
of the spiracles, and often these were the only specific characters
detected. Although they may provide generic characters, the spi-
racles do not provide much help in the recognition of higher cate-
gories. Their various features suggest an endless series of recom-
binations of a few characteristics. The spines of the atrium, the
spines of the subatrium, the peritreme, the collar, etc., may be large,
small, or absent, each apparently without regard to the condition of
the other structures.

The functional significance, if any, of the spiracular spines and
other spiracular ornamentation is unknown. Possibly these struc-
tures are of value in keeping pollen grains out of the openings. If
so, the variability might be correlated to some extent with vari-
ability in the kinds of pollen utilized by the bees. It is possibly
significant that the only bee larva known to me in which important
differences exist between anterior and posterior spiracles is the first

Michener: Comparative Studies of Bee Larvae 1005

stage larva of Coelioxys, in which the posterior spiracles are much
larger than the others. The first stage larva lives within the rather
liquid provisions of Megachile, not on the surface as does the Mega-
chile and as does the Coelioxys also after the first stage.

The elevations or tubercles of the larval body also vary, like the
spiracular characters, in an apparently meaningless manner. In
general the tubercles seem higher and more consistently present in
group I larvae than in group II larvae, but this is only a tendency.

As is well known (Brauns, 1926; Bayment, 1951) the larvae of
certain Ceratinini (some species of AUodape, Exoneura) have
peculiarly elongate and modified ventrolateral projections on the
anterior part of the body which are used in holding the small quan-
tities of pollen offered the larvae by these bees, which feed the
larvae progressively. Other species of AUodape are provided with
numerous body tubercles arranged in more than the usual series
(dorsolateral and ventrolateral) found in other bee larvae. Ap-
parently these serve to anchor the larvae in groups on the insides
of hollow stems where they are fed progressively.

A long known peculiarity of some parasitic bee larvae is the great
size of the head and mandibles of the first stage larva. It is in this
stage that larvae of these cockoo bees destroy the egg or larva of
their host. This condition occurs in quite unrelated parasites, for
example, Coelioxys (Megachilidae), Triepeohis (Apidae), and
Melecta (Apidae).

SOME GENEBAL PBOBLEMS OF LABVAL
EVOLUTION

Certain general problems concerning larval forms which have
been more or less evident to entomologists for many years are per-
haps worth brief consideration here. The problems involved are
incorporated into the following question: How does it happen
that larvae can evolve in quite different directions and at different
rates from adults, adapting themselves to different food require-
ments, habitats, and the like?

As to rates, it must of course be remembered that what is meant
is rates of phenotypic evolution; there can be no differences in rates
of true or genotypic evolution between larvae and adults.

It should be pointed out that mutations affect processes, and
only indirectly, characters (see Beadle, 1945, 1946). A given
process may influence only early development, only late develop-
ment, or both. A mutation that affects a process influencing only
a certain portion of the development might affect principally larval

1006 The University Science Bulletin

characters, since the cells which will develop into the adult struc-
tures are segregated into primordia which do not develop much
until larval life is well along, or it might affect only adult characters
if its influence is not felt until late in development. A mutation
may affect a process influencing characters of selective value in one
state ( e. g. the larva ) and other characters of little selective value
in another stage. It may influence characters visible phenotypically
and externally in one stage, but not in the other.

It is thus evident that the basis exists for the appearance and
subsequent selection of characters of one stage in the life history,
with little effect on other stages. Of course since larva and adult
are the same individual, selection will depend upon success in both
stages; it cannot act independently on either. Fundamentally the
situation is little different from that which makes it possible for the
young of animals without metamorphosis to have distinctive fea-
tures ( e. g. immature plumages of birds ) different from those of
the adult and subject to their own partially independent evolution-
ary development.

The segregation of larval from adult characters is certainly
exaggerated in holometabolous insects. Possibly this is due in part
to the fact that, at least in Drosophila and Tineola, larval and adult
structures are determined at different times during development.
Thus there is a time when minor injuries to the egg will cause
larval but not adult defects, a later time when they will cause adult
but not larval defects. ( For a brief summary of our knowledge in
this connection, see Wigglesworth, 1950. )

Rapid evolution of one stage, without comparable change in the
other, could easily result and is not fundamentally different from
another common phenomenon, namely rapid evolution of one struc-
ture or group of structures while the others remain little changed.
In this light it is easy to see that larval characters are merely more
characters, of no more systematic value and of no less, than adult
characters.

TECHNIQUES

All studies were based upon larvae preserved in alcohol. This is
best done by dropping live larvae into nearly boiling water for a
few minutes before they are transferred to 70% alcohol. The ma-
terial available, however, having been preserved by various col-
lectors and at different times and in different ways, is in many
different states of preservation and the drawings of whole larvae
are undoubtedly adversely affected by this, for some specimens were

Michexer: Comparative Studies of Bee Larvae 1007

badly shriveled. It should be pointed out also that last stage larvae
vary greatly in shape, size of the tubercles, and the like according
to their age. Two drawings of Emphor larvae are presented to
illustrate a perhaps extreme example of this point. For most species
only a few last stage larvae and these of unknown age, were avail-
able. There is no doubt, however, that in most instances the hiber-
nating form is the one illustrated.

In order to study the mandibles in detail they may be removed
from the head and cleared with caustic, or the whole head treated
in the same manner. Spiracles were studied by removing them with
a bit of adjoining integument and boiling in caustic. The parts so
treated can be stained with acid fuchsin if necessary, and were
examined in drops of glycerin so that, even under high magnification
they could be moved about for study. For preservation such parts
were placed in alcohol in small vials within the larger vial contain-
ing the rest of the larva.

Illustrations of heads were prepared from a study of the whole
head both before and after clearing in caustic.

Illustrations of the mandibles were prepared by removing a man-
dible from the head and examining it in glycerin. Orientation was
by means of the mandibular apodemes and articulations. In the
inner view the adductor apodeme is as nearly as possible directly
above the abductor while in the ventral (or dorsal) view the points
of articulation are one above the other and the apodemes are at
lateral extremities of the mandibular base.

Illustrations of spiracles were prepared with the aid of a com-
pound microscope. Because of the need for frequent reorientation
of the specimens, oil immersion was rarely used. Lateral views of
spiracles are optical sections, while facial views show structures
visible at all levels.

EXTERNAL MORPHOLOGY OF ANTHOPHORA
STANFORDIANA COCKERELL

The following morphological account, based on the mature larva,
is presented in order to provide a basis for the terminology used
elsewhere in this paper and as a contribution to the general fund
of knowledge on insect morphology. Anthophora was selected, not
only because its larvae were available in numbers but because it
is the genus utilized for a comparable study of adult morphology
(see Michener, 1944).

The larva is grublike, robust, curved, with no separation between
the thorax and the abdomen (figure 7). The color is yellowish
34—6490

1008 The University Science Bulletin

white, the spiracular peritremes and the harder parts of the head
brownish, the apices of the mandibles black.

The head (figures 8 to 11) is rounded, that part above (not in-
cluding) the maxillae slightly sclerotic, this sclerotized portion
which may be called the head capsule thickened and slightly in-
flexed along posterior margin. At each lower posterior angle of
the head capsule there is a deep invagination into the posterior
tentorial pit. The thickening along the posterior margin of the
head capsule (termed postocciput by Vance and Smith, 1933) is
directly connected with the posterior tentorial arms and continues
anteriorly as the hypostomal thickening along the lateral margins of
the head capsule to the bases of the mandibles. [These thickenings
are termed habenae by Starke (1949).] From each lower (i.e.
posterior) mandibular articulation the marginal thickening of the
head capsule bends dorsally as the pleurostomal thickening, ex-
tending around the posterior margin of the mandibular corium to
the anterior mandibular articulation where it meets with the thick-
ening which is the internal evidence of the epistomal suture. This
entire thickening (including hypostomal and pleurostomal thicken-
ings) may be called the marginal thickening of the head capsule.
The cleavage lines (see Snodgrass, 1947) or epicranial suture are
weak, A-shaped, the unpaired portion appearing rather thick, the
arms feeble, appearing as scarcely recognizable creases. Each
antenna is a blisterlike convexity with a darkened center. The
parietal bands are ill defined, slightly differentiated areas of cuticle
(termed optic plate by Nelson, 1924, temporal fossa by Vance and
Smith, 1933, and bandarella temporale by Grandi, 1934, etc.). The
epistomal suture, separating the clypeus x from the frons, arches
between the anterior mandibular articulations, which are in deep
pockets or precoilae (Ritcher, 1933) beneath the sides of the cly-
peus. The epistomal suture is feeble medially and is marked in-
ternally by a ridge (likewise weak medially) which is connected
laterally to the marginal thickening of the head capsule. The
anterior tentorial arms arise from this internal ridge above and
mesad of the precoilae. The labroclupeal suture is a mere fold.
The lahrum is a thick projection from the lower margin of the
clypeus which bears apically a pair of tubercles, the lahral tuber-
cles. Beneath the labrum is a transverse slit, the mouth, leading

1. Du Porte (1946) considers this structure as the frontoclypeus. However, I see
no reason for disregarding the evidence provided by muscular attachments emphasized by
Snodgrass (1947), and have therefore followed Snodgrass in the interpretation of the face.

Since this paper was in proof a subsequent paper (Du Porte, E. Melville and R. S. Bige-
low, 1953, Canadian Jour. Zool., vol. 31, pp. 20-29) greatly modifies Du Porte's earlier
( 1946) views.

Michener: Comparative Studies of Bee Larvae 1009

into the foregut, the floor of which, a short distance behind the
mouth, is marked by a transverse fold. Below the mouth is the
small convex hypopharynx. The salivary opening is a transverse
slit, shorter than the mouth, between two thin sclerotized projecting
lips.

The labium, hypopharynx, and to some extent the maxillae form
a single complex of fused or partially fused parts.

For morphological purposes it is assumed that the salivary open-
ing is a suitable landmark lying between the hypopharynx and the
prementum. From a study of Anthophora alone this would not be
questioned, since in this genus the lips of the salivary opening are
in a feeble transverse groove which seems to be the premental-
hypopharyngeal line. In most other bee larvae studied a more
distinct transverse groove exists in about the same position; how-
ever, the salivary opening is more apical in position on the labio-
hypopharyngeal lobe and not in or near the groove. The groove,
here called the hypopharyngeal groove, cannot be the premental-
hypopharyngeal line if the salivary opening is the true landmark.
Nonetheless, for descriptive purposes, the area basal to this groove
is called the hypopharynx, that distal to it the prementum. This is
done because the hypopharynx in this sense is frequently distinc-
tively spiculate while the area beyond the hypopharyngeal groove
is usually like the prementum in surface characteristics. Dobrovsky
(1951) calls the entire region the labium-hypopharynx without
attempting to differentiate the two components. The simpler ex-
pression "labial lobe" has been used in the descriptions which
follow.

The mandibles are distinctly sclerotic, truncate, the inner apical
surface concave, this concavity limited basally by an arcuate carina.
The broad membranous mandibular corium lies between the man-
dibular articulations.

The maxillae are fleshy, with no separation of cardines and
stipites. [Some bees (some megachilids, halictids, and Bombus)
have a small faintly sclerotic area on the outer surface of the maxilla
at its base ( posterior end ) which represents the cardo, and extend-
ing distad from this a weakly sclerotic ribbon described below as
the labio-maxillary rod.] Near the apex of each maxilla is a project-
ing, slightly sclerotic papilla, the maxillary palpus.

Various authors ( e. g. Kellogg, 1902, and Nelson, 1924 ) have cor-
rectly interpreted this structure. Ritcher (1933) in an important
paper on bumblebee larvae, challenges this view and on the basis
of ontogenetic evidence states that the papilla is not the palpus but

1010

The University Science Bulletin

polpus

stipes

palpus -

lacinio

cordo

palpus-

palpus

galea palpus

lacinia

— galea

locinia

Figs. 2-6. 2. Diagram of maxilla of larva of Tenthredinidae. 3. Diagram
of maxilla of larva of Sphecidae. 4. Diagram of maxilla of larva of Antho-
phorinae. 5. Ventral view of right maxilla of larva Bombus americanorum,
showing by broken lines the developing pupal maxilla within. 6. Ventral view
of right maxilla of an individual of Bombus americanorum as it is becoming a
pupa.

is the distal part of the galea. It is clear that he misinterpreted the
evidence before him, however, largely due to the fact that in the
adult and pupal Bombus the galea is a very large structure and the
palpus very small. Grandi (1934a) shows clearly, by a series of
developmental stages, that in Polistes, which has two maxillary
papillae, the outer one is the palpus. In those sphecoid wasps in
which one of the two papillae is reduced in size, it is the inner one;
thus one might presume that the remaining one in the bees would
be the outer one, or palpus. A re-examination of the developmental
stages in Bombus clearly establishes this surmise. As time for pupa-

Michener: Comparative Studies of Bee Larvae 1011

tion approaches, the larval maxilla is seen to contain two projecting
processes, the longest and most slender reaching the larval palpus,
the other shorter (fig. 5 ) and reaching toward the tip of the maxilla
but not toward the palpus. It was perhaps natural for Ritcher to
conclude that the longer process must represent the galea, which
is so long in the pupa and adult. This leaves the inner process un-
explained, however. The obvious conclusion is that the outer
process, in spite of its length, represents the palpus, the inner the
galea, and a weak convexity on the inner surface of the developing
pupal maxilla the lacinia. That this interpretation is correct is
shown in figure 6, a diagram of the maxilla of a pupa which had
recently emerged from the last larval skin. In this individual the
processes can readily be homologized with adult structures, and
almost equally readily, with the processes of the pupal maxilla seen
within the larval maxilla. The phylogenetic series of larval galeae
also supports this interpretation ( see figures 2 to 4 ) .

The upper margin of the maxilla, beneath the hypostomal thick-
ening of the head capsule, is curved inward to form a furrow. Nel-
son (1924) calls this the lateral furrow but it is more accurately
termed the hypostomal furrow. In many aculeate Hymenoptera
there is a sclerotic rod or thickening marking the union between
the maxilla and the postmentum (see Snodgrass, 1935, and Ritcher,
1933). The upper or posterior end of this rod is connected with
the hypostomal thickening. This rod is very feeble in Anthophora,
but is termed the labiomaxillary rod or thickening. This structure
is present in the tenthredinids, as a fragmentum of the stipes which
supports the lacinia, or perhaps as a basal prolongation of the lacinia
itself so that the latter contacts the cardo. Its reduction is clearly
shown in the series of maxillae ( figures 2 to 4 ) .

The labium is divided by a weak transverse furrow into two parts
which, following Snodgrass (1935), are termed the postmentum
and prementum. The prementum bears a pair of apical tubercles
which appear to represent labial palpi (see Ritcher, 1933).

The tentorium in larvae which are not preparing to molt consists
of posterior tentorial arms, united by a tentorial bridge, connected
to the anterior tentorial arms which give rise to the slender dorsal
tentorial arms. As shown in figure 14, the bridge, the dorsal arms,
and the posterior halves of the anterior arms disappear before the
pupal molt, probably before every molt since molting would seem
to be impossible with the tentorium intact. The portions which do
not disappear are more heavily sclerotic than the remainder of the
tentorium.

1012

The University Science Bulletin

caudal annulet

cephalic annulet

spiracle

anal opening

abductor apodeme of mandible dorsal tentorial arm

lobrum
labral tubercle

mouth

mandible

hypopharynx

salivary opening

anterior tentorial arm '
I '

epistomal ridge ] '

antenna /

V X I I

\ \

clypeus \

posterior thickening of head capsule

adductor apodeme of mondible
foregut

tentoriol bridge
solivOTium

maxtllo

hypostomal thickening

premenlum

postmentum

abductor opodeme of mandible

anterior tentoriol arm / W*!* b°nd . , . . . „rm

/ ' dorsal tentorial arm

frons * i /

I /

epistomal suture antenna | /
clypeus \ \

labroclypeal suture
labrum-^. ~~

labral tubercle

mandibular corium —

'/ adductor apodeme of mandible

mandible-

salivary opening -

posterior thickening of head capsule

—posterior tentorial orm
posterior tentorial pit

/ /

moxillory palpus j

premenlum

/ \

postmentum

salivcrium

hypostomal thickening

pleurostomal thickening

Figs. 7-9. Anthophora stanfordiana, mature larva; 7, lateral view; 8, head cut
on saggital line to show internal skeletal structures; 9, lateral view of head.

Michener: Comparative Studies of Bee Larvae 1013

clevage imo

frons

cdductor opodeme of mandible s

epistomol suture \

\ \ \

ciypeus^

obductor opodeme of mandible ^

lobroclypeol suture

labrum — _

labrol tubercle .

labium 7f

po/ielol bond

( '*) ~~J antenna

. — -onterlor tentorial arm

"7 _- — ' anterior tentorial pit
i^- — pleurostomol thickening
mandibular corium
■ — mandible

/ moxillo

maxillory palpus

/ labial palpus

solivory opening

adductor opodeme of mandible —
obductor opodeme of mandible —

mandibular corium —

maxilla
maxillary pa

maxillary palpus
\iHl;li<i- ■iiiw/

poster.or thickening of heed capsule

foramen magnum

foregut

posterior tentorio! arm

postmenfum

solivanum

prementum

mand

lobral tubercle

labial palpus

~ solivory opening

obductor opodeme

adductor opodeme

anterior tentoriol orm

posterior tentorial orm-

antenor tentoriol pit

i position of ontenno

dorsal tentoriol arm

\ posterior tentorial pit

tentorial bridge

Figs. 10-14. Anthophora stanfordiana, mature larva; 10, dorsal view of
head; 11, ventral view of head; 12, inner view of maxilla; 13, inner view of
mandible; 14, dorsal view of tentorium. (Shaded portion disappears before
the molt into pupal stage.)

1014

The University Science Bulletin

There are thirteen postcephalic segments, three thoracic and ten
abdominal. The intersegmental lines between the thoracic seg-
ments are absent laterally; they are weakened laterally on the
abdomen. The tergal regions of most of the segments are divided
by a transverse furrow into a cephalic annulet which is transversely
wrinkled and a caudal annulet which is not wrinkled and is convex
dorsolateral^, forming very weak dorsolateral tubercles. It is pro-
duced in some bee larvae to form distinct tubercles. In others there
are tubercles below the line of spiracles, termed ventrolateral tu-

otriai spines
i \

subatrium

primary tracheal opening

Fig. 15. Longitudinal sectional view of spiracle of Anthophora stanfordiana.

bercles. The last body segment bears an arcuate slit, the anal
opening.

There are ten pairs of spiracles, two thoracic and eight abdominal,
all round. The primary tracheal opening lies between the cup-
shaped, heavy-walled atrium and the annulated subatrium which
extends inward to a sharp constriction in the trachea. The primary
tracheal opening is guarded by long serrate or toothed spines form-
ing a collar. The atrium is lined with rows of small spines arising
from ridges. These spines together with the collar spines are called
atrial spines. The walls of the atrim extend above the general body
surface and flare slightly to form a sharp rim. The transparent
peritreme extends inward from the rim, narrowing the atrial opening.

Michener: Comparative Studies of Bee Larvae 1015

DESCRIPTIVE ACCOUNTS

Family Colletidae

This family contains rather diverse elements, both on the basis
of larval and of adult structures. As indicated elsewhere, the larvae
lack certain primitive features which they would be expected to
exhibit in view of the primitive mouthparts of the adults. Out-
standing among these is the reduction of the salivary opening from
a straight transverse slit guarded by sclerotic lips to small snout-
like projection (Caupolicana and Policana), a small curved slit
without lips (Hylaeus and Euryglossa), or an apparently closed
scar (Colletes). This reduction is doubtless correlated with the
fact that these larvae spin no cocoons. Presumably it is unnecessary
for them to do so, since their cells are lined with a smooth trans-
parent material secreted by the adult female. Perhaps for this
reason the antennal papillae are reduced, another specialized fea-
ture. Mandibles of colletids do not show the large apical teeth
which characterize wasp larvae but are instead acute in known
forms. Those of Caupolicana and Policana have the subapical mul-
tidentate inner cusp which appears to be a primitive feature among
bee larvae but the others lack the cusp and have the mandibles
apically attenuate. In a number of other features also Caupolicana
and Policana show a close relationship. (For example, the sub-
apical maxillary palpus, the short labrum compared to the clypeus,
and the spinose atrium and subatrium. ) For these reasons I ap-
prove of the relationships of these groups indicated in Moure's
(1945) classification based upon the adults.

The relationship of Euryglossa and Hylaeus, evident also in adults,
is supported by the similar mandibles and salivary openings of the
larvae and especially by the spiracles, which in these two genera
differ from other colletids studied in having atria produced above
the body surface.

Unfortunately larvae contribute nothing to the question of the
relationships between the Hylaeinae and Euryglossinae on the one
hand and the Colletinae, Paracolletinae, etc. on the other. These
groups are held together as the family Colletidae almost solely
because of the primitive wasplike mouthparts of the adults. There
seems to be no group of characters which will unite the larvae of the
Colletidae and distinguish them from the Halictidae and Andreni-
dae; neither are there characters which show that any of the colletids
are closely related to any other bees and that the family should be
split.

1016

The University Science Bulletin

Figs. 16-26. 16, Euryglossa fasciatella, larva; 17, HyJaeus sp?, larva; 18, lat-
eral view of head of same; 19, dorsal view of head of same; 20, 21, 22, ventral,
dorsal and inner views of mandible of same; 23, 24, inner and ventral views of
mandible of Euryglossa fasciatella; 25, spiracle of same; 26, spiracle of Hylaeus
sp.

Hylaeus sp?

(Figs. 17-22, 26)

The larva is rather slender, with the intersegmental lines con-
spicuous ventrally but indicated only by broad concavities dorsally
and laterally. The annulets of the segments are scarcely separated
from one another. Feeble dorsolateral tubercles are present on the

Michener: Comparative Studies of Bee Larvae 1017

dorsum high above the spiracles; except for these tubercles the body
is faintly transversely wrinkled. Rounded ventrolateral tubercles
are feebly in evidence.

Head capsule and mouthparts without setae; labrum and hypo-
pharynx finely spiculate; head capsule weakly sclerotized without
much constriction between head and thorax. Marginal thickening
of head capsule very weak, posterior tentorial pits small and ten-
torial arms slender; epistomal suture weak, its position indicated
by a depression arching between bases of mandibles; cleavage line
(and dorsal longitudinal median thickening of head capsule)
scarcely evident. Antennae mere convexities; clypeolabral suture
weak, labrum with small labral tubercles, as in Anthophora. Man-
dibles with attenuate, pointed, apices, their upper margins with a
row of small teeth. Maxillae with apices not bent inward, maxillary
palpi more than twice as long as broad; hypostomal furrow shallow;
salivary opening small, crescentic, at apex of labial lobe; labial palpi
longer than broad, weak furrow between postmentum and premen-
tum.

Body without spicules or setae. Spiracular atria without spines,
its walls projecting slightly beyond body surface, peritreme flat,
primary spiracular opening narrowed by a collar.

Mineral King, Tulare County, California, August, 1939 (Bohart).

Hylaeus variegata (Fabricius)

Stockhert ( 1922 ) describes the larva of this species briefly. So
far as can be determined, it is similar to that described above,
although the dorsolateral tubercles are probably sharper.

EnrygJossa fasciatella Cockerell

(Figs. 16, 23-25)

This larva is rather slender, similar in many ways to that of
Hylaeus, with the intersegmental lines conspicuous ventrally, rather
weak dorsally. The dorsolateral tubercles are much higher than in
Hylaeus and are transverse but reach neither the middorsal line
nor the level of the spiracles. The ventrolateral tubercles are
rather distinct, small and rounded.

Head capsule and mouthparts without setae, head capsule slightly
more sclerotized than in Hylaeus and less broadly attached to
thorax. Marginal thickening of head capsule present although
weak, posterior tentorial pits more conspicuous than in Hylaeus.
Epistomal suture indicated by a shallow trough arched between
bases of mandibles, thickening apparent in this trough only later-

1018 The University Science Bulletin

ally; antennae indicated by weak convexities; labroclypeal suture
indicated by distinct furrow, lower margin of labrum beneath the
small labral tubercles with a band of minute setae. Mandibles
pointed, more robust than in Hylaeus, with sharp serrate ridge on
inner surface apically. Maxillary palpi conical, about as long as
broad; salivary opening not clearly recognizable, not provided with
produced lips; labial palpi small, rather close together, conical,
broader than long; furrow between postmentum and prementum
inconspicuous.

Body without spicules or setae. Spiracles with atrium without
spines (but with broken ridges), projecting beyond body surface,
peritreme flat, primary tracheal opening narrowed by collar.

A single specimen is available without locality data but marked
14 months old. It is from the collection of the U. S. National
Museum.

Chilicola spp.

Claude-Joseph (1926) describes and figures the larvae of C
inermis Friese and C. friesei Herbst. His brief accounts, and figures
which show few details, indicate a larva similar to that of Hylaeus.

Colletes fulgidus Swenk

(Figs. 27, 30-33, 35)

The larva is similar to Hylaeus, a little more robust, with the
dorsolateral tubercles large though broad and rounded, extending
downward nearly to the level of the spiracles. The ventrolateral
tubercles are absent.

Head capsule as described for Hylaeus but tentorium virtually
absent,* epistomal suture represented by thickening of cuticle
arching between mandibular bases, labroclypeal suture weak,
strongly arched. Antennae represented by distinct swellings. Man-
dibles with attenuate apices finely serrate on both margins. Sali-
vary opening apparently closed but duct present; a furrow separates
hypopharynx from upper margin of prementum.

Spiracular atria shallow, without spines, with a few broken cir-
cular ridges; peritreme large; primary tracheal opening wide but
provided with a small collar.

Bodega Bay, Sonoma County, California, January 14, 1948 ( Mac-
Swain) and Montara, San Mateo County, California, September,
1940 (MacSwain).

* It is presumably present in feeding stages but, as indicated in the section on Antho-
phora morphology, the tentorium largely disappears before a molt.

Michener: Comparative Studies of Bee Larvae 1019

Figs. 27-35. 27, Colletes fulgidus, larva; 28, 29, inner and ventral views of
mandible of Colletes araucariae; 30, 31, inner and ventral views of mandible of
Colletes fulgidus; 32, 33, lateral and dorsal views of head of same; 34, spiracle
of Colletes araucariae; 35, spiracle of Colletes fulgidus.

1020 The University Science Bulletin

Colletes araucariae Friese

(Figs. 28, 29, 34)

The larva, first described by Claude-Joseph ( 1926 ) , is similar to
that of C. fulgidus but the labral tubercles more pronounced; the
mandibles are longer, crossing near their apices, and minutely spicu-
late apically on the inner surfaces. The spiracles have atria which
are broader and shallower, with more broken internal ridges. The
peritreme is smaller. There is no distinct collar around the primary
tracheal opening. The subatrium is very short.

Correo Nunoa, Chile (Claude-Joseph) (one specimen).

Colletes succinctus ( Linnaeus )

Mayet (1875) described and figured the larva of this species. His
figures are not very detailed, but there are no obvious differences
indicated between his species and C. fulgidus.

Colletes spp.

Claude-Joseph ( 1926 ) describes and figures larvae of C. laticeps
Friese and C. ciliatus Friese. The illustrations of the heads are not
detailed but show structures not widely different from those of
C. fulgidus. The body apparently bears no tubercles in C. laticeps
and only ventrolateral tubercles in C. ciliatus.

Lonchopria marginata Spinola

The larva of this species, described and figured by Claude-Joseph
(1926), seems similar to that of Colletes, at least in its attenuate
mandibular apices. The body of the larva apparently lacks all evi-
dence of tubercles.

Policana herbsti Friese

(Figs. 36-38, 44-46)

The larva first described by Claude-Joseph (1926) is rather ro-
bust, strongly tuberculate, with the dorsolateral tubercles transverse,
nearly meeting on the middorsal line, not extending downward to
the level of the spiracles at the sides. The ventrolateral tubercles
are conspicuous, rounded, present on the first eight abdominal seg-
ments, those of the first small. The intersegmental lines are deep,
strongly constricted dorsally and ventrally, but weak laterally. The
lines between the cephalic and caudal annulets of the segments are
absent except as they mark the anterior margins of the dorsolateral
tubercles, especially on the second to the fifth abdominal segments.

Head capsule more heavily sclerotized than in Colletes, marginal

Michener: Comparative Studies of Bee Larvae 1021

Figs. 36-47. 36, Policana herbsti, larva; 37, 38, lateral and dorsal views of
head of same; 39, Campolicana gayi, larva; 40, 41, lateral and dorsal views of
head of same; 42, 43, ventral and inner views of mandible of same; 44, 45,
ventral and inner views of mandible of Policana herbsti; 46, spiracle of same;
47, spiracle of Campolicana gayi.

1022 The University Science Bulletin

thickening distinct, posterior tentorial pits distinct and tentorial
arms presumably not as slender as in Colletes, epistomal suture as
in Hylaeus, that is weak but its position indicated by a depression
arching between bases of mandibles; unpaired portion of cleavage
line visible but arms of cleavage line absent, dorsal longitudinal
median thickening of head capsule not evident. Antennae repre-
sented by distinct convexities; clypeolabral suture distinct, slightly
arched upward medially; labrum with minute setae distally, labral
tubercles large and blunt. Apices of mandibles not attenuate and
sharply pointed as in Hylaeus and Colletes but broader, with inner
apical concavity poorly defined, dorsal margin strongly toothed,
this row of teeth continuous with those of mandibular cusp; con-
cavity with a few very small teeth; dorsal surface of mandible spicu-
late. Maxillae with apices not bent inward, maxillary palpi near
apices, about twice as long as broad; hypostomal furrow shallow:
salivary opening being near apex of labial lobe, the lobe as a whole
being very thick and considerably projecting; salivary opening nar-
row, only about one fourth as broad as distance between labial
palpi, but lips of the opening exceedingly long, forming a projection
that is nearly as long as wide and is distinctly longer than the labial
palpi; labial palpi considerably longer than broad; labium with a
distinct furrow between postmentum and prementum; outer sur-
faces of mouthparts with scattered minute setae, labiomaxillary
thickening rather distinct.

Body without spicules or setae. Spiracles with atrium shallow,
not projecting above level of body surface, with minutely spinose
broken ridges within; peritreme flat; subatrium spinose; primary
tracheal opening not narrowed by collar.

Correo Nunoa, Chile (Claude-Joseph) (one specimen).

Policana occidentalis (Haliday)

According to Claude-Joseph ( 1926 ) the larva is similar to that of
P. herbsti. His figures of both species, doubtless made from life,
show larval forms very different from that shown in the present
paper and based on one of Claude-Joseph's preserved specimens of
P. herbsti. He shows no dorsal tubercles, but very strong ventral
convolutions for both species.

Caupolicana gayi Spinola

(Figs. 39-43, 47)

The larva first described by Claude-Joseph ( 1926 ) is rather robust
with the intersegmental furrows quite deep, the dorsolateral tuber-

Michener: Comparative Studies of Bee Larvae 1023

cles high, transverse, nearly meeting middorsally and on the second
and following abdominal segments extending down to the level of
the spiracles, these elevated tubercles arising abruptly from the
surface of the body. The lines between the cephalic and caudal
annulets of the body segments are absent unless they are indicated
by the abrupt anterior faces of the dorsolateral tubercles. Folds
and slight elevations below these tubercles indicate ventrolateral
tubercles on the third to the ninth abdominal segments.

Head capsule rather distinctly sclerotized, its marginal thickening
distinct; posterior tentorial pits rather large and tentorial arms pre-
sumably not as slender as in Hylaeus; epistomal suture not easily
recognizable, its position indicated by a depression arching between
bases of mandibles, a slight broad thickening of cuticle probably
following this depression; unpaired portion of cleavage line present
but arms feeble or absent and dorsal longitudinal median thickening
of head capsule absent. Antennae present as distinct convexities;
labroclypeal suture distinct, feebly arched; labrum with rather
numerous spicules, some of them long and setalike, tubercles large
and broad. Mandibles much as in Policana but with fewer teeth,
no spicules, and a very strong toothed cusp, a distinct untoothed
space between its teeth and teeth of upper apical margin; median
portions of mandibles hidden from view in repose by overlapping
labral tubercles which practically contact maxillae. Maxillae with
long spicules, apices bent inward and downward, reaching labial
palpi; maxillary palpi not apical, more than twice as long as broad;
labial lobe very thick and elongate; salivary opening at apex of
labial lobe, almost circular, on a projecting snoutlike process, the
apex of which is dark; labial palpi about twice as long as broad;
labium with distinct furrow between postmentum and prementum.

Body without spicules or setae. Spiracles with atrium shallow,
not projecting above level of body surface, with spinose ridges
within, flat peritreme, weak collar at primary tracheal opening;
subatrium spinose within.

Correo Nurioa, Chile ( Claude-Joseph ) ( one specimen ) .

Family Halictidae

This family contains a relatively homogenous group of larvae.
Like those of the Colletidae, they combine certain primitive fea-
tures with features specialized by reduction. For example the
salivary opening in all known cases is a curved slit, provided with
no projecting lips. Such forms do not spin cocoons. However,
Systropha is reported to spin cocoons, and it is therefore presumed

35—6490

1024

The University Science Bulletin

that it has a transverse salivary opening with lips, as with other
cocoon-spinning bees and wasps. As in the Colletidae, the antennal
papillae are insignificant or absent. The palpi are reduced in
halictid larvae, even absent in some. The mandibles are of what is
believed to be the most primitive type found in bees, that is, with

Figs. 48-56. 48, Lasioglossum kincaidii, larva; 49, 50, inner and ventral views
of mandible of same; 51, 52, lateral and dorsal views of head of same; 53,
spiracle of same; 54, 55, inner and ventral views of mandible of Lasioglossum
spar sum; 56, spiracle of same.

Michener: Comparative Studies of Bee Larvae 1025

two large apical teeth (except in Augochlora and some Nomia)
and with a multidentate cusp on the inner surface, the region be-
tween this cusp and the apex concave.

The close relationship of Lasioglossum and Halictus, obvious in
any event from the adults, is shown in the larvae also by the basal
obliteration of the lines between the labium and the maxillae, by
the shallow spiracular atria, not projecting above the body surface,
with their spines minute or absent, and by the peritremes which are
often incomplete or absent.

The great diversity of species of Nomia, likewise evident from
adults, is shown among the larvae, which, for example, may have
acute or bidentate mandibular apices, spinose or spineless spiracular
atria, etc.

Claude-Joseph ( 1926 ) describes the larvae of a number of halic-
tine bees from Chile. So far as can be determined from his de-
scriptions and figures, all agree rather well with the better known
members of this group.

Lasioglossum (Evylaeus) kincaidii (Cockerell)

(Figs. 48-53)

The larva is rather robust with the intersegmental lines distinct
all around the body but with no clear division of the segments into
cephalic and caudal annulets. The dorsolateral tubercles are con-
spicuous and transverse, nearly meeting on the middorsal line and
extending downward on the sides nearly to the level of the spiracles.
Only on the last two or three abdominal segments are the dorso-
lateral tubercles low and rather inconspicuous. They are highest
and most sharply ridged on the thoracic segments.

Head capsule without setae, with spicules in temporal regions,
rather distinctly sclerotized, separated by distinct constriction from
thorax. Marginal thickening of head capsule weak; posterior ten-
torial pits inconspicuous from the outside but represented internally
by short and rather robust tentorial arms similar to those found in
specimens of Anthophora approaching the pupal stage; head with
two large rounded convexities above the level of the antennae so
that seen in lateral view it seems to have a "high forehead"; episto-
mal suture nearly absent between anterior tentorial pits, represented
by shallow, transverse depression, not indicated by an internal thick-
ening; cleavage line and dorsal longitudinal median thickening of
head capsule absent or nearly so. Antennae clearly defined, broadly
conical; labroclypeal suture weak, scarcely arched; labrum with a
pair of large tubercles rather near to one another, bearing minute

1026 The University Science Bulletin

setae arising from large sockets; sides of clypeus spiculate. Man-
dibles each with large tubercle on outer surface midway between
base and apex, this tubercle bearing one or two minute setae; apical
portions of mandibles attenuate, ending in two teeth; margins den-
tate; cusp multidentate, numerous small teeth basad from cusp.
Maxillae with apices not bent inward, maxillary palpi absent, a very
few scattered minute setae on outer surface of each maxilla in some
specimens; labial lobe scarcely exceeding maxillae; salivary opening
minute and inconspicuous, not marked by lips or other projecting
structures, located near apex of labial lobe; labial palpi absent; no
distinct furrow between postmentum and prementum.

Body with spicules on some areas. Spiracles rather shallow, with
broken rows of small spines; peritreme flat, incomplete; primary
tracheal opening without collar.

Montara, San Mateo County, California, June 12, 1940 (Mac-
Swain ) .

Lasioglosswn (Evylaeus) malachurus (Kirby)

Soika (1934) has described and figured the larva of this species.
It is similar to that of Lasioglosswn kincaidii but the antennae are
lower and flatter. The mandibles are more attenuate than in
L. kincaidii, bidentate at the apices, with one tooth much shorter
than the other, with a distinct multidentate cusp, and apparently
without small teeth on the margins of the mandibles. As with
L. kincaidii, the palpi are absent and the maxillae are blunt and
rounded.

Lasioglossum {Chloralictus) sp?

(Figs. 62, 63, 65)

This species is similar to L. kincaidii in general structure but
differs in numerous details. The dorsolateral tubercles are much
lower and less conspicuous than in L. kincaidii although transverse
as in that species. The head capsule is white and apparently not
at all sclerotized, the convexities above the antennae are lower and
more rounded than in L. kincaidii. The antennae are lower and
blunter than in L. kincaidii and the labral tubercles are mere broad
prominences. The mandibles are fundamentally as in kincaidii but
the attenuate apical portions are shorter and do not have teeth on
their upper margins, the lower margins have more teeth, the tubercle
on the outer surface of each mandible is weaker than in L. kincaidii.
The spicules and setae on the head and body are apparently absent.
The spiracles lack atrial spines and teeth, although the inner sur-

Michener: Comparative Studies of Bee Larvae 1027

face of the atrium is marked with curved lines; the peritreme is
absent.

Arlington, Virginia, collected under bark.

Larvae of another unidentified species of Chloralictus from North
Carolina were essentially similar to the form described above.

Lasioglossum (Chloralictus) sparsum (Robertson)

(Figs. 54-56)

Similar to the above described species of Chloralictus. The man-
dibular structure differs slightly, however, in the greatly reduced
number of teeth on the lower margin of the mandible, the more
transverse cusp, perhaps with heavier teeth, and the sharper apical
teeth. The spiracles differ in the presence of rows of teeth on the
inner surface of the atrium, the presence of a peritreme and of a
collar around the primary tracheal opening.

Lawrence, Kansas, April 24, 1951 (Michener).

Halictus (Halictus) sexcinctus (Fabricius)

Grandi (1937) has described 'and figured the larva of this species.

It is similar to that of Lasioglossum kincaidii but the antennae
are lower and flatter and the mandibular dentation appears different
in details. ( The cusp is not clearly shown although it is apparently
present.) The palpi are absent. The spiracular atria evidently
contain many more teeth than in L. kincaidii.

Packard ( 1897 ) gives brief descriptions of the larvae of Halictus
(Halictus) ligatus Say and H. (H.) parallelus Say. They seem to
agree in major features with the halictine bees described herein.

Halictus (Seladonia) tripartitus Cockerell

(Figs. 60, 61, 66, 67)

Larva similar to that of Lasioglossum kincaidii from which it
differs in minor features such as the larger number of teeth on the
lower margin of the mandible, the more slender mandibular apex
with the ventral apical tooth longer than the dorsal one, and the
larger number of seta-bearing tubercles on the outer surfaces of the
mandibles. The spiracular atrium is marked with anastomosing
circular ridges within, these ridges sometimes bearing numerous
small spines. The peritreme is absent and the primary tracheal
opening is narrowed by a small collar.

The presence or absence of atrial spines seems erratic and mi-
correlated with body segments.

Chino, Arizona, July 4, 1950 ( R. H. Beamer, J. G. Rozen ) .

1028

The University Science Bulletin

Augochlora (Augochlora) pura (Say)

(Figs. 57-59, 64)

The larva is similar in most respects to that of Lasioglossum kin-
caidii. It differs in the smaller and lower thoracic tubercles, so
that these are lower than the abdominal tubercles instead of higher
as in L. kincaidii, and in the presence of ventrolateral tubercles on
the second and following abdominal segments. The mandibles

Figs. 57-67. 57, Augochlora pura, larva; 58, 59, inner and ventral views of
mandible of same; 60, 61, ventral and inner views of mandible of Halictus
tripartitus; 62, 63, ventral and inner views of mandible of Lasioglossum ( Chlora-
lictus) sp?; 64, spiracle of Augochlora pura; 65, spiracle of Lasioglossum
(Chloralictus) sp?; 66, 67, spiracles of Halictus tripartitus.

Michener: Comparative Studies of Bee Larvae 1029

bear several setae on their outer surfaces; the apices of the mandibles
are simple and acute, the ventral margin not serrate, the cusp pro-
duced as a slender projection which may end in two large teeth
instead of the simple apex shown in the figure. The presence of a
distinct curved line outlining the distal mandibular concavity and
extending from the cusp to a point on the ventral margin of the
mandible is distinctive. The spiracles have the atrium slightly pro-
jecting above the body surface, its inner surface with rows of minute
spines; the peritereme is flat. The primary tracheal opening is nar-
rowed by a very small collar.

Short Mountain, Shenandoah, Virginia, June 6, 1941 (A. B. Gur-
ney); Lawrence, Kansas (Michener); York County, Pennsylvania,
June 16, 1951; Sand Gap, Kentucky, August 30, 1946 (Ritcher).

Among specimens presumably from a single nest from York
County, Pennsylvania, considerable variability occurs in details of
mandibular dentition. For example, in one specimen the cusp ends
in two distinct darkened teeth, while in the other the teeth of the
cusp are all small, as shown in the figure.

Nomia (Epinomia) nevadensis nevadensis Cresson

(Figs. 68, 71, 72, 76)

The larva is similar to that of Lasioglossum in many respects. The
intersegmental lines are distinct except laterally while the lines be-
tween cephalic and caudal annulets of the segments are absent.
The dorsolateral tubercles of the body are high and pointed, those
of the thorax forming distinct transverse ridges but those of the
abdomen not transverse but conical; these tubercles do not extend
down toward the level of the spiracles. The ventrolateral tubercles
are absent.

Head capsule and outer surfaces of mouthparts with scattered
minute setae, the former rather distinctly sclerotized, separated
from thorax by a distinct constriction. Marginal thickening of head
capsule narrow; cleavage line and dorsal longitudinal median thick-
ening of head capsule absent; head with a rounded convexity on
each side above antennae as in Lasioglossum and with a broad
longitudinal median depression of the dorsal surface extending
downward to level of anterior tentorial pits. Antennae distinct,
rounded conical; epistomal suture absent but its position roughly
indicated by a transverse depression between the anterior tentorial
pits, clypeolabral suture weak, the labrum with small apical tu-
bercles. Mandibles robust, with large tubercle on outer surface of
each, apices deeply bifid, cusp with many teeth of varying sizes;

1030

The University Science Bulletin

lower margin of mandible serrate subapically. Maxillae with apices
not bent inward; maxillary palpi robust, longer than broad; hy-
postomal furrow rather shallow; labial lobe exceeded by maxilla;
labial palpi much broader than long; salivary opening small, near
apex of labial lobe, not projecting, but about as wide as projection
in Nomia melandri.

Figs. 68-76. 68, Nomia nevadensis, larva; 69, 70, lateral and dorsal views of
head of Nomia melandri; 71, 72, inner and ventral views of mandible of Nomia
nevadensis; 73, 74, inner and ventral views of mandible of Nomia melandri;
75, spiracle of same; 76, spiracle of Nomia nevadensis.

Michener: Comparative Studies of Bee Larvae 1031

Body without spicules or setae; spiracles with thick-walled atria
bearing broken irregular ridges within, atria projecting slightly
above body surface and with rim at upper extremity; peretreme
nearly flat; primary tracheal opening narrowed by large collar.

Blythe, California, July 25, 1945 (Linsley and MacSwain).

Nomia (Acunomia) melandri Cockerell

(Figs. 69, 70, 73-75)

This species is similar to N. nevadensis but differs in having the
dorsolateral tubercles slightly lower, those of the abdominal seg-
ments somewhat transverse, more like those of the thorax although
not so strongly so as those of the thorax. The longitudinal median
depression in the head capsule is less conspicuous than in N. neva-
densis. The apices of the mandibles not bidentate but acute, the
cusp with fewer teeth. The maxillary palpi are shorter than in
nevadensis, being pigmented but only about as long as broad; the
salivary opening is on a small but distinct pigmented prominence.
There are distinct spicules on some areas of the body. The spiracles
are more heavily pigmented than in nevadensis, the atrium much
narrowed near the surface, its inner surface spinose. The atrium
projects slightly above the body surface and provided with a rim.
The peritreme is flat. The primary tracheal opening is narrowed
by a collar.

Fillmore, Utah, June 29, 1950 (Michener).

Nomia ruficornis Spinola

The larva of this species is described and figured by Soika ( 1932 ) .
It differs from N. nevadensis in not being gently curved but rather
being abruptly bent in the vicinity of the base of the abdomen.
The dorsolateral tubercles are larger, although not higher, than in
N. nevadensis, largest on the last thoracic and first two abdominal
segments. The antennae are low and rounded, much less pro-
tuberant than in N. melandri. The labrum has slightly more con-
spicuous labral tubercles than does N. melandri. The mandibles
are simple, not bidentate at their apices, and each has a multiden-
tate cusp, apparently with more teeth than in N. melandri. Each
also has a clearly defined inner apical concavity as in our species of
Nomia, and the surface of this concavity bears small scattered teeth.
The maxillary palpi are longer than broad, apical, not slightly sub-
apical as in N. melandri. The labial palpi are about as broad as
long.

1032

The University Science Bulletin

Figs. 77-85. 77, 78, Dorsal and lateral views of head of Andrena sp.; 79,
larva of same; 80, 81, ventral and inner views of mandible of same; 82, 83,
ventral and inner views of mandible of Andrena complexa; 84, spiracle of
Andrena sp.; 85, spiracle of Andrena complexa.

Michener: Comparative Studies of Bee Larvae 1033

Family Andrenidae

Like the last family, this one is relatively homogenous in larval
characters. Indeed most of its characters are shared with the
Halictidae. Like that family and the Colletidae, larvae of the
Andrenidae exhibit a mixture of primitive and specialized features.
Thus the salivary opening is reduced to a curved slit with no lips
and the antennal papillae are insignificant or absent. The palpi are
small, but on the basis of the small number of species examined, are
less often absent than in the Halictidae. Very often the maxillary
palpi are strikingly larger than the labial palpi. The mandibles
are acute (or sometimes bidentate) at their apices and exhibit the
presumably primitive feature of a multidentate cusp, sometimes
reduced to a few small teeth (see Nomadopsis fracta). The region
between this cusp and the mandibular apex is distinctly concave
in the genus Andrena but not in the Panurginae. In all species ex-
amined the spiracular atria lack spines. The atrial walls may
project above the body surface or not, even within the genus
Andrena.

As shown by the descriptions and figures, the Panurginae have
particularly large dorsolateral tubercles which are ordinarily conical,
not transverse as in most groups. These tubercles reach maximum
length in Perdita zebrata ( Cresson ) ( see Custer, 1929. )

Andrena sp.

(Figs. 77-81, 84)

The larva is similar to that of Lasioglossum kincaidii in many
respects. The intersegmental furrows are distinct and faint lines
between cephalic and caudal annulets of the abdominal segments
are visible. The dorsolateral tubercles are conspicuous, particu-
larly on the thorax where they are high and rounded; on the ab-
domen they are lower and more transverse, reaching downward
almost to the level of the spiracles. The ventrolateral tubercles are
absent.

Head capsule and mouthparts with a few minute setae; head cap-
sule shaped much as in Lasioglossum with a rounded elevation
above each antenna and with the antennae themselves arising from
rounded prominences. Marginal thickening of head capsule feeble
and inconspicuous but posterior tentorial pits distinct; epistomal
suture indicated by a weak depression, no thickening between the
anterior tentorial pits; cleavage line and longitudinal median thick-
ening of head capsule weak. Antennae represented by convexities;

1034

The University Science Bulletin

labroclypeal suture weak; labrum with two rather large tubercles.
Mandibles with apices attenuate and sharply pointed, upper and
lower margins finely serrate apically, the serrations continued along
the margin to the densely toothed cusp. Maxillae with apices not
bent inward; maxillary palpi large, slightly longer than broad; labial
lobe similar to that of Lasioglossum but with minute labial palpi.

Body without setae, with spicules on some areas ventrally. Spi-
racular atria with internal ridges; atrium not projecting above body
surface; peritreme present; collar around primary tracheal opening
present.

Tesla, San Joaquin County, California, May 23, 1946 (MacSwain).

Andrena (Ptilandrena) complexa Viereck

(Figs. 82, 83, 85)

The larva of this species seems to differ from that of A. sp? de-
scribed above in the somewhat larger dorsolateral tubercles of the
body segments and in having the spiracular atria projecting above
the body surface and the peritreme flat.

Berkeley, California (MacSwain).

Figs. 86-91. 86, 87, Inner and ventral views of mandible of Panurginus
melanocephalus; 88, spiracle of same; 89, 90, inner and ventral views of mandi-
ble of Andrena erythronii; 91, spiracle of same.

Michener: Comparative Studies of Bee Larvae 1035

Packard ( 1897 ) gives a brief description of the larva of Andrena
vicina Smith. It seems to agree in major features with the species
described herein.

Andrena (Leucandrena) erythronii Robertson

(Figs. 89-91)

The larva is similar to that of Andrena sp? described above but
has more distinct dorsolateral tubercles all along the body so that
those of the thorax are only slightly larger than those of the abdo-
men. The details of mandibular shape and dentition differ slightly
from those of Andrena sp?. The spiracular atrium is much deeper,
more constricted toward the surface, and projects above the body
surface.

Lawrence, Kansas, May 11, 1952 (Michener).

Panurgus banksianus (Kirby)

The larva is described and figured by Micheli (1931). In body
form it somewhat resembles Andrena, but the lines between the
cephalic and caudal annulets of the body segments are distinct on
all segments, and the caudal annulets are swollen to form large
dorsolateral tubercles, largest on the middle segments of the body,
and extending downward almost to the level of the spiracles. There
are also small ventrolateral tubercles. The head is shaped much
as in Nomadopsis and Calliopsis, with the low antennae on broad
protuberances. The clypeus and labrum are not clearly defined but
the latter bears large and acute labral tubercles. The mandibles
are slender and acutely pointed, much as in Calliopsis, but the
upper margins are more coarsely toothed, and the teeth spread out
to form a multidentate cusp area much larger than in Calliopsis
and nearer the base of the mandible. The apex of the maxilla is
broad and blunt as in Calliopsis, with the maxillary palpus arising
in the middle of apex. Maxillary and labial palpi are about as long
as broad.

Panurgus calcaratus Scopoli

Micheli (1936) describes and figures the larva of this species,
comparing it with that of P. banksianus. It is similar in body form
although with the dorsolateral tubercles higher. The upper lat-
eral portions of the head, above the antennae, are more convex,
as in Andrena. There are also minor differences in mandibular
dentition, etc. The spiracles are shown with a peritreme, with no
atrial teeth, but with atrial ridges.

1036 The University Science Bulletin

Panurginus melanocephalus ( Cockerell )

(Figs. 86-88)

The larva is similar to that of Nomadopsis fracta. The antennal
protuberances are larger. The labral tubercles are more distinct
and longer, the margin between them straight. The mandible is
rather slender, pointed, serrate on both margins, with a multidentate
cusp. The maxillary palpi are large, several times as large as the
labial palpi, longer than broad, distinctly pigmented. The spiracular
atria are produced above the body surface, without inner spines,
with nearly flat peritreme and distinct collar.

Berkeley, California, April 28, 1952 ( MacSwain ) .

Nomadopsis fracta Rozen

(Figs. 92-98)

The larva is rather robust and similar to that of Andrena but the
dorsolateral tubercles are all high and rounded, not transverse as in
Lasioglossum, Andrena, etc. The ventrolateral tubercles are absent.
The head capsule is essentially as in Andrena, agreeing with it in
the light sclerotization, the reduced antennae and other features;
the swellings above the antennae and those on which the antennae
are borne are smaller than in Andrena. The mandibules are slender,
thin upper margins serrate apically, the row of teeth in such a posi-
tion that they are out of sight when the mandible is seen from be-
neath. The cusp is nearly absent although there are a few small
teeth in its position. The maxillary palpi are broader than long.
The labial lobe bears the apical salivary opening which is not or but
little protruding and in some views almost circular. The spiracular
atrium projects above the body surface, with a rim and with feeble
internal ridges. The peritreme is nearly flat and the primary tra-
cheal opening is provided with a collar.

Mount Diablo, Contra Costa County, California, June 5, 1940
(Bohart and MacSwain). Additional specimens presumably of the
same species are from Oakley, Contra Costa County, California,
May, 1939 (Bohart and MacSwain).

Nomadopsis enphorbiae (Cockerell)

(Figs. 99, 106-108)

The larva is similar to that of Nomadopsis fracta except that the
dorsolateral tubercles are very much higher, those of some of the
body segments being twice as high as the width near the base,
those of the first two thoracic segments being distinctly more robust
than the others but nonetheless higher than basal width. The

Michener: Comparative Studies of Bee Larvae

1037

Figs. 92-98. 92, 93, Lateral and dorsal views of head of Nomadopsis fracta;
94, larva of same; 95, 96, 97, ventral, dorsal, and inner views of mandible of
same; 98, spiracle of same.

head is similar to that of N. fracta except that the antennae are much
more conspicuous and distinctly conical while the tubercles above
the antennae and the elevations on which the antennae arise are
less conspicuous. The mandible more robust, less attenuate, api-
cally than in N. fracta, cusp more evident, provided with numerous
teeth, marginal teeth in evidence seen from beneath, not hidden on

1038 The University Science Bulletin

upper surface as in N. fracta. The maxillary and labial palpi are
absent; the salivary opening is on a feeble prominence.

Riverside County, California, August 17, 1946 (MacSwain).

Spinoliella spp.

Claude-Joseph (1926) has described and figured larvae of S.
macidata (Spinola) and S. herbsti (Friese). Both species have
high dorsolateral tubercles but this is especially true of S. herbsti
in which the tubercles of the thorax are elongated and directed
backward. The mandibular apices are acute.

Calliopsis (Calliopsis) andreniformis Smith

(Figs. 100-105)

The larva is similar to that of Nomadopsis fracta but the dorso-
lateral tubercles of the thorax are a little higher and more angular
and ventrolateral elevations are feebly evident on the middle seg-
ments of body. The head is much as in Nomadopsis but the upper
parts are more gently rounded. The mandible bears teeth along its
dorsal margin and in an inner subapical patch or cusp. The spiracu-
lar atrium has thick walls without distinct ridges inside. It is not
produced above the body surface. The peritreme is large, and the
primary tracheal opening is provided with a collar.

Lawrence, Kansas, June, 1951 ( Michener ) .

Acamptopoeum submetallicum (Spinola)

The larva is briefly described and figured by Claude-Joseph
(1926). It has high dorsolateral tubercles as in Nomadopsis
euphorbiae. Unlike this species, the mandibular apices are bi-
dentate. The salivary opening appears to be narrow and quite
protuberant.

Perdita spp.

Custer (1929) briefly describes the larva of Perdita zebrata Cres-
son and P. opuntiae Cockerell, and also gives brief comments on
that of Nomadopsis australior (Cockerell). All of these bees have
the usual long dorsolateral tubercles of panurgine larvae, but they
are unusually attenuate, broadened and provided with bristles at the
apices in P. zebrata. This larva is even more remarkable for the
possession of a very long posterior median "spine" or tubercle.

Family Melittidae

Only a single species of this small family has been studied. It
agrees in most respects with the larvae of the families Colletidae,

Michener: Comparative Studies of Bee Larvae

1039

Figs. 99-108. 99, Nomadoptis euphorbiae, larva; 100, Calliopsis andreni-
fortnis, larva; 101, 102, lateral and dorsal views of head of same; 103, 104,
inner and ventral views of mandible of same; 105, spiracle of same; 106, 107,
ventral and inner views of mandible of Nomadopsis euphorbiae; 108, spiracle of

same.

36—6490

1040 The University Science Bulletin

Andrenidae, and Halictidae. The salivary opening is short and
transverse. The antennal papillae are absent, and unlike represent-
atives of the three families mentioned, the remnants of the antennae
are mere discs which are not themselves elevated nor are they borne
on elevations of the head surface. The palpi are reduced and the
mandibles are pointed, each with a multidentate cusp.

Hesperapis rufipes (Ashmead)

(Figs. 281-287)

The larva is moderately robust with the intersegmental lines
conspicuous, the lines between the cephalic and caudal annulets of
the segments absent or nearly so but posterior portions of each seg-
ment somewhat elevated on the dorsum. Ventrolateral tubercles
form a low ridge beneath the spiracles.

Head capsule scarcely sclerotic, without setae. Marginal thick-
ening of head capsule weak; epistomal suture not recognizable; dor-
sal longitudinal line of head capsule absent; parietal bands con-
spicuous. Antennae mere discs, not at all elevated; labroclypeal
suture distinct; labrum spiculate laterally. Mandibles each acutely
pointed, a series of small teeth along lower margin near apex, the
series along upper margin grading into the multidentate cusp; outer
surfaces of mandibles each with a single small seta. Maxillae not
clearly separated from labium except apically, not much bent in-
ward, palpi as long as broad. Labium short, rounded, exceeded by
maxillae, palpi broader than long; salivary opening small, nearly
round, in a slight depression, its upper margin produced as a slight
lip.

Spiracular atria without internal ridges or spines, produced above
body surface, with distinct rim and peritreme; primary tracheal
opening with collar.

Tanbark Flat, San Gabriel Mountains, Los Angeles County, Cali-
fornia, July 16, 1952 (MacSwain).

Family Megachilidae

The family is shown by larval characters, as well by those of the
adults, to be very homogeneous. The larvae exhibit certain clearly
primitive characters. Thus they differ from all other known bee
larvae except those of certain Allodape in the abundant setae on
the body. Other primitive features include the distinct antennal
papillae, the long maxillary and labial palpi, and the large trans-
verse salivary opening provided with large lips. As explained else-
where, it is believed that these primitive features may be reversions,

Michener: Comparative Studies of Bee Larvae

1041

not actual indications of primitiveness, for the mandibles are of a
specialized type, like those of many of the other long tongued bees,
having an inner apical concavity limited by a distinct transverse

115 \ M6

Figs. 109-118. 109, Trachusa perdita, larva; 110, 111, dorsal and lateral
views of head of same; 112, 113, inner and ventral views of mandible of same;
114, Stelis lateralis, larva; 115, 116, inner and ventral views of mandible of
same; 117, spiracle of Trachusa perdita; 118, spiracle of Stelis lateralis.

1042 The University Science Bulletin

ridge (absent in Stelis lateralis) on the inner surface. The apices
of the mandibles have two large teeth, a presumably primitive fea-
ture, except in Stelis lateralis in which they are acute, in Trachusa
in which they are truncate, and in Megachile (Chalicodoma) in
which one of the teeth is much reduced. The Anthidiini (except
Trachusa) are noteworthy for the presence of small teeth along
the mandibular margins apically. Such teeth are absent in most
Megachilini, although Coelioxys exhibits them. They are also
found in immature larvae of Megachile brevis (see Michener, in
manuscript ) , but are scarsely discernable in mature larvae. As such
teeth are found in short-tongued families of bees, it is assumed that
they are a primitive feature, supporting the view, based upon adult
morphology, that the Anthidiini are a more primitive group than
the Megachilini.

An interesting feature of megachilid larvae is that there are no
dorsolateral tubercles in the sense that these structures are under-
stood in other groups of bees. Dorsal protuberances when present
are middorsal or if they extend onto the sides of the body (as in
Ashineadiella ) they are continuous across the middorsal line.

Spiracles in all forms examined have atria extending above the
body surface. Atrial spines are common but not universal. It may
be remarked that among the few species examined the absence of
atrial spines distinguishes larvae of Megachile, subgenera Chelo-
stomoides and Chalicodoma, from other subgenera of Megachile.

Lithurge dubius (Herbst)

The larva is described by Claude-Joseph ( 1926 ) as fusiform with
the posterior segments more dilated than the anterior. The caudal
annulets of the segments are elevated. As in other megachilids, the
antennal papillae and the palpi are distinct.

Trachusa perdita Cockerell

(Figs. 109-113, 117)

The larva is very robust, with the intersegmental lines incon-
spicuous except ventrally where they are rather distinct and with
the lines between the cephalic and caudal annulets of the segments
present anteriorly but inconspicuous. The caudal annulets of the
dorsal segments in the middle portion of the body are somewhat
elevated, the elevations extending across the middorsal line and
highest middorsally, not produced to form distinct dorsolateral
tubercles. Rounded ventrolateral tubercles are present on the ab-
domen ( except for the apical segments ) and less strongly so on the
second and third thoracic segments.

Michener: Comparative Studies of Bee Larvae , 1043

Head capsule and mouthparts somewhat sclerotized, with nu-
merous distinct setae. Marginal thickening of head capsule dis-
tinct; posterior tentorial pits conspicuous; epistomal suture indi-
cated by a slightly arched line between anterior tentorial pits,
cuticle not thickened in this area; dorsal longitudinal thickening of
head capsule distinct, divided into two branches anteriorly which
are directed toward anterior tentorial pits. Antennae represented
by broad low convexities, each of which bears a small median
papilla somewhat longer than broad; labroclypeal suture distinct,
slightly arched; labrum with apex concave. Mandibles rather short
and robust, apices truncate, upper apical angle more rounded than
lower; inner surface of apical portion gently concave, the concavity
bounded by a weak ridge. Maxillae each with a basal swelling;
apices bent somewhat inward as in Anthophora; maxillary palpi
longer than broad; apical portions of maxillae bearing coarse setae;
hypostomal furrow deep, labiomaxillary rod distinct; labial lobe
rather thick, salivary opening at its apex, a transverse slit provided
with very large sclerotized lips; labial palpi more than twice as long
as broad, separated by a distance equal to width of salivary open-
ing; postmentum and prementum clearly separated, the latter pro-
vided with coarse bristles.

Body provided with numerous setae. Spiracles with large atrium,
inner walls of which are spinose, spines reduced to tubercles below;
atrium projecting above body surface; peritreme rather narrow, flat;
primary tracheal opening with delicate collar.

The Pinnacles, California, April, 1940 ( MacSwain ) .

Dianthidium sp.

(Figs. 120, 125, 126, 129)

The larva is similar to that of Trachtisa but even more robust, the
intersegmental lines very inconspicuous, the division of the segments
into cephalic and caudal annulets feeble. The ventrolateral por-
tions of the segments are slightly swollen. The head capsule is
somewhat more elongate than that of Trachusa but is similar in all
important respects. The setae of the head capsule and mouthparts
are slender, not bristlelike. The cleavage lines are more distinct
than in Trachusa. The antennal papilla is about twice as long as
broad. The apex of the mandible is bifid, the teeth pointed, the
lower tooth much longer than the upper, the margin between the
teeth and the lower margin of the mandible finely serrate, a long
slender tooth on the lower margin. The maxillary and labial palpi
are about twice as long as broad.

1044

The University Science Bulletin

129

Figs. 119-129. 119, Anthidiellum erhorni, larva; 120, Dianthidium sp?, larva;
121, 122, inner and ventral views of mandible of Anthidiellum notatum robert-
soni; 123, 124, inner and ventral views of mandible of Anthidiellum erhorni;
125, 126, inner and ventral views of mandible of Dianthidium sp?; 127, spiracle
of Anthidiellum erhorni; 128, spiracle of Anthidiellum notatum robertsoni; 129,
spiracle of Dianthidium sp?.

Michener: Comparative Studies of Bee Larvae 1045

The spiracular atria lack spines but bear internal ridges. The
primary tracheal opening is provided with a collar.

Mount Diablo, Contra Costa County, California, December, 1940
(MacSwain).

Dianthidium curvatum sayi Cockerell

(Figs. 130-133)

The larva is similar to that of D. sp. described above. It differs,
however, in the absence of serrations along the upper margins of
the mandibles, and in other details of mandibular dentition, particu-
larly in the presence of a slender sharp spine arising in the notch
between the mandibular teeth. The spiracles differ from those of
the other Dianthidium available in the presence of numerous groups
of atrial teeth.

Garden City, Kansas ( Roland Fischer ) .

Figs. 130-133. 130, 131, ventral and inner views of mandible of Dian-
thidium sayi; 132, spiracle of same; 133, inner ventral view of apex of man-
dible of same.

Anthidiellum erhorni ( Cockerell )

(Figs. 119, 123, 124, 127)

The larva is very robust, similar in shape to that of Dianthidium,
but with the intersegmental lines more distinct and the lines be-
tween the cephalic and caudal annulets of the segments more
clearly visible. The ventrolateral tubercles are large and distinct.
The head capsule is essentially as in Dianthidium. The mandibles
have bidentate apices with the notch between the teeth a narrow
sinus, the teeth obliquely truncate, their distal margins, and the
upper distal margin of the mandible, finely serrate. The apical con-
cavity of the mandible is deeper than in Trachusa or Dianthidium.
The spiracles are much as in Dianthidium but with a slight rim
around outer margin of the atrium and a smaller collar around the
primary tracheal opening.

1046 The University Science Bulletin

Vidal, San Bernardino County, California, September 6, 1947
(MacSwain).

Anthidielhtm notatum robertsoni (Cockerell)

(Figs. 121, 122, 128)

This larva may not be distinguishable from that of erhorni. How-
ever, in the two specimens examined there are differences of man-
dibular dentition, the notch between the two principal teeth being
broader and the serrations coarser in robertsoni. The spiracular
atria are more narrowed at the body surface, and the rim is more
distinct.

Antioch, Contra Costa County, California, September 10, 1947
(MacSwain).

Anthidium oblongatum Latreille

The larva of this species is described by Xambeu (1896) and
Maneval (1937). It is much like that of Megachile brevis in shape,
perhaps a little more robust anteriorly, with a similar series of some-
what confluent ventrolateral tubercles. The characters in general
are like those of most megachilids, the palpi and antennal papillae
being about three times as long as broad, the salivary opening being
transverse and apparently with lips. The labroclypeal suture is
strongly arched as usual in the family. The maxillary palpi arise
preapically on the maxillae. The mandibles are bidentate apically,
the teeth blunt, the notch between them rather shallow; the upper
margin of each mandible bears a slender tooth like the single long
slender tooth on this margin in Dianthidium sp?. Otherwise the
mandibles lack small teeth. The labrum bears a marginal series of
papillae.

Anthidium chilense Spinola

Claude-Joseph ( 1926 ) describes and figures the larva of this
species. It is similar in shape to that of Dianthidium, but has a
subspiracular ridge made up of fused ventrolateral tubercles. The
mandibular apices also appear to be similar to those of Anthidiellum
robertsoni.

Paranthidium (?) caturigense (Giraud)

Micheli (1935) and Maneval (1936a) describe and figure the
larva of this species. The larva is robust, not swollen posteriorly
as in many megachilids, with lines between the cephalic and caudal
annulets of the body segments evident dorsally, the dorsal tubercles
absent, the ventrolateral tubercles present. The usual features of

Michener: Comparative Studies of Bee Larvae 1047

a megachilid are present, such as the setose body, the long palpi and
antennal papillae, the large transverse salivary slit provided with
distinct lips, the arched labroclypeal suture, etc. The labrum is
armed with several rows of papillae apically. An unusual feature
according to Maneval is the bifid apex of each antenna but Micheli
shows a normal apex. The mandibles are bidentate, the teeth short,
not separated by a deep notch, the upper margin of the mandible
denticulate.

Paranthidium septemdentatum (Latreille)

A larva, supposedly of this species, was briefly described by
Xambeu (1896). Since he describes the maxillary palpi as biartic-
ulate and the mandibles as laterally ciliate, it seems likely that he
had the larvae of some other insect.

However, Grandi (1934a, 1935) describes and figures the larva
in detail. As with P. coturigense, the larva is not swollen posteriorly
as in many megachilids. The comments on P. coturigense apply
also to the present species, except that the deeply emarginate apex
of the labrum bears only a single irregular row of papillae, the
antennae are not bifid, the mandibular teeth are blunt, especially
the shorter one, and there are small denticles on the upper margin
of the upper (shorter) tooth. The spiracular atria bear small inner
teeth.

The placement of this species in Paranthidium is subject to ques-
tion. It does not fall in any described subgenus, and should per-
haps fall in the allied Paraanthidium (see Michener, 1948). Unlike
American Paranthidium, it uses both plant down and resin in its
nests.

Paranthidiellum lituratum (Panzer)

Enslin (1923) and Micheli (1934) give accounts of the larva of
this species. The larva is shaped much like that of Dianthidium
but is more slender anteriorly so that the swollen posterior portion
is more striking. The setose body, long antennal papillae, long
palpi, and broad salivary opening with lips are as in other Mega-
chilidae. The labroclypeal suture is more strongly arched than
shown in Trachusa, almost angulate medially. The lower margin
of the labrum bears an irregular transverse row of small papillae.
Unfortunately the mandible is not well illustrated nor fully de-
scribed, but it is bidentate, and evidently without the small teeth
found in Anthidiellum and Dianthidium.

1048 The University Science Bulletin

Stelis nasuta (Latreille)

The larva of this species was described by Maneval (1937). It is
very different from the larva of Stelis lateralis illustrated in the
present paper. The body form is similar but middorsal tubercles
are entirely absent and ventrolateral tubercles are present. Weak
lines separating the cephalic from the caudal annulets of the body
are present dorsally on the middle body segments. Like most
megachilids, the palpi and antennal papillae are elongate, the sali-
vary opening transverse and with lips, the labroclypeal suture
strongly arched, the sides of the labrum drawn down to form broad
and undefined labral tubercles. The margin of the labrum bears a
row of papillae. The maxillary palpi are preapical on the maxillae.
The mandibles, unlike those of Stelis lateralis, are bidentate at their
apices, each tooth obliquely truncate. They are said to be much
like those of S. ornatula.

Stelis ornatula Nylander

The larva of this species was described and figured by Micheli
(1935). Unlike S. lateralis, the larva is robust, swollen posteriorly,
the dorsal tubercles virtually absent and ventrolateral tubercles
present. Distinct lines separate the caudal from the cephalic
annulets of the body segments. The palpi and antennal papillae
are elongate as in most megachilids, the salivary opening is trans-
verse and provided with large lips, the labroclypeal suture is arched,
the sides of the labrum are produced to form distinct tubercles.
The margin of the labrum bears a broken row of papillae. The
maxillary palpi are preapical. The mandibles are bidentate, the
upper margin of each mandible minutely dentate apically.

Stelis (Microstelis) lateralis Cresson

(Figs. 114-116, 118)

The larva is similar in body form to that of Hoplitis (Alcidamea)
sp? described elsewhere in this paper, but with the middorsal tuber-
cles higher, that of the fourth abdominal segment particularly large.
The ventrolateral tubercles are absent. The head is similar to that
of other megachilids, but the mandible is pointed, neither truncate
nor bifid, with virtually no indication of an apical inner concavity
or of a cusp. The spiracular atrium is provided with scattered long
spines arising from short ridges. The atrium is produced above the
level of the body surface. The peritreme is flat, and the primary
tracheal opening is narrowed by a collar.

Lawrence, Kansas, June 27, 1950, from nest of Hoplitis {Alci-
damea) pilosifrons (Cresson) (Michener).

Michener: Comparative Studies of Bee Larvae 1049

Stelis minuta Lepeletier

Enslin ( 1925 ) described and figured the larva of this species. It
apparently does not have conspicuous dorsal tubercles but has
large rounded ventrolateral tubercles. The lines of division be-
tween caudal and cephalic annulets of the dorsal body segments
are shown as feeble. The mandibles, like those of S. lateralis, are
sharply pointed.

Dioxys cincta (Jurine)

Micheli (1936) described and illustrated the larva of this para-
site. It is peculiar for a megachilid in having only a few scattered
setae on the body. The head has only a few scattered but rather
long setae. It agrees with the majority of megachilids in the long
antennal papillae, long palpi, transverse salivary opening with large
lips, etc. The spiracles are described as having numerous very
minute teeth in the atrium. The labrAclypeal suture is strongly
arched; the labrum bears a small cluster of papillae subapically at
each side. The mandible is bidentate, the teeth rather small and
blunt. There are no minute teeth or serrations. Micheli shows
no defined inner apical concavity, but he fails to show the transverse
line defining the concavity basally in certain other bees which almost
certainly have such a line ( e. g. Hoplitis lepeletieri ) . Probably
there is no such line in Dioxys, but it may be present.

It seems likely that the larva described by Grandi ( 1934a ) from
a nest of Megachile (Chalicodoma) muraria is a Dioxys.

Heriades (Heriades) crenulatus Nylander

The larva of this species is fully described and figured by Grandi
(1934a). The body shape is typical of megachilids, the posterior
portion being swollen. The lines separating the cephalic from the
caudal annulets of the dorsal body segments are evident and extend
downward below the spiracles as shown for Megachile brevis. The
caudal annulets are swollen dorsally but not so strongly as in
Ashmeadiella. Feeble ventrolateral swellings are present. Other
characters also are like those of most megachilids. The body bears
numerous setae, the antennal papillae and the palpi are elongate,
the salivary opening is transverse and provided with very long lips,
and the labroclypeal suture is strongly arched. The labrum is
unusually narrow, with the apex emarginate; it lacks papillae, or
they are represented by small pegs. The mandibles are bidentate,
the teeth acute, the inner apical concavity distinct. The spiracular
atria contain a few blunt teeth.

1050

The University Science Bulletin

Figs. 134-141. 134, Ashmeadiella sp?, larva; 135, lateral view of head of
Osmia lignaria; 136, 137, inner and ventral views of mandible of same; 138,
139, ventral and inner views of mandible of Ashmeadiella sp?; 140, spiracle of
Osmia lignaria; 141, spiracle of Ashmeadiella sp?.

Michener: Comparative Studies of Bee Larvae 1051
Ashmeadiella sp.

(Figs. 134, 138, 139, 141)

The larva is essentially as in Trachusa but the caudal annulets of
the dorsal body segments are much more strongly elevated, the ele-
vations continued across the dorsum and extending down laterally
to the elevations of the ventrolateral regions of the body which are
not formed into distinct tubercles but are elevated to produce an
irregular ridge. The head capsule is essentially as in Trachusa.
The papilla of each antenna is about twice as long as broad and the
maxillary and labial palpi are distinctly longer than broad. The
mandible is bidentate at its apex, the teeth rounded, the upper tooth
slightly shorter than the lower. The spiracular atria lack spines but
have high sharp ridges within; the collar surrounding the primary
tracheal opening is thick.

Mount Diablo, Contra Costa County, California, December 24,
1940 (MacSwain) (a single specimen removed from the nest of
Pseud omasaris ) .

Hoplitis (Alcidamea) sp.

(Figs. 142-145, 149)

In fundamental structure this larva is similar to that of Trachusa
but it is much more slender, the lines of separation between the
cephalic and caudal annulets of the body segments are even less
conspicuous and the ventrolateral tubercles are inconspicuous. The
posterior portions of the middle body segments, and to a lesser
extent of the last thoracic to the sixth abdominal segments bear
middorsal unpaired elevations or tubercles. The head capsule dif-
fers from that of Trachusa in the absence of a cuticular thickening
along the branches of the cleavage line. The antennae are more
conspicuously conical than in Trachusa, the apex of each cone bear-
ing a papilla as in Trachusa. The mandibles are deeply bifid at
their apices, the upper tooth shorter than the lower, the inner apical
concavity weak. The maxillary palpi are over twice as long as
broad. The spiracular atria are deep, thick walled, nearly smooth
inside, much produced above the body surface; the peritreme is
narrow and flat.

Mineral King, Tulare County, California, August 10, 1939 (Bo-
hart). From their large size it is clear that they are either H.
sambuci Titus or H. uvulalis (Cockerell).

Larvae of H. (Alcidamea) parvula (Dufour and Perris) and H.
(Tridentosmia) tridentata (Dufour and Perris) were described
briefly by Dufour and Perris ( 1840 ) .

1052

The University Science Bulletin

1 r s w *

, ,, Q* »

v..

Figs. 142-150. 142, Hoplitis (Alcidamea) sp?, larva; 143, lateral view of
head of same; 144, 145, inner and ventral views of mandible of same; 146,
antenna and surrounding region (lateral view) of Proteriades xerophila; 147,
148, inner and ventral views of mandible of same; 149, spiracle of Hoplitis
(Alcidamea) sp?; 150, spiracle of Proteriades xerophila.

Hoplitis (Hoplitis) adunca (Panzer)

The larva of this species is described and figured by Grandi
(1935). Grandi suggests that this larva may have been that of a
parasite, Dioxys cincta Jurine, but Micheli (1936) describes the
larva of Dioxys and shows clearly that Grandi's "Osmia adunca" is
correctly identified.

Michener: Comparative Studies of Bee Larvae 1053

The body is more robust and more curled than that of Hoplitis
(Alcidamea) , and lacks dorsal protuberances as well as ventro-
lateral tubercles. The cephalic and caudal annulets of the body are
distinct dorsally. Most structures are typical of megachilids, for
example, the elongate antennal papillae and palpi, the arched labro-
clypeal suture, the transverse salivary opening with lips, and the
setae on the body surface. The mandibles are bidentate with the
teeth rounded, the inner concavity well defined. The labral margin
is provided with a broken row of papillae.

Hoplitis lepeletieri (Perez)

The larva of this species is described and figured by Micheli
(1933). It is more robust than that of Hoplitis (Alcidamea), being
more like the larva of Megachile in form although not so robust as
usual in that genus. H. lepeletieri exhibits the usual megachilid
features. The labroclypeal suture is strongly arched and the lowest
margin of the labrum bears an irregular broken row of small
papillae. The mandibles are bidentate, the upper tooth shorter
than the lower, in ventral view looking much like those of Osmia
lignaria, but with the upper margin provided with several small
blunt teeth.

Hoplitis loti (Morawitz)

The larva of this species is described by Micheli (1931). It has
very much the shape of Hoplitis (Alcidamea), although Micheli
shows it in a more curled position. All of its features are typical
of the Megachilidae in general. The mandibles are bidentate, the
teeth shorter than in Alcidamea and blunt as in Proteriades. The
anterior margin of the labrum bears a broken row of papillae absent
in the similar Hoplitis leucomelaena.

Hoplitis leucomelaena (Kirby)

The larva of this species is described by Enslin (1925) and by
Micheli ( 1930 ) . The larva has the body form of the Hoplitis
(Alcidamea) described in the present paper except that it is more
slender with the intersegmental lines more deeply constricted on
the venter. All its features described and figured by Micheli are
typical of Megachilidae in general. The mandibles are bidentate
with the teeth shorter than those figured for Hoplitis (Alcidamea)
but not blunt as are those of Proteriades.

1054 The University Science Bulletin

Diceratosmia siibmicans (Morawitz)

The larva of this species is briefly described and figured by
Maneval (1939). It exhibits the usual megachilid features. The
antennal papillae and palpi are over three times as long as broad.
The maxillary palpi are preapical on the maxillae. The labrum
bears an apical row of papillae. The salivary opening is transverse
with unusually long lips. The mandibles are bidentate, the apical
teeth narrowly rounded, the notch between them deep. There is
no evidence in Maneval's description and figures of the well-defined
inner apical concavity of most megachilids.

Osmia (Osmia) lignaria Say

(Figs. 135-137, 140)

The larva is similar to that of Trachnsa except that the inter-
segmental lines are even less conspicuous and the lines between the
cephalic and caudal annulets of the segments are also less con-
spicuous, the caudal annulets not being elevated as in Trachusa.
The ventrolateral tubercles are absent. The head capsule is essen-
tially as in Trachusa; the swellings on which the antennae are
located are even lower, being essentially indistinguishable, and the
antennal papillae are higher, being more than twice as long as
broad. The mandibles are bidentate at their apices, the upper
tooth being smaller and shorter than the lower; the inner apical
concavity is shallow. The maxillary palpi are longer than broad
and the maxillae and labium are not provided with bristlelike
hairs but only with ordinary fine setae. The spiracles have atria
spinose within and produced well above body surface. The peri-
treme is narrow and the primary tracheal opening lacks a collar.

Pittsburg, California, May, 1940 (MacSwain).

Osmia (Osmia) rufa (Linnaeus)

The larva of this species was described and figured by Losinski
(under the name O. bicornis Linnaeus) in 1911. No conspicuous
specific characters are evident but the structures in general are
similar to those of O. ligriaria.

Osmia (Helicosmia) aurulenta (Panzer)

Marechal (1926) has described and illustrated the larva of this
species. The mature larva is more slender than that of Trachusa,
and differs from that of O. lignaria in having the lines between
cephalic and caudal annulets of the segments distinct dorsally, the
caudal annulets elevated dorsally. The ventrolateral tubercles of
the body are present. The usual megachilid characters of long

Michener: Comparative Studies of Bee Larvae 1055

antennal papillae, long palpi, transverse salivary opening with lips,
and setose body are present. The mandibles are bidentate, the
upper tooth apparently being short and rounded in the mature larva
although pointed in younger ones.

Proteriades xerophila (Cockerell)

(Figs. 146-148, 150)

The larva is similar to that of Trachusa but somewhat more
slender and with less distinct ventrolateral tubercles. The head
capsule is essentially as in Trachusa with the position of the cleavage
line being indicated by slight depressions and slight thickenings and
the position of the epistomal suture between the anterior tentorial
pits being indicated by a slight depression. The antennal papillae
are much larger than in Trachusa, being more than twice as long as
broad. The mandibles are bifid, the teeth blunt, the lower one
longer than the upper. The maxillary as well as the labial palpi
are more than twice as long as broad. The spiracles have very
thick-walled atria which have scattered large spines within and are
produced above the body surface. The peritreme is narrow and
the primary tracheal opening is much narrowed by a thick collar.

Little Lake, Inyo County, California, February 29, 1940 ( Linsley,
Bohart, and MacSwain ) .

Megachile sp.

(Figs. 154, 155, 158)

The larva is similar to that of Trachusa, but the caudal annulets
of the body segments are less elevated (although weak middorsal
elevations are present) and the ventrolateral regions of the body,
instead of being formed into separate tubercles, are elevated to
form a broad ridge on each side below the level of the spiracles.
The head capsule is much as in Trachusa, the antennal papillae
being more than twice as long as broad. The apices of the man-
dibles are bifid, the lower tooth longer than the upper, the inner
apical concavity of each mandible deep and delimited by a distinct
and sharp ridge which forms a conspicuous inner shoulder on the
mandible. The maxillary and labial palpi are similar in appearance,
both more than twice as long as broad. The maxillae and labrum
bear setae similar to those found elsewhere on the head and mouth-
parts, not coarse and bristlelike. The spiracular atria are spinose
within, the spines being arranged in short rows, each row on a short
ridge; the spines are lower toward the primary tracheal opening.
The peritreme is rather narrow.

New York City, August, 1928 (Lillian Clum).
37—6490

1056

The University Science Bulletin

Figs. 151-158. 151, Megachile brevis, larva; 152, 153, inner and ventral
views of mandible of same; 154, 155, inner and ventral views of mandible of
Megachile sp?; 156, spiracle of Megachile (Chelostomoides) sp? from Panama;
157, spiracle of Megachile brevis; 158, spiracle of Megachile sp?.

Michener: Comparative Studies of Bee Larvae 1057
Megachile (Litomegachile) brevis Say

(Figs. 151-153, 157)

This species is similar to that of Megachile sp. described above
but the mandibular teeth are less flaring; the teeth of the atrium
are more numerous, the subatria of the spiracles are spinose within,
swollen and only irregularly annulate.

Lawrence, Kansas (White, Michener, Fischer, LaBerge, Wille).

Megachile (Megachile) centuncularis (Linnaeus)

Buysson ( 1902 ) makes a brief comment on the larva of this spe-
cies, comparing it with that of M. lagopoda; Packard (1897) de-
scribes the general characteristics of a larva supposed to be this
species; and Grandi (1934a) gives a full description and figures.
The larva is similar to that of M. sp. and M. brevis, described in the
present paper. The mandibles are more like those of M . brevis, but
have the lower tooth even more acute. The atria contain groups of
small teeth, as in M. brevis but perhaps smaller.

Megachile (Eutricharaea) argentata (Fabricius)

The larva of this species is described and figured by Grandi
(1931). It differs from that of M. albiseta only by very minor
characters.

Megachile nigriventris Schenck

The larva of this species is described and figured in some detail
by Micheli (1937). It is similar to that of M. brevis in most re-
spects. The most striking characteristic of the larva of nigriventris
is that the head bears a pair of conspicuous tubercles well above the
antennal bases. The apices of these tubercles are rounded, not
pointed as they are in Centris, the only other bee larva known to
have comparable tubercles. The mandible is similar to that of
M. brevis but with the upper tooth conspicuously acute, the upper
margin smooth. The body is less plump than in M. brevis, with the
lines between cephalic and caudal annulets of the segments con-
spicuous, the latter distinctly more swollen than the former. The
spiracles are not figured or adequately described.

Megachile albiseta ( Klug )

Grandi (1931) describes and figures the larva of this species in
detail. As with M. nigriventris, the body is less plump than in
such forms as M. brevis, not noticeably swollen posteriorly, with
the caudal annulet of each segment considerably swollen. Ventro-

1058 The University Science Bulletin

lateral tubercles are present. The elongate palpi and antennal
papillae, the transverse salivary opening with lips, and the setose
body are characteristic of megachilids. The mandibles are biden-
tate, with an inner apical concavity.

Megachile spp.

Buysson (1902) very briefly describes the larva of M. lagopoda
(Linnaeus), pointing out such conspicuous characters as the bi-
dentate mandibles. He then states that larvae of the following
are similar to that of M. lagopoda: M. maritima (Kirby), pyrenaica
Lepeletier, and buyssonii Perez.

Another group of species was described by Claude-Joseph
(1926). These are M. pollinosa Spinola, saulcyi Guerin-Meneville,
rancaguensis Friese, and euzona Perez. All have bidentate man-
dibles and the other usual characters of the genus.

Megachile (Chalicodoma) muraria (Retzius)

The larva of this species is well described and figured by Grandi
(1934a). Like the adult, it is somewhat unusual for a Megachile,
having mandibles which are not bidentate but narrowed apically
with a suggestion of a subapical tooth represented by a small shoul-
der. As in the subgenus Chelostomoides, the spiracular atria lack
spines. As would be expected, it resembles most megachilids in the
principal characters of the group, such as the setose body, the long
antennal papillae and palpi, and the large transverse salivary open-
ing with lips.

Megachile (Chalicodoma) pyrenaica alpina Morawitz

The larva is described by Micheli (1935) who notes only very
minor differences between this species and M. muraria.

Megachile (Chelostomoides) sp?

(Fig. 156)

This species is much like the above, differing in the less elevated
caudal annulets of the body segments (in the single specimen
studied) and in the spiracular details, the atrium being narrowed
toward the surface and the subatrium having few annulations.

Between Pacora and Chepo, Panama, February 25, 1924 ( Snyder,
Zetek, and Molino). The larva was in the outer layer of a nest of
Amitermes niedius. The adult bee was identified by S. A. Rohwer
as Megachile howardi Cockerell. This species is a member of the
subgenus Sayapis, but the female originally associated with it is

Michener: Comparative Studies of Bee Larvae

1059

Megachile (Chelostomoides) zoptlana Cresson (see Mitchell, 1937).
Identification was probably based on a female, and it seems likely
that the larva in question is a Chelostomoides, perhaps M. zaptlana.

Megachile (Chelostomoides) sp?

(Figs. 162-164)

The larva is shaped approximately as in M. brevis but the lines
between the cephalic and the caudal annulets of the body seg-

Figs. 159-166. 159, Coelioxys 8-dentata, larva; 160, 161, ventral and inner
views of mandible of same; 162, 163, ventral and inner views of mandible of
Megachile (Chelostomoides) sp. from California; 164, spiracle of same; 165,
lateral view of head of first stage larva of Coelioxys 8-dentata; 166, spiracle of
mature larva of same.

1060 The University Science Bulletin

ments are conspicuous, and the caudal annulets are elevated dorsally
and dorsolaterals. The ventrolateral tubercles are absent. The
head structures are essentially as in M. brevis. The mandibular
teeth are shorter and more rounded than in that species while
the basal limit of the mandibular concavity, is distinct but not
strongly elevated as in M. brevis. Unlike M. brevis, the spiracular
atrium lacks spines and the primary tracheal opening is narrowed
by a collar.

Five miles northwest of Blythe, California, April 2, 1941, in old
Colletes burrow (Linsley, MacSwain).

Coelioxys 8-dentata Say

(Figs. 159-161, 165, 166)

The larva is similar to that of Megachile but differs by having
longer setae on the body, numerous long setae on the outer sur-
faces of the mandibles, and a projection, variable in size, extending
downward from the hypostomal thickening immediately behind
the mandible. The spiracular atria have very long spines, progres-
sively shorter toward the primary tracheal opening. The preatrium
is spinose or tuberculate within, swollen, only irregularly annulate,
and longitudinally folded.

Lawrence, Kansas, various dates (Michener and White), in nests
of Megachile brevis Say.

Xylocopinae

The evidence of relationship of the Ceratininae and the Xylo-
copinae on the basis of adult characters has been rather tenuous.
It is therefore interesting that the larval characters of Xylocopa
and Ceratina in general support the relationship. Common char-
acters include the absence of body tubercles (although there are
greatly enlarged and modified ventrolateral tubercles in Allodape
and Exoneura), the bidentate mandibles (acute in Allodape) with
small marginal teeth, the great reduction of the salivary opening
which is hidden from outside view, being far back on the upper
surface of the labial lobe, and the distinct antennal tubercles. In
the attenuate mandible which has lost its inner apical concavity,
Ceratina is more specialized than Xylocopa.

Among the most remarkable bee larvae are those of Allodape
(Brauns, 1926, Masi, 1930; Yasumatsu, 1938) and Exoneura (Ray-
ment, 1949, 1949a, 1951; Erickson and Rayment, 1951). They are
progressively fed and have various modifications of the body tuber-
cles important for the progressive feeding utilized by members of

Michener: Comparative Studies of Bee Larvae

1061

Figs. 167-172. 167, Xt/locopa virginica, larva; 168, 169, lateral and dorsal
views of head of same; 170, 171, inner and ventral views of mandible of same:
172, spiracle of same.

1062 The University Science Bulletin

these genera. Unfortunately few of the finer details of structure
have been recorded for members of these genera, the most com-
plete information available being that provided by Yasumatsu. It
is interesting that at least certain Allodape larvae have setae on the
body like those of the Megachilidae and more abundant than those
of Bombus.

Xylocopa (Schonherria) virginica (Linnaeus)

(Figs. 167-172)

The larva, which was briefly described by Packard (1897), is
rather slender with intersegmental lines conspicuous but not deep,
the lines between the caudal and cephalic annulets of the segments
absent, the dorsolateral and ventrolateral tubercles also absent.

Head capsule somewhat sclerotized with a distinct constriction
separating it from thorax; head capsule and mouthparts provided
with scattered minute setae; epistimal suture represented by a
rather distinct groove between anterior tentorial pits; cleavage line
not evident but dorsal longitudinal median thickening of head
capsule present. Antennae each with distinct papilla which is
longer than broad arising from a broad convexity; labroclypeal
suture distinct; labrum with apical margin subtruncate, labial tuber-
cles absent. Mandibles each with a tubercle or projection on outer
side midway between base and apex, apices bidentate, upper tooth
much longer than lower, its upper margin coarsely serrate; lower
tooth small and slender, its inner margin feebly serrate. Maxillae
with apices extending a short distance beyond maxillary palpi which
are longer than broad; labium with salivary opening small, circular,
on dorsal surface of labial lobe, without lips or projections; labium
with a distinct ventral tubercle just anterior to line between post-
mentum and prementum; hypostomal furrow deep.

Body without spicules or setae. Spiracles with atrium and pre-
atrium both densely spinose within, the former produced above
body surface, with rim; preatrium short, not annulate, thrown into
longitudinal folds; peritreme flat.

Geitch, Virginia, July 7, 1914 ( T. D. Snyder ) ( A single specimen ) .

Xylocopa (Xylocopa) violacea (Linnaeus)

The larva of this species was briefly described and figured by
Lucas ( 1868 ) and fully so by Grandi ( 1934a ) . The larva is similar
to that of X. virginica, differing in the more slender upper mandib-
ular tooth which bears a distinct shoulder at the base of the lower
mandibular tooth. Minor apparent differences in the arrangement
of the minute setae may or may not exist in reality.

Michener: Comparative Studies of Bee Larvae

1063

Xylocopa aestuans (Linnaeus)

Dover (1924) described and figured the larva of this species.
So far as can be determined, the species agrees with the two better
known Xylocopa species whose larvae are described above.

Ceratina (Zadontomerus) dupla Say

(Figs. 173-179)

The larva, which was crudely figured by Packard (1897), is slen-
der with the intersegmental lines rather distinct but the lines be-
tween cephalic and caudal annulates of segments absent or visible
only on some of the middle segments of the body. The dorsolateral
and ventrolateral tubercles are absent.

Figs. 173-179. 173, Ceratina dupla, larva; 174, 175, ventral and inner
views of mandible of same; 176, 177, lateral and dorsal views of head of same;

178, dorsal view of labiomaxillary complex showing salivary opening of same;

179, spiracle of same.

1064 The University Science Bulletin

Head capsule and mouthparts with scattered exceedingly minute
setae; head capsule rather weakly sclerotized, a slight constriction
separating it from the thorax; marginal thickening of head capsule
weak and posterior tentorial pits inconspicuous; epistomal suture
indicated by a very feeble depression and feeble thickening of the
integument extending between the anterior tentorial pits; cleavage
line feebly indicated, dorsal longitudinal median thickening of head
capsule absent. Antennae distinct, each represented by a rather
high prominence; labroclypeal suture distinct; labrum with apex
subtruncate, labral tubercles absent. Mandibles with apices bi-
dentate, upper tooth longer than lower, upper and lower margins
of mandible serrate apically; cusp absent, inner apical concavity not
distinct. Maxillae with apices not bent inward as in Anthophora,
maxillary palpi broader than long; hypostomal furrow deep;
labium with salivary opening small, round, on dorsal side of labial
lobe; division between prementum and postmentum distinct.

Body without spicules or setae. Spiracles with atrium with a few
feeble ridges, without spines, not projecting above level of body sur-
face; peritreme present; collar around primary tracheal opening
present and thick.

Washington, D. C, June IS, 1930 (J. E. Barr).

Ceratina (Ceratina) cucurbitina (Rossi)

Grandi (1935) has described and illustrated this species. It is
similar to C. duplo, with the antennae and palpi perhaps slightly
shorter. The small denticles of the mandibles appear to be some-
what differently arranged than in C. dupla.

Ceratina (Ceratina) callosa (Fabricius)

The larva of this species is described and figured by Micheli
( 1936 ) . It is essentially like C. dupla, apparently with the an-
tennal papillae and the palpi slightly longer. The mandibles evi-
dently have the small teeth of the margins smaller than in dupla,
indeed Micheli's outer view does not show them at all.

Manuelia gayatina (Spinola)

Claude-Joseph ( 1926 ) describes and figures the larva of this spe-
cies. It is similar to that of Ceratina, the body lacking tubercles.
The antennal papillae are unusually large. The mandibles are
slender apically as in Ceratina and terminate in a group of teeth,
but are not bidentate.

Michener: Comparative Studies of Bee Larvae 1065

AUodape ccratinoides Gribodo

The larva of this species is briefly described and figured by
Brauns ( 1926 ) . The head is of the usual shape with a few very long
hairs on the vertex. Antennae are not illustrated, perhaps they are
much reduced. The dorsolateral tubercles are absent while the
ventrolateral ones anteriorly are elongated to form "arms" in which
pollen is held during feeding. The dorsum of the larva is provided
with numerous setae.

AUodape strandi Masi

Masi ( 1930 ) described and figured the larva of this species. It is
similar to that of A. ccratinoides but differs in details of the "arms",
etc. It, too, has setae on the body.

AUodape marginata Smith

Yasumatsu (1938) has described and figured the larva of this
species. It is somewhat similar in general form to A. ccratinoides
but the ventrolateral tubercles ("arms") are absent. The head is
greatly broadened, the lateral projecting parietal regions being
provided with long curved hairs. The antennae are low. The palpi
are virtually absent. The mandibles are attenuate apically, acute,
with three to six small inner subapical teeth. The body is provided
with setae. As in A. ccratinoides, the last two body segments are
unusually elongate.

AUodape pringlei Cameron

The larva of this species is described and figured by Brauns
(1926). The head is of the ordinary shape, without long setae.
The antennal papillae are long. The labrum has distinctly pro-
duced, broad labral tubercles. No setae are shown on the body but
numerous small tubercles are present both dorsally and ventrally.

AUodape sp.

A distinctive type of AUodape larva described by Brauns ( 1926 )
( his third type ) has only two tubercles on each side of the body.

Exoneura spp.

The larvae of several species of Exoneura have been recently
described, as follows: E. simillima Bayment, frogattii Cockerell,
fultoni Cockerell, obliterata Cockerell (Bayment, 1949); concava
Bayment, sub-holmesi Bayment, roddi Bayment, montana Bayment,
variabilis Bayment, excavata Cockerell, and apposita Bayment ( Bay-

1066 The University Science Bulletin

ment, 1949a); rufitarsis Rayment, sitb-baculifera Rayment, richard-
soni Rayment, hamulata Cockerell (Rayment, 1951); and illustris
Erickson and Rayment, pictifrons Alfken (Erickson and Rayment,
1951). Unfortunately the numerous descriptions and figures are
not detailed, emphasis being placed on the armlike projections, ap-
parently modified ventrolateral tubercles, which are highly variable
in size, position, and number, much as in AUodape. No details of
the mouthparts are known. The antennal papillae are usually very
long, several times as long as broad. Presumably this is correlated
with the somewhat active life of the larvae in a common chamber
where they are fed progressively. One species ( siib-baculifera ) has
a conspicuous frontal projection on the head.

Subfamily Anthophorinae

That the bees which were placed in the Anthophorinae in a pre-
vious work (Michener, 1944) constitute a diverse, although probably
monophyletic, unit is evident enough from adult morphology. The
diversity is emphasized by larval morphology, and as pointed out
in the section on phylogeny, a rearrangement of the classification
may be indicated. However, at least until larvae of the Exoma-
lopsini can be studied, or until adult characters are found, such a
rearrangement will not be formalized.

Larvae of Nomada and Neopasites, especially the latter, share
most of their characters with the Halictidae and Andrenidae. For
example, the salivary opening is reduced, without lips, the antennal
papillae are reduced or absent, and the mandibles are acute at their
apices with small teeth on the margins and an inner multidentate
cusp. No inner apical concavity marked by a ridge is present.
Epeohis and Triepeolns are essentially similar except that the small
teeth of the mandibles are much reduced and the cusp is absent,
the distal portions of the mandibles being attenuate. Nomada and
the epeolines show the most extensive labiomaxillary reduction and
fusion seen in bee larvae. In all four of the above mentioned genera
the spiracular atria are produced above the body surface.

The other genera of Anthophorinae are quite different from those
discussed above in that the salivary opening is a slit, provided with
lips which are usually large but are much reduced in Anthophora.
The antennal papillae are distinct except in Anthophora. Each
mandible possesses an inner apical concavity demarked basally by
a distinct line or ridge. Small teeth are reduced or absent and the
multidentate cusp is absent, or in Melissodes indicated by a mere
remnant. The mandibular apices are usually blunt, often subtrun-

Michener: Comparative Studies of Bee Larvae 1067

cate, but may be nearly acute, as in some species of Anthophora,
or bidentate, as in Melissodes and Emphor.

The view that MeJecta arose from ancestors of Anthophora, as
indicated by several adult characters (see Michener, 1944), not
from Anthophora itself as suggested by Griitte (1935) and others,
is supported by larval characters. For example, Melecta has long
antennal papillae, larger palpi, a larger salivary opening, and larger
lips for this opening than Anthophora. All of these are presumably
primitive features among the bees of this group.

Exomalopsis caerulea Friese

Claude-Joseph (1926) briefly describes and figures the larvae of
this bee. It appears to have the caudal annulets of the body seg-
ments strongly elevated down to the level of the spiracles. The
antennal papillae and the palpi are elongated. The labrum is
strongly emarginate apically so that the labial tubercles are promi-
nent. The mandibles are robust, oblique or beveled apically; un-
fortunately nothing is known of their inner surfaces. The salivary
opening is transverse but short, not as long as the distance between
the labial palpi.

Neopasites sp?

(Figs. 180-185)

The larva is somewhat slender, the intersegmental lines absent
laterally and weak dorsally, the lines between the cephalic and the
caudal annulets of the segments absent unless the anterior margins
of dorsolateral tubercles represent them. The dorsolateral tubercles
are low, transverse, nearly meeting on the middorsal line. The
ventrolateral tubercles are absent.

Head capsule scarcely sclerotized, with a few minute scattered
hairs on clypeus and mouthparts; marginal thickening of head
capsule weak; epistomal suture indicated by weak concavity, no
thickening between anterior tentorial pits; cleavage line indicated
on upper part of head, arms not indicated; longitudinal median
thickening of head capsule absent. Antennae prominent convexi-
ties; head capsule above antennae with broad convexities suggestive
of those of the halictids; labrum without distinct tubercles although
distal margin has feeble convexities laterally which bear minute
setae arising from large pits. Mandible with apex acute, bearing
two large teeth on upper margin and some small teeth on both
margins; cusp distinct, bearing several large teeth. Maxillae with-
out distinct palpi; labial palpi mere convexities; salivary opening a
short slit without lips at apex of labial lobe.

1068

The University Science Bulletin

Spiracular atria without spines or ridges, produced above level of
body surface, strongly rimmed; peritreme large; primary tracheal
opening narrowed by distinct collar.

Lawrence, Kansas, June, 1951 ( Michener); described from a
single specimen obtained from a nest of Calliopsis andreniformis
and identified by comparison with a shed larval skin from which a
Neopasites pupa emerged. As no adult was obtained specific identi-
fication proved impossible.

Figs. 180-185. 180, Neopasites sp?, larva; 181, 182, dorsal and lateral views
of head of same; 183, 184, inner and ventral views of mandible of same; 185,
spiracle of same.

Michener: Comparative Studies of Bee Larvae

1069

Nomada (Nomada) fowled Cockerell

(Figs. 187-189)

This species agrees with Triepeohis in nearly all details, including
the body shape, the large and conspicuous spiracles, and the an-
atomical peculiarities of the head capsule. The following differ-
ences exist: The hypopharynx and labium are even more receding,
much overhung by the labrum; the external portion of the hypo-
pharynx is nearly round, not transverse, the salivary opening, there-
fore, is farther from the mouth than in Triepeolus. The mandibles
are thick basally, pointed and curved inward apically, their upper
and lower margins coarsely serrate apically. (The details of serra-
tion vary in the two mandibles of one individual. ) The cusp is
indicated by a group of teeth. The maxillary palpi are about twice
as long as their basal widths.

The body bears spicules in certain dorsal areas. The spiracular
atria lack spines. The atria are produced above the body surface,

Figs. 186-189.

186, Nomada fowleri, larva; 187, 188, ventral and inner views
of mandible of same; 189, spiracle of same.

1070 The University Science Bulletin

each with a rim. The peritreme is flat and the primary tracheal
opening is much narrowed by a collar.

Berkeley, California, June 1, 1946, from the nest of Andrena
complexa Viereck (MacSwain).

Packard ( 1897 ) gives a brief description of the larva of Nomada
imbricata Smith? It seems to agree in major features with the
species described above.

Triepeolus sp?

(Figs. 190-194, 197)

The larva is unusual in shape, being broadest at about the third
abdominal segment and tapering posteriorly from this point. The
intersegmental lines are distinct but the segments are not divided
into cephalic and caudal annulets. The dorsolateral tubercles are
entirely absent, the abdominal segments are swollen laterally, below
the level of the spiracles, and the integument is finely and trans-
versely wrinkled.

Head capsule and mouthparts with a very few scattered small
setae; head capsule weakly sclerotized, attached to thorax broadly,
without much constriction between head and thorax; marginal thick-
ening of head capsule very weak, posterior tentorial pits small and
inconspicuous, tentorium reduced to exceedingly delicate strands
(strands incomplete in specimens studied but perhaps complete in
younger specimens); epistomal suture faint, not represented by an
internal ridge; anterior tentorial pits weak, located near anterior
mandibular articulation; dorsal longitudinal median thickening of
head capsule absent; arms of cleavage line absent; parietal bands
present. Antennae mere convexities; labroclypeal suture weak,
labrum a thick rounded lobe bearing a pair of very distinct though
small tubercles. Mandibles thick basally, with attenuate and sclero-
tized apices which are feebly serrate on upper margins. Maxillae
reduced to a mere lobe on each side, much exceeded by labium,
indistinguishably fused to labium posteriorly and not separated by
a furrow but merely by the pigmentation of the hypostomal thick-
ening from the head capsule; apices of maxillae mere broad con-
vexities, separated from labium only by shallow furrow; maxillary
palpi slightly pigmented, about as broad as long; labium a broadly
rounded, thick lobe, exceeded by hypopharynx, the external portion
of which is a transverse convex area separated from labium by shal-
low furrow; salivary opening small, lunate (not slit-shaped), sur-
rounded by feebly projecting dark lips; labial palpi as broad as
long; furrow between postmentum and prementum absent; labio-
maxillary rod absent.

Michener: Comparative Studies of Bee Larvae

1071

Figs. 190-198. 190, Triepeolus sp? larva; 191, 192, lateral and dorsal views
oi head of same; 193, 194, inner and ventral views of mandible of same; 195,
196, inner and ventral views of mandible of Epeolus sp?; 197, spiracle of
Triepeolus sp? 198, spiracle of Epeolus.

38—6490

1072 The University Science Bulletin

Body without spicules or setae. Spiracles large, conspicuous,
pigmented, on slight protuberances; atria clearly defined, pig-
mented, shallow, densely lined with long spines, no particularly
large spines around primary tracheal opening, atrium projecting
well beyond level of body surface, this outer portion sloping inward;
peritreme flat; few small spines in what is presumed to be the distal
portion of the subatrium; no collar around primary tracheal opening.

Marsh Creek Canyon, Contra Costa County, California, Feb-
ruary 18, 1947, in nests of Melissodes sp? (MacSwain, Bohart).

Epeolus sp.

(Figs. 195, 196, 198)

Like Triepeolus sp? described above but smaller with the apical
portions of the mandibles shorter and thicker. The spiracles have
that portion of the peritreme which projects above the level of the
body surface erect, not sloping, and the primary tracheal opening
is provided with a collar. The subatrium lacks spines.

Bodega Bay, Sonoma County, California, January 14, 1948, in
nests of Colletes fulgidus Swenk (MacSwain).

Epeolus tristis Smith

This species is briefly described and figured by Mayet (1875).
The information available is not detailed, but the body form, large
protruding spiracles, slender mandibular apices, and fused maxillae
and labium agree with the Triepeolus and Epeolus available for
study.

Epeolus gaiji Spinola

Claude-Joseph ( 1926 ) describes and illustrates this species. The
larva is more slender than that of other known species of the genus,
but the head structure seems like that of North American Epeolus
and Triepeolus. The mandibles are much attenuate, like those of
the species of Triepeolus described above.

Isepeolus luctuosus (Spinola)

The larva is described and figured by Claude-Joseph ( 1926 ) . It
has a more tuberculate body than Epeolus, and the palpi are very
long. The salivary opening is a broad transverse slit, formed by
large lips, unlike Epeolus. The mandibular apices are attenuate.

Hemisia cineraria (Smith)

Claude- Joseph ( 1926 ) describes and figures the larva of this
species. It is shaped about in Anthophora, with the caudal an-

Michener: Comparative Studies of Bee Larvae 1073

nulets of the first five body segments having transverse dorsolateral
tubercles. Maxillary palpi appear to be distinct and subapical; the
labial palpi are absent. The mandibles are narrowly truncated at
their apices. The salivary opening is rather large and transverse.

Melissodes sp?

(Figs. 199-205)

The larva is about the same shape as that of Anthophora, with
the intersegmental lines weak, and the lines between the cephalic
and caudal annulets of the body even weaker except on the thorax.
The dorsolateral tubercles are weak posteriorly, being indicated by
broad convexities, extending down to level of spiracles and highest
only a little above that level, but become progressively higher
anteriorly, being quite conspicuous on the thoracic segments. The
fine transverse wrinkles on the body are very faint.

Head capsule white, unsclerotized, with a few scattered minute
setae on clypeus and mouthparts; marginal thickening of head cap-
sule delicate, narrower than in Anthophora; epistomal suture weaker
than in Anthophora, represented by a broad transverse sulcus, indi-
cated internally by a feeble, broad thickening; arms of cleavage line
very short or absent; longitudinal median thickening of head capsule
extending to epistomal suture or nearly so. Antennae conical, with
median, acutely pointed papilla; labrum much shorter than clypeus,
labial tubercles large, broad, rounded. Mandibles broad seen from
above or below, narrow apically seen from side of head, apices
minutely bidentate; cusp oblique, upper margin of mandible which
extends to cusp minutely denticulate, inner apical concave area of
mandible large; mandibles thus similar to those of Emphor; inner
upper surfaces of mandibles with dense group of hairs; outer sur-
faces of mandibles, maxillae and labium with scattered minute hairs.
Maxillae with palpi subapical, apices not much bent inward as in
Anthophora, palpi long and slender, a small rounded tubercle below
each; labium thick, salivary opening at apex of labial lobe; salivary
opening large, as broad as distance between labial palpi, with
projecting lips which are very large and apparently armed with
large teeth as in Emphor; labial palpi over twice as long as broad.

Body neither spiculate nor setose. Spiracular atria with spines
and a few ridges; atria not projecting above body surface, peritreme
large, collar around primary tracheal opening absent.

Marsh Creek Canyon, Contra Costa County, California, February
18, 1947 (Bohart, MacSwain).

1074

The University Science Bulletin

Figs. 199-205. 199, Melissodes sp?, larva; 200, 201, 202, dorsal, ventral,
and inner views of mandible of same; 203, 204, lateral and dorsal views of head
of same; 205, spiracle of same.

Tetralonia spp.

Claude-Joseph (1926) briefly describes and illustrates larvae of
T. chilensis Herbst, tristrigata Spinola, and melanura Spinola. All
are rather like the larva of Melissodes. Unfortunately details such
as mandibular structure are not recorded.

Michener: Comparative Studies of Bee Larvae 1075
Diadasia enevata (Cresson)

(Figs. 206-211)

The larva is more slender than that of Anthophora, frequently
more curved than in the specimen figured, the dorsolateral tubercles
of the body feebly represented by broad, ill-defined convexities.
Both the cephalic and the caudal annulets of the body segments
are transversely wrinkled.

Head capsule with scattered minute hairs. Marginal thickening
of head capsule weaker than in Anthophora, epistomal suture
weaker than in Anthophora, represented internally only by weak
broad thickening; arms of cleavage line absent; parietal bands
scarcely recognizable. Antennae with a slender papilla arising
from the broad feeble convexity; labral tubercles absent. Mandi-
bles short and robust, obliqely truncate, with apical concavity mar-
gined by a ridge, much as in Anthophora stanfordiana; outer sur-
faces of mandibles, maxillae, and labium with scattered minute
hairs. Maxillae with apical portions bent inward as in Anthophora;
maxillary palpi long and slender, far from apices of maxillae; labium
thick, salivary opening at apex of labial lobe; salivary opening
larger than in Anthophora, as broad as distance between labial
palpi, with thin projecting lips which are much longer than in
Anthophora; labial palpi dark, twice as long as broad; labium only
feebly divided into postmentum and prementum; labiomaxillary
rod slightly more heavily sclerotized than in Anthophora.

Body minutely spiculate, especially densely so dorsally, without
setae; spiracles with atrium broad and shallow, without spines or
ridges, wide open since peritreme is reflexed; primary tracheal
opening narrowed by a large collar.

Delta, Utah, June 27, 1950 (Bohart, Michener).

Melitoma chilensis Spinola

The larva, as described and figured by Claude-Joseph (1926),
is similar to that of Diadasia, with no evidence of tubercles on the
body. The mandibles are beveled at the apices and the salivary
opening is transverse.

Emphor bo7nbiformis (Cresson)

(Figs. 212-218)

The larva is more slender than that of Anthophora or Diadasia,
immature specimens being particularly slender. In immature lar-
vae the intersegmental lines are weak, absent laterally, and there

1076

The University Science Bulletin

Figs. 206-211. 206, Diadasia enevata, larva; 207, 208, lateral and dorsal
views of head of same; 209, 210, ventral and inner views of mandible of same;
211, spiracle of same.

Michener: Comparative Studies of Bee Larvae

1077

Figs. 212-218. 212, 213, Emphor homhiformis, last stage larvae of different
ages; 214, 215, lateral and dorsal views of head of same; 216, 217, ventral and
inner views of mandible of same; 218, spiracle of same.

is no division of the body segments into cephalic and caudal annu-
lets. In prepupae the intersegmental lines are more distinct and
the anterior body segments are somewhat swollen just below the
level of the spiracles. The dorsolateral tubercles are absent but
each abdominal segment bears a transverse median dorsal eleva-

1078 The University Science Bulletin

tion, possibly produced by confluence of dorsolateral tubercles.
Except for these elevations, most of the body surface is finely trans-
versely wrinkled in the prepupa.

Head capsule with scattered minute setae. Marginal thickening
of head capsule distinct, hypostomal and pleurostomal portions
particularly dark; epistomal suture weaker than in Anthophora,
represented internally by weak, broad, thickening; arms of cleav-
age line absent or short; parietal bands distinct. Antennae with
slender papilla and broad basal convexity; labral tubercles absent
or nearly so, distal half of labrum densely setose. Mandibles ro-
bust, with a few minute setae on outer surfaces; apices bidentate,
cusp very oblique seen in inner view (as in Melissodes), connected
by a ridge or carina to upper apical tooth, thus partially enclosing
a subapical inner concavity. Outer surfaces of maxillae with scat-
tered small hairs, apical portions of maxillae extending inward as
in Anthophora, maxillary palpi long and slender, far from apex of
maxilla; labium thick, the salivary opening at apex of labial lobe;
salivary opening larger than in Anthophora, as broad as distance
between labial palpi, with thin projecting lips which are larger
than in Anthophora; labial palpi pale, longer than broad; line be-
tween postmentum and prementum weak; labiomaxillary rods more
distinct than in Anthophora.

Body neither spiculate nor setose; spiracles with atria devoid of
spines but with weak ridges, atrium much narrowed near body
surface, projecting beyond body surface to form strong rim; peri-
treme flat; primary tracheal opening with collar.

Hattiesburg, Mississippi, August 20, 1944 (Michener). Some
half grown larvae are from Washington, D. C, July 9, 1935 ( Sand-
house and Brown).

Anthophora stanfordiana Cockerell

(Figs. 7-15, 219-222, 225)

Since the larva of this species is rather fully described from a
morphological standpoint in an earlier part of this paper, it seems
unnecessary to repeat characteristics here. Certain features not
mentioned in that description are as follows:

Labroclypeal suture angulate upward medially; labral tubercles
low, separated by a long, nearly straight, papillate margin. Man-
dibles robust, truncate at apices, with distinct inner apical con-
cavity. Maxillae with palpi weak, broader than long, far from
apices of maxillae; salivary opening much smaller than in such forms
as Emphor, shorter than distance between labial palpi, on dorsal

Michener: Comparative Studies of Bee Larvae

1079

226

Figs. 219-226. 219, 220, Lateral and dorsal views of head of Anthophora
stanfordiana; 221, 222, inner and ventral views of mandible of same; 223, 224,
inner and ventral views of mandible of Anthophora urhana; 225, spiracle of
Anthophora stanfordiana; 226, spiracle of Anthophora urhana.

1080 The University Science Bulletin

surface of labial lobe rather than at its apex, provided with small
projecting lips; labial palpi much broader than long.

Montara, California, May, 1940 (MacSwain); Bodega Bay, So-
noma County, California, January 14, 1948 (MacSwain); Sommers-
ville and Nortonville, Contra Costa County, California, February
18, 1947 (Bohart, MacSwain, Hurd).

Anthophora urbana Cresson

(Figs. 223, 224, 226)

This species differs from A. stanfordiana in the slightly higher
dorsolateral tubercles of the body. In this respect it resembles
A. edwardsii. The mandible is narrower apically than in A. stan-
fordiana and the upper margin is feebly crenulate, suggesting the
serrations found in some species. The spiracles have fewer rows
of atrial spines than in A. stanfordiana, the inner ones being smaller,
while the toothed collar spines nearly meet in the center, closing
the primary tracheal opening. The shape of the atrium is distinc-
tive, for it is narrowed toward the surface (fig. 226) so that much
of the inner wall is invisible from the surface. The atrial wall is
more produced above the body surface than in A. stanfordia7ia and
lacks a rim.

Montara, California, May 18, 1940 (MacSwain) (a single speci-
men).

Anthophora edwardsii Cresson

(Figs. 228, 234, 235)

The mature larva differs significantly from A. stanfordiana in the
slightly more convex and higher dorsolateral tubercles of the body
and in the spiracles (figure 235). The atrial spines, except for those
which cover the primary tracheal openings, are shorter than many
of those of stanfordiana, while the collar spines do not meet to
close the primary tracheal opening; the atrial walls form no rim
but converge above the level of the body surface.

One mile south of Pittsburg, California, May 21, 1940 (Mac-
Swain) and five miles north of Madera, California, March 29, 1941
(Linsley, MacSwain).

Anthophora abrupta Say

(Figs. 232, 233)

The mature larva differs from that of A. stanfordiana in the low
dorsolateral tubercles, in which it resembles A. edwardsii and A.
urbana. The mandible is very broadly truncate, the upper surface
spiculate medially. The spiracles resemble those of A. urbana but

Michener: Comparative Studies of Bee Larvae 1081

Figs. 227-237. 227, spiracle of Anthophora linsleyi; 228, spiracle of Antho-
phora edwardsii; 229, spiracle of Anthophora furcata syringae; 230, 231, inner
and ventral views of mandible of same; 232, 233, ventral and inner views of
mandible of Anthophora abrupta; 234, 235, inner and ventral views of mandible
of Anthophora edwardsii; 236, 237, inner and ventral views of mandible of
Anthophora linsleyi.

1082 The University Science Bulletin

are not narrowed at the surface, the atria being shaped as in A.
stanfordiana.

Alexandria, Virginia, July 20, 1935 (Sandhouse) (a single speci-
men).

Anthophora linsleyi Timberlake

(Figs. 227, 236, 237)

The mature larva is smaller than that of A. stanfordiana and has
slightly more convex dorsolateral tubercles, much as in A. edwardsii.
The mandibles are more slender than in stanfordiana, narrowly
truncated at the apices and the upper margins apically are coarsely
serrate (figure 236); the upper surfaces, medially, are spiculate.
The spiracular atrium has the usual large spines nearly covering
the primary tracheal opening, but the atrial walls above the spines
have fewer spines than in A. stanfordiana.

Twenty miles east of Bakersfield, California, March 29, 1941
(Linsley and MacSwain).

Anthophora personata (Illiger)

Semichon (1925) briefly describes this species. It evidently
agrees in its main features with the various species described above.

Anthophora incertus Spinola

Claude-Joseph ( 1926 ) describes the larva which is evidently simi-
lar to that of North American species. The mandibles are truncate,
as in A. stanfordiana.

Anthophora villostda Smith

The larva is described by Torikata (1931). It apparently re-
sembles the various species described above.

Anthophora (Clisodon) furcata syringae (Cockerell)

(Figs. 229-231)

This species is most similar to A. linsleyi, with which it agrees
in the serrate upper apical margins of the mandibles. The mandible
is more robust, however, than that of A. linsleyi, with a translucent,
rounded, subapical projection on the lower margin, and with the
apex rounded, not truncate. Specimens at hand are somewhat
shriveled so that the height of the dorsolateral tubercles cannot be
determined but they appear to be about as in A. linsleyi, slightly
higher than in A. stanfordiana. The spiracles are essentially like
those of edwardsii.

Mineral King, Tulare County, California, August, 1939 (Bohart).

Michener: Comparative Studies of Bee Larvae

1083

Melecta (Melectomorpha) calif ornica Cresson

(Figs. 238-242)

The body form of the larva, to judge by the two shriveled speci-
mens available, is similar to that of Anthophora stanfordiana. The
dorsolateral tubercles are transverse, nearly meeting middorsally, and
are slightly higher and more conspicuous than in Anthophora. Both
the cephalic and the caudal annulets of the body, except for the
dorsolateral tubercles, are transversely wrinkled.

Head capsule rather strongly sclerotized, with scattered minute
hairs. Marginal thickening of head capsule strong; epistomal suture
weak, represented externally by shallow transverse depression, in-
ternally by a cuticular thickening; arms of cleavage line absent.

Figs. 238-242. 238, 239, lateral and dorsal views of head of Melecta cali-
jurnica; 240, 241, ventral and inner views of mandible of same; 242, spiracle of
same.

1084 The University Science Bulletin

Antennae with rather large papillae; labroclypeal suture particularly
distinct; labral tubercles broad and rounded so that entire apex of
labrum is broadly emarginate, apical portion of labrum rather
densely setose. Mandibles tapering from broad bases to blunt
apices, upper apical margin feebly serrate, apical portion of man-
dible concave on inner surface, this concavity not margined by such
distinct ridges as in Antlwphora stanfordiana; outer surfaces of
mandibles, maxillae, and labium with scattered minute hairs. Max-
illae short and broad, much exceeded by labium, apices not bent
inward as in Antlwphora but maxillary palpi slightly preapical,
these palpi large, twice as long as basal width; salivary opening at
apex of labial lobe; salivary opening not as broad as distance be-
tween labial palpi, lips long and distinctly sclerotic, much larger
than in Antlwphora; hypopharynx well defined; labial palpi dark,
longer than broad; division between postmentum and prementum a
deep furrow; labiomaxillary rod rather distinct.

Body without spicules or setae, a perpendicular flat area above
anal opening. Spiracular atria with weak internal ridges, walls
slightly projecting above level of body surface, with rims; peritreme
flat; primary tracheal opening surrounded by thick collar.

Specimens are from Montara, California, May, 1940, in nests of
Antlwphora (probably stanfordiana) (MacSwain).

Melecta (Melecta) armata Panzer

This species is briefly described but not figured by Semichon
(1925). Evidently it does not differ greatly from that of M. cali-
fornica.

Subfamily Apinae

The larvae of this subfamily exhibit considerable diversity, due
perhaps to reductions of structures correlated with social life. The
antennal papillae are largest in Centris, small in Bombtis and Psi-
thyrus, virtually absent in Melipona, Trigona, and Apis. The palpi
are present in all genera although largest in Centris. The mandibles
lack a cusp. They have an inner apical concavity, clearly defined, in
Centris, Bombus, and Psithyrus. In Melipona such a concavity is
present although less conspicuous, in Trigona it is inconspicuous or
absent, while in Apis it is entirely absent. The mandibular apices
are bidentate, with the upper apical tooth much smaller than the
lower, in Bomhas and Psithyrus; in Centris the upper tooth is re-
duced to a mere tubercle well away from the blunt apex of the
lower tooth; the remaining genera have the mandibular apex blunt
with no evidence of two teeth but in Melipona and Trigona with

Michener: Comparative Studies of Bee Larvae 1085

coarse setae or long spicules. The mandibles of Melipona are rather
slender, those of Trigona conspicuously attenuate; those of Apis
are more reduced than in any other bee larva known. In all the
Apinae the salivary opening is a transverse slit provided with large
lips. The larva of Apis is one of the few forms examined in which
the spiracular peritreme is entirely absent. (It is also absent in
some Lasioglossum, incomplete in Trigona and other Lasioglossum ) .
The larvae of Centris, Bomhus, Psithyrus, and Melipona, differ from
those of other bees in having the dorsolateral tubercles of the tho-
racic segments represented by small points. Centris is remarkable
for having a pair of similar points on the head. Trigona has such
points on most of the body segments, arising from the summits of
broad rounded tubercles. It is surprising that there are a few scat-
tered setae on the body in Bombus. They are shown clearly by
Grandi ( 1934a ) and are present in B. americanorum.

Centris (Enplusia) violacea (Blanchard)

(Figs. 243-247)

The only specimen available is one which was killed as it was
molting into the pupal stage so that details as to larval form cannot
be established. It is obvious, however, that the intersegmental lines
are rather inconspicuous. No lines are visible separating the
cephalic from caudal annulets of the segments. Each of the thoracic
segments bears a pair of minute, dark, and slightly sclerotized
pointed dorsal or dorsolateral tubercles. A pair of similar tubercles
occurs on the upper part of the head capsule.

Head capsule lightly sclerotic, the thickening of the posterior
margin of the head capsule virtually absent but remainder of mar-
ginal thickening of head capsule distinct; posterior tentorial pits
apparently inconspicuous, the posterior tentorial arms short and
robust; epistomal suture not represented by a thickening between
anterior tentorial pits but probably represented by a depression in
fresh or properly preserved larvae. Antennae each with clearly
defined papilla arising from broad convexity; labroclypeal suture
weakly evident in shed skin; labrum with apex emarginate medially,
no distinct labral tubercles, apical portion with numerous minute
setae. Mandibles short and robust, heavily sclerotized, apices
bluntly rounded; inner surfaces each with a broad apical concavity
margined by a ridge, upper margin with a large tooth midway the
length of the concavity. Apical portions of maxillae rather small,
not bent inward but with a small apical projection immediately

1086

The University Science Bulletin

Figs. 243-247. 243, 244, lateral and dorsal views of head of Centris violacea;
245, 246, ventral and inner views of mandible of same; 247, spiracle of

same.

mesad of base of each maxillary palpus, the latter nearly three times
as long as broad; labial lobe with salivary opening at apex, a trans-
verse slit provided with rather large sclerotized lips, width of slit
equal to distance between labial palpi which are about twice as long
as broad; labium and maxillae both with rather numerous setae;
hypostomal furrow very deep.

Body without setae or spicules. Spiracle with atrium shallow,
without spines or ridges, scarcely projecting above body surface but

Michener: Comparative Studies of Bee Larvae 1087

with weak rim; peritreme flat; primary tracheal opening narrowed
by collar.

Brasil (W. E. Kerr).

The name Centris is here used for the group usually called
Eulaema.

Bombus (Fervidobombus) americanorum Fabricius

(Figs. 248-253)

The larva is robust with the intersegmental lines weak, absent on
the sides, and the lines between the caudal and the cephalic annu-
lets of the body segments weak. The dorsolateral tubercles are
absent except for minute conical ones on the thoracic segments,
similar to those of Centris. Head capsule rather distinctly sclerotic,
conspicuously constricted in front of thorax, marginal thickening
of head capsule rather weak but posterior tentorial pits distinct,
epistomal suture represented by an indistinct thickening arching
between anterior tentorial pits, longitudinal median thickening of
head capsule present and reaching epistomal suture. Antennae
represented by convexities, each with minute low median papilla;
labroclypeal suture distinct laterally but weak medially; labrum
deeply emarginate medially and bearing exceedingly minute setae.
Mandibles short and rather heavily sclerotized, apices bluntly
rounded with a blunt subapical tooth on upper margin; apical por-
tions deeply concave on inner surface, concavity except basally
delimited by a sharp ridge. Maxillae not bent inward apically,
maxillary palpi about twice as long as broad; labial lobe with sali-
vary opening at apex, this opening marked by weakly sclerotic
lips, the opening nearly as broad as distance between labial palpi,
the latter nearly twice as long as broad; hypostomal furrow deep;
outer surfaces of maxillae and labium with scattered setae.

Body with spicules and a very few scattered setae. Spiracles
with atrium rather shallow, inner wall with spines, mostly in groups,
collar spines very long and many of them toothed at apices; pre-
atrium not annulate; peritreme flat.

Lawrence, Kansas, August 20, 1950 (Michener).

This species has also been described by Ritcher (1933).

Bombus (Fervidobombus) fervidus (Fabricius)

This species agrees with B. americanorum except in details of the
spiracular atria (see Ritcher, 1933). Its general features were also
described by Packard ( 1897 ) .
39—6490

1088

The University Science Bulletin

h\^

;iUii^x-_N

Figs. 248-254. 248, Bombus americanorum, larva; 249, 250, dorsal and
lateral views of head of same; 251, 252, inner and ventral views of mandible
of same; 253, spiracle of same; 254, spiracle of Bombus vosnesenskii.

Bombus (Separatobombus) griseocollis (DeGeer)

This species was described by Ritcher (1933) under the name
separatus Cresson. It resembles B. americanorum except in certain
details of the spiracular atria.

Michener: Comparative Studies of Bee Larvae 1089

Bombus (Bombus) terricola Kirby

This species resembles B. americanorum except in certain details
of the spiracular atria (see Ritcher, 1933).

Bombus (Pratobombus) vosnesenskii Radoszkowski

(Fig. 254)

Specimens at hand differ from those of Bombus americanoriim in
showing feeble intersegmental lines across the pleural regions of
the body 1 as well as on the sternal and tergal regions. Spiracles
with spines isolated, not in groups as in americanoriim, no large
collar spines, collar region not dilated as in americanoriim.

Berkeley, California, March 22 and June 4, 1946 (MacSwain);
Hat Creek, Lassen County, California, June 4, 1941 (Linsley and
Michener ) .

The larva of this species runs to the vicinity of B. impatiens and B.
bimaculatus in Ritcher 's ( 1933 ) key.

Bombus (Pratobombus) impatiens Cresson

This species resembles B. americanoriim except in the details of
spiracular atria (see Ritcher, 1933).

Bombus (Pratobombus) bimaculatus Cresson

This species resembles B. americanorum except in the details of
spiracular atria (see Ritcher, 1933).

Bombus (Pratobombus) perplexus Cresson

This species resembles B. americanorum except in the details of
spiracular atria ( see Ritcher, 1933 ) .

Bombus (Pratobombus) vagans Smith

This species resembles B. americanorum except in the details of
spiracular atria (see Ritcher, 1933).

Bombus (Agrobombus) agrorum pascuorum Scopoli

This species was described and illustrated by Grandi (1937).
It resembles B. americanorum except for details of the spiracular
atria.

Bombus (Agrobombus) senilis Smith

Sakagami ( 1951 ) has illustrated the larva of this species. It
resembles the majority of Bombus larvae. The spiracular atria are

1. The larvae are all rather small, and almost equally distinct lines are visible in small
individuals of B. americanorum. The lines across the pleural regions of the body are not
as distinct as in the subgenus Bombias.

1090 The University Science Bulletin

said to be similar to those of B. fervidus. The illustration of the
mandible shows the teeth more acute than in other Bombus, but
not as acute as in Psithyrus.

Bombus (Mastrucatobombus) mastrucatus Gerstaecker

The head and mandibles are well figured by Moczar ( 1938 ) but
unfortunately the spiracular details are not illustrated.

Bombus silvarum Linnaeus

The larva of this species is described and figured by Grandi
(1934a). It evidently resembles B. americanorum in most of its
characters, differing, however, in the details of the spiracular atria.
The intersegmental lines are rather clearly illustrated in the pleural
regions of the body by Grandi, perhaps they are more distinct
than in B. americanorum. The mandibles have the lower apical
margin minutely denticulate, unlike other known Bombus.

Bombus (Bombias) auricomus Robertson

The larva of this species was described by Ritcher (1933). It
is the only known larva of the subgenus Bombias. It closely re-
sembles B. americanorum but differs from it and all other known
Bombus larvae in the greatly reduced atrial spines and in the pres-
ence of intersegmental lines across the pleural regions of the body.

Psithyrus variabilis (Cresson)

The larva of this species is similar to that of Bombus ameri-
canorum but differs in having the apical and subapical mandibular
teeth acute, falcate ( see Ritcher, 1933 ) . The details of the spiracu-
lar atria also differ from those of any known species of Bombus, but
do not seem to provide a generic character. The atrial spines are
long, as in most known species of Bombus, not short or absent as in
the subgenus Bombias.

Melipona quadrifasciata quadrifasciata Lepeletier

(Figs. 255-259)

The larva is robust, similar in shape to that of Bombus, exhibiting
weak intersegmental lines and even weaker lines between the
cephalic and caudal annulets of the body segments but showing
somewhat distinct ventrolateral tubercles below the level of the
spiracles on the sides of the body, particularly on the abdominal
segments. The thoracic segments have minute conical dark and
slightly sclerotized dorsal or dorsolateral tubercles similar to those
found in Centris and Bombus.

Michener: Comparative Studies of Bee Larvae

1091

Figs. 255-265. 255, 256, lateral and dorsal views of head of Melipona
quadrifasciata; 257, 258, inner and ventral views of mandible of same; 259,
spiracle of same; 260, spiracle of Melipona marginata; 261, 262, ventral and
inner views of mandible of same; 263, spiracle of Melipona variegatipes; 264,
265, ventral and inner views of mandible of same.

1092 The University Science Bulletin

Head capsule weakly sclerotized with a distinct constriction be-
tween it and thorax. Marginal thickening of head capsule feeble,
head essentially as in Bomb us but anterior margin of clypeus con-
vexly truncated; lower portion of labrum with numerous scattered
minute setae; outer surfaces of maxillae and labium with scattered
minute setae, the former spiculate above. Apical portions of man-
dibles more slender than in Bombus, apices bluntly pointed without
subapical dorsal tooth; inner apical concavity rather shallow but
distinct; mandibles abundantly spiculate in certain areas, some
large spicules or small spines on upper margin subapically. Max-
illary and labial palpi slightly longer than broad; labium with sali-
vary opening broad, formed by distinct lips about as broad as
distance between labial tubercles. Spiracular atria thin walled, shal-
low, not produced above body surface, inner surfaces with ridges;
primary tracheal opening without a collar, peritreme (in specimens
examined) slightly inflexed.

Brasil (W.E.Kerr).

Melipona variegatipes Gribodo

(Figs. 263-265)

The larva is similar to that of M. quadrifasciata although in the
poorly preserved specimens at hand the ventrolateral swelling or
row of tubercles obvious in quadrifasciata is not visible. The head
capsule and mouth parts seem to be similar to those of quadri-
fasciata except that the apical portions of the mandibles are broader
and fringed with small spines while elsewhere the mandibles are
not or scarcely spiculate. The spiracular atria are heavier walled
and more spherical than in quadrifasciata, with fewer ridges, and
the annulations of the preatrium are poorly developed.

Montserrat, British West Indies, March 3, 1894 (Hubbard).

Melipona marginata Lepeletier

(Figs. 260-262)

This species agrees with the description of M. quadrifasciata ex-
cept that the labroclypeal suture is distinct throughout its entire
length (this appears to be a matter of individual variation). The
mandible is more robust than in quadrifasciata with spicules or
teeth on the upper portion subapically but margin otherwise smooth.
The spiracular atrium has very few ridges.

Specimens are from Brasil collected by Kerr.

Michener: Comparative Studies of Bee Larvae

1093

Trigona (Partamona) cupira Smith

(Figs. 266-270, 273)

The larva is rather robust with the intersegmental lines weak,
though distinctly recognizable, and with the lines separating the
caudal and the cephalic annulets of the segments also weak but

Figs. 266-274. 266, Trigona cupira, larva; 267, 268, inner and ventral views
of mandible of same; 269, 270, lateral and dorsal views of head of same; 271,
272, ventral and inner views of mandible of Trigona corvina; 273, spiracle of
Trigona cupira; 274, spiracle of Trigona corvina.

1094 The University Science Bulletin

recognizable. The dorsolateral tubercles are present, of moderate
size and rounded, high on the body and not extending downward
toward the tubercles. All these tubercles except for those of the
last two abdominal segments terminate in small conical papillae
which are largest on the thoracic segments where they correspond
to similar tubercles or papillae found on the thorax in Melipona,
Bombus, and Centris. The head is similar to that of Melipona
quadrifasciata, the apex of the labrum being shallowly and broadly
emarginate. The antennae are broad convexities without distinct
papillae. The labroclypeal suture is continuous. The maxillary
and labial palpi are small, about as long as broad. The mandibles
have slender and attenuate apices, slightly curved inward, without
margined concave spaces on the inner surfaces. The apices bear
a comb of teeth and the lower margins are spiculate. The body
is spiculate. The spiracular atria are shallow, with internal ridges
but no spines. They are not produced above the level of the body
surface. The peritreme is incomplete. The primary tracheal open-
ing lacks a collar.

Specimens are from Juan Mina, Canal Zone, May 4, 1945 ( Mich-
ener ) .

Trigona (Trigona) corvina Cockerell

(Figs. 271, 272, 274)

The specimens at hand are poorly preserved but in so far as can
be determined the body form closely resembles that of T. cupira.
As in that species, much of the body is densely spiculate. The
mandibles are more robust and are provided with minute denticles
or spines on both the upper and the lower margins near and around
the apices, and are considerably spiculate. The spiracles are fun-
damentally like those of T. cupira although they differ in details,
such as the very short preatrium, which may or may not be of
specific importance.

Juan Mina, Canal Zone, April 27, 1945 ( Michener ) .

Apis mcllifera Linnaeus

(Figs. 275-280)

The larva is robust with the intersegmental lines conspicuous
and the lines between the cephalic and caudal annulets of the seg-
ments visible. The caudal annulets are elevated laterally, the ele-
vations being continuous with rather distinctly elevated ventro-
lateral tubercles below the level of the spiracles. The thorax and
even the first abdominal segment have transverse dorsolateral tuber-
cles which are progressively smaller toward the rear.

Michener: Comparative Studies of Bee Larvae

1095

Figs. 275-280. 275, Apis mellifera, larva; 276, 277, lateral and dorsal views
of head of same; 278, 279, ventral and inner views of mandible of same; 280,
spiracle of same.

1096

The University Science Bulletin

Head capsule scarcely sclerotic, separated by slight constriction
from thorax; setae on head capsule and mouthparts very short and
sparse. Marginal thickening of head capsule feeble; epistomal
suture not clearly recognizable between anterior tentorial pits;
dorsal longitudinal thickening of head capsule present posteriorly.
Antennae distinct, dome shaped, without papillae; labroclypeal
suture distinct; apex of labrum broadly concave and bearing minute
setae and elongate spicules. Mandibles feebly sclerotic, bluntly
pointed, without concavity on inner surface apically. Maxillae with
apices not bent inward, maxillary palpi longer than broad; labium
exceeding maxillae; salivary opening at its apex, the opening a

Figs. 281-287. 281, 282, dorsal and lateral views of head of Hesperapis
rufipes; 283, larva of same; 284, 285, ventral and inner views of mandible of
same; 286, lateral view of salivary opening and salivarium of same; 287, spiracle
of same.

Michener: Comparative Studies of Bee Larvae 1097

transverse slit margined by slightly sclerotic but large lips, the slit
equal in length to distance between bases of labial palpi; labial palpi
longer than broad; hypostomal furrow deep; prementum and post-
mentum distinctly separated by a furrow.

Spiracular atria shallow, without internal ridges or spines; peri-
treme absent; primary tracheal opening without collar; preatrium
very short and scarcely annulate.

The larva of this species has been described and figured in a
general way by many workers. The first detailed figures of the
head structures are those of Grandi ( 1934, 1934a ) .

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THE UNIVERSITY OP KANSAS

SCIENCE BULLETIN

Vol. XXXV, Pt. II] September 10, 1953 [No. 9

A Revision of the North American Species of Typhlocyba
and its Allies (Homoptera, Cicadellidae ) x

By Paul J. Christian 2

Abstract: This revision includes descriptions of the Typhlocybid genera
Henribautia, Ribautiana, Ossiannilssonola, Typhlocyba, Empoa, Edwardsiana,
and a new genus Mcateeana (Homoptera, Cicadellidae), with redescriptions of
all of the known North American species of each, keys to the species, new illus-
trations, new host records, new distribution records, and descriptions and illus-
trations of the following twenty-two new species: Henribautia beameri;
Ribautiana multispinosa, R. parapiscator; Ossiannilssonola bangsoni, O. knulli,
O. mcateei, O. rossi; Empoa acericola, E. caryata, E. elmata, E. latifasciata,
E. platana; Typhlocyba alabamaensis, T. attenuata, T. inftata, T. medleri, T.
transviridis; Edwardsiana dejecta, E. delongi, E. nigripennis, E. projecta, E.
pseudocommissuralis.

Typhlocyba jacobii nom. nov. for T. duplicata Jacobi, nee T. duplicata Mc-
Atee is proposed, and Ossiannilssonia Young and Christian, nee Lambers, is
replaced by Ossiannilssonola nom. nov.

Edwardsiana plebeja and E. frustrator are recorded for the first time as
occurring in North America.

New synonymy is as follows : Ribautiana unca ( McAtee ) ( = Typhlocyba
surda DeLong and Johnson); Ossiannilssonola australis (Walsh) (= Typhlocyba
nicarete McAtee), O. danae (McAtee) (= Typhlocyba eurydice McAtee),
O. appendiculata (Malloch) (= Typhlocyba gillettei var. sellata McAtee);
Typhlocyba rubriocellata Malloch (= Typhlocyba escana Ross and DeLong),
T. persephone McAtee ( = Typhlocyba lancifer McAtee ) ; Empoa albicans
Walsh ( = Typhlocyba cijmba var. pollens McAtee ) ; Edwardsiana australis
Froggatt replaces the name Typhlocyba froggatti (Baker).

TABLE OF CONTENTS

PAGE

Introduction 1104

Historical Account 1 105

Acknowledgments 1107

Genitalic Characteristics 1110

Biological Notes 1110

Technique 1113

1. Contribution number 804 from the Department of Entomology of the University of
Kansas, Lawrence.

2. Present address, Department of Biology, University of Louisville, Louisville, Kentucky.

(1103)
40—6490

1104 The University Science Bulletin

PAGE

Systematic Account 1114

Introduction to the Typhlocyba Complex 1114

Key to the Genera of the Typhlocyba Complex 1114

The Genus Henribantia 1115

Key to the Species of Henribautia 1116

The Genus Ribautiana 1119

Key to the Species of Ribautiana 1120

The Genus Mcateeana 1 130

The Genus Ossiannilssonola 1132

Key to the Species of Ossiannilssonola 1133

Color Key to the Species of Ossiannilssonola 1135

The Genus Typhlocyba 1 160

Key to the Species of Typhlocyba 1161

The Genus Empoa 1187

Key to the Species of Empoa 1190

The Querci Group 1 192

The Casta Group 1 196

The Gillettei Group 1199

The Vestita Group 1204

The Spinosa Group 1205

The Albicans Group 1206

The Genus Edwardsiana 1208

Key to the Species of Edwardsiana 1209

Discussion of the Commissuralis Complex 1228

Glossary of Terms 1230

Literature Cited 1232

Index to Genera and Species 1236

Plates 123S

INTRODUCTION

The purpose of this revision is to define more clearly each of
the genera of the Typhlocyba Complex which contain species oc-
curring in North America, and to present clear descriptions, illus-
trations, keys, and new synonymy for the known North American
species of these genera of leaf hoppers.

The genera of the Typhlocyba Complex included are as follows:
Typhlocyba, Empoa, Edwardsiana, Ribautiana, Henribautia, Os-
siannilssonola and a new genus Mcateeana.

New descriptions and illustrations of the male genitalia are given
for nearly all of the previously described species, for two European
species not previously recorded from North America, and for
twenty-two new species.

New host records, biological notes, and locality records have been
included under each species subsequent to its description, and for
many species these add greatly to our knowledge of the habits and
distribution of the species.

Revision of the Species Typhlocyba 1105

HISTORICAL ACCOUNT

The first revision of the species now placed in the Typhlocyba
Complex was made by W. L. McAtee in 1926, who gave descrip-
tions of the known North American species, all of which were then
included in Typhlocyba, and described twenty-three new species
and sixteen new varieties, with figures of the male genitalia for each
species. The descriptions of two European species were also in-
cluded although no North American specimens of these species had
been seen. Much of the work included in this revision was accurate,
but due to insufficient material to work with, lack of proper equip-
ment, or to insufficient time, a number of inaccuracies occurred.

Since McAtee's revision a number of papers, which have added to
our knowledge of the genus Typhlocyba, have appeared in various
parts of the world. In 1928, Edwards moved the British species of
Typhlocyba into the genus Anomia Fieber, and in 1929 Jazykov
(Zachvatkin) proposed the genus Edwardsiana with Cicada rosae
Linnaeus as the type species. Ribaut redescribed the French spe-
cies of the Ulmi and Rosae Groups in 1931, with the description of
several new species. The Oriental species of Typhlocyba were
described by Matsumura in 1931-32, but without descriptions of the
internal male genitalia. Wagner reviewed the species for Northern
Germany in 1935, while in the same year Ossiannilsson reviewed
the Typhlocyba species occurring in Sweden. In 1936 Ribaut re-
described all of the French species of Typhlocyba with illustrations
of the male genitalia for each. Jacobi described a number of spe-
cies of Typhlocyba from Lombok Island and adjacent islands in
1941.

The British species were brought up to date by China in 1943,
who redescribed those species not considered by Ribaut ( 1936 ) .
In 1946 Dlabola recorded four species new to Bohemia. The genus
Ribautiana was proposed in 1945 (1947) by Zachvatkin, along with
the descriptions of new species of Edwardsiana. In 1949 Linnavouri
described several species which had not been previously recorded
for Finland. In 1950 China listed the British species, giving recog-
nition to the genera Edwardsiana and Ribautiana, and in the same
year Dlabola ( 1950 ) revised the leafhoppers from central Europe
in Melichar's Collection, listing the species of Typhlocyba in the
collection and indicating previous misdeterminations.

While this work was going on in other parts of the world, work
was continuing on the North American species as well. New spe-
cies were described by a number of workers: one by DeLong

1106 The University Science Bulletin

(1926), one by Osborn (1928), one by DeLong and Davidson
(1934), one by Ossiannilsson (1936), six by DeLong and Johnson
(1936), one by Medler (1942), three species and one subspecies
by Beamer (1943), six by Knull (1944), two by Knull (1945), and
five by Ross and DeLong ( 1949 ) . In 1949 a list of synonyms and
references for another European species, of economic importance
to cultivated prune, was made by Oman ( 1949a ) . Two new intro-
ductions of European species were reported recently, one in 1949 by
Ross and DeLong, and the other by Andison, 1950.

With this increase in the number of North American species,
some of which were known to be synonyms of previously described
species and others whose identity was held in question, it was im-
perative that a revision be made in order to clear up the identity
of all of the known species. While undertaking the revision of this
genus the author has tried to use those methods which would permit
the most accurate observations, and has included illustrations of
those structures which best characterize the species.

Since the work on this genus was begun, the genera Henribautia
Young and Christian, and Ossiannihsonia Young and Christian,
have been segregated from Typhlocyba (see Young, 1952). The
study of additional material subsequent to the completion of that
paper has led the author to regard the species sexnotata, previously
placed in the genus Ossiannihsonia, to have characters which jus-
tify the establishment of a new genus in which it has been placed.

Further consideration of the species of Typhlocyba, which on the
basis of aedeagal structure appeared to be intermediate between
the species in the Rosae Group and other species of Typhlocyba,
has led the author to believe that the absence of atrial processes in
such species as T. tortosa, T. persephone, T. niobe, and T. sollisa is
due to a fusion of these with the aedeagal shaft, as is partly seen in
T. athene. In species of the Rosae Group there is no indication
that atrial processes have ever existed. When considered from this
aspect, species of the Rosae Group are seen as having the aedeagus
fundamentally different enough to set them off from Typhlocyba
as a separate genus (Edwardsiana Zachvatkin). The group of
species near and including querci Fitch has, on the basis of the
structure of the aedeagus and pygofer, also been recognized as a
genus distinct from Typhlocyba and has been segregated as the
genus Empoa Fitch.

Revision of the Species Typhlocyba 1107

Ossiannilssonola, nom. nov.

The name replaces Ossiannilssonia Young and Christian in Young,
Bull. Univ. Kansas Sci. Bull, vol. 35, pt. 1, July 1, 1952, p. 97 (nee
Lambers, Ent. Tidskr., vol. 73, no. 1-2, May 10, 1952, p. 41).

Typhlocyba jacobii nom. nov.

The name Typhlocyba duplicata Jacobi (Jacobi, 1941, nee.
Typhlocyba duplicata McAtee, 1926) is here replaced by the name
Typhlocyba jacobii. See Ossiannilssonola duplicata (McAtee)
p. 1146.

During the time spent working on this revision, it was possible
for the author to study type specimens and determined specimens
in the following collections: the Illinois State Natural History Sur-
vey Collection, Urbana, Illinois; the Colorado Agricultural and
Mechanical College Collection, Ft. Collins, Colorado; the U. S.
National Museum Collection, Washington, D. C; and the private
collection of Dr. D. M. DeLong, Columbus, Ohio. In addition to
these, type material in the Snow Entomological Collections of the
University of Kansas was available for study, and type specimens
were loaned for study by the Canadian National Collection, and by
Mrs. J. N. Knull from her personal collection.

ACKNOWLEDGMENTS

The author wishes to make acknowledgment of particular as-
sistance given him by the following persons during his work on this
revision:

Dr. R. H. Beamer of the University of Kansas, who brought the
author's attention to the problem, and under whose direction the
research has been carried out; who has assisted in bringing to-
gether preserved material, and in developing suitable techniques
for collecting and studying material; who has accompanied the
author on collecting trips, and has made particular efforts during
the summers of 1948, 1949, and 1950 to collect material for the
author's use; and who has made available from his files and from
his personal experience information which has been of great assist-
ance in the preparation of this revision.

Dr. P. W. Oman, Systematic Entomologist for the U. S. Depart-
ment of Agriculture, Bureau of Entomology and Plant Quarantine,
who first suggested this problem and has shown continued interest
in the progress being made by giving helpful suggestions, by mak-
ing comparisons of specimens with types, and by making available
for the author's study a number of paratypes and a large number

1108 The University Science Bulletin

of undetermined specimens from the U. S. National Museum Col-
lections.

Dr. D. A. Young, Jr., Systematic Entomologist for the U. S. De-
partment of Agriculture, Bureau of Entomology and Plant Quaran-
tine, who has worked jointly with the author in the solution of the
generic problems concerned, who by numerous suggestions has
immeasurably helped to facilitate the work on this revision, and
through whose assistance it was made possible for the author to
study the material at the U. S. National Museum.

Dr. Frej Ossiannilsson of Uppsala, Sweden, for the loan of Euro-
pean specimens of Edwardsiana bergmani (Tullgren), E. candidula
(Kirschbaum), E. frustrator (Edwards), E. prunicola (Edwards)
for study; and for the gift of specimens of E. rosae (Linnaeus),
Typhlocyba quercus (Fabricius), and E. hippocastani (Edwards)
for deposition in the Snow Entomological Collections of the Uni-
versity of Kansas.

Dr. H. H. Ross of the Illinois State Natural History Survey, who
has made available for study a number of holotype specimens and
a large number of determined and undetermined specimens, most
of which have been recently collected in Illinois, in the Great
Smoky Mountains, Tennessee, and in British Columbia.

Mr. I. E. Davis, Entomologist for the Oregon State Department
of Agriculture, Division of Plant Industry, who has made available
for study pinned material and a large number of specimens in
alcohol, especially sorted out for the author from material collected
on pear psylla boards distributed throughout the state of Oregon.

Dr. D. M. DeLong of the Ohio State University, who has made
available for study all of the material in his personal collection,
including a number of holotype specimens.

Mrs. J. N. Knull of Columbus, Ohio, for the loan of type speci-
mens, for permission to use the original illustrations of Ribautiona
sciotoensis, for the gift of specimens of a number of species for
deposition in the Snow Entomological Collections of the University
of Kansas, and for a number of distribution records and helpful
suggestions.

Dr. J. T. Medler of the University of Wisconsin, for the gift of
his personal collection of specimens of species covered by this
revision, to be deposited in the Snow Entomological Collections of
the University of Kansas.

Dr. T. O. Thatcher of the Colorado Agricultural and Mechanical
College, for assistance while studying material in the College col-
lection, for the loan of a number of determined and undetermined

Revision of the Species Typhlocyba 1109

specimens including type specimens of Ossiannihsonola flavomar-
ginata, and for data for numbered specimens from the list of Gillette
numbers.

Dr. C. D. Michener of the University of Kansas, for helpful sug-
gestions concerning problems confronted in the querci and com-
missuralis complexes, and for suggestions and opinions regarding
certain nomenclatorial problems.

Mr. R. L. McGregor of the University of Kansas Department of
Rotany, who has accompanied the author on several collecting trips,
who has provided information regarding the locations of certain
host plants, and who has identified a number of the host plants.

Acknowledgment is also made for the loan of material borrowed
from the following collections, and to those who have made this
material available: U. S. National Museum Collection; Illinois
State Natural History Survey Collection; Oregon State Department
of Agriculture Collection; Colorado Agricultural and Mechanical
College Collection; Kansas State College of Agriculture and Ap-
plied Science Collection, D. A. Wilbur; Iowa State College of Agri-
culture and Mechanical Arts Collection, H. H. Knight; University
of California Collection, A. T. McClay and P. Hurd; California
Academy of Sciences Collection, E. S. Ross; Museum of Compara-
tive Zoology at Harvard College, J. Requaert; Pomona College Col-
lection, M. D. McCarthy; North Carolina Department of Agriculture
Collection, D. L. Wray; University of New Hampshire Collection,
J. G. Conklin; University of Minnesota Collection, C. E. Mickel and
H. E. Milliron; Rhode Island State College Collection, H. Knutson;
University of Delaware Collection, H. E. Milliron; Utah State Agri-
cultural College Collection, G. F. Knowlton; Oregon State College
Collection, C. H. Martin; and to the following individuals for ma-
terial from their personal collections: Dr. Frej Ossiannilsson, Mrs.
J. N. Knull, Dr. D. M. DeLong, Dr. R. A. Flock, Dr. D. A. Young, Jr.,
and Mr. R. R. Dreisbach.

Others who have assisted in a lesser degree, in supplying records
and in giving helpful suggestions and encouragement are as follows:
M. E. Neary, Nova Scotia Department of Agriculture; W. E. China,
Rritish Museum of Natural History, London, England; H. Andison,
Dominion Entomological Laboratory, Victoria, Rritish Columbia;
J. A. Rerly, Clemson College, South Carolina; G. E. Wallace, Carne-
gie Museum, Pittsburgh, Pennsylvania; C. A. Wilson, Mississippi
State College; J. A. Wilcox, New York State Science Service; H. C.
Severin, South Dakota State College; L. R. Penner, University of
Connecticut; L. Kuitert, Agricultural Experiment Station, Gaines-

1110 The University Science Bulletin

ville, Florida; W. J. Gerhard, Chicago Natural History Museum; and
J. R. Eyer, New Mexico College of Agriculture and Mechanical Arts.

GENITALIC CHARACTERISTICS

The genitalic structures of the male have been found important
in distinguishing genera as well as species of the Typhlocyba Com-
plex. The characteristic genitalic structure has been found to be
generally constant for each species, and has proven to be reliable for
use in the determination of species.

Although the form of the styles, connective, and male plates
is generally uniform for all of the species of some genera, in other
genera these structures are sometimes specifically distinct.

The shape of the aedeagus shows the greatest degree of modifica-
tion, and has proven to be the most useful structure for grouping
the species comprising different genera. The form of the aedeagus
is equally useful in distinguishing between species within a genus.

The shape of the pygofer, though not as useful for generic dis-
tinction because of its wide diversity of form in some genera, is in
these genera almost as important as the aedeagus for helping to
distinguish between species within the genus.

For a concise summary of the relations of the male genitalic struc-
tures to each other, in the Cicadellidae, the reader is referred to
Oman (1949b), pp. 22-23.

BIOLOGICAL NOTES

A number of interesting observations of the habits of some of the
species collected have been made by the author while collecting.

Population shifts during the day were noticed for several species,
and appeared to be caused by difference in temperature, difference
in light intensity, or a combination of both of these factors. During
the mornings in early June, adults could usually be found in greater
abundance on those branches upon which the sun was shining, while
they were seldom found on shaded branches. Shortly before noon,
as the temperature began to rise, the adults were found to have
shifted to the shaded lower branches. In the warm afternoons they
were found in greatest abundance on branches on the shaded sides
of the trees. Species living on small trees and shrubs were found
in greater abundance along the edges of woods where the sun shone
on the host plants, while in densely shaded areas only a few speci-
mens could be found.

When disturbed from the branches, it was found that most species
would fly to dark objects such as the bark on the trunk of trees,

Revision of the Species Typhlocyba 1111

stones, or to the ground. Species living in thickets were found to
fly deeper into the thicket, or toward the ground when disturbed.
In one instance, while collecting on Quercus macrocarpa, it was
found to be more profitable to beat the lower branches of the trees
with the net handle and to aspirate the insects from the tree trunks
than to attempt to use the bag in the dense foliage. When return-
ing to the leaves after having been disturbed, the insects usually
alight on the under surfaces of the leaves. In collecting species
which are not abundant or are not easily disturbed, when the
branches are within easy reach it is sometimes possible to aspirate
the insects from the undersurfaces of the leaves before beating. A
large number of specimens were taken from Acer saccharum in this
way, being easily seen from beneath, appearing as dark spots on
the translucent light green leaves. This method was found to be
most effective for collecting Edwardsiana candidula from the
fastigiate form of Populus alba, after beating had proven ineffective.

On warm humid nights the collection of Typhlocybid leafhoppers
at light was found to be particularly good. While collecting in
Milwaukee, Wisconsin, on June 27, 1950, a warm humid night when
thousands of specimens of species of Empoa were seen, it was found
that certain colors of neon lights were more attractive than others.
Few specimens were found at yellow, some at red, more at white,
while most specimens were found around green and blue lights.
Blue light was apparently more attractive than green.

Serious host injury was observed in only a few cases, the greatest
injury seen being that done by T. hockingensis on a species of
Viburnum used in Milwaukee County parks as an ornamental shrub.
Damage was particularly heavy in some areas, with leaves nearly
white from feeding injury, while in other areas only slight injury
or none was seen. The host plant was quite abundant, and usually
planted in hedgelike rows, which made it possible to easily see the
irregularity of injury. Similar irregularity of host injury was ob-
served on Quercus alba, caused by several species of leafhopper,
but principally by Ossiannilssonola berenice in association with
O. danae and O. australis. Noticeable injury was seen on Acer
saccharum caused by an association of several species, but primarily
by Typhlocyba niobe and T. persephone.

This factor of irregularity of population distribution makes it
possible to collect large numbers of specimens in one place while
in another place under apparently the same conditions specimens of
the same species are rare.

Males were found to begin reaching maturity several days be-

1112 The University Science Bulletin

fore females of the same species, so that early collections frequently
contained a large majority of males. Within one week after the
first males were taken, about equal numbers of both sexes were
found, but after two more weeks the majority of specimens found
were females. Observations made on Typhlocyba modesta showed
that this pattern was not followed, but that a ratio of nearly equal
numbers of both sexes was maintained for several weeks before the
population became predominantly female in composition. Observa-
tions on Typhlocyba persephone showed a rapid change to an all
female population within about two weeks, and after four weeks it
was impossible to find even a single female specimen in places
where there had previously been an abundance of specimens.

The only species observed by the author to have a second genera-
tion were Typhlocyba modesta, T. hockingensis, and T. melite.
Other species have been observed to have more than one genera-
tion a year by other workers, but populations of some of these
species were so small that it was impossible to be certain whether
a second generation had developed. Some of the species observed
could not possibly have had a second generation on the host that
the first generation was observed to feed on, because the leaves of
the host withered and dropped before a second generation could
have developed.

During the process of maturing, adults of some of the species of
Ossiannilssonola and Empoa pass progressively through several
color stages, with the color markings appearing to increase in ex-
tent and in intensity so that specimens killed at different stages of
this process are marked to greater or lesser degree. Another factor
influencing coloration which appears to be independent of the ma-
turing process controls the ultimate extent of color on the mature
adult. A more complete discussion of coloration is given in the
description of the genus Empoa, and problems which concern other
species are discussed in the color descriptions of these species.

Because of progressive coloration, and because male and female
insects begin emerging at different times, male and female speci-
mens taken at the same time often appear to be differently marked,
the females being usually much lighter in color than the males with
which they are taken. When fully colored specimens of both sexes
are compared there is no color distinction between them.

Eye color has sometimes been referred to in the descriptions of
light colored species. This color is caused by the migration of dark
color pigment in the iris cells of the compound eye ( Wigglesworth,
1947, pp. 113-115). When the insect is in a dark place the dark pig-

Revision of the Species Typhlocyba 1113

ment moves toward the surface of the eye, but when exposed to the
light the pigment moves away from the surface of the eye. Because
of this movement of pigment it has been possible for the author to
collect specimens of one species showing both light and dark colored
eyes by collecting on both shaded and sunny sides of a single tree,
or by collecting at different times of the day. Specimens collected
at night are frequently dark eyed when they come to the light, but
the eye color becomes lighter as they remain at the light for some
time.

TECHNIQUE

A particular type of net was found to be more efficient in collect-
ing the insects studied than were the types commonly used for col-
lecting leafhoppers. Since all of the species collected live on trees
or bushes, a durable net bag was needed. A heavy canvas beating
net was found to be impractical because it could not be moved
rapidly enough, did not collapse quickly enough, and did not let
in sufficient light. After testing several kinds of net bags, a fine-
meshed nylon bag was found to be best suited for this type of col-
lecting. Because most of the species collected were light colored,
it was found that a bag dyed navy blue permitted them to be seen
more easily, and they appeared to be less active than in a light
colored bag. The bag was also pointed to restrict these rather
active inser ts to a smaller space from which they could be aspirated
more easily.

Most of the species studied were particularly fragile, and in order
to remove the abdomen for dissection without detaching the speci-
men from the point, or injuring its wings, the paper point with the
insect attached was removed from the pin and placed inverted on
a piece of sheet cork. This exposed the abdomen in ventral aspect
so that it could easily be separated from the thorax by a slight
pressure with the point of the pin at its base.

Less heavily sclerotized specimens and those used for illustration
were stained with acid fuchsin to show differences in sclerotization
and to bring out obscure details.

Dissections were held stationary for drawing by means of a small
amount of petroleum jelly placed on the slide before adding the
glycerine. Dry slide mounts of the wings were used for making
illustrations of the wings.

Drawings of the pygofer, plates, and head represent a magnifica-
tion of 60 times actual size; drawings of the aedeagus, styles, con-
nective of the male, and eighth abdominal sternite of the female,
120 times; drawings of the wings 56 times.

1114 The University Science Bulletin

SYSTEMATIC ACCOUNT
Introduction to the Typhlocyba Complex

Most of the species included in the genus Typhlocyba prior to
the segregation of the genera included in this revision were placed
in this genus on the basis of wing venation as found in the type
species, Typhlocyba quercus (Fabricius), the hind wing having
two open apical cells, the fore wing with the inner and outer apical
cells short and not attaining the wing apex, and the third apical cell
of the fore wing triangular and usually stalked. Some of the species
included in the genus differ from the type species in having the
third apical cell of the fore wing quadrate and not stalked. The
North American species with rare exceptions agreed with the type
species in wing venation.

In recent interpretation of the genus Typhlocyba the structure of
the male genitalia has been regarded as more significant than wing
venation as a generic character, resulting in the segregation of
groups of species from Typhlocyba as genera (Zachvatkin, 1939,
1947; Young, 1952). This change in generic concept has brought
about a need for restudying the species of Typhlocyba to determine
their generic status on the basis of the male genitalia. It is the
opinion of the author that when this has been done for the species
from other regions, additional new genera will be added to the
Typhlocyba Complex.

The following systematic account includes only those genera of
the Typhlocyba Complex which occur in North America.

Key to the Genera of the Typhlocyba Complex

1. Crown with median length equal to median length of pronotum

or nearly so; ocelli present; pygofer without group of mac-
rosetae near basal angle of male plate. (Pi. LXXIII, fig. 1),

Henribautia p. 1115

Crown with median length much less than median length of pro-
notum; ocelli rarely present; pygofer usually with macrosetae
near outer basal angle of male plate 2

2. Aedeagal shaft reduced to a flattened membranous structure oc-

curring between a pair of arms formed by longer forcipate
atrial processes; plate without macroseta at outer basal angle.
( Pi. IV, fig. 1 ) Ossiannilssonola p. 1132

Aedeagal shaft strongly sclerotized or absent, not so enclosed;
plate with macroseta at outer basal angle 3

3. Aedeagal shaft absent, atrial processes fused at base, branched

near apex. (Pi. I, fig. 4) Mcateeana p. 1130

Aedeagal shaft present, heavily sclerotized, atrial processes when
present not branching 4

Revision of the Species Typhlocyba 1115

4. Mesal margin of style with distinct preapical angular protuber-

ance. ( PI. II, fig. 1 ) Ribautiana p. 1119

Mesal margin of style without distinct preapical angular pro-
tuberance 5

5. Aedeagal shaft with three pairs of apical processes and three

broad, thin plates arising from anterior margin.

( PI. LXXXVIII ) Empoa p. 1187

Aedeagal shaft with less than three pairs of apical processes or
none, rarely with one thin plate arising from anterior margin . . 6

6. Aedeagal shaft with two pairs of apical processes, frequently one

or both pairs branched; without atrial processes; aedeagal
apodeme a slender arm nearly two thirds the length of shaft,

Edwardsiana p. 1208
Aedeagal shaft with one pair or without apical processes; usually
with atrial processes present; aedeagal apodeme usually later-
ally broadened, less than two thirds the length of shaft,

Typhlocyba p. 1160

Genus Henribautia Young and Christian

(PI. LXXIII, figs. 1, 2, 3)

Henribautia Young and Christian, in Young, Bull. Univ. Kansas Sci. Bull., vol.
35, pt. 1, 1952, pp. 96-97.

Type of the genus, Typhlocyba nigricephala Beamer, by original
designation.

Fore wings. — Inner and outer apical cells short, not attaining
wing apex; second apical cell much broader at apex than at base;
third apical cell petiolate; wing apex somewhat oblique; apical half
of wing having a number of transparent areas scattered along the
cross veins and in apical cells. (PI. LXXIII, fig. lh).

Hind wings. — Vein IV branching from vein 2V near its mid-
length; submarginal vein absent at wing apex; both apical cells open
apically; posterior branch of R fused with apical portion of vein
M1+2; basal half whitish hyaline, apical half fumose with veins
brown. (PI. LXXIII, fig. lh).

Genital capsule. — Male plate, in ventral aspect, broadened on
basal half, abruptly narrowed near middle of length to narrow up-
turned apex; in lateral aspect with single macroseta near outer basal
angle of male plate, and a few small setae on apical half; pygofer,
in lateral aspect with numerous small setae scattered over ventral
half of disc, completely without macrosetae, dorsal posterior half of
disc covered with numerous awl-shaped spines, posterior margin
slightly inrolled, differentially sclerotized area near base of plate,
posterodorsal margin with row of small setae; anal hooks wanting.

Internal male genitalia. — Style narrow, gradually tapered and
curved laterad on apical half, with preapical triangular protuber-

1116 The University Science Bulletin

ance on inner margin, with one or few setae on outer, and several
alveoli on inner margin near middle of length; connective triangular
to Y-shaped, the stem broad, aedeagal articulation subterminal;
aedeagus without preatrial arm, apodeme simple, well developed,
shaft slender, elongate, with paired apical processes.

Female. — With posterior margin of eighth abdominal sternite as
in H. nigricephala. (PL LXXXVII, figs. 2a, b).

Head well produced medially, conical, median length of the
crown greatly exceeding length next the eye particularly in female,
nearly, or quite as long as, but narrower than pronotum, crown
sharply rounded to the slightly convex face, contour divergent from
the line of dorsum; ocelli present, situated on the margin between
crown and face, distant from compound eyes; pronotum with lat-
eral margins divergent posteriorly, posterior margin scarcely emar-
ginate; pleural portion much broader than the ocellocular area;
head, pronotum, and scutellum black to dark brown, face yellow
ventrad of antennal pits, abdomen black to dark brown, male plates
yellow with apices brown.

The known distribution of this genus is the southern United
States from Mississippi westward to New Mexico.

Key to the Species of Henribautia

1. Vertex of he:xl brownish black with only ocelli white; fore wings

with basal half white to yellow-orange and meeting black-
brown apical half in a transverse line, a small bilobed black

mark along scutellum nigricephala p. 1116

Vertex of head brownish black with a narrow white line extend-
ing between eye and ocellus; fore wings with black-brown
markings extending along commissural margin or with light
markings meeting black-brown markings on apical half in an
oblique line 2

2. Vertex of head with narrow white line extending between ocelli;

fore wing with white to yellow markings on basal half extend-
ing to or nearly to commissural margin hubbardi p. 1117

Vertex of head with narrow white line not extending between
ocelli; fore wing with white to yellow markings only on lateral
third of basal two thirds along costal margin beameri p. 1118

Henribautia nigricephala (Beamer)
( PI. LXXIII, fig. 1 )

Typhlocyba nigricephala Beamer, Canadian Ent., vol. 75, no. 7, 1943, pp. 131-

' 133.
Henribautia nigricephala, Young, Bull. Univ. Kansas Sci. Bull., vol. 35, pt. 1,
1952, p. 97.

Resembling H. hubbardi, but differs in having dark brown mark-
ings on outer half of fore wing with anterior margin almost trans-

Revision of the Species Typhlocyba 1117

verse, and in lacking narrow white band between ocelli and eyes.

Length.— 2.5-3.0 mm.

Color. — Fore wings, with dark brown markings on clavus border-
ing scutellum roughly bilobed; milky-white to golden-yellow on
basal half of wing, dark brown along scutellum; anterior margin of
dark brown apical marking almost transverse.

Specimens from Arizona are dark brown to black, while speci-
mens from other localities are lighter brown, nearly maroon.

Genital capsule. — Male pygofer, in lateral aspect, with posterior
margin forming two evenly rounded lobes, the dorsal lobe bearing
a few microsetae and projecting beyond ventral lobe posteriorly,
ventral margin slightly lobed.

Internal male genitalia. — Shaft of aedeagus with inferior pair of
apical processes directed basad to slightly below middle of shaft,
superior pair short, thornlike, slightly curving laterodorsad.

The only known host plant is Rhamnus californica, a large series
having been collected from it by Dr. R. H. Reamer, July, 1950, at
Granite Dells, Arizona. The name of the host plant from which the
type series was taken, which differed from the above host, is not
known.

Specimens have been seen from the following localities : Arkansas:
Fouke, December 21; Mississippi: Shuqualak, July 16; Louisiana:
Calcasieu County, August 16; Caddo County, August 19; Colfax,
December 23; Vinton, December 25; Texas: Orange County, Au-
gust 14; Arizona: Santa Catalina Mountains, July 14; Santa Rita
Mountains, July 17, August 18; Yarnell, July 27; Oak Creek Canyon,
August 9; Granite Dells, July 5.

The specimens taken in Arizona differ from those from the other
states, in being darker in color, and in being one-fifth to one-sixth
larger in size in all dimensions, though identical in form and propor-
tions with those from other places. After more is known about the
biology of this species, these forms may prove to be distinct species.
At present the author considers it best to continue to regard these
as extremes in variation of a single species.

Types. — Holotype male, allotype female, and numerous paratypes
of both sexes, in the Snow Entomological Collections of the Univer-
sity of Kansas.

Henribautia hubbardi (McAtee)
(PI. LXXIII, fig. 3)

Erythroneura hubbardi McAtee, Florida Ent., vol. 8, no. 3-4, 1924, p. 35.
Typhlocyba hubbardi, Beamer, Canadian Ent., vol. 66, no. 1, 1934, p. 18.
Henribautia hubbardi, Young, Bull. Univ. Kansas Sci. Bull., vol. 35, pt. 1, 1952,
p. 97.

1118 The University Science Bulletin

Resembling H. nigricephala, but differs in having dark brown
markings on outer half of fore wing with anterior margin strongly
oblique, and in having a narrow white band on vertex of head be-
tween ocelli and eyes.

Length.— 2.75-3.0 mm.

Color. — A narrow white line between eyes on vertex across ocelli;
fore wings with dark brown markings on clavus along scutellum
evenly rounded; brown markings continuing along commissural
margin on inner half of clavus, joining anterior and posterior mark-
ings in some specimens; if not joined, with anterior edge of posterior
markings crossing wing obliquely; with a narrow V-shaped white
band bordering dark areas and enclosing a triangular golden-yellow
area which borders on costal margin near middle; male plate light
brown on lateral margins.

Genital capsule. — Male pygofer with posterior lobes not as deeply
notched between as in H. nigricephala.

Internal male genitalia. — Aedeagus with apical processes parallel
to shaft, superior and inferior pairs of almost equal length, fused at
base near point of attachment to shaft, superior pair strongly di-
vergent, inferior pair directed basad parallel to shaft.

Specimens have been seen from the following localities in Arizona:
Chiricahua Mountains, July 8, August 7, September 6; Santa Rita
Mountains, June 16, July 16; Huachuca Mountains, June 11, July 9,
August 1, October 31.

Types. — Holotype female, in the U. S. National Museum collec-
tion; allotype male, in the Snow Entomological Collections of the
University of Kansas. Types have been seen by the author.

Henribautia beameri sp. nov.
(PI. LXXIII, fig. 2)

Resembling H. nigricephala, but differs in having more extensive
black markings on wing and in having inferior pair of apical proc-
esses of aedeagus sharply angled at the middle and directed mesad
toward the shaft.

Length. — 2.5 mm.

Color. — Vertex of head with narrow yellow line extending be-
tween eye and ocellus on each side, yellow color of face sometimes
extending anteriorly between ocelli to anterior third of crown; fore
wings dark brown over most of their surface, with a bright yellow
stripe along basal two thirds of costal margin outlined by a narrow
white band.

Revision of the Species Typhlocyba 1119

Genital capsule. — Male pygofer, in lateral aspect, with posterior
margin medially produced posteriorly as a single lobe.

Internal male genitalia. — Aedeagus with inferior pair of apical
processes diverging laterocephalad from shaft for two thirds their
length, then directed toward shaft, apices meeting or almost meet-
ing near basal third of shaft, superior pair of processes short, ap-
pearing as a distal continuation of base of inferior processes, proc-
esses somewhat S-shaped from right lateral aspect.

The host of this species is thought to be Condalia spathulata.

Types. — Holotype male, allotype female, and numerous male and
female paratypes, Oracle Junction, Arizona, April 29, 1948, R. H.
Reamer and L. D. Reamer. One male paratype, Tucson, Arizona,
April 28, 1948, R. H. Reamer. Types in the Snow Entomological
Collections of the University of Kansas.

Genus Ribautiana Zakhvatkin

(Pis. LXXIV and LXXV)

Ribautiana Zakhvatkin, Rev. Ent. URSS, vol. 28, no. 3-4, (1945), 1947, p. 113.

Type of the genus, Cicada idmi Linnaeus, by original designation.

Fore wings. — Inner and outer apical cells short, not attaining wing
apex; second apical cell much broader at apex than at base; third
apical cell petiolate, wing apex smoothly rounded; with dark mark-
ings at apices of the veins which form distal margins of inner and
outer apical cells, dark areas in apices of inner three basal cells.
(PI. LXXIV, fig. 1).

Hind wings. — Vein IV branching from vein 2V near its mid-length;
submarginal vein absent at wing apex; both apical cells open api-
cally; posterior branch of R fused with apical portion of vein M1+2;
usually whitish hyaline, fuscous on apical half in R. ulmi. (PL
LXXIV, fig. 1).

Genital capsule. — Male plate, in ventral aspect, abruptly nar-
rowed near middle its length to narrow upturned divergent apex,
with single macroseta near outer basal angle and two smaller setae
near lateral margin near middle of plate; in lateral aspect, with a
number of scattered microsetae, some of which tend to form a sub-
marginal row along lateral margin; pygofer, in lateral aspect, with
group of macrosetae just dorsad of outer basal angle of plate, with
numerous microsetae in region caudad and dorsad of these, and a
small row on inrolled margin directed mesad; posterior margin
scarcely inrolled, not differentially sclerotized, not well differenti-

41—6490

1120 The University Science Bulletin

ated, merging with ventral margin; pygofer hooks wanting; anal
hooks wanting.

Internal male genitalia. — Style elongate, slender, gradually taper-
ing and curved laterad or dorsad towards apex, inner margin with
preapical triangular protuberance and several alveoli, outer margin
with a few long setae; connective Y-shaped or triangular; aedeagal
articulation terminal or subterminal; aedeagus without preatrial
arm; aedeagal apodeme well developed, simple; atrial processes or
basal shaft processes present, and usually one or more pairs of
processes more distad on the shaft.

Female. — With posterior margin of eighth abdominal sternite as
in R. piscator (PI. LXXXVII, figs. 3a, 3b) except in R. ulmi (PI.
LXXXVII, figs. 4a, 4b).

Head in dorsal aspect, narrower than pronotum, scarcely pro-
duced, median length not greatly exceeding the length next the eye,
female longer than male, anterior margin of crown smoothly
rounded; in lateral aspect contour of face convex and divergent
from the line of dorsum; ocelli absent; pronotum short, with lateral
margins greatly divergent posteriorly, and with posterior margin
shallowly emarginate; width of pleural portion greatly exceeding
width of ocellocular area.

The genus has a Holarctic distribution.

Key to the Species of Ribautiana

1. Length 3.5-4.0 mm.; fore wings greenish-yellow; abdomen black

on venter; aedeagus without processes on shaft. (Pi. LXXIV,

fig. 1) ulmi p. 1121

Length 3.25 mm. or less; fore wings white to orange-yellow; ab-
domen yellow on venter; aedeagus with processes on shaft ... 2

2. Scutellum dark brown or black 3

Scutellum light colored 4

3. Aedeagus with two pairs of apical processes, shaft three times as

long as atrial processes. (PI. LXXV, fig. 1 ) luculla p. 1127

Aedeagus with three pairs of apical processes, median pair
branched, shaft twice as long as atrial processes. (Pi. LXXV,
fig. 5) sciotoensis p. 1128

4. Aedeagus with a pair of processes arising near middle of shaft ... 5
Aedeagus without processes arising near middle of shaft 6

5. Shaft of aedeagus slender, apex gradually curving ventrocaudad,

with two or three pairs of short apical processes. ( Pi. LXXIV,

fig. 4) unca p. 1129

Shaft of aedeagus stout, apex not curving ventrad, with a pair of
apical processes directed cephalad, and a single unpaired
process directed dorsad from posterior margin. (Pi. LXXIV,
fig. 2 ) tenerrima p. 1122

Revision of the Species Typhlocyba 1121

6. Inferior pair of apical processes of aedeagus with numerous spine-

like projections 7

Inferior pair of apical processes of aedeagus smooth 8

7. Length of shaft of aedeagus beyond apical processes equal or

nearly equal to length of inferior processes. (PI. LXXV, fig.

4) mtdtispinosa p. 1125

Length of shaft beyond apical processes of aedeagus less than
half the length of inferior processes. ( Pi. LXXV, fig. 3 ) ,

piscator p. 1124

8. Apical half of aedeagal shaft nearly straight, inferior pair of proc-

esses only slightly curved. (PI. LXXIV, fig. 3) . . parapiscator p. 1123
Apical half of aedeagal shaft curved into nearly a complete circle,
inferior pair of apical processes following curve of shaft. ( PL
LXXV, fig. 2) foliosa p. 1126

Ribautiana ulmi (Linnaeus)
(PL LXXIV, fig. 1)

Cicada ulmi Linnaeus, Systema naturae, Regnum animale. ed. 10, 1758, p. 439.

( Engelmann reprint, 1894, p. 439. )
Typhlocyba ulmi, Burmeister, Handbuch der Entomologie, 1835, vol. 2, p. 107.
Anomia ulmi, Fieber, Katalogue der europaischen Cicadinen, 1872, p. 15.
Empoa ulmi, Van Duzee, Check List of Hemiptera (excepting the Aphididae,

Aleurodidae and Coccidae) of America North of Mexico, 1916, p. 77.
Ribautiana ulmi, Zakhvatkin, Rev. d'Ent. URSS, vol. 28, nos. 3-4, (1945), 1947,

p. 113.
Eupteryx ocellata Curtis, British Entomology, vol. 14, art. 640, 1837, p. 2.

Resembling R. tenerrima, but easily distinguished by its larger
size, darker color, and absence of processes on aedeagal shaft.

Length. — 3.5-4.0 mm.

Color. — Head, pronotum, and scutellum pale greenish-yellow,
female with two oval black spots on vertex of head between eyes,
another on middle of anterior margin of pronotum also present on
male, spots faint or absent on teneral specimens; fore wings with
basal half evenly colored light yellowish-green to olive-green; ab-
domen with dorsum and venter black, with a narrow yellow band on
posterior margin of each segment, pygofer black.

Genital capsule. — Male pygofer, in lateral aspect, with posterior
margin produced mesally in an evenly rounded lobe.

Internal male genitalia. — Connective distinctly Y-shaped; aedea-
gus with pair of broadly flattened atrial processes directed dorso-
laterad for half the length of aedeagal shaft; shaft without proc-
esses, slender, gradually tapering to slightly bifid apex, recurved on
outer fourth; aedeagal apodeme more than one third as long and
almost twice as broad as shaft in lateral aspect.

A large series of specimens was taken on Uhnus fnlva in Mil-
waukee, Wisconsin, by the author, and a short series is on hand
from Salem, Oregon, taken on Ulmus glabra camperdownii, by Mr.

1122 The University Science Bulletin

John E. Davis. Other specimens seen are labeled only "Uhnus",
or "Elm".

Specimens have been seen from the following North American
localities: Ontario: Vineland, July, August, September, October;
Jordan, August 11, 15; Massachusetts: Cambridge, October 24, 30,
November 1; Wood's Hole, July 10, September 1; Arlington, Sep-
tember 10; Nova Scotia: King's County, July; Rhode Island: Kings-
ton, September; New York: Ithaca, October 6; New Jersey: Bound
Brook; Ohio: Columbus, October 13; Missouri: St. Louis, April 25;
Wisconsin: Milwaukee, June 27-July 7; Texas: ?; Utah: Logan,
August 26; British Columbia: Vernon; Victoria, August 8, October
12; Washington: Seattle; Shelton, July 24; Tacoma, June 29; Puyal-
lup, June; Oregon: Salem, September 26; Orenco, November 1;
California: Mountain View, Milbrae, October 3; Mill Valley, Marin
County, October 3; Berkeley, September 1, 8, 12; San Jose, October
24.

Other specimens have been seen from the following European
localities: Germany: Halle, September 23; France: Paris, October
28; Russia: Samara, July 15; England: Cambridge, October 17
Oxfordshire, June 17; Scotland: Edinburgh, August 19; Finland
Helsinki, September 25; Bohemia: Perimov, September 24; Sweden
Uppland, Solna, October 26; Skane, Ven, August 2; Gotland, Roma
Kl, July 3.

Ribautiana tenerrima (Herrich-Schaffer)
(PI. LXXIV, fig. 2)

Typhlocyba tenerrima Herrich-Schaffer, Faunae Insectorum Germanicae initia;

oder Deutschlands Insecten gesammelt und herausgegeben von D. G. W. F.

Panzer. Fortgesetzt von G. A. W. Herrich-Schaffer, vol. 124, 1834, p. 10.
Typhlocyba rubi Hardy, Trans. Tyneside Nat. Club, vol. 1, 1850, p. 417.
Typhlocyba misella Boheman, Handlingar. Kongliga Svenska Vetenskaps

Akademien, 1851, p. 122.
Anomia tenerrima, Fieber, Katalogue der europaischen Cicadinen, 1872, p. 15.
Empoa tenerrima, Van Duzee, Trans. San Diego Soc. Nat. Hist., vol. 2, no. 1,

1914, p. 57.
Ribautiana tenerrima, Zakhvatkin, Revue d'Ent. URSS, vol. 28, no. 3-4. ( 1945),

1947, p. 113.

Resembles R. piscator and other similarly marked species in ex-
ternal appearance, but differs in having aedeagal shaft straight, not
curving posteriorly at apex, and in having a pair of short slender
basally fused processes arising on posterior margin near middle,
another pair on anterior margin, and an unpaired process on pos-
terior margin at apex.

Length. — 3.0 mm.

Color. — Head, pronotum, and scutellum pale white to light yellow,
without dark markings; fore wings white to yellow; abdomen black

Revision of the Species Typhlocyba 1123

on basal half of dorsum of each segment, venter light yellow, or
with narrow black basal band on each segment, basal half of pygofer
black.

Genital capsule. — Male pygofer, in lateral aspect, subquadrate,
ventral angle projecting slightly beyond dorsal angle.

Internal male genitalia. — Connective triangular; aedeagus with
pair of atrial processes elongate, slightly flattened on distal half,
length exceeding aedeagal shaft, gradually reduced to acute apices;
aedeagal shaft expanded on basal third, laterally compressed shortly
before apex, with a pair of short slender basally fused processes
arising on posterior margin near middle, another pair at apex di-
rected laterocephalad from anterior margin, and with unpaired
dorsally directed process arising from posterior margin at apex;
aedeagal apodeme one sixth length of shaft, as broad as narrowest
width of shaft in lateral aspect.

The description of this species made by McAtee (1926) was
based on a specimen of R. cruciata Ribaut, according to Ribaut
(1931a, p. 287). The first reliable report of the occurrence of this
species in North America was made by H. Andison (1950) from
specimens collected on Loganberry at Rrentwood, British Columbia,
June 20, 1947, and illustrations for this species have been made from
one of these. No other North American specimens of this species
are known to the author, and previous records of the occurrence of
this species apparently apply to other species in the genus.

Determination of this species is based on figures of male genitalia
as illustrated by Ribaut (1936, p. 117). A pair of specimens from
Sweden, determined by Dr. Ossiannilsson as this species, agree with
the North American specimens seen. Other specimens from Europe,
determined as tenerrima are: a pair of specimens from Budapest,
determined by Horvath, which agree with the figures of Typhlocyba
scalaris Ribaut, (Ribaut, 1936), and a specimen from France, de-
termined by Signoret, which has male genitalia which agree with
T. debilis Douglas as illustrated in the same paper.

The approved common name for this species is "The Bramble
Leafhopper" (Muesbeck, 1950, p. 138).

Ribantiana parapiscator sp. nov.

(PI. LXXIV, fig. 3)

Typhlocyba piscator McAtee, Proc. U. S. Nat. Mus., vol. 68, art. 18, 1926, p. 7
(part).

Resembling R. piscator in external appearance, but with aedeagal
shaft forming nearly a right angle near middle and with inferior
pair of apical processes smooth.

1124 The University Science Bulletin

Length. — 2.75 mm.

Color. — Dorsum pale milky white to light orange-yellow; abdo-
men black on basal half of dorsum of each segment, venter light
yellow.

Genital capsule. — Male pygofer, in lateral aspect, subquadrate
with dorsal angle projecting beyond ventral angle.

Internal male genitalia. — Aedeagus with atrial process elongate,
evenly tapered from base to acute apices, sinuate, almost as long as
shaft; aedeagal shaft elongate, slender, bent posteriorly at middle
in almost a right angle, with two pairs of slender, smooth, acute
processes at apex, superior pair appearing as a continuation of shaft
in lateral aspect, slightly curved ventrad, length two thirds that of
inferior pair which curve dorsad toward apices of atrial processes.

This species has been collected by the author from Ostrija vir-
giniana in abundance, both in Douglas County, Kansas, and in Mil-
waukee County, Wisconsin, along with R. multispinosa.

Types. — Holotype male and twenty-seven male paratypes, Doug-
las County, Kansas, June 7, 1950, P. J. Christian; additional para-
type males: twenty-eight, Douglas County, Kansas, May 30, 1949,
R. H. Beamer; two, Douglas County, Kansas, May 30, 1949, P. J.
Christian; one, June 1, 1949, Douglas County, Kansas, P. J. Chris-
tian; twenty-six, Milwaukee, Wisconsin, June 26-July 5, 1950, P. J.
Christian; one, Ames, Iowa, September 20, 1940, D. R. Lindsay;
one, Brandenburg, Kentucky, September 14, 1941, D. A. Young;
one, Vineland Station, Ontario, September 21, 1940, W. L. Putman;
one, Ames, Iowa, September 28, 1894, a paratype of Ribautiana
piscator bearing the same data as the allotype female of that species.

Holotype and paratypes in the Snow Entomological Collections
of the University of Kansas; the Brandenburg, Kentucky, paratype
in the U. S. National Museum Collection; the Vineland Station,
Ontario, paratype in the Canadian National Collection; and the
Ames, Iowa, specimen of the type series of R. piscator, in the Iowa
State College Collection.

Ribautiana piscator (McAtee)
(PI. LXXV, fig. 3)

Typhlocyba piscator McAtee, Proc. U. S. Nat. Mus., vol. 68, art. 18, 1926, p. 7

(part).
Ribautiana piscator, Young, Bull. Univ. Kansas Sci. Bull., vol. 35, pt. 1, 1952,

p. 100.

Resembling R. multispinosa, but differing in having length of
aedeagal shaft distad of superior pair of processes, shorter than
these.

Revision of the Species Typhlocyba 1125

Length.— 2.75-3.0 mm.

Color. — Head, pronotum, scutellum and light portions of fore
wing pale milky-white to light yellow; abdomen black on basal half
of dorsum of each segment, venter yellow.

Genital capsule. — Male pygofer, in lateral aspect, somewhat
quadrate, dorsal angle rounded, projecting beyond ventral angle
which is less distinct.

Internal male genitalia. — Aedeagus, with pair of atrial processes
three fourths as long as shaft, elongate, gradually reduced to acute
apices, sinuate; shaft broadly curved posteriorly, with two pairs
of apical processes of about equal length, superior pair slender,
smooth, directed ventrolaterad toward apices of atrial processes, in-
ferior pair fused basally, a stout spine on mesal margin at basal
third, numerous small spines scattered over apical two thirds.

Most of the specimens seen have the inferior pair of processes
more spinose than in the specimen figured which is nearer in ap-
pearance to the holotype than any of the other specimens seen.

A series of twenty specimens, eleven males and nine females, were
collected by the author from Carpinus caroliniana in Milwaukee,
Wisconsin, July 7, 1950.

Tyj)c. — Holotype male, Elizabeth, Illinois, in the Illinois State
Natural History Survey Collection; allotype female, in the Iowa
State College Collection, bears the same data as a paratype male
which is a specimen of Ribautiana parapiscator, and is probably
also a specimen of that species.

Ribautiana multispinosa sp. nov.
(PL LXXV, fig. 4)

Resembling R. piscator, but differing in having length of aedeagal
shaft distad of superior pair of processes longer than these.

Length. — 2.75-3.0 mm.

Color. — Head, pronotum, scutellum and light areas of fore wings
pale milky-white to light yellow; abdomen black on basal half of
dorsum of each segment, venter yellow.

Genital capsule. — Male pygofer, in lateral aspect, somewhat
quadrate, dorsal angle rounded, projecting beyond ventral angle.

Internal male genitalia. — Aedeagus with pair of atrial processes
elongate, evenly tapered from base to acute apices, sinuate, almost
half as long as aedeagal shaft; shaft elongate, slender, bent pos-
teriorly on outer third, forming a half circle in lateral aspect, apex
curving toward middle of shaft; two pairs of processes on apical
sixth of shaft, superior pair slender, smooth, directed laterocaudad

1126 The University Science Bulletin

toward apices of atrial processes, half as long as inferior pair which
are broadly flattened laterally forming an oval in caudal aspect and
armed with numerous short stout spines which vary somewhat in
number and location on different specimens.

This species has been collected by the author from Ostrya vir-
giniana in association with R. parapiscator, in Milwaukee, Wis-
consin, and in Douglas County, Kansas.

Types. — Holotype male, July 2, 1950, Milwaukee, Wisconsin, P. J.
Christian; paratype males: eight, May 30, one, June 5, 1949, Douglas
County, Kansas, R. H. Beamer; one, June 1, 1949, eight, June 7,
1950, Douglas County, Kansas, P. J. Christian; one, June 28, three,
July 1, two, July 2, one, July 3, three, July 5, 1950, Milwaukee, Wis-
consin, P. J. Christian; one, July 4, 1949, Cheboygan County, Michi-
gan, H. B. Hungerford. An abnormal male of this species, August 1,
1949, Cheboygan County, Michigan, H. B. Hungerford, is not made
a paratype. Types in the Snow Entomological Collections of the
University of Kansas.

Ribautiana foliosa (Knurl)

(PI. LXXV, fig. 2)

Typhlocyba foliosa Knull, Ohio J. Sci., vol. 45, no. 3, 1945, pp. 104, 107,

pis. 1-2.
Ribautiana foliosa, Young, Bull. Univ. Kansas Sci. Bull., vol. 35, pt. 1, 1952,

p. 100.

Resembling R. piscator, but differing in having apical half of
aedeagal shaft forming nearly a circle, and in having the inferior
pair of shaft processes smooth and strongly curved, following the
curvature of the shaft.

Length. — 3.0 mm.

Color. — Head, pronotum, and scutellum pale milky-white to
light orange yellow; abdomen black on basal half of dorsum of each
segment.

Genital capsule. — Male pygofer, in lateral aspect, somewhat quad-
rate, dorsal angle rounded, projecting beyond ventral angle.

Internal male genitalia. — Aedeagus with atrial processes broadly
attached at base, distinctly S-shaped in left lateral aspect, gradually
reduced to sharp apices, half as long as aedeagal shaft, sinuate in
posterior aspect; aedeagal shaft swollen at base and broadly flat-
tened on outer third, with two pairs of processes near apex, inferior
pair foliaceous, curved ventrolaterad in a semicircle, with apices
directed toward apices of atrial processes, superior pair of processes
forming almost a complete circle with the distal half of shaft, apices
directed ventrolaterad toward middle of shaft in lateral aspect,
shaft sharply reduced to a short sinuate apex beyond processes.

Revision of the Species Typhlocyba 1127

A number of specimens of this species have been taken by the
author from Fagus grandifolia. Two other specimens are on hand
labeled "host Carpinus caroliniana", while another specimen from
the same locality is labeled "Fagus grandifolia". One male speci-
men from New Hampshire, determined by McAtee as R. piscator,
and a male paratype have been seen. Specimens seen have been
from the following localities: Ontario: Vineland Station, September
19; Manitoba: Birtle, August 6, 10, September 2; New Hampshire:
Durham, June 23; Virginia: Mountain Lake, July 2; Tennessee:
Gatlinburg, June 24, 25; Ohio: Hocking County, September 16;
Delaware County, October 10; Wisconsin: Milwaukee, July 5; Min-
nesota: Itaska County, July 26-28.

Types. — Holotype, allotype, and paratypes in the collection of
Mrs. J. N. Knull; six paratypes in Ohio State University Collection.

Ribantiana luculla (Medler)
(PI. LXXV, fig. 1)

Typhlocyba luculla Medler, Minnesota Agr. Exp. Sta. Tech. Bull., no. 155, 1942,

pp. 139-140, pi. 9.
Ribautiana luculla, Young, Bull. Univ. Kansas Sci. Bull., vol. 35, pt. 1, 1952,

p. 100.

Resembling R. sciotoensis in outward appearance, but with aedea-
gal shaft dorsoventrally flattened, with only two pairs of apical proc-
esses on shaft.

Length. — 3.25 mm.

Color. — Head, pronotum, and light portions of fore wing pale
yellowish white; scutellum fuscus; fore wing with band across distal
ends of three inner basal cells more pronounced, cross veins darker
near wing margins, third apical cell fuscus; abdomen black on dor-
sum, yellow on venter.

Genital capsule. — Male pygofer, in lateral aspect, somewhat
quadrate, dorsal angle rounded, projecting beyond ventral angle.

Internal male genitalia. — Aedeagus, with pair of atrial processes
one half width and one third length of shaft in lateral aspect, very
gradually reduced to acute apices, sinuate; aedeagal shaft slender,
greatly attenuated, width in lateral aspect one third width from
caudal aspect, inferior pair of processes at outer third of shaft with
apices directed toward apices of atrial processes, slightly broader
but only half as long as atrial processes, superior pair of processes
as long and stout as inferior pair, directed dorsolaterad, arising at
half their length from apex of shaft.

The host plant for this species is believed to be Corylus ameri-
cana, on the basis of a female specimen taken on that host, which

1128 The University Science Bulletin

agrees in external appearance with a male paratype of this species.

The following specimens have been seen: one female, La Crosse,
Wisconsin, August 7; one female, Merrillan, Wisconsin, August 5;
one female, Thornton, Illinois, September 7; and a series of twelve
unpinned male and female specimens from Illinois, in the Illinois
Natural History Survey Collection.

Types. — Holotype male and two paratype males, in the Univer-
sity of Minnesota Collection; one male paratype in the Snow Ento-
mological Collections of the University of Kansas.

Ribautiana sciotoensis (Knull)
(PL LXXV, fig. 5)

Typhlocyba sciotoensis Knull, Ohio J. Sci., vol. 45, no. 3, 1945, pp. 103, 107,

pi. 2.'
Ribautiana sciotoensis, Young, Bull. Univ. Kansas Sci. Bull., vol. 35, pt. 1, 1952,

p. 100.

Resembling R. lucaUa in external appearance, but with third
apical cell white, and with three pair of apical processes on aedeagal
shaft, the median pair branching.

Length. — 3.0 mm.

Color. — Head and pronotum chalky white with a yellowish
tinge; scutellum dark brown, darker in median basal area; fore wing
with a pronounced dark brown band over white crossveins, extend-
ing farther anteriorly than posteriorly.

Genital capsule. — Male pygofer, in lateral aspect, with posterior
margin distinct, ventral angle projecting beyond dorsal angle.

Internal male genitalia. — Aedeagus, with pair of atrial processes
narrow, reaching to two thirds length of shaft, separated at base by
three times width of shaft, directed dorsocaudad parallel to each
other, apex of shaft slightly broadened and strongly curved ventrad,
bearing three pairs of apical processes the superior pair arising
before apex, incurving, median pair branched, directed laterad,
inferior pair curved sharply ventrad, apices directed toward apex
of shaft in lateral aspect; aedeagal shaft sharply curved, narrowing
from broad base to complex apex, forming an incomplete semi-
circle, somewhat swollen at juncture of processes. Illustrations for
this species are adapted from the original description by permission
of the author.

The following specimens have been seen: one paratype, Scioto
County, Ohio, June 17, and one female from Salamanca, New York,
July 24.

Types. — Holotype, allotype, and male paratype in the collection
of Mrs. J. N. Knull.

Revision of the Species Typhlocyba 1129

Ribautiana unca (McAtee)

(PI. LXXIV, fig. 4)

Typhlocyba unca McAtee, Proc. U. S. Nat. Mus., vol. 68, art. 18,^1926, p. 8.
Ribautiana unca, Young, Bull. Univ. Kansas Sei. Bull., vol. 35, pt. 1, 1952,

p. 100.
Typlilocyba surda DeLong and Johnson, Ent. News, vol. 47, no. 4, April, 1936,

pp. 101-102. (new synonymy).
Ribautiana surda, Young, Bull. Univ. Kansas Sei. Bull., vol. 35, pt. 1, 1952,

p. 100.

Resembling R. piscator in external appearance, but with a pair of
processes arising from near middle of aedeagal shaft, apical proc-
esses short and without numerous spines.

Length. — 3.0 mm.

Color. — Head, pronotum, and scutellum pale milky-white to light
orange yellow.

Genital capsule. — Male pygofer, in lateral aspect, somewhat quad-
rate, dorsal angle rounded, projecting beyond ventral angle.

Internal male genitalia. — Aedeagus, with atrial processes elongate,
nearly as long as shaft, evenly tapered from base to acute apices,
sinuate; shaft elongate, slender on apical half, with a pair of dorso-
laterally directed processes arising at or slightly below middle, apex
curved toward apices of atrial processes, bearing at apex from one
to three pairs of short processes which show a marked degree of
variability. ( PL LXXIV, figs. 4d, e, f, g. )

R. surda (DeLong and Johnson) is based upon a specimen of
this species with the apical processes broadened toward the base,
and is considered synonymous with R. unca (McAtee).

A series of specimens from Glen Haven, Colorado, differ from
other specimens seen in having the median pair of processes arising
from the base at a point lower than the aedeagal apodeme.

Numerous specimens of this species have been collected by Dr.
R. H. Reamer and the author from Corylus americana. Other speci-
mens have been seen with the following host labels: "vine maple,"
"on filbert," "alder," and "on beech."

Specimens have been seen from the following localities: Maine:
Fryeburg, August 20; Massachusetts: Holliston, September 4; New
Hampshire: Durham, September 9; New York: Cranberry Lake,
July 25; Salem, June 27, July 27; Minetto, July 23, September 16;
Pennsylvania: Hartstown Rog, September 12, 13, 14; Snowshoe,
August 22; Northeast, July 4; Ontario: Ottawa, June 24, Trenton,
September 30; Tennessee: Gatlinburg, June 29; Great Smoky Moun-
tain National Park, September 1; Virginia: Glencarlyn, June 12;
Michigan: September 23, Gogebic, August 18; Wisconsin: Mil-

1130 The University Science Bulletin

waukee, June 26-July 5; Lake Geneva, June 21; Minnesota: Itaska
County, July 16, 26, 27; Anoka County, September 21; Illinois:
Salem, September 22; Manitoba: Deepdale, August 1; Missouri:
Goodman, May 28; Kansas: Douglas County, May 27, 30, June 1, 7;
Jefferson County, June 15; Colorado: Glen Haven, August 3; Ore-
gon: Woodburn, October 2; Selma, June 14; California: Marin
County, August 3; Giant Forest, July 28; Fort Dick, July 13; Leona
Hights, Alameda County, August; Wild-Cat Canyon, San Pablo,
Contra Costa County, May 16.

Types. — Holotype male, and paratype males and females, in the
U. S. National Museum Collection; allotype female and paratypes
of both sexes in the Iowa State College Collection.

Mcateeana gen. nov.
(PL LXXIII, fig. 4)

Type of the genus, Empoa querci var. sexnotata Van Duzee.

Fore wings. — Inner and outer apical cells short, not attaining
wing apex; second apical cell much broader at apex than at base;
third apical cell petiolate; wing apex smoothly rounded. (Pi.
LXXIII, fig. 4h).

Hind wings. — Vein IV branching from vein 2V near its midlength;
submarginal vein absent at wing apex; both apical cells open api-
cally; posterior branch of R fused with apical portion of vein M1+2.
(PI. LXXIII, fig. 4h).

Genital capsule. — Male pygofer, in lateral aspect, with a small
group of short macrosetae dorsad of outer basal angle of male plate
and a few microsetae caudad of these, with posterior margin slightly
inrolled and bearing a few microsetae near posterior angle on mesal
surface, with awl-shaped spines scattered on posterior third, pygofer
hooks wanting; male plate in ventral aspect, broadened at base,
widest at basal fourth, gradually reduced to upturned apex, in lateral
aspect with single macroseta near outer basal angle, a row of micro-
setae on dorsolateral margin of apical two thirds, others along
ventrolateral margin on apical half.

Internal male genitalia. — Style elongate, slender, gradually taper-
ing and curved laterad or dorsad, abruptly curved ventrad before
apex, inner margin with several alveoli on outer margin, with a few
setae near middle, and a single large seta basad of apical curve;
connective Y-shaped; aedeagal articulation subterminal; aedeagus
with preatrial arm well developed, aedeagal apodeme well de-
veloped, shaft not present, gonopore at base of fused portion of
apically branching atrial processes.

Revision of the Species Typhlocyba 1131

Female. — With posterior margin of eighth abdominal sternite as
shown in PI. LXXXVII, figs. 5a, b.

The head in dorsal aspect narrower than pronotum, moderately
produced, its median length not greatly exceeding length next the
eye, anterior margin of crown smoothly rounded; face convex, di-
vergent from the line of the dorsum; ocelli present; pronotum short,
with lateral margins greatly diverging caudally, posterior margin
shallowly emarginate.

The following species is the only representative of this genus
known.

Mcateeana sexnotata (Van Duzee) (new combination)

(Pi. LXXIII, fig. 4)

Empoa querci var. sexnotata Van Duzee, Trans. San Diego Soc. Nat. Hist.,

vol. 2, no. 1, 1914, p. 57.
Typhlocyba querci var. sexnotata, McAtee, Canadian Ent., vol. 51, no. 8, 1919,

pp. 225-226.
Typhlocyba sexnotata, McAtee, Proc. U. S. Nat. Mus., vol. 68, art. 18, 1926,

p. 34.
Ossiannilssonia sexnotata, Young, Bull. Univ. Kansas Sci. Bull., vol. 35, pt. 1,

1952, p. 99.

Length. — 3.5 mm.

Color. — Head and pronotum yellowish-white; scutellum orange-
red, disc yellow; fore wings whitish hyaline, with inner half of clavus
and most of brachial cell light orange-yellow, with a large brown
spot in apex of brachial cell and two round black spots on clavus,
one at middle, the other near apex of scutellum, with commissural
vein black near apex of scutellum in some specimens, distal ends
of second and third basal cells, and apical cells fumose; abdomen
with dorsum black, apical margin of each segment yellow, venter
yellow, male plates yellow.

Internal male genitalia. — Aedeagus with a pair of stout, forcipate,
basally fused atrial processes bearing two pairs of slender, pos-
teriorly directed, apical branches, the superior pair directed dorso-
caudad, each pair crossing apically; aedeagal apodeme directed
cephalad for three fourths length of preatrial arm.

Specimens labeled "sycamore", and others labeled "Ribes" are on
hand. All specimens seen by the author are from California.

Types. — Holotype female, in California Academy of Sciences Col-
lection; neoallotype male, Gilroy, California, August, 1942, R. H.
Smith, here designated, and one male paraallotype, San Luis Obispo,
August, 1942, R. H. Smith, in the Snow Entomological Cellections
of the University of Kansas. Additional paraallotype males: one,
Cucamonga, California, December 21, 1917, in the Cornell Uni-

1132 The University Science Bulletin

versity Collection; one, Niles Canyon, California, July 15, 1916,
E. P. Van Duzee, in the California Academy of Sciences Collection.

Genus Ossiannilssonola nov. nom.

(Pis. LXXVI-LXXX)

Ossianniksonia Young and Christian, in Young, Bull. Univ. Kansas Sci. Bull.,
vol. 35, pt. 1, 1952, pp. 97-99.

Type of the genus, Typhlocyba berenice McAtee, by original
designation.

Fore icings. — Inner and outer apical cells short, not attaining
wing apex; second apical cell much broader at apex than at base;
third apical cell petiolate; wing apex smoothly rounded.

Genital capsule. — Male plate, in ventral aspect, gradually nar-
rowed on outer margin towards apex, occasionally with small lobe
on lateral margin before apex (O. appendiculata and O. phryne),
without single macroseta near outer basal angle of male plate,
with a dorsal submarginal row of setae and a row of microsetae
along ventrolateral margin on apical half; pygofer, in lateral aspect,
with posterior margin distinct, not rounded to ventrocaudal margin,
rectilinear in form; pygofer wall frequently with sclerotized bars
occurring within the limits of pygofer proper, or prolonged dorso-
caudad or caudad as short pygofer hooks; a group of macrosetae
usually present on pygofer just dorsad of outer basal angle of male
plate, with numerous small awl-shaped cuticular projections over
dorsal posterior half of pygofer, and a row of microsetae on disc
extending dorsocaudad from the group of macrosetae; anal hooks
wanting.

Internal male genitalia. — Style elongate, slender, gradually curved
laterad or dorsolaterad apically, usually without triangular pre-
apical projection on inner margin (exceptions, O. danae, O. bang-
soni, and O. flavomarginata), with elongate slender setae on outer
margin, with a few alveoli on inner margin; connective triangular or
Y-shaped; aedeagus with preatrial arm distinct, aedeagal apodeme
short, well developed or not, shaft occurring as a flattened mem-
branous structure between basal portions of a pair of forcipate
atrial processes.

Female. — With posterior margin of eighth abdominal sternite
usually broadly evenly rounded as in O. berenice (PI. LXXXVII,
figs. 6a, b), except in O. jlavomarginata where it is strongly incised

Revision of the Species Typhlocyba 1133

laterally forming two lateral lobes and a slightly emarginate median
lobe (PI. LXXXVII, figs. 7a, b).

Head, including the eyes, narrower than pronotum, only slightly
produced medially on rounded anterior margin, median length of
the crown not greatly exceeding length next the eyes, face strongly
convex to slightly below antennal insertions, then slightly convex
to tip of clypellus; ocelli present or absent, when present, situated
on round margin between crown and face, nearer the eyes than to
each other; pronotum short, but much longer than head, lateral
margins greatly divergent caudally, posterior margin shallowly
concave; width of pleural portion greatly exceeds width of ocel-
locular area.

The species are usually pale in color, with dark markings which
are occasionally extensive.

Most of the species of this genus live on species of Quercus, but
Ulmus, Crataegus, Acer, and Primus are also host genera. None of
the known North American species has been recorded from out-
side of this continent, but from the figures of Typhlocyba callosa
Then, by Ribaut ( 1936 ) , it appears that this species could belong to
this genus, and should this be true, the range of this genus would
be holarctic.

Key to the Species of Ossiannilssonola

1. Male plate distinctly forked near apex 2

Male plate not forked 3

2. With black parenthesis-shaped marks on pronotum; fore wing

with narrow black lines along commissural margin to cross-
veins; pygofer with posterior angle strongly produced in a

broad rounded lobe. (Pi. LXXIX, fig. 2a) phryne p. 1152

With light colored pronotum, pygofer with posterior margin
nearly vertical, only slightly produced on dorsal angle. (Pi.
LXXIX, fig. la) appendiculata p. 1150

3. Styles with apical third sharply curved mesad, a small triangular

protuberance on lateral margin. (Pi. LXXX, fig. 2e),

flavomarginata p. 1157
Styles with apical third not curved mesad 4

4. Styles appearing truncate at apex, with small projection near apex, 5
Styles evenly tapering to an acute apex 6

5. Triangular projection on lateral margin of style near apex; pygofer

with two vertically arranged, posteriorly directed hooks on

dorsal posterior margin. (Pi. LXXVIII, fig. 1) bangsoni p. 1145

Triangular projection on mesal margin of style near apex; pygofer
with two horizontally arranged, mesally directed hooks on dor-
sal posterior margin. (Pi. LXXVIII, fig. 5) danae p. 1149

6. Atrial processes forked near apex 7

Atrial processes not forked 8

1134 The University Science Bulletin

7. Pygofer with posterior margin nearly vertical, dorsal angle form-

ing a broad, blunt, dorsomesally directed process. (PL

LXXVI, fig. 1 ) berenice p. 1136

Pygofer with posterior margin obliquely slanted toward base of
plate, dorsal angle forming a moderately rounded projection
with two mesally directed, spinelike processes. (PL LXXVI,
fig. 2), hermione p. 1137

8. Atrial processes tumid, apical third sharply reduced, slender,

thornlike 9

Atrial processes not tumid, gradually tapering to apex 11

9. Atrial processes greatly broadened at base in posterior aspect,

lateral margins nearly parallel; fore wing with entire basal half
bright crimson red, without black markings. (PL LXXVII,

fig. 3 ) tunicaruhra p. 1141

Atrial processes not greatly broadened at base in posterior aspect,
lateral margins divergent toward apex; fore wings white to
yellow with black spots in apices of inner three basal cells .... 10

10. Posterior margin of tumid portion of atrial processes forming

nearly a right angle near middle, in lateral aspect. (PL

LXXVII, fig. 4) australis p. 1142

Posterior margin of tumid portion of atrial processes forming an
evenly rounded semicircle, in lateral aspect. (PL LXXVII,
fig. 2 ) clymene p. 1140

11. Ventral angle of pygofer directed caudally as a small thumblike

lobe. (PL LXXVI, fig. 3) volans p. 1138

Ventral angle of pygofer without lobe, or with lobe directed
ventrad when present 12

12. Atrial processes gradually curved dorsocaudad 13

Atrial processes abruptly bent caudad 14

13. Dorsal angle of pygofer projecting caudad beyond ventral angle

as a broad quadrate process bearing two mesally directed di-
verging spines. (PL LXXVIII, fig. 2) duplicata p. 1146

Dorsal angle of pygofer not projecting caudad as a quadrate
process, but as two short fingerlike processes which cross each
other. (PL LXXVII, fig. 1) antigone p. 1139

14. Dorsal angle of pygofer forming a broad caudally directed hook,

lower half of the posterior margin S-shaped from the left side,

ventral angle broadly rounded 15

Dorsal angle of pygofer not forming a hook, posterior margin
straight, ventral angle somewhat angulate 17

15. Pygofer hook large, half as deep as broad; atrial processes of

aedeagus broadly flattened beyond membrane, appearing

forked in posterior aspect. (PL LXXIX, fig. 3) rossi p. 1153

Pygofer hook small, one third to one fourth as deep as broad;
atrial processes not appearing forked in posterior aspect 16

16. Base of aedeagus greatly broadened laterally, abruptly reduced

before bases of atrial processes to one third its greatest width;
with sclerotized plate extending over bases of atrial processes;
pygofer with two acute mesocaudally directed spines on dorsal
posterior margin. (PL LXXX, fig. 3) quadrata p. 1155

Revision of the Species Typhlocyba 1135

Base of aedeagus of almost uniform width; bases of atrial proc-
esses not covered by a sclerotized plate; pygofer with only
slight projections on dorsal posterior margin. (Pi. LXXIX,
fig. 4) knulli p. 1154

17. Atrial processes widely separated at base, apices finely serrate.

(PI. LXXVIII, fig. 3) serrula p. 1147

Atrial processes not widely separated at base, apices smooth 18

18. Dorsal angle of pygofer excavated; fore wings milky-white, with

two dark crossbands, anterior band interrupted by white

scutellum. (Pi. LXXVII, fig. 5) hinei p. 1144

Dorsal angle of pygofer not excavated; fore wings with black
spots only in apices of inner three basal cells 19

19. Dorsal angle of pygofer directed dorsad as a broad acutely-angled

hook, ventral angle produced as a short caudally directed hook.

(PI. LXXX, fig. 1) troza p. 1156

Dorsal angle of pygofer not directed dorsad as a broad acutely
angled hook, but mesocaudad as a short broad lobe; ventral
angle rounded, not projecting caudad. (Pi. LXXVIII, fig. 4),

mcateei p. 1148

Many of the species of this genus can be distinguished on the basis
of color markings. The following key based on fully colored speci-
mens will permit the determination of female specimens of a num-
ber of species, but closely colored species which are not easily dis-
tinguished have of necessity been grouped together.

Color Key to the Species of Ossiannilssonola

1. Without black or brown markings on fore wings 2

With black or brown markings, or traces of these on fore wing ... 6

2. Fore wings with crimson, or light red markings 3

Fore wings white or orange-yellow 4

3. Anterior half of fore wings bright crimson-red, white on remain-

der of dorsum tunicarubra p. 1141

Entire fore wing colored fight red; head, pronotum, and scutellum

bright yellow flavomarginata type III p. 1157

4. Dorsum white to cream colored flavomarginata type V p. 1158

Dorsum orange-yellow 5

5. Length 3.25 mm flavomarginata type IV p. 1158

Length 3.5-3.75 mm serrula p. 1147

6. Dark markings extensive, not restricted to three spots anterior to

cross veins 7

Dark markings not extensive, restricted to three spots, one in

each of inner three basal cells just anterior to cross veins 13

7. Dark markings in the form of two transverse bands 8

Dark markings not as two transverse bands 10

8. Anterior band near base of fore wings, interrupted by white

scutellum hinei p. 1144

Anterior band near middle of fore wing 9

42—6490

1136 The University Science Bulletin

9. Posterior band solid color volans p. 1138

Posterior band made up of six distinctly separated spots, ground
color of dorsum milky-white, resembling forms of Empoa, casta
group duplicata p. 1146

10. Dark markings covering entire dorsum flavomarginata type II p. 1157

Dark markings less extensive 11

11. Dark markings forming a brown saddle over fore wings anterior

to cross veins, sometimes limited in extent to only the apical
third of inner three basal cells and a stripe along claval suture,

appendiculata (in part) p. 1150
Dark markings otherwise 12

12. With dark parenthesis-shaped markings on pronotum which con-

tinue posteriorly to cross veins as a black commissural line,

phryne p. 1152
With dark markings in apical cells along cross veins and apical

veins berenice (in part) p. 1136

13. Ground color orange-yellow,

antigone, berenice (in part), quadrata, rossi, and kmdli
Ground color white to light yellow 14

14. With a flavescent stripe along commissural suture 15

Without such a stripe 16

15. With row of black spots slanting obliquely toward apex of wing,

flavomarginata type I p. 1157
With row of black spots nearly transverse australis (in part) p. 1142

16. Ground color cream, dark brown spots filling most of outer fourth

or fifth of inner three basal cells, irregular in intensity,

appendiculata (in part) p. 1150
Ground color white, dark spots smaller, of even intensity,

danae, ch/mene, hcrmione, troza, mcateei,
bangsoni, australis (in part)

Ossionnilssonola berenice (McAtee) (new combination)

(PI. LXXVI, fig. 1)

Typhlocyba berenice McAtee, Proc. U. S. Nat. Mus., vol. 68, art. 18, 1926, p. 38.
Ossiannilssonia berenice, Young, Bull. Univ. Kansas Sci. Bull., vol. 35, pt. 1,
1952, p. 99.

Resembling O. hermione in shape of the aedeagus, but differing
in lacking spinelike hooks on mesal margin of dorsal angle of
pygofer, and in having posterior margin of pygofer vertical.

Length. — 3.25-3.5 mm.

Color. — Head, pronotum, and scutellum yellow to orange-yellow;
fore wings light yellow to uniform deep orange-yellow anterior to
cross veins, lighter apically, veins sometimes red-orange, wing sub-
hyaline to cross veins, hyaline apically, with a transverse row of
three black spots in apices of inner three basal cells separated from
veins by a narrow yellow margin; apical cells smoky along veins and
outer margins, forming indistinct spots in apical cells three and

Revision of the Species Typhlocyba 1137

four; abdomen with dorsal segments black medially, yellow to
orange-yellow laterally, venter yellow, male plate yellow. Black
spots sometimes reduced or missing.

Genital capsule. — Male pygofer, in lateral aspect, with posterior
margin vertical, dorsal angle forming a broad blunt dorsally directed
projection, ventral angle nearly a right angle, a few macrosetae on
posterior margin near dorsal angle.

Internal male genitalia. — Aedeagus, with atrial processes elon-
gate, slender, apices almost meeting medially, in posterior aspect
outlining a rectangle, a single pair of slender subapical processes
arising from the posterior margin and curving mesocaudad, cross-
ing at outer fourth; aedeagal apodeme one fourth length of atrial
processes, directed cephalad on apical third in lateral aspect.

Several thousand specimens of this species have been collected by
the author from Quercus alba in Milwaukee, Wisconsin, in associa-
tion with O. australis and O. danae.

Specimens have been seen from the following localities: Ontario:
Toronto, August 8; Massachusetts: Wood's Hole, July 10; Con-
necticut: New Haven, July 4; New York: Ithaca, July 8; District of
Columbia: Rock Creek, June 19; Virginia: Arlington, October 12;
Mountain Lake, July 17; North Carolina: Franklin, August 17;
Illinois: Thornton, September 7; Wisconsin: Milwaukee, June 26-
July 7; Minnesota: Ramsay County, August 30; St. Paul, June 27.

Types. — Holotype male and one male paratype in the U. S. Na-
tional Museum Collection, Washington, D. C; a female specimen
from Milwaukee, Wisconsin, June 30, 1950, taken in copula with a
male of this species, here designated neoallotype, in the Snow Ento-
mological Collections of the University of Kansas.

Ossiannilssonola hermione (McAtee) (new combination)

(PI. LXXVI, fig. 2)

Typhloct/ba hermione McAtee, Proc. U. S. Nat. Mus., vol. 68, art. 18, 1926,

pp. 38-39.
Ossiannilssonia hermione, Young, Bull. Univ. Kansas Sci. Bull., vol. 35, pt. 1,

1952, p. 99.

Resembling O. berenice in shape of aedeagus, but differing in
having two spinelike hooks on mesal margin of dorsal angle of
pygofer, and in having ventrocaudal margin oblique.

Length. — 4.0 mm.

Color. — Head, pronotum, and scutellum white to yellowish-white;
fore wings whitish-hyaline with three black spots anterior to cross
veins, apical cells slightly fumose; abdomen white, male plates
white.

1138 The University Science Bulletin

Genital capsule. — Male pygofer, in lateral aspect, with ventro-
caudal margin obliquely slanted from base of plate to dorsal angle,
ventral angle directed ventrad in a lobe, dorsal angle appearing as
a moderately rounded projection with two vertically arranged tooth-
like processes directed mesad from inner margin, upper process
one third length of lower.

Internal male genitalia. — Aedeagus with atrial processes elon-
gate, slender, gradually reduced toward apex, curving ventromesad
and continuing anteriorly, a pair of short subapical processes arising
from dorsal surface; aedeagal apodeme a short apically enlarged
shaft directed cephalodorsad.

The following specimens have been seen by the author: holotype
male, Washington, D. C, July 2; paratype male, Madison, Wis-
consin, August 11; paratype male, Bluemont, Virginia, July 1; one
male, Northeast, Pennsylvania, July 7.

Types. — Holotype male and paratype males, in the U. S. National
Museum Collection.

Ossiannilssonola volans (McAtee) (new combination)

(Plate LXXVI, fig. 3)

Typhloei/ba qaerci var. volans McAtee, Canadian Ent., vol. 51, no. 8, 1919,

pp. 225-226.
Typhlocyha gillettei var. volans, McAtee, Proc. U. S. Nat. Mus., vol. 68, art. 18,

1926, p. 28.

Resembling O. hinei in color markings, but differing in having
anterior band at middle of fore wing; ventrocaudally directed
thumblike lobe on pygofer distinguishing it from other species in
the genus.

Length. — 3.75-4.0 mm.

Color. — Head, pronotum, and scutellum light yellow; fore wing
light yellow overlaid with variable dark brown markings which
usually form two broad bands, one over the cross veins forming a
broad inverted V, the other V-shaped, arising at the middle of
clavus and slanting obliquely forward to costal margin; some speci-
mens from the Great Smoky Mountains taken on Primus pennsyl-
vanicus differ from this pattern by having a longitudinal band con-
necting these transverse bands, others by having only traces of this
band; specimens collected from Ulmus fulva differ by having the
anterior band transverse and posterior band nearly transverse; apical
cells fumose; abdomen with black markings on dorsum, venter
yellow, plates yellow.

Genital capsule. — Male pygofer, in lateral aspect, with dorsal
angle a broad rounded lobe projecting caudally beyond ventral

Revision of the Species Typhlocyba 1139

angle, without hooks, ventral angle produced caudally into a thumb-
like lobe, a number of macrosetae scattered submarginally along
dorsal two thirds of posterior margin.

Internal male genitalia. — Aedeagus, with atrial processes elon-
gate, slender, unbranched, in caudal aspect describing an oval,
apices sharply directed ventromesad and crossing near tip, aedeagal
apodeme reduced to a short anteriorly directed arm.

Several hundred specimens have been collected by the author in
Milwaukee, Wisconsin, and in Lawrence, Kansas, from Ulmus fulva.
Additional specimens seen are from the following localities: On-
tario: Vineland Station, August 2; New York: Monroe, July 10; Ba-
tavia, July 15; Indian Lake, Sabael, August 25; Pennsylvania: Harts-
town Bog, June 26; Tennessee: Great Smoky Mountain National
Park, July 30, September 1; Michigan: Agricultural College, July 7;
Wisconsin: Marshfield, August 20; Milwaukee, June 26-July 7; Utah:
Richfield, July 15; Kansas: Douglas County, June 9, August; Ore-
gon: Portland, August 1; Independence, June 21; Azalea, Septem-
ber; McMinville, August 15; California: Placer County, June 16.

One pair of paratype specimens of Typhlocyba gillettei var. casta
McAtee, bearing the label "Douglas Co. Ks. August 1923 W. Robin-
son" are specimens of this species like those collected on Ulmus
fulva.

Types. — Mr. M. E. Neary, Entomologist for the Nova Scotia De-
partment of Agriculture and Marketing, informs me that the holo-
type female of this species was destroyed by fire in June, 1946.

Because of the lack of material from the type locality, and the
diversity of color pattern found in the specimens on hand, the
author considers it best not to erect a neotype for this species until
specimens from the type locality are available.

Ossiannilssonola antigone (McAtee) (new combination)

(PL LXXVII, fig. 1)

Typhlocyba antigone McAtee, Proc. U. S. Nat. Mus., vol. 68, art. 18, 1926,

pp. 35-36.
Ossiannilssonia antigone, Young, Bull. Univ. Kansas Sci. Bull., vol. 35, pt. 1,

1952, p. 99.
Typhlocyba eurydice var. distincta McAtee, Proc. U. S. Nat. Mus., vol. 68,

art. 18, 1926, p. 38. (new synonymy)

Resembling O. berenice externally, but distinguished by having
two fingerlike processes at dorsal angle of pygofer, and in not
having atrial processes of aedeagus branching near apex.

Length. — 3.5 mm.

Color. — Head, pronotum, and scutellum pale yellow to orange-

1140 The University Science Bulletin

yellow; fore wing pale yellow to orange-yellow, subhyaline to cross
veins, apex hyaline with indistinct black spots in apices of inner
three basal cells, veins in some specimens orange-red, in others
almost white; abdomen yellow, plates yellow.

Genital capsule. — Male pygofer, in lateral aspect, with posterior
margin sinuate, dorsal angle produced in two short fingerlike proc-
esses directed mesocaudad and crossing each other, ventral angle
forming an angular lobe projecting caudally as much as dorsal
processes, without macrosetae dorsad of outer basal angle of plate,
with two or three macrosetae on dorsal posterior margin.

Internal male genitalia. — Aedeagus with atrial processes short,
stout, half as long as base in lateral aspect, with apices directed
toward each other and slightly caudad, describing almost a com-
plete circle in caudal aspect; aedeagal apodeme short, as broad as
long in lateral aspect.

Only a few specimens of this species have been seen. The author
has taken a single pair from Qiiercus alba in Jefferson County, Kan-
sas, in association with O. titnicarabra, and a single male from the
same host in Milwaukee, Wisconsin. Other specimens seen are from
the following localities: Connecticut: New Haven, July 4; Mary-
land: Beltsville, June 23; Delaware: Wilmington, June 20; Virginia:
Glencarlyn, June 12, 16, 20; Arlington, June 14; Mountain Lake, July
23; Illinois: Monticello, June 11; Wisconsin: Milwaukee, July 3;
Kansas: Jefferson County, June 15.

The holotype and six paratype males have been seen.

Types. — Holotype male, in the Illinois State Natural History
Survey Collection; paratypes in the U. S. National Museum, and
DeLong Collections; a female specimen collected by the author in
Jefferson County, Kansas, June 15, 1950, here designated neoallo-
type, in the Snow Entomological Collections of the University of
Kansas.

The holotype specimen of Typhlocyba eurydice var. discincta
McAtee, from Beltsville, Maryland, has been dissected and found
to be a specimen of O. antigone.

Ossiannilssonola clymene (McAtee) (new combination)

(PI. LXXVII, fig. 2)

Typhlocyba clymene McAtee, Proc. U. S. Nat. Mus., vol. 68, art. 18, 1926,

pp. 36-37.
Ossiannilssonia clymene, Young, Bull. Univ. Kansas Sci. Bull., vol. 35, pt. 1,

1952, p. 99.

Resembling O. australis in external appearance and in the shape
of the aedeagus, but differing in having the ventral angle of the

Revision of the Species Typhlocyba 1141

pygofer rounded apically and slightly projecting caudad, and in
having ventral margin of atrial processes of aedeagus rounded, not
angular.

Length — 3.5-3.75 mm.

Color. — Head, pronotum, and scutellum white to light yellow;
fore wings pale yellowish-white with a transverse row of three
black spots in apices of inner three basal cells, apical cells hyaline;
abdomen white, male plates white.

Genital capsule. — Male pygofer, in lateral aspect, with posterior
margin nearly vertical, dorsal and ventral margins nearly parallel,
dorsal and ventral angle evenly rounded, not produced, a row of
macrosetae along dorsal half of posterior margin.

Internal male genitalia. — Aedeagus with atrial processes short,
great* y enlarged to outer third, sharply reduced on outer third to
slender elongate acute apices directed ventromesad and posteriorly
and crossing at apex, processes appearing cordate in caudal aspect;
aedeagal apodeme slender, as long as apical portion of atrial proc-
esses, directed anteriorly; aedeagus with posterior margin slightly
sinuate, posterior margin of atrial processes smoothly rounded, not
angulate.

A short series of three females and four males from Vineland
Station, Ontario, collected from Querciis alba by W. L. Putman,
July 8, 1931, and the holotype, McLean, New York, July 14, 1919,
are the only specimens known to exist. The aedeagus of the holo-
type has been lost.

Types. — Holotype male, in U. S. National Museum Collection,
Washington, D. C. A female specimen from the above mentioned
series, here designated neoallotype, in the Canadian National Col-
lection, Ottawa, Canada.

Ossiannilssonola tunicarubra (Gillette) (new combination)

(PI. LXXV, fig. 3)

Typhloci/ba tunicarubra Gillette, Proc. U. S. Nat. Mus., vol. 20, no. 1138, 1898,

pp. 752-753.
ErytJironeura tunicarubra, Van Duzee, Cheek List of Hemiptera (excepting

the Aphididae, Aleurodidae and Coecidae) of America North of Mexico,

1916, p. 77.
Ossiannilssonia tunicarubra, Young, Bull. Univ. Kansas Sci. Bull., vol. 35, pt. 1,

1952, p. 99.

Resembling O. clymene and O. australis in structure of male
genitalia, but easily distinguished from these species by having
anterior half of fore wings bright crimson red, and bases of atrial
processes of aedeagus enlarged in caudal aspect.

Length. — 3.5-4.0 mm.

1142 The University Science Bulletin

Color. — Head, pronotum, and scutellum bright lemon-yellow;
fore wings from base to short distance before cross veins crimson
red, forming a sharply distinct transverse line posteriorly, remainder
of wing yellow, apical cells hyaline; abdomen, dorsum black with
outer margins of segments yellow, venter bright lemon-yellow, plates
yellow.

Genital capsule. — Male pygofer, in lateral aspect, with posterior
margin vertical, dorsal margin horizontal, ventral margin obliquely
slanting dorsad, dorsal and ventral angles evenly rounded, not pro-
duced except for small tooth on mesal margin near dorsal angle, a
row of macrosetae along dorsal half of posterior margin.

Internal male genitalia. — Aedeagus with atrial processes short,
greatly enlarged laterad on basal third, slightly reduced on median
third, greatly reduced to slender, elongate, acute apices directed
ventromesad on outer third and crossing at apex, in caudal aspect
almost quadrate in outline; aedeagal apodeme slender, as long as
apical third of atrial processes, directed ventrocephalad, posterior
margin in lateral aspect strongly sinuate, margin of atrial processes
smoothly rounded, not angulate.

A short series of specimens of this species were collected from
Quercus alba by the author in Jefferson County, Kansas, June 15,
1950, including a number of teneral females which show only a
slight tinge of pink on the anterior half of the fore wings, and three
males of intermediate color intensity. Final instar nymphs collected
with these bear wing pads which are also colored pink.

Specimens have been seen from the following localities: New
York: Conesus Lake, July 16; Ithaca, July 22; Pennsylvania: Point
Royal, July 24; Virginia: Mountain Lake, July 24; Tennessee: Knox-
ville, June 24; Clarksville, July 5; Ohio: Delaware County, July 4,
8, 2, 27, August 3; Michigan: Agricultural College, July 15; Wis-
consin: Lake Geneva, July 21; Iowa: Ames, July 26; Kansas: Doug-
las County, August; Jefferson County, June 15; Atchison, July 8.

Types. — Holotype female, in the U. S. National Museum Collec-
tion, Washington, D. C; a male specimen compared with holotype,
collected by the author in Jefferson County, Kansas, June 15, 1950,
here designated neoallotype, is in the Snow Entomological Collec-
tions of the University of Kansas.

Ossiannilssonola australis (Walsh) (new combination)

(PI. LXXVII, fig. 4)

Enjthroneura australis Walsh, The Prairie Fanner, (n. s.), vol. 10, no. 10, Sep-
tember 6, 1862, p. 149.
Typhlocyba australis, Woodworth, Psyche, vol. 5, no. 157-159, 1889, p. 214.

Revision of the Species Typhlocyba 1143

Empoa australis, Van Duzee, Check List of Hemiptera (excepting the Aphid-
idae, Aleurodidae and Coccidae) of America North of Mexico, 1916, p. 77.

Typhlocyba nicarete McAtee, Proc. U. S. Nat. Mus., vol. 68, art. 18, 1926,
p. 36. (new synonymy)

Ossiannilssonia nicarete, Young, Bull. Univ. Kansas Sci. Bull., vol. 35, pt. 1,
1952, p. 99.

This species is not Empoasca australis Froggatt (1918), which
was subsequently moved into Typhlocyba ( Myers, 1921 ) , renamed
Typhlocyba froggatti (Baker, 1925), as a secondary homonym, and
has recently been moved to Edwardsiana ( China, 1950 ) . Synonymy
of that species is given on p. 1220 of this paper.

Resembling O. clymene externally in lightly colored specimens,
and in shape of aedeagus, but differs in having the ventral angle
of the pygofer slightly angular and reduced cephalad, and in having
ventral margin of atrial processes of aedeagus angular near middle.

Length. — 3.5 mm.

Color. — Head, pronotum, and scutellum light yellow to yellow;
fore wings show marked progressive coloration from whitish-hyaline
with only veins yellow, to orange-yellow or deep orange-yellow with
longitudinal red band along commissural margin from base of wing
to cross veins on inner half of clavus, with a transverse row of three
sharply defined black spots in apices of inner three basal cells;
abdomen with dorsum black, outer margin of each segment yellow,
male plates yellow.

Genital capsule. — Male pygofer, in lateral aspect, with posterior
margin slightly convex, with dorsal angle evenly rounded, ventral
angle reduced forming a slight projection on ventral margin, a row
of macrosetae along dorsal half of posterior margin.

Internal male genitalia. — Aedeagus with atrial processes short,
stout, tumid on basal two thirds, sharply reduced on outer third to
a slender elongate acute apex directed ventromesad posteriorly, in
caudal aspect somewhat cordate in outline; aedeagal apodeme
slender, as long as outer third of atrial processes, directed an-
teriorly; posterior margin of aedeagus nearly straight to middle of
atrial processes, then abruptly curved dorsocephalad at nearly a
right angle.

Numerous specimens of this species have been collected by the
author from Quercus alba in Milwaukee, Wisconsin. Another
large series of this species from Illinois was seen in the Illinois
Natural History Survey Collection. Quercus macrocarpa has also
been recorded as a host plant.

The original description of Erythroneura australis Walsh (1862)
(nee. australis Froggatt, 1918) indicates that this species belongs

1144 The University Science Bulletin

in the genus Ossiannilssonola. In the absence of a type specimen
the author has concluded, after considering all of the species in
this genus which might fit the original description, that the majority
of specimens of the species Typhlocyba nicarete McAtee ( 1926 ) ,
which is abundant in Illinois, the type locality of australis, fit the
limited description more nearly than do specimens of any other
species seen.

Specimens have been seen from the following localities: New
York: Salem, July 27, 28; Pennsylvania: Greenberg, August 17;
Virginia: Blacksburg, Glencarlyn, June 12, 16, 20; Mountain Lake,
July 8-26; near District of Columbia, June 15; Washington, D. C;
Tennessee: Clarksville, July 11, 29; Tullahoma, August 3; Ohio:
Delaware County, August 27; Wooster, July 5; Shawnee Forest,
June 9; Minnesota: St. Paul, June 16, 22, August 6, 11; Wisconsin:
Milwaukee, June 27; Polk County; Illinois: Tinley Park, September
8; Olmsted, July 15; Urbana; Pere Marquette State Park, August 12
Decatur, August; Bell Smith Springs, July 16; Bellwood, June 21
Missouri: Goodman, May 28; Arkansas: Siloam Springs, May 26
Oklahoma: LeFlore County, May 24; Louisiana: Ida, June 6.

Types. — The holotype male of Typhlocyba nicarete McAtee,
White Heath, Illinois, June 24, 1916, here designated neotype of
Ossiannilssonola australis (Walsh), in the Illinois State Natural
History Survey Collection; a female specimen taken in copula
with a male of this species, July 3, 1950, Milwaukee, Wisconsin, by
the author, here designated neoallotype, in the Snow Entomological
Collections of the University of Kansas.

Ossiannilssonola hinei (Knull) (new combination)
(PI. LXXVII, fig. 5)

Typhlocyba hinei Knull, Ohio. J. Sci., vol. 44, no. 6, 1944, p. 272.
Ossiannilssonia hinei, Young, Bull. Univ. Kansas Sci. Bull., vol. 35, pt. 1, 1952,
p. 99.

Resembling O. volans in outward appearance, but with anterior
crossband at base of fore wings interrupted by white scutellum,
and with dorsal angle of pygofer excavated in lateral aspect.

Length. — 3.5-4.0 mm.

Color. — Head, pronotum, and scutellum chalky white; fore wings
chalky white, subhyaline, with two transverse chocolate-brown
bands, one at base of wings interrupted by white scutellum, the
other over cross veins formed by a row of four spots in apices of
basal cells, veins between spots white, size of spots progressively
reduced from inner to outer cell, apical cells whitish-hyaline; ab-
domen yellow, pygofer yellow.

Revision of the Species Typhlocyba 1145

Genital capsule. — Male pygofer, in lateral aspect, with posterior
margin vertical on lower two thirds, dorsal third shallowly exca-
vated, dorsal margin broadly produced, with dorsal angle ending in
a short, mesally directed spine, ventral angle forming an angular
ventrally directed lobe, several submarginal macrosetae near middle
of posterior margin.

Internal male genitalia. — Aedeagus with atrial processes elongate,
slender, gradually tapering toward apex, curved ventromesad cau-
dally and crossing at apices, in posterior aspect nearly describing a
circle; aedeagal apodeme broadly joined to base, enlarged apically
into a smoothly rounded club.

According to Mrs. J. N. Knull (1944) this species occurs on
Quercus alba in association with O. tunicarnbra.

Specimens have been seen from the following localities: Ohio:
Delaware County, June 25, July 2, 4, 5; Tennessee: Clarksville, July
1; North Carolina: Franklin, August 17.

Types. — Holotype female and female paratypes, in the collection
of Mrs. J. N. Knull, Columbus, Ohio; one paratype female, in the
Herbert Osborn Collection of the Ohio State University; a male
specimen, Delaware County, Ohio, July 5, 1945, D. J. and J. N.
Knull, here designated neoallotype, three paraallotype males from
Delaware County, Ohio, July 2, 5, 1945, and July 4, 1947, D. J. and
J. N. Knull, in the Knull Collection; five paraallotype males, Dela-
ware County, Ohio, July 5, 1945 and July 4, 1947, in the Snow Ento-
mological Collections of the University of Kansas.

In his revision of the genus Typhlocyba, (1926), McAtee makes
reference to a collection record by Mrs. Annie Trumbull Slosson
(1906) from Mount Washington, New Hampshire, of T. nitidula
( Fabricius ) , a synonym of T. bifasciata Boheman, and on the basis
of this record described it as doubtfully occurring in North America,
suggesting that the Slosson record might be based on one of the
varieties of Typhlocyba gillettei or T. cymba. It is the opinion of
the present author that this record was more likely based upon a
specimen of O. hinci which was described subsequently, since no
specimens of T. bifasciata Boheman (nee. bifasciata Gillette and
Baker, 1895) have been seen from North America.

Ossiannilssonola bangsoni sp. nov.
(PI. LXXVIII, fig. 1)

Resembling O. troza in external appearance, but differs by bear-
ing two short posteriorly-directed spinelike processes on dorsal
angle of pygofer.

Length. — 3.75 mm.

1146 The University Science Bulletin

Color. — Head, pronotum, and scutellum yellowish-white; fore
wing yellow with veins lemon yellow, subhyaline to cross veins, apex
hyaline and slightly fumose, a transverse row of three black spots
in apices of inner three basal cells.

Genital capsule. — Male pygofer, in lateral aspect, with posterior
margin convexly rounded, dorsal angle produced into a short acute
caudally directed hook, a blunt triangular process arising from the
mesal margin near dorsal third of posterior margin and directed
mesocaudad, ventral angle reduced to a small slightly angular
ventrally directed lobe, without macrosetae near outer basal angle
of plate, a few macrosetae on posterior margin along concavity
between processes.

Internal male genitalia. — Style with apex appearing somewhat
truncate, with small triangular projection on lateral margin near
apex, aedeagus with base greatly enlarged, atrial processes shorter
than base, gradually narrowed from base to apex, directed ventro-
mesad posteriorly, nearly meeting at apex, in lateral aspect with
ventral margin of basal two thirds produced medially forming nearly
a right angle, apical third abruptly recurved ventrad; aedeagal
apodeme reduced to a broad triangular anteriorly directed process
in lateral aspect.

This species is known from two male specimens both of which
were taken at light, and was doubtfully referred to as O. quadrata
(DeLong and Johnson) by Medler (1942).

Types. — Holotype male, Berea, Kentucky, July 4, 1941, J. S. Bang-
son, in the U. S. National Museum Collection; one paratype male,
Ramsey County, Minnesota, U. Farm Light, July 8, 1921, collected
by Wm. E. Hoffman, in the University of Minnesota Collection.
Since its description by Medler, and prior to its being sent to the
author, the body of the paratype was lost from the point, but the
dissection of the abdomen is complete, and in view of the clear
description of the specimen by Dr. Medler which agrees in every
particular with that of the holotype, it has been made a paratype
even though it is not now complete.

Ossiannilssonola duplicata (McAtee) (new combination)

(PI. LXXVIII, fig. 2)

Typhlocyba duplicata McAtee, Proc. U. S. Nat. Mus., vol. 68, art. 18, 1926,
pp. 16-17. (nee. T. duplicata Jacobi, 1941) (Typhlocyba jacobii, nom.
nov. for Typhlocyba duplicata Jacobi, see p. 1107.

Ossiannilssonia duplicata, Young, Bull. Univ. Kansas Sci. Bull., vol. 35, pt. 1,
1952, p. 99.

Resembling Empoa casta in color pattern, but with ground color

Revision of the Species Typhlocyba 1147

milky white, and without macroseta near base of plate; dorsal angle
of pygofer produced as a broad distinctly quadrate process.

Length. — 3.0 mm.

Color. — Head, pronotum, and scutellum milky white to light
yellowish white; fore wings milky white, with brown markings
forming two transverse bands, one near middle of wing variable in
width from a trace of a line in partly teneral specimens to a band
one fourth the width of wing, the other band over cross veins made
up of six spots, one in each of the inner three basal and first, sec-
ond, and fourth apical cells; abdomen, dorsum black with apical
half of each segment white to yellow; venter yellow, plates light
yellow.

Genital capsule. — Male pygofer, in lateral aspect, with posterior
margin sinuate on lower two thirds, dorsal third produced dorso-
caudad beyond ventral angle as a broad quadrate process bearing
two mesally directed diverging spines, ventral angle directed ven-
trad in a broad lobe slightly constricted near apex, a few macrosetae
along posterior margin of quadrate process.

Internal male genitalia. — Aedeagus with atrial processes elongate,
broad at base, gradually reduced to acute apices, outer margins
quadrate from posterior aspect, apices directed mesad toward each
other, widely separated, in lateral aspect sinuate, curved anteriorly
near middle, apex directed dorsad; aedeagal apodeme similar to
that of O. hinei, enlarged apically.

Several hundred specimens of this species were collected by the
author from Crataegus sp. in Milwaukee, Wisconsin, June 26-July 4.
The type series, from Toronto, Ontario, August 8, the only other
specimens known, have been seen by the author.

Types. — Holotype male, allotype female, and female paratypes,
in the U. S. National Museum Collection.

This species shows the same type of color variability due to vary-
ing degrees of maturity, as O. appendiculata, O. volans, species of
Enipoa, and heavily marked species of Typhlocyba.

Ossiannilssonola serrula (Ross and DeLong) (new combination)

(PI. LXXVIII, fig. 3)

Typhlocyba serrula Ross and DeLong, Ohio J. Sci., vol. 49, no. 3, 1949, pp. 117-

118.
Ossiannilssonia serrula, Young, Bull. Univ. Kansas Sci. Bull., vol. 35, pt. 1, 1952,

p. 99.

Resembling Typhlocyba niobe and T. persephone in outward ap-
pearance, but distinguished from these by its larger size and absence
of macrosetae on outer basal angle of male plate; distinguished

1148 The University Science Bulletin

from other species of Ossiannilssonola by lacking brown markings
anterior to cross veins in inner three basal cells, and by having
apices of atrial processes of aedeagus serrate.

Length. — 3.5-3.75 mm.

Color. — Head, pronotum, and scutellum light yellow; fore wings
sulfur yellow, yellow-orange to nearly red-orange in some speci-
mens, subhyaline to shortly before cross veins, without black or
brown spots anterior to cross veins, areas occupied by spots in other
species hyaline; apical cells hyaline; apical veins sulfur yellow;
hind wing with veins yellow; abdomen and plates entirely yellow.

Genital capsule. — Male pygofer, in lateral aspect, with posterior
margin straight, slightly slanting toward base of plate; dorsal angle
rounded, wi*h a short, acute, dorsomesally directed spine, another
similar spine on mesal margin, slightly lower; ventral angle a broad
evenly rounded ventrally projecting lobe, a few slightly submarginal
macrosetae near dorsal posterior margin.

Internal male genitalia. — Aedeagus with atrial processes elongate,
slender, apices almost meeting mesally, in posterior aspect diverging
to outer third, then broadly curved mesad toward each other, finely
serrate at apex, in lateral aspect apical third bent ventrocaudad,
processes joining base of aedeagus at an acute angle; aedeagal apo-
deme short, curving caudad, closely appressed to base of aedeagus.

A large series of specimens has been collected by the author
from Acer saccharum in association with the similarly colored spe-
cies Typhlocyba niobe, T. perscphone, T. athene, and Edwardsiana
lethierryi. Specimens have been seen from Gatlinburg, Tennessee,
June 21, 28; Milwaukee, Wisconsin, June 28-July 5; and the type
series from Waynesburg, Pennsylvania, July 17; and North Bloom-
field, Pennsylvania, July 16.

Types. — Holotype male, allotype female, and female paratypes,
in the DeLong Collection, Columbus, Ohio; one male paratype, in
the Illinois State Natural History Survey Collection.

Ossiannilssonola nicateei sp. nov.
(PL LXXVIII, fig. 4)

Resembling O. danae in external appearance, but with apex of
style acute, not truncate; pygofer with dorsal angle a broad blunt
projection.

Length. — 3.5 mm.

Color. — Head, pronotum, and scutellum pale white to light yel-
low; fore wing with a transverse row of three black spots in apices

Revision of the Species Typhlocyba 1149

of inner three basal cells, apical cells slightly fumose; abdomen
white, male plates white.

Genital capsule. — Male pygofer, in lateral aspect, with posterior
margin nearly transverse, dorsal angle produced into a broad blunt
mesally directed projection, ventral angle somewhat angular, api-
cally rounded, a few macrosetae on dorsocaudal margin.

Internal male genitalia. — Style with acute apex; aedeagus with
atrial processes slender, elongate, sharply bent ventromesad cau-
dally, crossing near apex, in caudal aspect quadrate, in lateral as-
pect evenly curving dorsad to middle then sharply bent ventro-
caudad; aedeagal apodeme short, indistinct.

This species was first recorded under the name Typhlocyba eurtj-
dice by McAtee (1926), from a male specimen from Odenton,
Maryland, July 12, 1914, which was made a paratype of eurydice.
The holotype, allotype, and paratype females of eurydice from
Beltsville, Maryland, June 23, 1916, seen by the author, are speci-
mens of O. danae agreeing with the holotype of that species. On
the basis of specimens compared with the type, Ross and DeLong
(1949) published a description of eurydice which agrees with the
ho'otype. The original descriptions of both species occur on p. 37
(McAtee, 1926), but since the description of O. danae precedes that
of O. eurydice, the latter is here recognized as a synonym of
O. danae (McAtee).

Types. — Holotype male, Odenton, Maryland, July 12, 1914, W. L.
McAtee, (paratype of T. eurydice) in the U. S. National Museum
Collection, Washington, D. C; allotype female, and one pair of
paratypes, Bell Smith Springs, Illinois, July 16, 1947, L. J. Stannard,
from Quercus alba, and two male paratypes, Palos Park, Illinois,
June 22, 1949, Ross and Stannard, from Quercus alba, in the Illinois
State Natural History Survey Collection; one female and two male
paratypes, "Ohio PL," Pennsylvania, July 19, 1919, D. M. DeLong,
and one pair of paratypes, Northeast, Pennsylvania, July 4, 1918,
D. M. DeLong, in the Collection of D. M. DeLong, Columbus, Ohio.
One male, from Berea, Kentucky, June 28, 1941, J. S. Bangson, is
recognizable as this species, but is in too poor condition to be
made a paratype.

Ossiannilssonola danae (McAtee) (new combination)

(PL LXXVIII, fig. 5)

Typhlocyba danae McAtee, Proc. U. S. Nat. Mus., vol. 68, art. 18, 1926, p. 37.
Typhlocyba eurydice McAtee, op. cit., pp. 37-38. (new synonymy)
Ossiannilssonia danae, Young, Bull. Univ. Kansas Sci. Bull., vol. 35, pt. 1,

1952, p. 99.
Ossiannilssonia eurydice, loc. cit.

1150 The University Science Bulletin

Resembling O. mcateei in external appearance, but with apex of
style appearing truncate; pygofer with dorsal angle acute and with
two acute horizontally arranged spines visible in caudal aspect.

Length. — 3.5 mm.

Color. — Head, pronotum, and scutellum milky white; fore wings
milky white, subhyaline nearly to cross veins, with a transverse
row of three indistinct brown spots in apices of inner three basal
cells; abdomen white; male plate white.

Genital capsule. — Male pygofer, in lateral aspect, with posterior
margin obliquely slanted from dorsal angle to base of plate, dorsal
angle produced in an acute mesally-directed spine, a similar spine
arising from the mesal margin, ventral angle a large tongue-shaped
lobe arising from mesal margin near dorsal third of pygofer, a few
macrosetae on posterior margin of dorsal angle.

Internal male genitalia. — Style with a sharp subapical triangular
projection on mesal margin; aedeagus with atrial processes elongate,
slender, sharply bent ventromesad posteriorly, crossing near apex,
strongly divergent on basal half, apical halves bent at right angles
toward each other, directed ventrocaudad; aedeagal apodeme elon-
gate, slender, curving dorsocephalad.

A large series of specimens of this species has been collected by
the author from Quercus alba in Milwaukee, Wisconsin, in associa-
tion with O. berenice and O. australis.

The synonymy of this species is discussed under O. mcateei,
pages 1148-1149.

Specimens have been seen from the following localities: New
York: Conesus Lake, July 16; Maryland: Beltsville, June 23; Vir-
ginia: Glencarlyn, June 12, 16, 20, 23; Wisconsin: Milwaukee, June
30-July 7; Illinois: Palos Park, June 22.

Types. — Holotype male, allotype female, and a pair of paratypes
from Conesus Lake, New York, in the Snow Entomological Collec-
tions of the University of Kansas; nine paratypes from Glencarlyn,
Virginia, in the U. S. National Museum Collection. All type speci-
mens have been seen.

Ossiannilssonola appendiculata (Malloch) (new combination)

(PI. LXXIX, fig. 1)

Typhlocyba appendiculata Malloch, Canadian Ent, vol. 52, no. 4, 1920, p. 95.
Ossiannilssonia appendiculata, Young, Bull. Univ. Kansas Sci. Bull., vol. 35,

pt. 1, 1952, p. 99.
Typhlocyba gillettei var. sellata McAtee, Proc. U. S. Nat. Mus., vol. 68, art. 18,

1926, p. 29. (new synonymy)
Typhlocyba querci var. sellata, Young, Bull. Univ. Kansas Sci. Bull., vol. 35,

pt. 1, 1952, p. 103.

Revision of the Species Typhlocyba 1151

Resembling O. phryne in having male plate bilobed near apex,
but differing by not having black or brown markings on head, pro-
notum, or seutellum; pygofer with posterior margin almost vertical.

Length. — 3.5 mm.

Color. — Head, pronotum, and seutellum light yellow; fore wings
with ground color light yellow, subhyaline to near cross veins;
with variable amounts of brown color forming rather indistinct
patterns, lightly colored specimens with brown markings in outer
fourth of inner three basal cells forming almost a solid band, with
a dash of brown along basal half of claval suture, color irregular
in intensity, progressively increasing in area, darkly marked speci-
mens with most of the wing anterior to cross veins brown, a yellow
semicircle along basal third of costal vein and another along middle
of costal vein extending inward to middle of third basal cell, apices
of third and fourth basal cells yellow; apical cells hyaline, slightly
fumose.

Genital capsule. — Male pygofer, in lateral aspect, with posterior
margin almost vertical, slightly projecting on dorsal angle, smoothly
rounded, ventral margin forming nearly a right angle with posterior
margin, a number of macrosetae submarginally arranged near dorsal
posterior margin; plates with a short laterally directed lobe at outer
third (PI. LXXIX, fig. If).

Internal male genitalia. — Style with apex more elongate than
usual (PL LXXIX, fig. le); adeagus with atrial processes elongate,
slender, sinuate, gradually tapering from base to acute apices which
are sharply directed mesad on outer third, crossing at apex medially,
in caudal aspect nearly quadrate, in lateral aspect slightly sinuate;
aedeagal apodeme slender, elongate, slightly enlarged apically,
directed cephalad.

A large series of specimens has been taken from Quercus macro-
carpa in Milwaukee, Wisconsin, and several other series from the
same host taken by H. H. Ross and L. J. Stannard in Illinois, are at
hand.

Specimens have been seen from the following localities: New
York: Batavia, July 4, August 12; Norris, August 15; New York City
Botanic Garden, July 7; Pennsylvania: Hartstown Bog, August 13;
Washington D. C. June 18, 29; Tennessee: Great Smoky Mountain
National Park, June 12; Ohio: Barberton, August 13; Illinois: Cook
County — Palos Park, June 22; Urbana, July 9, 14; Tinley Park, Sep-
tember 8; Thornton, September 7; Iowa: Ames, June 19, September
11; Kansas: Manhattan, June 8; Wisconsin: Madison, July 19; Mil-

43—6490

1152 The University Science Bulletin

waukee, June 26-July 4; Minnesota: Ramsey County, July 20, 26;
August 3, 25.

Both the holotype female and female paratype of McAtee's
Typhlocyba gillettei var. sellata have been seen, and are identical
in color marking with heavily pigmented specimens of this species.

Types. — Holotype male, allotype female, and one paratype male
in the Illinois State Natural History Survey Collection. Types
have been seen by the author.

Ossiannilssonola phryne (McAtee) (new combination)

(PI. LXXIX, fig. 2)

Typhlocyba phryne McAtee, Proc. U. S. Nat. Mus., vol. 68, art. 18, 1926,

pp. 34-35.
Typhlocyba phryne var. subpulchra McAtee, loc. cit.
Ossiannilssonia phryne, Young, Bull. Univ. Kansas Sci. Bull., vol. 35, pt. 1,

1952, p. 99.

Resembling O. appendiculata in the shape of male plate, but
differing by having a black line along commissural margin and
parenthesis-shaped black lines on scutellum, pronotum, and some-
times extending onto head; pygofer strongly produced caudally on
dorsal half as a broad apically rounded lobe.

Length. — 3.5 mm.

Color. — Head, in very darkly marked specimens, with two choco-
late brown longitudinal bars on either side of midline from anterior
margin of eye to pronotum, ecdysial line dark, remaining portions
yellow, entirely yellow in most specimens; pronotum, with two
parenthesis-shaped chocolate brown longitudinal bands continuing
posteriorly from basal margin of head broadening caudally and
nearly meeting medially at base, with remaining portions yellow,
almost entirely yellow in lightly marked specimens with only a
faint trace of chocolate brown color; scutellum, with lateral angles
chocolate brown, yellow medially; fore wings, with chocolate brown
stripes along commissural margin to cross veins covering inner half
of clavus, chocolate brown spots in apices of inner three basal cells
forming a transverse band; size of spots progressively smaller to-
wards costal margin, lighter areas anterior to cross veins subhyaline,
yellowish-white; apical cells hyaline, fumose; abdomen, chocolate
brown on basal half of median third of dorsal segments, laterally
deep orange yellow, dorsal third of pygofer chocolate brown, venter
yellow, plates yellow.

Genital capsule. — Male pygofer, in lateral aspect, with posterior
margin greatly produced caudally on dorsal half, ventral angle
forming a right angle, with one or usually two macrosetae just

Revision of the Species Typhlocyba 1153

dorsad of outer basal angle of plate, a row of macrosetae along
posterior margin on dorsal half; male plate elongate, dorsoventrally
compressed near middle, forked shortly before apex into two thumb-
like processes, one directed dorsad, the other laterad, setae mostly
restricted to outer fourth. ( PI. LXXIX, fig. 2a ) .

Internal male genitalia. — Aedeagus with atrial processes elongate,
slender, apices directed mesocaudad, separated at apex, in caudal
aspect nearly forming a circle; aedeagal apodeme reduced to a
broad triangular anteriorly directed process.

A large series of specimens of this species collected by H. H.
Ross and L. J. Stannard, from Quercus macrocarpa, in Illinois, has
been seen by the author.

Specimens have been examined from the following localities:
New York: Norris, August 15; Ontario: Ottawa, June 30, Septem-
ber 27; Illinois: Western Springs, June 21; Tinley Park, September
8; Wisconsin: Milwaukee, June 27; Minnesota: St. Paul, June 22-23,
August 12, 18, 25.

Types. — Holotype female, in the Illinois State Natural History
Survey Collection; another female holotype of var. subpulchra, in
the Iowa State College Collection; a male specimen collected by the
author, June 27, 1950, Milwaukee, Wisconsin, here designated neo-
allotype, two paraallotijpe males, June 21, 1949, Western Springs,
Illinois, Stannard and Ross, and three paraallotijpe males, June 16,
1949, Tinley Park, Illinois, Ross and Stannard, in the Snow Ento-
mological Collections of the University of Kansas; additional para-
allotijpe males: five, June 21, 1949, Western Springs, Illinois, Stan-
nard and Ross; eight, September 8, 1949, Tinley Park, Illinois, Ross
and Stannard, in the Illinois State Natural History Survey Collection;
one, August 15, 1942, Norris, New York, R. E. Olson, in Cornell
University Collection; one, August 12, 1938, St. Paul, Minnesota,
A. A. Granovsky, in the University of Minnesota Collection; one,
September 8, 1904, Ottawa, Ontario, W. Metcalfe, in the Colorado
Agricultural and Mechanical College Collection.

Ossiannilssonola rossi sp. nov.
(PI. LXXIX, fig. 3)

Resembling O. quadrata and O. knulli in external appearance
and in general form of pygofer and aedeagus, but distinguished
from these species by having the dorsal angle of pygofer greatly
produced into a large hook, and in having the apices of the atrial
processes appearing forked in caudal aspect.

Length. — 3.5 mm.

1154 The University Science Bulletin

Color. — Head, pronotum, and scutellum light yellow; fore wings,
light yellow, subhyaline to near cross veins, apex hyaline and
slightly fumose, with a transverse row of three clearly defined black
spots in apices of inner three basal cells; abdomen black on dorsum,
venter light.

Genital capsule. — Male py gofer, in lateral aspect, with posterior
margin distinctly S-shaped on lower half, dorsal angle produced
posteriorly in a broad ventrally hooked lobe, curving mesad, and
appearing as a broad hook in caudal aspect as well, ventral angle
rounded, reduced, a dense brush of macrosetae on dorsal lobe.

Internal male genitalia. — Aedeagus with atrial processes elongate,
slender, apices distant from each other the width of base, in caudal
aspect subquadrate, slightly diverging toward apex, sharply bent
mesad on apical fourth, with apices flattened and twisted dorso-
caudad, appearing forked; base of aedeagus roughly S-shaped from
left side; aedeagal apodeme reduced to a slight projection on an-
terior margin.

Type.— Holotype male, Thornton, Illinois, June 22, 1949, H. H.
Ross and L. J. Stannard, on Quercus bicolor, in the Illinois State
Natural History Survey Collection.

Ossiannilssonola knulli sp. nov.
(PI. LXXIX, fig. 4)

Resembling O. quadrata in external appearance, but with dorsal
angle of pygofer less prominent, and with base of aedeagus not
broadly inflated below bases of atrial processes.

Length. — 3.0-3.25 mm.

Color. — Head, pronotum, and scutellum bright sulfur-yellow; fore
wings sulfur-yellow, subhyaline to cross veins, apex hyaline, slightly
fumose, with a transverse row of three black spots in apices of inner
three basal cells, sometimes one or two missing; abdomen yellow,
male plates yellow.

Genital capsule. — Male pygofer, in lateral aspect, with posterior
margins somewhat S-shaped on lower half, dorsal angle directed
caudally as a broad ventrally hooked lobe one fourth as long as
broad, hook more prominent from caudal aspect, ventral angle
broadly rounded, a group of macrosetae on dorsal posterior margin.

Internal male genitalia. — Style abruptly reduced to an acute apex
(PI. LXXIX, fig. 4), with atrial processes elongate, slender, slightly
curved ventromesad caudally, crossing near apex; base of aedeagus
roughly S-shaped in lateral aspect; aedeagal apodeme reduced to a

Revision of the Species Typhlocyba 1155

slight projection on anterior margin; base of aedeagus slender in
caudal aspect, not enlarged as in O. quadrata.

A large series of both male and female specimens has been col-
lected by the author, from a single tree of Quercus borealis in Law-
rence, Kansas.

Types. — Holotype male and numerous paratypes of both sexes,
Douglas County, Kansas, June 8, 1951; allotype female, two male
and twenty-one female paratypes, Douglas County, Kansas, June
10, 1950; ten female paratypes Douglas County, Kansas, June 12,
1950, P. J. Christian, in the Snow Entomological Collections of the
University of Kansas; one male paratype collected by Nathan Banks,
North Mountain, Pennsylvania, July 4, in the Museum of Compara-
tive Zoology; a pair of paratypes collected by E. D. Ball, Woods
Hole, Massachusetts, July 7, 1925, in the U. S. National Museum
Collection; one female, June 12, and pair of paratypes June 20, 1951,
Dale Bray, Wilmington, Delaware, in the University of Delaware
Agricultural Experiment Station Collection.

Ossiannilssonola quadrata (DeLong and Johnson)
(new combination)

(PI. LXXX, fig. 3)

Typhlocyba quadrata DeLong and Johnson, Ent. News, vol. 47, no. 4, April,

1936, pp. 102-104.
Ossiannilssonia quadrata, Young, Bull. Univ. Kansas Sei. Bull., vol. 35, pt. 1,

1952, p. 99.

Resembling O. knulli in outward appearance, but with dorsal
angle of pygofer more prominent, hook terminating in strong, acute,
ventrally-directed spine, base of aedeagus laterally inflated below
bases of atrial processes.

Length. — 3.5 mm.

Color. — Head, pronotum, and scutellum pale yellow; fore wings
yellow, subhyaline to cross veins, with a transverse row of three
black spots in apices of inner three basal cells; apex fumose, hyaline.

Genital capsule. — Male pygofer, in lateral aspect, with lower
half of posterior margin less distinctly S-shaped than in O. rossi,
dorsal angle produced caudally into a short broad process curving
ventrad as a sharp elongate spine and with another shorter spine
directed mesad near dorsal margin, a few submarginal macrosetae
on dorsal angle, ventral angle broadly rounded but less prominent
than in preceding species.

Internal male genitalia. — Style with apex less abruptly reduced
than in O. knulli (Pi. LXXIX, fig. 4e); aedeagus with atrial processes
elongate, slender, directed ventromesad caudally; aedeagal apo-

1156 The University Science Bulletin

deme absent; base of aedeagus greatly inflated laterally, sharply
constricted before bases of atrial processes, with a sclerotized plate
extending over bases of processes into membrane.

Type. — This species is known from one male specimen collected
at Kane, Pennsylvania, August 19, 1928, by D. M. DeLong, and is in
the collection of the collector. Illustrations for this species have
been drawn from the type specimen.

Ossiannilssonola troza (Ross and DeLong) (new combination)

(PL LXXX, fig. 1)

Tt/phlocijba troza Ross and DeLong, Ohio J, Sci., vol. 49, no. 3, 1949, pp. 116-

' 118.
Ossiannilssonia troza, Young, Bull. Univ. Kansas Sci. Bull., vol. 35, pt. 1, 1952,
p. 99.

Resembling O. bangsoni in external appearance, but with dorsal
angle of pygofer produced as a broad, acute, dorsomesally-directed
arm.

Length. — 3.5-3.75 mm.

Color. — Head, pronotum, and scutellum white to light yellow;
fore wings, light yellow to yellow, subhyaline to cross veins, with
a transverse row of three black spots in apices of inner three basal
cells, apex hyaline, slightly fumose; abdomen yellow, male plates
yellow.

Genital capsule. — Male pygofer, in lateral aspect with posterior
margin nearly transverse, dorsal angle produced into a large broad
dorsomesally directed hook, ventral angle slightly produced caudally
in a short hook, a row of macrosetae on posterior margin of dorsal
hook.

Internal male genitalia. — Aedeagus with atrial processes elongate,
slender, apices separated, in caudal aspect nearly quadrate in out-
line, widely separated near basal third, slightly converging to outer
third which is abruptly curved mesodorsad caudally; aedeagal
apodeme short, enlarged apically, directed dorsad.

In addition to the holotype, a short series collected by R. H.
Beamer and the author, from Quercus muhlenbergii, and one male
taken on Quercus pahistrus by the author, in Douglas County, Kan-
sas, June 11-24, have been seen.

Types. — Holotype male, in the Illinois State Natural History Sur-
vey Collection; a female specimen collected by R. H. Beamer, Doug-
las County, Kansas, June 11, 1949, taken with males of this species,
here designated neoallotype, in the Snow Entomological Collections
of the University of Kansas.

Revision of the Species Typhlocyba 1157

Ossiannihsonola flavomarginata (Gillette and Baker)
(new combination)

(PI. LXXX, fig. 2)

Typhlocyba flavomarginata Gillette and Baker, Bull. Colorado Agr. Exp. Sta.,

no. 31, Tech. ser. no. 1, 1895, pp. 111-112.
Typhlocyba flavomarginata var. vesta McAtee, Proc. U. S. Nat. Mus., vol. 68,

art. 18, 1926, p. 18.
Typhlocyba flavomarginata var. scoria McAtee, op. cit., pp. 18-19.
Typhlocyba flavomarginata var. media McAtee, op. cit., p. 19.
Ossiannussonia flavomarginata, Young, Bull. Univ. Kansas Sci. Bull., vol. 35,

pt. 1, 1952, p. 99.

Specimens of Form I which have a red longitudinal stripe on
clavus along commissural margin, resemble in external appearance
deeply colored species of O. australis, but are easily distinguished
from this species and other species in the genus by having styles
with apical third sharply curved mesad, and a triangular projection
on lateroventral margin.

Length. — 3.25-4.0 mm.

Color. — The color of this species is highly variable resulting in
five recognizable forms, but with numerous intermediates of all
degrees between these. Perhaps a study of the genetic constitution
of this species will be necessary before the diversity of color pattern
will be understood.

Color descriptions of the five most distinct patterns, designated
by Roman numerals I to V, are as follows:

Form I. — Head light yellow; pronotum yellow with a red triangle
with apex on disc and base on posterior margin; scutellum with
outer angles red, forming an inverted V with the triangle on pro-
notum; fore wings with red commissural band extending from base
to cross veins on inner half of clavus, with a dark spot in each of
inner three basal cells appearing as a band obliquely slanted cau-
dally toward costal margin, remaining portions of wing anterior to
cross veins subhyaline, yellow, apex hyaline, fumose; abdomen with
dark brown median band on dorsum broader on basal segments,
sides yellow, venter and male plates yellow.

Form II. — Head, yellowish-white; pronotum yellow, light brown
from disc to posterior margin; scutellum light brown; fore wings
light to dark brown, hyaline, without spots; abdomen with median
band on dorsum dark brown, sides bright yellow, venter and male
plates yellow.

Form III. — Head, pronotum, and scutellum bright yellow; fore
wings pale red throughout, subhyaline to cross veins, hyaline be-
yond; abdomen and male plates yellow.

1158 The University Science Bulletin

Form IV. — Head yellow-orange; pronotum yellow-orange, disc
orange; scutellum yellow-orange; fore wings deep orange through-
out, subhyaline to cross veins, hyaline beyond; abdomen and male
plates yellow.

Form V. — Head, pronotum, scutellum, and all of fore wings white
to ivory, without dark markings.

All degrees of color intergradation can be found between forms
I and II, I and V, and II and V. Some intergrades occur between
III and IV, and there is a gradation through teneral specimens of
III and IV, toward V. The holotype of McAtee's var. media is like
form I, the holotype of var. scoria is like form III, and the holotype
of var. vesta is like form V. The lectotype of var. flavomarginata
is intermediate between forms II and V, but nearer form II.

Genital capsule. — Male pygofer, in lateral aspect, with posterior
margin nearly vertical, ventral margin extended dorsad obliquely
nearly to dorsal angle; dorsal angle broadly rounded, ventral pos-
terior margin slightly recurved, with a slight, papillose, ventral lobe,
a number of macrosetae on dorsal posterior margin; male plate in
ventral aspect, wedge-shaped, lateral and mesal margins nearly
straight, in lateral aspect slightly curved dorsad, with a broad dor-
solateral excavation on outer three fourths.

Internal male genitalia. — Styles with outer third of apical portion
abruptly curving mesad, a small triangular hooklike projection on
lateroventral margin ( PI. LXXX, fig. 2e ) ; aedeagus with atrial proc-
esses broadly flattened laterally on basal two thirds, reduced on
apical third to slender mesoventrally curving acute apices, directed
caudally; aedeagal apodeme in lateral aspect broad, elongate, api-
cally enlarged, directed cephalad.

Female. — With posterior margin of eighth abdominal sternite
strongly incised laterally forming two lateral lobes and one slightly
emarginate median lobe (Pi. LXXXVII, figs. 7a, b).

Two large series collected by R. H. Beamer in Sapello, New
Mexico, and in Raton, New Mexico, from Quercus sp., have been
seen by the author, and short series have been seen from the follow-
ing localities: Arizona: Grand Canyon, August 1, 3, 11; Flagstaff,
August 1, 5; Oak Creek Canyon, July 9, 31, August 9, 14; Coconino
County, August 13; Huachuca Mountains, July 8, August 1, 2;
Cochise County, July 29; New Mexico: Ruidoso, August 10, Octo-
ber 10, 15; Pecos, Sapello, July 24; Raton, July 26; Jamez Springs,
July 22; Taos Pass, August 13; Cimarron, August 13; Colfax County,
August 21; Cowles, July 18; Luna, July 25; Soroco County, August

Revision of the Species Typhlocyba 1159

18; Colorado: LaVeta Pass, July 28; Colorado Springs, July 19;
Palmer Lake, October 9; Trinidad, July 13, August 19; Manitou,
September 29; Cimarron, August 22; Cerro Summit, August 21;
Durango, August 7; Salida, October 8; Alder, August 25; Rabbit
Ear Pass, August 18; Royal Gorge, July 3; Glenwood Springs,
August 17; Garden of the Gods; Colorado City; Utah: Freedom,
August 24, Fruitland, July 16; Keetley, August 16; Park City, August
16; Grant, June 25, July 4, 14, 15, 22, 24, August 6; Farmington,
June 12; Panguitch, September 4; Alton, July 30, August 14; Salt
Lake City, July 3, 13, August 29; Emery, August 16; Cove Fort,
August 14; Cedar City, August 13; Pintura, August 11; Salina,
August 13, 26; Fish Lake, September 2; Richfield, July 14; Mount
Carmel Highway, June 18.

Types. — Specimens seen bearing machine printed cotype labels
are as follows: two specimens, "Colo 2280" in the Snow Ento-
mological Collections of the University of Kansas; four specimens,
"Colo 2280" and one specimen, "Colo 2266" in the Colorado Agri-
cultural and Mechanical College Collection; and two specimens,
"Colo 2266" and "Colo 2270" in the U. S. National Museum Collec-
tion. Two female specimens each bearing a handwritten label
"type" on white paper, have also been seen, one "Colo 1375" in the
U. S. National Museum Collection, and the other "Colo 1780" in the
Colorado Agricultural and Mechanical College Collection. These
numbers refer to the collecting numbers of either Baker or Gillette
and have been checked on the lists of each of these men, for the data
which the numbers represent.

The data for number 1780 on Gillette's list which fits that given
for the three females in the original description of T. flavomarginata
G. & B., is "9-29-'94, at Manitou, Colo. C.P.G., Quercus undulosa".
The data for each of the other numbers seen indicates that the
specimens bearing these can not be more than metatypes.

In 1926 McAtee selected a lectotype from three females which he
thought were the type series of flavomarginata, and designated a
fourth specimen, a male with a cotype label on it, as neoallotype,
recognizing that it was not a cotype since only females were in the
type series. These specimens were given the U.S.N.M. catalogue
number 3456, and placed in the U. S. National Museum Collection.
Only three of these four specimens are now in this collection, and
of these, two, "Colo 2270" a female, and "Colo 2266" a male, bear
cotype label U.S.N.M. 3456. The male is unquestionably the neo-
allotype set up by McAtee while the other specimen is only a

1160 The University Science Bulletin

metatype. The third specimen, "Colo 1375" which on Baker's list
is "May, Ft. Collins", may have been the specimen selected by Mc-
Atee as lectotype, since it does not bear a U.S.N.M. cotype label as
the others do, although it does not have a lectotype label either.
There is also the possibility that the fourth specimen seen by McAtee
may have been the one selected as lectotype, but this specimen is
now missing.

Since the only remaining specimen which could have been
selected as lectotype, "Colo 1375", is not one of the type series,
and since only one specimen of the type series is now known to exist,
the author proposes that this specimen, "Colo 1780" in the Colorado
Agricultural and Mechanical College Collection, be made lectotype
of Typhlocyba flavomarginata Gillette and Baker.

Genus Typhlocyba Germar
(Pis. LXXXI-LXXXVII )

Typhlocyba Germar, Rev. Ent. Silbermann, vol. 1, 1833, p. 180.

Anomia Fieber, Verh. Zool.-bot. Ges. Wien, vol. 16, 1866, p. 509. (type,
Cicada quercus Fabricius, 1794, by subsequent designation of Evans, 1947,
Trans. Roy. Ent. Soc. Lond., vol. 98, p. 200). Type of the genus, Cicada
quercus Fabricius, by subsequent designation of Woodworth, 1889.

Fore icings. — Inner and outer apical cells short, not attaining
wing apex; second apical cell much broader at apex than at base;
third apical cell petiolate; wing apex smoothly rounded. (PL
LXXXI,fig. If).

Hind wings. — Vein IV branching from 2V near its midlength;
submarginal vein absent at wing apex; both apical cells open api-
cally; posterior branch of B fused with apical portion of vein M1+2
(PL LXXXI, fig. If).

Genital capsule. — Male plate gradually curved dorsad apically,
reduced near middle but enlarged apically, with one or rarely two
macrosetae near outer basal angle, submarginal row of microsetae
near outer basal angle, submarginal row of microsetae parallel to
lateral margin near middle and extending over apical half of length,
a few other irregularly arranged microsetae, apex usually black;
pygofer, in lateral aspect, of various forms with or without sclero-
tized barlike thickenings which when present may or may not be
extended as caudal or dorsocaudal pygofe/ processes, with ventro-
caudal margin occasionally inrolled, often with group of macrosetae
just dorsad of outer basal angle of male plate, almost always with
group of small submarginal setae along dorsocaudal margin and
with numerous microsetae arranged over disc; apex of ventral lobe
or ventral hook usually black.

Revision of the Species Typhlocyba 1161

Internal male genitalia. — Style elongate, slender, usually either
smoothly curved laterad or dorsad apically, with mesal preapical
protruberance poorly developed or absent in broad aspect (which
may be dorsal or caudal); style with setae in various arrangements;
connective massive, the aedeagal articulation subterminal; aedeagus
with preatrial arm well developed or not, apodeme usually well
developed; aedeagal shaft with paired ventral processes arising
from atrium or base of shaft, sometimes fused to shaft, with one or
two pairs of apical processes and occasionally with processes along
length of shaft.

Head in dorsal aspect narrower than pronotum, longer medially
than next the eye, anterior margin of crown broadly rounded; pro-
notum short and broad, lateral margins strongly divergent caudally,
posterior margin smoothly, shallowly convex, pleural portion
broader than ocellocular area.

The species, for the most part, are slender and delicate in appear-
ance, usually pale white or yellow, occasionally with darker mark-
ings.

The genus is holarctic in distribution.

The following key to the North American species of the genus is
based primarily on characteristics of the male genitalia.

Key to the Species of Typhlocyba

1. Aedeagus with atrial processes arising from base of shaft 2

Aedeagus without atrial processes, or with processes fused to

shaft 20

2. Atrial processes fused with aedeagal shaft on basal fourth 3

Atrial processes arising from near, or at the base of aedeagal

shaft 4

3. Aedeagal shaft with elongate apical processes (PI. LXXXI, fig.

3b, c) athene p. 1166

Aedeagal shaft simple, without apical processes (PI. LXXXV,

figs. 2b, c) Cassiopeia p. 1181

4. Aedeagal shaft with processes 5

Aedeagal shaft without processes 7

5. Aedeagal shaft with two pairs of processes (PI. LXXXIV, fig.

3b, c ) shawneeana p. 1178

Aedeagal shaft with one pair of processes 6

6. Aedeagal shaft processes apical, broad at base, short and stout,

slightly curving laterad. (Pi. LXXXI, fig. 2b, c) oneka p. 1165

Aedeagal shaft processes slightly subapical, elongate, slender,

curving strongly laterodorsad. (Pi. LXXXI, fig. 4b, c) . arsinoe p. 1166

7. Aedeagal shaft flattened laterally and apically into a broad thin

semitransparent plate 8

Aedeagal shaft not flattened laterally and apically 12

1162 The University Science Bulletin

8. Posterior margin of pygofer projecting ventrad as a long sharp

hook; face of males usually orange-red. (Pi. LXXXII, fig. 4),

pomaria p. 1170
Posterior margin of pygofer not projecting ventrad as a long
sharp hook; face never orange-red 9

9. Dorsal angle of pygofer forming a prominent caudally directed

process 10

Dorsal angle of pygofer not forming a prominent caudally di-
rected process 11

10. Dorsal process of pygofer narrow, acute, heavily sclerotized. ( Pi.

LXXXIII, fig. 3) surcula p. 1175

Dorsal process of pygofer a broad continuation of pygofer, not

heavily sclerotized. (Pi. LXXXIII, fig. 1) attenuata p. 1172

11. Posterior half of pygofer nearly quadrate; fore wing white, with-

out markings. (Pi. LXXXIII, fig. 4) andromache p. 1176

Posterior half of pygofer with margin strongly rounded; fore wing
with a broad brown band over cross veins, often with a red
spot in middle. (Pi. LXXXIII, fig. 2) rubriocellata p. 1174

12. Apex of male plate produced laterally as a short, heavily sclero-

tized, beaklike process (Pi. LXXXI, fig. 1); fore wings with
five red to orange-brown spots, and with brown markings in

cells bordering cross veins quercus p. 1163

Apex of male plate smoothly rounded; fore wings without red
or orange markings 13

13. Atrial processes of aedeagus broadly flattened. (Pi. LXXXIV,

fig. 1 ) melite p. 1176

Atrial processes of aedeagus not broadly flattened 14

14. Aedeagal shaft arising greatly elevated above bases of atrial

processes 15

Aedeagal shaft arising between or slightly above bases of atrial
processes 16

15. Atrial processes of aedeagus fused on basal fourth, or basal half;

style with thumblike projection on lateral margin near middle;
pygofer with a short, acute, dorsally directed hook. (PI.

LXXXII, fig. 1 ) modesta p. 1167

Atrial processes of aedeagus not fused on basal fourth, style with-
out thumblike projection near middle; pygofer with a long,
sharp, ventrally directed process on ventral angle. (Pi.
LXXXII, fig. 3) hockingensis p. 1169

16. Fore wings with a transverse brown or black band 17

Fore wings without dark transverse band 18

17. Fore wings with band distad of cross veins; shaft of aedeagus

with posterior margin concave; dorsal angle of pygofer forming
a broad, angular, laterally-concave process. (Pi. LXXXIV,

fig. 2) alahamaensis p. 1177

Fore wings with band covering middle third; shaft of aedeagus
convex on posterior margin; dorsal and posterior margins of
pygofer nearly continuous, not produced. (Pi. LXXXIV,
fig. 4 ) transviridis p. 1179

Revision of the Species Typhlocyba 1163

18. Aedeagus with atrial processes not exceeding shaft in length;

pygofer with a small ventrally directed lobe near middle of

posterior margin. (Pi. LXXXII, fig. 2) medleri p. 1169

Aedeagus with atrial processes exceeding shaft in length by one
third their length 19

19. Aedeagal shaft anterior to atrial processes throughout its length;

posterior margin of pygofer nearly straight, without short ven-
tral tooth. (PI. LXXXV, fig. 3) crassa p. 1182

Aedeagal shaft posterior to or even with atrial processes; pygofer
with posterior margin angled near middle, extended ventrad
as a short tooth. (Pi. LXXXV, fig. 1) putmani p. 1180

20. Aedeagus with two pairs of shaft processes, one pair arising near

middle of shaft. (Pi. LXXXI, fig. 3) athene p. 1166

Aedeagus with only one pair of shaft processes 21

21. Aedeagal shaft processes arising at or before middle of shaft, shaft

greatly reduced beyond processes 22

Aedeagal shaft processes apical 24

22. Aedeagal shaft processes arising before middle of shaft, pygofer

with ventral angle evenly rounded, not produced. (Pi.

LXXXV, fig. 2) Cassiopeia p. 1181

Aedeagal shaft processes arising at middle of shaft; pygofer with
ventral posterior angle produced laterad or ventrad as a short
process 23

23. Aedeagal shaft processes closely appressed to apical portion of

shaft, basal half of shaft stout, broad in lateral aspect. (PL

LXXXVI, fig. 4) niobe p. 1183

Aedeagal shaft processes distinctly separated from shaft, from
base; basal half of shaft slender, slightly wider than processes
in lateral aspect. (Pi. LXXXVI, fig. 1) sollisa p. 1182

24. Apical processes of aedeagus broadened in middle, half as long

as shaft; pygofer without a lobe at dorsal angle. (Pi.

LXXXVI, fig. 3 ) persephone p. 1184

Apical processes of aedeagus short, less than one fourth length of
shaft, appearing as a continuation of shaft 25

25. Apical processes of aedeagus nearly straight, shaft broadly in-

flated on distal half in caudal aspect; pygofer without lobe on

dorsal angle. (Pi. LXXXVII, fig. 1 ) infata p. 1186

Apical processes of aedeagus sinuate, shaft of uniform width
throughout in caudal aspect; pygofer with a lobe on dorsal
angle. (Pi. LXXXVI, fig. 2) tortosa p. 1185

Typhlocyba quercus (Fabricius)

(PI. LXXXI, fig. 1)

Cicada quercus Fabricius, Genera Insectorum, Rhyngota, 1777, p. 298.
Tettigonia quercus, Germar, Magazin der Entomologie, vol. 4, 1821, p. 73.
Typhlocyba quercus, Herrich-Schaffer, Deutschlands Insecten, vol. 124, 1834,

pp. 1-15.
Typhlocyba fasciata Tollen, Stett, Ent. Zeit, vol. 12, 1851, p. 73.
Anomia quercus, Fieber, Katalogue der Europaischen Cicadinen, 1872, p. 15.

This species is easily distinguished from other species in the

1164 The University Science Bulletin

genus by having the apex of the plate laterally produced as an
acute beaklike process, and by having fore wing with five orange-
red spots forming a rough W.

Length. — 3.0-3.5 mm.

Color. — Head, with dorsum bearing an orange-red, inverted V-
shaped mark extending from margins of eyes to near middle of disc;
pronotum, with an orange-red band along anterior margin, a large
round orange spot on disc, remaining portions milky-white to light
yellow; scutellum, with lateral angles orange-red, median band
white to yellow; fore wing, with ground color white, with three
evenly spaced red to orange-red spots on clavus along commissural
margin, and two spots of the same color on inner two basal cells
forming a rough W, connected to costal margin by two oblique
brown lines; apical and cross veins white, apical and basal cells with
light brown areas bordering cross veins; abdomen, dorsum black
with outer margins of segments yellow, venter yellow, basal seg-
ments darker on basal half; pygofer, dorsum black, venter yellow.

Genital capsule. — Male pygofer, in lateral aspect, with posterior
margin continuous with ventral margin, evenly rounded to dorsal
angle which is produced dorsad as a heavily-sclerotized, darkly-
pigmented, sharply pointed hook, without macrosetae dorsad of
outer basal angle of plate; plate, in ventral aspect, gradually reduced
toward apex, produced laterad as a short, heavily sclerotized, beak-
like process, in lateral aspect slightly curving dorsad, with one or
two macrosetae near outer basal angle, a number of scattered
microsetae in a submarginal row along lateral margin, a small group
of macrosetae at apex. (PI. LXXXI, fig. la).

Internal male genitalia. — Style with apical portion reduced, nearly
equal in length to basal apodeme, a row of setae on outer margin
and several alveoli on inner margin near apex; connective, almost
quadrate; aedeagus, with atrial processes elongate, slender, grad-
ually curving laterocephalad, length nearly equal to that of shaft;
aedeagal shaft without apical processes, width nearly uniform from
base to apex, slightly curving dorsad, anterior margin recurved
at apex, with lightly sclerotized areas on sides near apex; aedeagal
apodeme as wide as, but less than half as long as shaft in lateral
aspect.

Female. — With posterior margin of eighth abdominal sternite
nearly transverse, slightly sinuate, lateral fourth sharply directed
cephalad (PI. LXXXVII, figs. 8a, b).

A series of twenty-seven specimens collected from cultivated

Revision of the Species Typhlocyba 1165

cherry, in Vancouver, British Columbia, by Dr. H. H. Ross, July 15,
1948, and August 4, 1950, are the only North American specimens
seen by the author. European specimens seen are: a pair from
Poland, determined by J. Nast, 1937; one male from Sweden, labeled
"Primus", determined by F. Ossiannilsson, 1948; and three speci-
mens in the U. S. National Museum Collection, bearing only the
number 191.

Typhlocyba oneka Knull

(PI. LXXXI, fig. 2)
Typhlocyba oneka Knull, Ohio J. Sci., vol. 44, no. 6, 1944, p. 270.

Resembling T. tortosa and T. inflata in having an apical pair of
shaft processes, but differing in having a pair of slender atrial
processes.

Length. — 3.5 mm.

Color. — "Pale yellowish white, elytra semihyaline, below cream-
colored, eyes pale", [from original description].

Genital capsule. — Male py gofer, in lateral aspect, with posterior
margin slightly convex, dorsal angle produced dorsad as a broad
apically rounded projection, ventral angle slightly produced ventro-
caudad as a small lobe, a few macrosetae dorsad of outer basal
angle of male plate, a large group of macrosetae along middle of
posterior margin extending inward toward disc, posterior margin
deeply inrolled; plate with apex spatulate.

Internal male genitalia. — Style elongate, slender, gradually taper-
ing to acute apex, curving ventrolaterad, with a large patch of setae
of ventral surface near middle extending from mesal to lateral mar-
gins, several alveoli on mesal margin near middle; connective, Y-
shaped, with posterior margin produced medially; aedeagus, with
atrial processes setaeform, two thirds length of shaft, slightly di-
verging apically; aedeagal shaft elongate, slender, with apical proc-
esses broadly attached at base, sharply reduced to acute laterally
directed apices; aedeagal apodeme as broad as base of aedeagal
shaft, strongly curving dorsocephalad.

Only the dissected abdomen of one male specimen has been seen
by the author. The reported host for this species is Corylus. The
known distribution for this species: Minnesota: Itasca County, July
27; New York: Cranberry Lake, July 15.

Types. — Holotype male, allotype, and paratypes, in the Herbert
Osborn Collection of Ohio State University.

1166 The University Science Bulletin

Typhlocyba athene McAtee

(PI. LXXXI, fig. 3)

Typhlocyba athene McAtee, Proc. U. S. Nat. Mus., vol. 68, art. 18, 1926, p. 31.

Resembling Typhlocyba niobe and T. persephone externally, but
differing in having the dorsal angle of pygofer produced as a slender
acute hook, and by having a pair of atrial processes which are fused
to shaft to middle, and a pair of apical processes.

Length. — 3.0-3.5 mm.

Color. — Head, pronotum, and scutellum pale to deep yellow; fore
wings deep yellow; abdomen yellow.

Genital capsule. — Male pygofer, in lateral aspect, with posterior
margin slightly convex, dorsal angle produced as a sharp, elongate,
ventrocaudally directed hook, ventral angle produced as a short
acute hook directed ventrad, one or a few macrosetae dorsad of
outer basal angle of male plate, a large group of macrosetae dorsad
of outer basal angle of male plate, a large group of macrosetae on
middle of posterior margin extending inward toward disc; male
plate with apex spatulate.

Internal male genitalia. — Styles and connective like those of
Typhlocyba oneka (PI. LXXXI, fig. 2d, e); aedeagus, with atrial
processes fused to shaft to near middle, gradually tapering to acute,
dorsolaterally directed apices; aedeagal shaft broad at base, slightly
reduced toward apex, with a pair of laterally diverging apical proc-
esses; aedeagal apodeme broad as shaft, nearly vertical, with a short
anteriorly directed arm at apex.

A series of twenty-eight male specimens has been collected from
Acer saccharum, by the author, in Milwaukee, Wisconsin, taken in
association with T. niobe, T. persephone, Ossiannilssonola serrula,
and Edwardsiana lethierryi. It has not been possible to separate
the female specimens from those of some of these species.

Specimens have been seen from the following localities: New
Hampshire: Lee, July 7; Connecticut: New Haven, October 16;
Pennsylvania: Northeast; Kane; Illinois: Urbana, June 7; Wisconsin:
Milwaukee, June 27, 28, 30, July 1.

Types. — Holotype male, in the Illinois State Natural History Sur-
vey Collection.

Typhlocyba arsinoe McAtee

(PL LXXXI, fig. 4)

Typhlocyba arsinoe McAtee, Proc. U. S. Nat. Mus., vol. 68, art. 18, 1926,
pp. 31-32.

Resembling T. pomaria in external appearance, but differing in

not having face of male orange-red, in having a pair of apical proc-

Revision of the Species Typhlocyba 1167

esses on shaft of aedeagus, and in having ventral angle forming a
small apically rounded lobe.

Length. — 3.25-3.5 mm.

Color. — Head, pronotum, and scutellum white to light yellow;
fore wings white to deep orange-yellow; abdomen yellow; apex of
ventral angle of pygofer black.

Genital capsule. — Male pygofer, in lateral aspect, with posterior
margin slightly concave, dorsal angle slightly produced, ventral
angle strongly produced as a short, rounded ventrocaudally directed
lobe which curves laterad, macrosetae near anterior margin of
ventral lobe distant from outer basal angle of plate, microsetae pos-
terior to these, a few scattered along posterior margin; male plate,
with apex spatulate.

Internal male genitalia. — Styles and connective as in T. oneka
( PI. LXXXI, fig. 2d, e ) but with fewer setae on style; aedeagus with
atrial processes elongate, slender, gradually tapering to acute apices,
nearly attaining apex of aedeagal shaft in length, closely appressed
to ventral margin of shaft to middle its length, then diverging
laterad; aedeagal shaft slightly laterally compressed, a single pair
of apical processes arising subapically on posterior margin and
curving sharply laterodorsad; aedeagal apodeme directed dorso-
caudad on basal two thirds, cephalad on apical third in lateral
aspect.

Specimens have been seen from the following localities: Ontario:
Vineland Station, June 22; New Hampshire: Durham, August 30;
Michigan: Lake Gogebic, August 18; Wisconsin: Milwaukee, June
27, July 3, 5; Amery, August 13; Illinois: Muncie, July 23; Oakwood,
June 14; Massachusetts: Boston, August 31.

The host species is Tilea americana.

Types. — Holotype male and allotype female, in the U. S. National
Museum Collection.

Typhlocyba modesta Gibson

(PI. LXXXII, fig. 1)

Typhlocyba modesta Gibson, Canadian Ent., vol. 49, no. 5, 1917, p. 184.

Resembling Edwardsiana candidula in outward appearance, and
T. hockingensis in shape of aedeagus, but distinguished from these
and other species by having aedeagus with atrial processes fused
medially on basal fourth or half, and by having on dorsal angle of
pygofer a short, thornlike, dorsally directed hook.

44_6490

1168 The University Science Bulletin

Color. — Dorsum uniform ivory white to pale yellow; fore wings
subhyaline to cross veins, apical cells hyaline and slightly fumose;
abdomen ivory white.

Genital capsule. — Male pygofer, in lateral aspect, with posterior
margin produced on dorsal half as a broad arm, bearing a short,
acute, dorsally directed hook and a small ventral lobe, ventral angle
a smoothly rounded lobe, a group of short macrosetae just dorsad
of outer basal angle of male plate, and a patch of macrosetae on
dorsal arm near margin; male plate with apex spatulate.

Internal male genitalia. — Style with lateral margin bearing a short
thumblike lobe covered with setae, inner and outer margins with
a row of microsetae, several alveoli on inner margin near outer
third; connective broadly attached to base of aedeagus; aedeagus
with atrial processes fused to each other on basal fourth or half,
continuing dorsad nearly parallel to each other, slightly diverging
laterad at apex, elongate, slender, scarcely tapering until shortly
before apex; aedeagal shaft greatly elevated above atrial processes,
arising from anterior margin of base of aedeagus near dorsal ex-
tremity; base of aedeagus broadened laterally near apex; aedeagal
apodeme having both anterior and posterior arms.

Female. — With posterior margin of eighth abdominal sternite
having a distinct median notch between two smoothly rounded lobes
(PI. LXXXVII, fig. 11).

A large series has been taken by the author in Douglas County,
Kansas, and a short series in Milwaukee, Wisconsin, on Acer
saccharinum. Specimens have been seen from the following locali-
ties: New York: Minetto, August 15; Virginia: Mountain Lake,
July 11, 12, 15, 18, 23; Falls Church, May 30, August 4, 11; Massa-
chusetts: Holiston, July; Kentucky: Louisville, July 27; Kentucky
Ridge State Forest, June 11; North Carolina: Brevard, June 20;
South Carolina: Clemson College; Georgia: Pemiscot County, Sep-
tember 26; Ohio: Wooster, July 8; Barberton, August 11; Indiana:
Kosciusko County, July 8; Lafayette, August 21, 30; Illinois: Deca-
tur, August; Urbana, July 13; Wisconsin: Milwaukee, June 27-July
5; Minnesota: St. Paul, June 17, August 17; Iowa: Ames, June 8, 19,
September 6; Davenport, September 8; Muscatine, June 6, 8, 9,
August 22; County #88, September 23, August 1; Missouri: Charles-
ton, June 7; Kansas: Manhattan, June 8, 14; Douglas County, May
29 to August 14; Colorado: Ft. Collins, June 24, July 3, 14.

Types. — Holotype, allotype, and female paratype, in U. S. Na-
tional Museum Collection, have been seen.

Revision of the Species Typhlocyba 1169

Typhlocyba medleri sp. nov.
(PI. LXXXII, fig. 2)

Resembling T. surcula in external appearance, but distinguished
from this species by not having pygofer hook on dorsal angle, and
having aedeagal shaft a slender cylindrical tube of nearly uniform
diameter.

Length. — 3.5-3.75 mm.

Color. — Dorsum milky-white to cross veins, subhyaline, apical
cells entirely fumose, hyaline; abdomen milky-white, apex of male
plate black.

Genital capsule. — Male pygofer, in lateral aspect, with posterior
margin nearly straight, slanting obliquely ventrocaudad on dorsal
half, dorsal angle not produced, forming an apically rounded right
angle; ventral angle reduced to a small, rounded, ventrally directed
lobe arising near middle of posterior margin of pygofer, ventral
half of posterior margin forming an evenly rounded ventral lobe, a
group of macrosetae just dorsad of outer basal angle of male plate,
a row of macrosetae along dorsal half of posterior margin; male
plate with apex spatulate.

Internal male genitalia. — Connective broadly attached to aede-
agus; aedeagus with atrial processes elongate, slender, diverging
from each other from base, strongly curving dorsad; base of aede-
agus forming a U-shaped, posteriorly opened plate with atrial proc-
esses broadly attached to arms; aedeagal shaft slender, elongate,
arising from between arms of base, nearly uniform in diameter
throughout its length; aedeagal apodeme short, apically enlarged.

Types. — Holotype male, allotype female, paratype male, and
four paratype females, Milwaukee, Wisconsin, June 26, 1950; two
female paratypes, Milwaukee, Wisconsin, June 29, 1950, P. J. Chris-
tian, in the Snow Entomological Collections of the University of
Kansas.

Typhlocyba hockingensis Knull

(PI. LXXXII, fig. 3)
Typhlocyba hockingensis Knull, Ohio J. Sci., vol. 44, no. 6, 1944, p. 270.

Resembling T. pomaria and T. modesta, but differing from these
by having the dorsal angle of pygofer without hooks, and forming
a right angle; aedeagus with atrial processes distant from aedeagal
shaft, asymmetrical, the left process curving cephalad and crossing
over right process.

Length. — 3.25-3.5 mm.

Color. — Dorsum pale yellowish-white to light yellow; fore wings

1170 The University Science Bulletin

subhyaline to cross veins, apical cells faintly fumose, hyaline; ab-
domen light yellow.

Genital capsule. — Male pygofer, in lateral aspect, with posterior
margin slightly convex, dorsal angle slightly rounded forming nearly
a right angle, ventral angle produced ventrad in a long sharp hook,
a small group of macrosetae just posterior to lateral basal angle of
male plate, a few macrosetae on ventral half of posterior margin;
male plate with apex spatulate.

Internal male genitalia. — Aedeagus with atrial processes elongate,
slender, as broad as shaft in lateral aspect, asymmetrical, left proc-
ess crossing over right anteriorly near middle; shaft arising from
dorsal anterior margin of base, parallel to and half the length of
atrial processes; aedeagal apodeme reduced or absent.

A large series of this species was collected in Milwaukee, Wis-
consin, from a species of Viburnum used as an ornamental shrub
in the city parks. Serious injury was noted in some areas while
in other areas injury was only slight. A short series was collected
from blackberry bushes, Goodman, Missouri, and another short
series from Ulmus fulva, in Douglas County, Kansas. Specimens
have been seen from the following localities: Alaska: Ft. Yukon,
July 15; British Columbia: Hope, August 1; Ontario: Vineland
Station, July 1; Beamsville, June 20; New Hampshire: Alton, Oc-
tober 2; Jackson, September 18; Durham, July 8, 14; Maryland:
Ashton, September 5; Pennsylvania: Hartstown Bog, September 14;
Virginia: Mountain Lake, September 2; Kentucky: Louisville, June
7, July 28; Tennessee: Great Smoky Mountains National Park, Sep-
tember 1; Ohio: Shawnee Forest, June 9; Minnesota: St. Paul, Jane
16; Marshall County, June 24; Wisconsin: Lake Geneva, September
1; Milwaukee, June 26-July 5; Missouri: Goodman, May 28; Kansas:
Douglas County, May 28, 30, June 9-30, July 1, September 24.

Types. — Holotype male, allotype female, and paratypes, in the
Collection of Mrs. J. N. Knull, Columbus, Ohio; paratypes in the
Ohio State University Collection, in the Canadian National Col-
lection, and in the Snow Entomological Collections of the University
of Kansas.

Typhlocyba pomaria McAtee

(PI. LXXXII, fig. 4)

Typhlocyba pomaria McAtee, Proc. U. S. Nat. Mus., vol. 68, art. 18, 1926,

pp. 29-31.
Empoa pomaria, Stear, Pennsylvania Acad. Sci., vol. 2, 1928, pp. 54-58.

Resembling T. hockingensis in shape of pygofer, but distinguished
by having aedeagal shaft arising from between atrial processes;

Revision of the Species Typhlocyba 1171

pygofer with dorsal angle reduced and not forming a right angle;
face of male usually red-orange.

Length. — 3.25-3.5 mm.

Color. — Dorsum light yellowish-white to yellow; fore wings sub-
hyaline to cross veins, apical cells hyaline, fumose; abdomen light
yellow.

Genital capsule. — Male pygofer, in lateral aspect, with posterior
margin nearly straight, obliquely slanting posteriorly, dorsal angle
broadly rounded, ventral angle produced ventral in a long sharp
hook as in T. hockingensis, a group of macrosetae just dorsad of
outer basal angle of plate, a patch of macrosetae on posterior margin
near middle; male plate with apical lobe rounded, only slightly en-
larged.

Internal male genitalia. — Aedeagus with atrial processes elongate,
broadly attached at base, tapering to acute sinuate apices; aedeagal
shaft arising from between atrial processes, broadened laterally and
apically as a thin plate, gonopore subapical on ventrocaudal surface;
base of aedeagus and apodeme reduced.

Female. — With posterior margin of eighth abdominal sternite
sinuate, slightly produced medially, with lateral third curving
strongly dorsocephalad (PI. LXXXVII, figs. 12a, b).

This species occurs on species of Mains and on Ulmus americana.
The approved common name for this species is "The White Apple
Leafhopper" (Muesbeck, 1950, p. 138) and is referred to by this
name in the numerous papers concerning its biology and control
which have been published since its description. It probably occurs
in all of the regions of North America where its host plants are
found. Specimens have been seen from the following localities:
Alaska: Fairbanks, July 30; Matanuska, July 21; Ft. Yukon, July 15;
British Columbia: Vancouver, August 5, 8; Quebec: Newaygo, July
30; Nova Scotia: Smith's Cove, October 4; King's County; Ontario:
Vineland Station, August 17, 29, September 9, 14, October 10; Sim-
coe, October 6; Trenton, June 17; Chelsea, June 20-25; Ottawa, June
18, August 22, October 1; Fenwick, October 10; St. David's, June
26; Massachusetts: Greenfield, August 24; Holliston, July, Septem-
ber 16-22; New Hampshire: Wilton County, September 30; Durham,
September 13, August 21, 27; Vermont: Mount Mansfield, July 26;
New York: Ithaca, June 8, 12, 17; Pennsylvania: Procter, July 21;
Hartstown Bog, June 26, September 13; Northeast, June 15, July 2,
3; Westchester, July 6; Maryland: Lakeland, September 12; Virginia:
Arlington, May 30, June 11, 15, September 19, 29; Leesburg, August
17, September; Copeley, October 6; Kentucky: Louisville, October

1172 The University Science Bulletin

12; Tennessee: Gatlinburg, June 14; Great Smoky Mountains Na-
tional Park, September 1; Indiana: Noble County; Illinois: Cook
County, June 22; Thornton, September 7; R. Canyon State Park,
July 10; Olney, September 21; Minnesota: St. Paul, August 25;
Winona County, July 1; Itaska County, August 19; Michigan: Au-
gust 18; South Dakota: Brookings, June 7, 8, 25, July 1; Wisconsin:
Milwaukee, June 26-July 7; Brule, August 19; Rib Mountain State
Park, August 27; Madison, June 24; Cramoor, August 11; Lake
Geneva, June 16-24, July 4, 9, 12; Iowa: Ames, September 3 through
22; Davenport, June 10, September 3; Muscatine, June 4-15; Mis-
souri: Oregon County, May 28; Kansas: Douglas County, May 27-
June 24, September 20-21; Riley County, September 21; Arkansas:
Bentonville, September 1; Colorado: Glen Haven, August 1; Ft.
Collins, June 14, 20, July 3, September 4, 10, 23; Utah: Providence,
October 8; Richfield, June 15, August 7; Logan, June 2-27, Septem-
ber 14, 22; Provo, September 17; Lake View, July 17; New Mexico:
Espanola, June 18; Idaho: Twin Falls, June 14; Moscow, June 19;
Parma, July 25, September 22; Idaho Falls, July 27; River Dale,
August 12, September 1; Washington: Dishman, July 7; Oregon:
Salem, October 22; Hood River, July 17, August 20; Yoncalla, July
12; The Dalles, June 8, 10, 18; Azales, September, October; Peoria,
October; Lancaster, October; Umatilla, October; Junction City,
October; Gold Hill, October; 5 miles north of Coberg, October;
Durfur; Green Springs Mountain; Ashland; Jacksonville; Clackomas;
California: Mount Shasta Canyon, June 29.

Types. — Holotype, allotype, and paratypes, in the U. S. National
Museum Collection; paratypes, in the Snow Entomological Collec-
tions of the University of Kansas, in the Illinois State Natural History
Survey Collection, and in the Iowa State College Collection. The
male paratype from Cabin John Bridge, Maryland, June 17, 1915,
Roberts, is a male of Edwardsiana rosae.

Typhlocyba attenuata sp. nov.
(PL LXXXIII, fig. 1)

Resembling T. putmani, Edwardsiana dejecta, and Empoa albi-
cans in outward appearance, and T. rubriocellata in shape of aede-
agus, distinguished from these species by having dorsal posterior
angle of pygofer produced in an acute, lightly sclerotized, attenuate
hook.

Length. — 4.0-4.25 mm.

Color. — Dorsum pale white to yellow; fore wings subhyaline to

Revision of the Species Typhlocyba 1173

cross veins, apex hyaline, not fumose; abdomen yellow, ventral angle
of pygofer black on apex.

Genital capsule. — Male pygofer, in lateral aspect, with dorsal
angle produced posteriorly in an attenuate hook, ventral angle
acute, produced ventrad as a small, apically rounded tubercle, a
group of macrosetae just posterior to outer basal angle of plate, a
large patch of short macrosetae along middle of posterior margin
extending inward submarginally on disc; male plate with apex
rounded, not spatulate.

Internal male genitalia. — Style broad, elongate, slightly enlarged
subapically, with setae on outer margin extending nearly to apex,
several alveoli on mesal margin near middle, scattered setae basad
of these; connective broadly triangular; aedeagus with atrial proc-
esses elongate, slender, nearly straight, as long as shaft, fused shortly
before base, diverging laterad to dorsally curved apices, a small
ventral process at base; aedeagal shaft arising from anterior margin
of dorsal half of base of aedeagus, expanded laterally and apically
as a thin, lightly sclerotized plate, medially more heavily sclerotized,
apical third slightly broader and distally rounded, gonoduct open-
ing subapically on ventrocaudal surface; base of aedeagus forming a
broad pillar supporting and elevating shaft above atrial processes.

Female. — With posterior margin of eighth abdominal sternite
broadly evenly rounded, not produced or incised (PL LXXXVII,
fig- 9).

A large number of specimens of this species have been collected
on Aesculus glabra var. sargentii in Douglas County, Kansas.

Types. — Holotype male, allotype female, and numerous para-
types of both sexes, Douglas County, Kansas, May 28, 1949, R. H.
Beamer; additional paratypes: one male, Rosedale, Kansas, June 23,
1924, E. P. Breaky; six females, Atchison County, Kansas, July 10-
16, E. P. Breaky and R. H. Beamer; six females, Douglas County,
Kansas, June 25, 1945, R. H. Beamer; ninety-nine males and females,
Douglas County, Kansas, May 27, 1949, R. H. Beamer and P. J.
Christian; sixty-seven males and females, Douglas County, Kansas,
May 29-31, 1949, R. H. Beamer and P. J. Christian; two females,
June 18, 22, 1949, Douglas County, Kansas, P. J. Christian; three
females, May 29, 1950, ten females, June 3, 1950, one male and four
females, June 5, 1950, one male and two females, June 7, 1950, two
males and two females, June 8, 1950, Douglas County, Kansas, P. J.
Christian. Types in the Snow Entomological Collections of the
University of Kansas.

1174 The University Science Bulletin

Typhlocyba rubriocellata Malloch
(PI. LXXXIII, fig. 2)

Typhlocyba rubriocellata Malloch, Bull. Brooklyn Ent. Soc, vol. 15, nos. 2 and

3, 1920, p. 48.
Typhlocyba rubriocellata var. clara McAtee, Proc. U. S. Nat. Mus., vol. 68,

art. 18, 1926, p. 21.
Typhlocyba escana Ross and DeLong, Ohio J. Sci., vol. 49, no. 3, 1949, pp. 117-

118. {new synonymy)

Resembling T. attenuata in shape of aedeagus, but easily dis-
tinguished by having a broad brown band across apical veins, and
usually with a bright crimson-red spot on fore wing.

Length. — 3.5-4.0 mm.

Color. — Dorsum with ground color yellowish-white; fore wings
with a solid dark brown band over cross veins covering ends of
cells bordering on cross veins, with bright crimson-red spot of varia-
ble size in middle of each fore wing along middle of inner half of
clavus and over part of basal half of inner two basal cells, entirely
absent in some specimens, fore wings subhyaline to cross veins,
hyaline beyond dark band, slightly fumose; abdomen yellowish-
white.

Genital capsule. — Male pygofer, in lateral aspect, with posterior
margin strongly convex, dorsal posterior margin directed dorsad,
ventral angle forming a broadly rounded lobe interrupted by a
slight ventral hook, posterior margin deeply inrolled; male plate
with apex rounded but not spatulate.

Internal male genitalia. — Style, connective, and aedeagus very
similar to those of the preceding species; aedeagus with atrial proc-
esses shorter than shaft, elongate, slender; aedeagal shaft arising
from dorsal two thirds of base, broadly expanded, not constricted
subapically, more broadly expanded laterally; base of aedeagus
broadened laterally, a short anteriorly directed apodeme on anterior
margin at apex.

Female. — With posterior margin of eighth abdominal sternite
broadly evenly rounded, as in T. attenuata (PI. LXXXVII, fig. 9).

A large series of specimens has been seen from the type locality,
collected on Aesculus sp. Specimens have been seen from the
following localities: Illinois: Urbana, July 12, 14, 20; Ohio: Colum-
bus, June 1; New York: Monroe, July 10; Kentucky: Kentucky
Ridge State Forest, June 11; Tennessee: Gatlinburg, June 24, 28,
July 12, 20, 21; Great Smoky Mountains, June 18, September 1.

Types. — Holotype female, in the Illinois State Natural History
Survey Collection; a male from Urbana, Illinois, July 14, 1946, R. H.

Revision of the Species Typhlocyba 1175

Beamer, here designated neoallotype, in the Snow Entomological
Collections of the University of Kansas; one paraallotype male,
Great Smoky Mountains, Tennessee, June 18, 1939, C. P. Alexander,
in the U. S. National Museum Collection.

Typhlocyba surcula DeLong and Johnson

(PL LXXXIII, fig. 3)

Typhlocyba surcula DeLong and Johnson, Ent. News, vol. 47, no. 4, 1936,
p. 103.

Resembling T. medleri externally, but with an acute pygofer hook
directed posteriorly from dorsal angle.

Length. — 3.25 mm.

Color. — Dorsum pale yellowish-white; fore wings subhyaline to
cross veins, apical cells hyaline, faintly fumose; abdomen yellowish-
white.

Genital capsule. — Male pygofer, in lateral aspect, with posterior
margin slightly convex ventrad of dorsal processes, dorsal process
broad basally but reduced strongly to an acute, heavily sclerotized,
posteriorly directed apex, ventral angle forming a small round
ventrally directed lobe; male plate with apex not spatulate.

Internal male genitalia. — Style produced laterad in a broad, an-
gular projection near point of attachment to connective, apex
strongly curving ventrolaterad, setae scattered over ventral surface
from lateral to mesal margins at middle, a few setae on lateral
margin extending more distad, several alveoli on mesal margin at
outer third; connective with anterior median lobe strongly pro-
duced; aedeagus, similar to preceding species but with atrial proc-
esses slender from base, shaft arising slightly above bases of proc-
esses; apex transparent, acutely pointed, sclerotized portions making
it appear bifurcate; aedeagal apodeme V-shaped in dorsal aspect,
arising from lateral arms of base.

One specimen of this species was taken on Quercus muhlenbergii
in Douglas County, Kansas, and other specimens have been col-
lected in Milwaukee, Wisconsin, from mixed trees and bushes not
including the above species.

Specimens have been seen from the following localities: Minne-
sota: St. Paul, June 27; Wisconsin: Milwaukee, June 29, July 2, 7;
Wisconsin Rapids, July 27; Cranmoor, July 27; Illinois; Kansas:
Douglas County, June 11.

Types. — Holotype and paratype males, in the collection of D. M.
DeLong, Columbus, Ohio.

1176 The University Science Bulletin

Typhlocyba andromache McAtee

(PI. LXXXIII, fig. 4)

Typhlocyba andromache McAtee, Proc. U. S. Nat. Mus., vol. 68, art. 18, 1926,
p. 32.

Resembling T. surcula in external appearance and in shape of
aedeagus, but easily distinguished by lacking dorsal hook on pygo-
fer, and by having posterior margin nearly straight.

Length.— 3.0-3.25 mm.

Color. — Dorsum pale yellowish-white; fore wings subhyaline to
cross veins, apical cells hyaline, slightly fumose.

Genital capsule. — Male pygofer, in lateral aspect, with posterior
margin nearly straight, dorsal angle acute, slightly less than a right
angle, ventral angle slightly more than a right angle and forming a
small ventrally directed hook; male plate with apex only slightly
enlarged.

Internal male genitalia. — Aedeagus with atrial processes broad,
elongate, exceeding shaft in length, diverging laterodorsad from
base, slightly sinuate, reduced to acute apices on outer fourth;
aedeagal shaft arising from base slightly above bases of processes,
laterally broadened, margins foliaceous, broadest at gonopore, nearly
transparent and extending beyond gonopore as a thin plate with
apex acute; base of aedeagus reduced, aedeagal apodeme directed
dorsad and cephalad.

The following specimens have been seen: six males, Itasca County,
Minnesota, July 26, 1939, J. T. Medler, and the holotype male,
Salem, New York, June 27, 1924, on birch, E. D. Ball.

Typhlocyba melite McAtee

(PI. LXXXIV, fig. 1)

Typhlocyba melite McAtee, Proc. U. S. Nat. Mus., vol. 68, art. 18, 1926, p. 32.

Resembling T. arsinoe in external appearance, but differing in
having atrial processes of aedeagus greatly broadened on basal two
thirds, and with a short hook at middle of posterior margin of
pygofer.

Length.— 3.5-3.75 mm.

Color. — Dorsum pale yellowish-white to yellow; fore wings sub-
hyaline to cross veins, apical cells hyaline, not fumose; abdomen
yellow.

Genital capsule. — Male pygofer, in lateral aspect, with posterior
margin slightly convex, dorsal angle reduced, dorsal and posterior
margins almost continuous, a small ventrocaudally directed hook

Revision of the Species Typhlocyba 1177

near middle of posterior margin, ventrocaudal margin broadly
rounded ventrad of hook; male plate with apex spatulate.

Internal male genitalia. — Connective Y-shaped with anterior and
posterior medial lobes narrowed; aedeagus with atrial processes
broadly flattened on basal two thirds, slightly narrowed before base,
outer third gradually tapering to acute apex; aedeagal shaft arising
from ventrad of processes and between them, slender, laterally
flattened, elongate, gradually tapering to apex, strongly recurved
just before apex and appearing bifid due to differential sclerotiza-
tion, posterior margin strongly convex; aedeagal apodeme short,
sharply curving ventrocephalad.

A large series of specimens has been collected by the author from
gooseberry ( Ribes sp. ) in Milwaukee, Wisconsin, and another series
from pawpaw (Asimina triloba) in Douglas County, Kansas. Speci-
mens have been seen from the following localities: Ontario: Vine-
land Station, June 30, July 4, 8; Massachusetts: Boston, July 27; New
York: Sea Cliff, August; Hudson Valley, June 14; Kentucky: Ken-
tucky Ridge State Forest, June 11; Ohio: Columbus, June 15; Minne-
sota: St. Paul, June 15, 16, 19; Ramsay County, May 30; Wisconsin:
Milwaukee, June 27-July 4; Illinois: Western Springs, June 21; Iowa:
Ames, May 25; Missouri: Kansas City, May 29; Kansas: Douglas
County, June 2-July 5, September; Colorado; Washington: Kalama,
July 4.

Types. — Holotype male, and paratype, in U. S. National Museum;
allotype and paratypes of both sexes in the Museum of Comparative
Zoology.

Typhlocyba alabamaensis sp. nov.
(PI. LXXXIV, fig. 2)

Resembling T. medleri externally, but with dark apical markings
more intense along cross veins, aedeagus differs from those of other
species in having ventrocaudal margin concave rather than convex.

Length. — 3.5 mm.

Color. — Dorsum light yellow to cross veins; fore wings subhy aline
to cross veins, apical cells dark brown, hyaline, more intensely
colored along cross veins.

Genital capsule. — Male py gofer, in lateral aspect, with posterior
margin sinuate, dorsal angle produced in a broad, laterally concave,
dorsocaudally directed, apically rounded process, ventral angle di-
rected ventrocaudad as a broad apically rounded lobe arising from
mesad of ventral lobe, a group of macrosetae just dorsad of outer
basal angle of male plate extending to middle of disc, scattered

1178 The University Science Bulletin

microsetae posterior to these and along dorsal margin near base of
anal tube; male plate with apical lobe scarcely enlarged.

Internal male genitalia. — Connective with a transverse bar at
apex of anterior lobe, appearing T-shaped as in T. crassa; aedeagus
with atrial processes broad, elongate, diverging laterally from base,
curved sharply mesad near apex, curving dorsocaudad in lateral
aspect, gradually reduced to acute apex; aedeagal shaft arising from
between processes, half as long as atrial processes, broadly attached
to base in lateral aspect, laterally compressed, posterior margin
forming a sharp ventrally concave keel gradually diminished toward
apex; base of aedeagus shield shaped in posterior aspect, with aede-
agal apodeme V-shaped and attached to divergent arms of base.

Types. — Holotype male and male paratype, La Place, Alabama,
near Tuskeege, June 9, 1917, in the Cornell University Collection.

Typhlocyba shawneeana Knull
( PI. LXXXIV, fig. 3 )
Typhlocyba shawneeana Knull, Ohio J. Sci., vol. 44, no. 6, 1944, p. 270.

Resembling T. andromache in external appearance and in shape
of pygofer, but easily distinguished from that species by having two
pairs of shaft processes.

Length. — 3.5-4.0 mm.

Color. — Dorsum pale yellow; fore wings subhyaline to cross veins,
apex hyaline.

Genital capsule. — Male pygofer, in lateral aspect, with posterior
margin nearly straight, almost vertical, dorsal and ventral angles
sharp right angles, slightly produced, a few macrosetae just dorsad
of outer basal angle of male plate, a group of macrosetae on dorsal
half of posterior margin of pygofer extending slightly inward on
disc; male plate with apex spatulate.

Internal male genitalia. — Aedeagus with atrial processes elongate,
broad, enlarged at base, gradually reduced on outer third to acute
apices, exceeding shaft in length; aedeagal shaft bearing two pairs
of processes of about equal length, one pair at apex directed dorso-
caudad, widely diverging, the other pair at outer fifth curving mesad
caudally from lateral margins of shaft, shaft greatly broadened,
margins foliacious, outer fifth reduced to half width at base, directed
dorsad, slightly convex on basal half; aedeagal apodeme broad,
directed cephalad.

This species has been collected by the author from witch hazel
(Hamamelis virginiana) in Milwaukee, Wisconsin. Although not
abundant, a number of specimens were collected by aspirator from

Revision of the Species Typhlocyba 1179

the under surfaces of those leaves which showed considerable feed-
ing injury. Nymphs and freshly emerged males were found on the
host.

Specimens have been seen from the following localities: Ohio:
Fairfield, June 16; Tennessee: Great Smoky Mountain National
Park, September 1, on witch hazel; Wisconsin: Milwaukee, July 5, 7.

Types. — Holotype and paratype males, in the collection of Mrs.
J. N. Knull, Columbus, Ohio; a female collected by the author, July
7, 1950, Milwaukee, Wisconsin, here designated neoallotype, in the
Snow Entomological Collections of the University of Kansas.

Typhlocyba transviridis sp. nov.
(PI. LXXXIV, fig. 4)

Resembling T. Cassiopeia in being light green when fresh and
later changing to yellow; easily distinguished from other species by
having a broad, brown, transverse band covering median third of
fore wings.

Length.— 3.25-3.5 mm.

Color. — Fresh specimens bright green with a broad greenish-
brown transverse band on median third of fore wings, color later
changing to yellow with a broad yellowish-brown band, veins across
band sometimes light colored, apical cells light colored, hyaline;
abdomen bright green changing to yellow.

Genital capsule. — Male pygofer, in lateral aspect, with posterior
margin continuous with dorsal margin, dorsal angle greatly re-
duced, ventral angle forming a broad caudally directed lobe with
a small ventrally directed hook on ventral margin, a group of macro-
setae just dorsad of outer basal angle of male plate, numerous
microsetae scattered on ventral lobe over disc and along dorsal
margin, a large group of macrosetae extending along most of the
posterior margin and inward on disc; male plate not enlarged at
apex.

Internal male genitalia. — Style with lateral margin bearing two
lobes near middle, a few scattered setae on lateral and mesal mar-
gins; connective as in T. Cassiopeia (PI. LXXXV, fig. 2); aedeagus
with atrial processes elongate, slender, gradually reduced to acute
apices, slightly divergent on outer sixth to apex, strongly curving
dorsocephalad from base, parallel to shaft; aedeagal shaft arising
between processes and continuing between them throughout its
length, two thirds as long as processes and of equal width; base of
aedeagus a broad dorsal arm bearing a short anteriorly directed
apodeme at apex.

1180 The University Science Bulletin

A large series of specimens of this species has been collected by
the author from Tilea americana (linden) in Milwaukee, Wis-
consin.

Types. — Holotype, allotype, six male and fourteen female para-
types, July 2, 1950, Milwaukee, Wisconsin, P. J. Christian; addi-
tional paratypes: one male, June 30, 1950; one female, July 1, 1950;
three females, July 3, 1950; four males, July 4, 1950; eight males
and four females, July 5, 1950; two females, July 7, 1950, Milwaukee,
Wisconsin, P. J. Christian. Types in the Snow Entomological Col-
lections of the University of Kansas.

Typhlocyba putmani Knull
(PI. LXXXV, fig. 1)
Typhlocyba putmani Knull, Ohio J. Sci., vol. 44, no. 6, 1944, p. 269.

Resembling T. attenuata, Empoa albicans, and Edwardsiana
dejecta in size and color; distinguished by having aedeagal shaft
flattened laterally and lying ventrocaudad of atrial processes
throughout its length.

Length. — 4.0-4.25 mm.

Color. — Dorsum pale yellow without dark markings; fore wings
subhyaline to cross veins, apical cells hyaline; abdomen pale yellow.

Genital capsule. — Male pygofer, in lateral aspect, with posterior
margin slightly angular, dorsal angle reduced, ventral angle pro-
duced ventrad as a short acute tooth, a vertical row of macrosetae
just dorsad of outer basal angle of male plate, a patch of macro-
setae at middle of posterior margin; male plate with apex spatulate.

Internal male genitalia. — Style with a short hook on lateral mar-
gin near attachment to connective, a row of setae along lateral and
mesal margins, several alveoli beyond these on mesal margin at
outer third; connective triangular; aedeagus, with atrial processes
broadly attached at base, compressed laterally, gradually tapering
to acute recurved apices, exceeding shaft in length by outer third;
aedeagal shaft elongate, slender, laterally flattened, gradually re-
duced to outer fourth, more abruptly reduced to apex, arising
ventrad of processes and continuing posterior of them throughout
its length; aedeagal apodeme similar to that of the preceding species.

A large series of this species has been collected by the author
from Cornus stolonifera, Milwaukee, Wisconsin, in association with
Edwardsiana dejecta, where noticeable injury could be seen on
hedges of the host plants in the city parks. Specimens have been
seen from the following localities: New York: Glen Cove, Long
Island, July 8; Virginia: Mountain Lake, July 21; West Virginia:

Revision of the Species Typhlocyba 1181

Great Cacapon, July 4; Tennessee: Great Smoky Mountains Na-
tional Park, September 1; Wisconsin: Milwaukee, July 1, 3, 4;
British Columbia: Vancouver, August 4.

Types. — Holotype male, allotype, and paratypes, Vineland Sta-
tion, Ontario, in the Canadian National Collection.

Typhlocyba Cassiopeia Knull
(PI. LXXXV, fig. 2)
Typhlocyba Cassiopeia Knull, Ohio J. Sci., vol. 44, no. 6, 1944, p. 269.

Resembling T. transviridis in shape of aedeagus and pygofer, but
lacking brown color markings; atrial processes fused to aedeagal
shaft on basal third and closely appressed to shaft concealing it
throughout most of its length.

Length. — 3.25-3.5 mm.

Color. — Entire body light green when fresh, color changing to
pale yellow, legs with apex of tibia remaining green longer than
rest of body.

Genital capsule. — Male pygofer, in lateral aspect, with posterior
margin evenly convex, dorsal angle reduced, dorsal and caudal
margins nearly continuous, ventral angle forming an evenly rounded
lobe; male plate not enlarged apically.

Internal male genitalia. — Aedeagus with atrial processes arising
from basal third of shaft, slender, elongate, gradually reduced to
acute apices, slightly divergent, directed dorsad, closely following
shaft and exceeding it in length; shaft slender, sometimes difficult
to see in less heavily sclerotized specimens; base of aedeagus short
with apodeme reduced.

The green color of the legs is useful in separating freshly collected
specimens of this species from those of other light colored species,
but after a month or two this color distinction is lost.

A large series of this species has been collected by the author
from Primus virginiana (P. serotina), in Milwaukee, Wisconsin.
Paratype males from Vineland Station, Ontario, have been seen.
Specimens have been seen from the following localities: Massa-
chusetts: Holliston, July; Ontario: Vineland Station, June 27, July
4; Minnesota: St. Paul, June 22, Itasca County, July 12; Wisconsin:
Milwaukee, June 26-July 7; Illinois: Cook County, June 22; Iowa:
Ames, June 17; Kansas: Douglas County, August 19, 24 ( a paratype
of T. pomaria McAtee); Washington: Mount Rainier, July 27;
Oregon: Jacksonville.

Types. — Holotype male, allotype, and paratypes, in the collection
of Mrs. J. N. Knull; paratypes, in the Canadian National Collection.

1182 The University Science Bulletin

Typhlocyba crassa DeLong and Johnson

(PI. LXXXV, fig. 3)

Typhlocyba crassa DeLong and Johnson, Ent. News, vol. 47, no. 4, 1936,
pp. 102, 104.

Resembling T. putmani in external appearance and in structure
of male genitalia, but with posterior margin of pygofer straight,
aedeagal shaft slender and of uniform diameter throughout its
length, and atrial processes curving ventrad at base forming a bow
which projects between male plates.

Length. — 3.5-3.75 mm.

Color. — Dorsum pale yellowish-white; fore wings subhyaline to
cross veins, apical cells hyaline.

Genital capsule. — Male pygofer, in lateral aspect, with posterior
margin straight, nearly vertical, dorsal margin convex, dorsal angle
rounded, ventral angle produced in a broad, acutely angled, ven-
trally-directed lobe, several macrosetae dorsad of outer basal angle
of male plate, a patch of macrosetae along dorsal half of posterior
margin; male plate with apex spatulate.

Internal male genitalia. — Connective broadened anteriorly as a
transverse plate (PI. LXXXV, fig. 3e); aedeagus, with atrial proc-
esses usually attenuate, twice length of shaft, directed ventrad at
base, curved dorsocaudad to outer third, then curved dorsocephalad
diverging laterally on outer fourth, median third appearing broad-
ened in lateral aspect; shaft slender, elongate, of nearly uniform
diameter from base to outer sixth where it is abruptly reduced be-
yond gonopore to a very acute apex; aedeagal apodeme forming a
massive anteriorly directed arm.

The recorded host for this species is Prnnns serotina.

Specimens have been seen from the following localities: Ontario:
Vineland Station, July 4; Pennsylvania: Hartstown Bog, June 16,
September 30; Wisconsin: Milwaukee, July 7; Colorado: Glen
Haven, August 1; Wyoming: Yellowstone Park, August 15.

Types. — Holotype male, allotype female, and three female para-
types, in the collection of Dr. D. M. DeLong, Columbus, Ohio.

Typhlocyba sollisa Ross and DeLong
( PI. LXXXVI, fig. 1 )

Typhlocyba sollisa Ross and DeLong, Ohio J. Sci., vol. 49, no. 3, 1949, pp. 116-
117.

Resembling T. niobe and T. persephone externally, but distin-

Revision of the Species Typhlocyba 1183

guished from these species by the aedeagal shaft having processes
subapical and distinctly separate from shaft from base.

Length. — 3.5 mm.

Color. — Head, pronotum, and scutellum yellow; fore wings yel-
low-orange, subhyaline to cross veins, apex hyaline, veins red-
orange.

Genital capsule. — Male pygofer, in lateral aspect, with dorsal
angle produced in a slight hook, ventral angle produced latero-
ventrad as a small tubercule, a single macroseta dorsad of outer basal
angle of male plate, patch of macrosetae near middle of posterior
margin; male plate, with apex spatulate.

Internal male genitalia. — Aedeagus with a pair of lateral proc-
esses arising near middle of shaft and continuing parallel to it to
apex, then curving sharply laterad, broadly attached to shaft;
aedeagal shaft greatly reduced distad of processes, slender, shorter
than processes; base of aedeagus slender, directed dorsad, a short,
broad, anteriorly directed apodeme at apex.

The only specimen known is the holotype, taken at Grand Tower,
Illinois, May 30, 1935, by Ross and Mohr.

Type. — Holotype male, in Illinois State Natural History Survey
Collection.

Typhlocyba niobe McAtee

(PL LXXXVI, fig. 4)
Typhlocyba niobe McAtee, Proc. U. S. Nat. Mus., vol. 68, art. 18, 1926, p. 20.

Resembling T. persephone and T. sollisa externally, but distin-
guished from these by having aedeagal shaft broad on basal half and
greatly reduced beyond shaft processes, shaft processes closely ap-
pressed to shaft, ventral angle of pygofer with pronounced hook.

Length. — 3.5 mm.

Color. — Dorsum light sulfur-yellow to deep yellow-orange; fore
wing subhyaline to cross veins, apex hyaline; abdomen sulfur-yellow.

Genital capsule. — Male pygofer in lateral aspect, with posterior
margin slightly convex, dorsal angle slightly pointed, ventral angle
produced ventrocaudad as a broad hooked lobe.

Internal male genitalia. — Aedeagus without atrial processes, with
a pair of lateral processes arising near middle of shaft broadly at-
tached to shaft and gradually diminishing to acute apices; aedeagal
shaft greatly reduced distad of processes, continuing parallel to
these nearly to apex where they diverge laterodorsad, basal half
broadened dorsoventrally; base of aedeagus directed dorsocaudad
with a short anteriorly directed apodeme.

45—6490

1184 The University Science Bulletin

A large series of this speries has been taken in Milwaukee, Wis-
consin, by the author, on Acer saccharum and Acer platanoides in
association with T. persephone and several similarly colored species.
Specimens have been seen from the following localities: Michigan:
Agricultural College, July 5; Wisconsin: Milwaukee, June 26-July 6;
Illinois: Andres, June 17; Iowa: Ames, June 20, 23.

Types. — Holotype male and paratypes of both sexes, in the U. S.
National Museum Collection; allotype female and one pair of para-
types, in the Iowa State College Collection.

Typhlocyba persephone McAtee
(PI. LXXXVI, fig. 3)

Typhlocyba persephone McAtee, Proc. U. S. Nat. Mus., vol. 68, art. 18, 1926,

p. 11.
Typhlocyba lancifer McAtee, op. cit., pp. 19-20. (new synonymy).

Resembling T. niobe and T . sollisa in external appearance, but
distinguished by having aedeagal shaft processes apical.

Length. — 3.5 mm.

Color. — Dorsum light sulfur-yellow to deep orange-yellow, some
specimens red-orange; abdomen sulfur-yellow.

Genital capsule. — Male pygofer, in lateral aspect, with posterior
margin slightly convex, dorsal angle not produced but slightly
angular, ventral angle produced as a broad lobe slightly hooked at
apex, a few macrosetae dorsad of outer basal angle of male plate,
a row of macrosetae along dorsal three fourths of posterior margin;
male plate with apex spatulate.

Internal male genitalia. — Style with apical portion beyond alveoli
strongly reduced, apex curving ventrad; connective strongly pro-
duced anteriorly in a slender, elongate, median lobe; aedeagus with-
out atrial processes, with shaft slender, elongate, gradually tapering
toward apex in lateral aspect, apical processes broadly inflated near
middle, directed dorsolaterad forming a distinct angle with shaft;
base of aedeagus broad, directed dorsocaudad, with aedeagal
apodeme arising from anterior margin at apex and directed cephalad.

Female. — With posterior margin of eighth abdominal sternite
strongly curving dorsocephalad from midline, slightly produced
medially as a rounded lobe (PI. LXXXVII, figs. 10a, b).

A large series of specimens of this species has been taken by
Dr. R. H. Beamer and the author, on Acer saccharum, in Douglas
County, Kansas, in an unmixed population. Another large series
was taken by the author from Acer saccharum and A. platinoides,
in Milwaukee, Wisconsin, in association with Typhlocyba niobe and
several other similar species.

Revision of the Species Typhlocyba 1185

Specimens have been seen from the following localities: Massa-
chusetts: Hampshire County, June; New Hampshire: Durham,
July 5, September 7; Vermont: Mansfield, July 24; Ontario: Vine-
land Station, June 22; New York: Minetto, June 13; Tennessee
Gatlinburg, June 14; Wisconsin: Milwaukee, June 27-July 5; Illinois
Urbana, June 4; Kansas: Douglas County, May 26-June 24; Iowa
Ames, June 20.

This species was described from a male specimen with apical
processes of the aedeagus broken off, but in all other respects like
the holotype of T. lancifer. Since the description of T. persephone
precedes that of T. lancifer, the latter is considered a synonym of
the former.

Types. — Holotype male, allotype and paratype females, in the
U. S. National Museum Collection; all paratype males of this species
seen by the author are specimens of T. niobe. The allotype and
female paratype specimens may also be this species, but at present
the author has not been able to distinquish between females of
these two species so that they are being regarded as specimens of
T. persephone until a sufficiently reliable method is found for dis-
tinguishing between them.

Typhlocyba tortosa Ross and DeLong

(PI. LXXXVI, fig. 2)

Typhlocyba tortosa Ross and DeLong, Ohio J. Sci., vol. 49, no. 3, 1949, pp 115-
116.

Resembling T. niobe in external appearance, but distinguished by
having pygofer with ventral angle smoothly rounded, and in having
apical processes short and only slightly subapical.

Length. — 3.0 mm.

Color. — Dorsum pale white to light yellow; abdomen yellow.

Genital capsule. — Male pygofer, in lateral aspect, with posterior
margin convex at middle, dorsal angle produced as a short, broad,
ventrally hooked process, ventral angle broadly rounded, with
macrosetae dorsad of outer basal angle of plate, a large group of
macrosetae on posterior margin at middle extending inward on
disc; male plates with apex spatulate.

Internal male genitalia. — Style broadly curved ventrolaterad,
slender, elongate, with ventral surface covered with setae at middle
from lateral to median margins, several alveoli on mesal margin;
connective as in T. persephone (PI. LXXXVI, fig. 3); aedeagus
without atrial processes, shaft slender, elongate, gradually reduced
toward apex, extending dorsad; apical processes short, parallel,

1186 The University Science Bulletin

sinuate, continuing in same direction as shaft; base of aedeagus
forming a broad, caudally concave plate, with apodeme directed
cephalad from dorsal margin.

A large series of specimens of this species collected by Dr. R. H.
Beamer and the author, from Ostrya virginiana, Douglas County,
Kansas, May 30, June 1, 7, a shorter series from the same host,
Milwaukee, Wisconsin, June 28, 29, July 2, 7, collected by the
author, and a single male, Wonalancet, New Hampshire, July 1,
have been seen.

Types. — Holotype male from Oakwood, Illinois, June 14, in the
Illinois State Natural History Collection; a female specimen col-
lected by the author in Douglas County, Kansas, June 7, 1950, in
association with males of this species, here designated neoallotype,
is in the Snow Entomological Collections of the University of
Kansas.

Typhlocyba inflata sp. nov.

(PI. LXXXVII, fig. 1)

Resembling T. tortosa in outward appearance and in form of male
genitalia, but differing in having distal half of aedeagal shaft inflated
laterally, and in lacking lobe on dorsal angle of pygofer.

Length.— 3.25-3.5 mm.

Color. — Dorsum pale whitish-yellow to light yellow; abdomen
light yellow.

Genital capsule. — Male pygofer, in lateral aspect, with posterior
margin convex on lower third; dorsal angle rounded apically, not
forming a lobe; ventral angle broadly rounded, a short laterally-
directed hook at apex, several short macrosetae dorsad of outer
basal angle of plate, a large group of macrosetae on posterior mar-
gin at middle extending inward on disc; male plate with apex
spatulate.

Internal male genitalia. — Style broadly curving ventrolaterad,
slender, elongate, with ventral surface covered with setae at middle
from lateral to median margins, several alveoli on mesal margin;
connective broadly triangular, median anterior lobe strongly pro-
duced; aedeagus without atrial processes, shaft slender, elongate,
broadly inflated on apical half in caudal aspect; apical processes
short, parallel, appearing as a continuation of shaft in lateral aspect;
base of aedeagus forming a laterally flattened dorsal arm, with
apodeme at apex reduced.

Types. — Holotype male, allotype female, one male and two fe-
male paratypes, Shelton, Washington, July 24, 1949, R. H. Beamer

Revision of the Species Typhlocyba 1187

in the Snow Entomological Collections of the University of Kansas;
one male paratype, Wanakena, New York, August 1, 1917, C. J.
Drake, in the Collection of Dr. D. M. DeLong.

Genus Empoa Fitch
(PI. LXXXVIII)
Empoa Fitch, Ann. Rpt. New York St. Cab. Nat. Hist., vol. 4, 1851, p. 63.

Type, Empoa querci Fitch, 1S51, by subsequent designation of
Van Duzee (Check List of the Hemiptera, 1916, p. 77).

Fore wings. — As in genus Typhlocyba.

Hind wings. — As in genus Typhlocyba (PI. LXXXI, fig. If).

Genital capsule. — Male plate gradually curving dorsad apically,
reduced near middle, gradually tapering to rounded apex, with one
macroseta near outer basal angle, a submarginal row of microsetae
parallel to lateral margin near middle and extending over apical half
of length; pygofer in lateral aspect with dorsal margin nearly hori-
zontal, dorsal angle prominent, apex rounded, ventral angle re-
duced, lacking group of macrosetae dorsad of outer basal angle of
male plate, a few scattered microsetae on disc and a group of
macrosetae on posterior margin.

Internal male genitalia. — Style elongate, slender, gradually re-
duced to an acute apex, curving lateroventrad apically, with nu-
merous setae on ventral surface at basal third, a row of setae extend-
ing along lateral margin to middle, several alveoli on mesal margin
near middle, apex attenuate beyond these; connective elongate,
triangular in ventral aspect; aedeagus without atrial processes, aede-
agal shaft with posterior margin strongly curving dorsad from base,
with three platelike enlargements on anterior margin, with three
pairs of apical processes; aedeagal apodeme a broad dorsally di-
rected arm arising at base of aedeagal shaft.

Female. — With posterior margin of eighth abdominal sternite
slightly sinuate, strongly resembling that found in Typhlocyba
pomaria (PI. LXXXVII, figs. 12a, b).

Head in dorsal aspect narrower than pronotum, longer medially
than next the eye, anterior margin of crown broadly rounded, face
with clypellus conspicuously gibbous; pronotum short and broad,
lateral margins strongly divergent posteriorly; posterior margin
smoothly, shallowly convex, pleural portion broader than ocellocular
area.

All of the species of the genus have black or brown color mark-
ings ranging in extent from only three spots along the cross veins,
to covering nearly the entire dorsum. The ground color is usually

1188 The University Science Bulletin

white to light yellow, but may be bright orange-yellow or even light
pink to rose. The dark color pattern is variable within a species,
and sometimes two or three sharply distinct forms may occur in one
population. Variability of coloration appears to be the result. of the
action of two factors or groups of factors, one causing the gradual
increase in intensity of color observed in maturing adults, the other
controlling the ultimate extent of color pattern, shade of color, and
intensity of color.

As a result of the action of the first factor, specimens of Empoa
gillettei and similarly marked species pass successively through
several color stages in the process of maturing. Extremely teneral
specimens show traces of spots in the apices of the inner three basal
cells, and as the intensity of color increases, an apical band will
gradually become evident. As the apical band deepens in intensity
a pale transverse median band begins to appear and increases in
intensity while the scutellum and dorsum of the abdomen begin to
show signs of dark color. The color of the scutellum continues
deepening until it becomes dark brown or black.

This same sequence is observed in the maturing of Empoa casta
and similarly marked species but usually terminates at the point
when only the apical and median bands have become dark. Speci-
mens of Empoa apicata and similarly colored species have only the
apical band dark so that this process of color change is less notice-
able. Specimens of Empoa vestita, and darkly marked specimens
of vennsta and scripta show a tendency toward a more uniform de-
velopment of color, the apical portion of the wings darkening
slightly earlier than the rest of the wing. In Empoa albicans the
dark color appears to become evident in the scutellum before it can
be seen in the wings. Color in the wings first appears in the apex
of the clavus and gradually spreads to include all of the clavus and
adjacent areas of the inner two or three basal cells.

The second factor, one which influences the final extent of color
pattern and intensity of color, appears to be responsible for the
occurrence of polymorphic species such as E. venusta, E. scripta, E.
albicans, and to a lesser degree E. casta, E. aureotecta, and E. ves-
tita.

Because of the similarity of partly colored specimens of species
of the Gillettei Group to more fully colored specimens of species
of the Casta Group, and because of similarity of partly colored
specimens of species of the Casta Group to specimens of species of
the Apicata Group, species in one group may be confused with
species of another group. In addition to intergroup similarity,

Revision of the Species Typhlocyba 1189

specimens of species within each group are frequently very similar
to each other when not fully mature. Variability of color pattern
within these closely similar species is such that it is impossible to
be certain to which species these marginal specimens belong in the
absence of reliable host records, although usually the majority of
mature specimens are easily distinguishable.

Although it is sometimes difficult to separate some of these species
on the basis of color alone, population behavior observed has been
such as would be expected of distinct species. Each species studied
in the field was found to be closely associated with a certain host
and was usually found on this host even in widely separated locali-
ties. Host relationship studies were made on E. vestita showing
that fourth and fifth instar nymphs when transferred from their
preferred host, Ulmus fulva, and restricted to either U. pumila or
U. americana were able to develop normally. Under natural condi-
tions where branches of U. fulva and U. americana were found
interlaced, the nymphs showed exclusive preference for U. fulva.
The nymphs of E. vestita were easily distinguished from those of E.
elmata, which usually occurs on U. americana, because they are
olive-green to greenish-brown in color and prefer to feed on the
upper surfaces of the leaves, while nymphs of E. elmata and other
species in the genus are white and prefer to feed on the lower sur-
faces of the leaves.

Population size was found to vary greatly for species on different
hosts within the same area and under the same climatic conditions.
Collections of a number of species on the University of Kansas
campus in 1949 shewed that populations of E. vestita and E. casta
were very large, populations of E. acericola, E. apicata, E. caryata,
and E. elmata to be small, and the population of E. querci very
small, while no specimens of E. venusta could be found although
collections were made on its host.

Changes in the size of population from 1949 to 1950 were found
to differ for some of the species studied, while other species popula-
tions appeared to remain constant. Only a slight decrease in
population was noted for E. vestita, while E. casta showed a very
marked decrease in the size of its population. The populations of
E. acericola, E. caryata, E. elmata, E. apicata, and E. querci did
not show any marked degree of change in population.

Population size was found to differ for species collected both
in Douglas County, Kansas, and in Milwaukee County, Wisconsin.
Populations of E. vestita, E. caryata, and E. acericola, were much
larger in Douglas County than in Milwaukee County. Population

1190 The University Science Bulletin

size of E. casta and E. querci were much larger in Milwaukee
County than in Douglas County, while E. apicata was in equal
abundance in both places and E. venusta was only found in Mil-
waukee County.

These studies have led the author to regard the following species
as distinct although population studies could not be made for all of
the species. In some cases where only a few specimens of a distinct
color form were on hand the author has preferred to refrain from
describing these as distinct species until sufficient material can be
seen and adequate studies have been made to show host relation-
ships.

This genus is Nearctic in distribution.

Because of the great similarity of structure of the male genitalia,
only Empoa spinosa and E. albicans have been found to have geni-
talia distinctly different enough for recognition on the basis of
these characteristics. The rest of the species have male genitalia
which may be characterized by the following description:

Genital capsule. — Male pygofer, in lateral aspect, with posterior
margin nearly vertical, dorsal angle rounded apically, ventral angle
reduced, ventral and posterior margins nearly continuous.

Internal male genitalia. — Aedeagal shaft with median plate on
apical half of anterior margin produced as a hook which arises from
the base of plate rather than from middle as seen in Empoa spinosa,
shape of hook variable within a species; median pair of apical shaft
processes of aedeagus strongly curving mesad and crossing near
middle, second pair of apical processes shorter than median pair
and diverging laterodorsad, lateral pair of apical processes nearly
straight, curving laterad and of nearly equal length to median pair.

Key to the Species of Empoa

1. Median plate on anterior margin of aedeagal shaft smoothly

rounded, not forming a hook (Pi. LXXXVIII, fig. lb); fore
wing without spots or apical band of color along cross veins,

albicans p. 1206
Median plate on anterior margin of aedeagal shaft forming a
hook; fore wing with spots or apical band along cross veins ... 2

2. Hook on anterior margin of shaft of aedeagus arising from the

middle of plate (Pi. LXXXVIII, figs. 2b, c); median apical
pair of processes of aedeagus slender, slightly curving mesad

and crossing on outer fourth spinosa p. 1205

Hook on anterior margin of shaft of aedeagus arising from base
of plate (PI. LXXXVIII, figs. 3b, c); median apical pair of
processes of aedeagus broadly flattened, sharply curving mesad
and crossing near middle 3

Revision of the Species Typhlocyba 1191

3. With pronotum brown or black (host Ulmus fulva) vestita p. 1204

With pronotum light colored 4

4. With scutellum brown or black 5

With scutellum neither brown nor black but sometimes deep

yellow 14

5. With only two transverse brown or black bands on fore wing .... 7
With brown or black markings more extensive 6

6. With dark markings on fore wing covering most of basal half of

wing, but not connecting with apical band (host Tilea ameri-

cana) venusta p. 1201

With dark markings on fore wing extending from base of wing to
cross veins, joining apical band with median band (host possi-
bly Salix sp. ) scripta p. 1202

7. Length 3.75-4.0 mm., median band on fore wing one half to one

third as broad as width of wing; apical band of fore wing as
broad or broader than median band ( host Alnus sp. ) ,

latifasciata p. 1203
Length 3.75 mm. or less, median band usually less than one third
as broad as width of wing; apical band narrower than median
band 8

8. Fore wing with brown or black spot absent from first basal cell,

or reduced to a faint trace along cross vein 9

Fore wing with brown or black spot present in first basal cell,
usually two thirds as long as width of cell 10

9. Length 3.0-3.5 mm., ground color yellow, subhyaline to hyaline

( host Carya ovata ) caryata p. 1198

Length 3.25-3.75 mm., ground color milky white, opaque (host

Acer saccharinum) acericola p. 1200

10. Fore wing with apical band of five spots, in inner three basal and

inner two apical cells; median band frequendy not reaching
costal margin; sometimes restricted to clavus; scutellum pale

yellowish brown (host Platanus occidentalis) platana p. 1199

Fore wing with apical band of more than five spots, median band
extending to costal margin; scutellum chestnut-brown, choco-
late-brown, or yellow-brown 11

11. Scutellum pale yellowish brown to yellow, usually light colored;

median band of fore wing narrow, one fourth to one fifth as
broad as fore wing (hosts Quercus alba, Q. macrocarpa, and

other white oak species) casta p. 1196

Scutellum usually dark colored, chestnut to dark chocolate brown;
median band of fore wing usually one third to one fourth as
broad as fore wing 12

12. Fore wing with median band narrow, one fourth to one fifth as

broad as fore wing, margins nearly straight (host Alnus sp.),

gillettei p. 1200
Fore wing with median band broad, one fourth to one third as
wide as fore wing, margins irregular 13

13. Fore wing with median band broad, nearly one third as broad as

fore wing, broadest near middle, narrowing toward margins of

wing (host Tilea americana) venusta p. 1201

1192 The University Science Bulletin

Fore wing with median band less broad, usually only one fourth
as broad as fore wing, usually somewhat irregular in outline,
but of nearly uniform width throughout (host ? Salix sp.),

scripta p. 1202

14. Fore wing with two transverse bands 18

Fore wing with one transverse band near apex 15

15. Length 2.75-3.0 mm. (host Ostrya virginiana) apicata p. 1195

Length 3.25-3.75 mm. (host Quercus or Ulmus) 16

16. Fore wing without brown spot in first basal cell, with only three

brown spots, in second and third basal and first apical cells

(hosts Quercus borealis and Q. palustrus) querci p. 1193

Fore wing with brown spot in first basal cell 17

17. Fore wing with distinct lunate white markings anterior to brown

spots in inner three basal cells and fourth apical cell; ground
color pale rose to bright red orange (host Quercus sp.),

aureotecta p. 1193
Fore wing without distinct white markings; ground color a uni-
form orange-yellow or whitish-hyaline 18

18. Fore wing opaque yellowish-white to orange-yellow, with four to

five brown spots in inner three basal and in inner two apical
cells; without any traces of a median band (host Ulmus ameri-

cana ) elmata p. 1194

Fore wing hyaline to subhyaline, pale yellow to whitish, some-
times with faint traces of a median band, with four to eight
spots along cross veins making up apical band (hosts Quercus
alba, Q. macrocarpa and other species of the white oak group),

casta p. 1196

19. Fore wing with median band nearly uniform in width and parallel

sided; apical band with brown spot usually absent from inner
basal cell, scutellum frequently light colored (host Carua

ovata ) caruata p. 1 198

Fore wing with median band irregular in width, color pale yel-
lowish brown; brown spot present in inner basal cell of fore
wing 20

20. Fore wing with apical band composed of five spots, in inner three

basal and inner two apical cells; median band frequently not
reaching costal margin, sometimes limited to clavus, band one
fourth to one third as wide as wing; ground color opaque milky
white to bright yellow (host Platanus occidentalis) . . . platana p. 1199
Fore wing with apical band usually composed of four to eight
black spots along cross veins; median band very narrow, in
most cases one fourth to one fifth as wide as wing; wing
hyaline to whitish subhyaline, rarely opaque (hosts Quercus
alba, Q. macrocarpa and other species of the white oak group),

casta p. 1196
The Querci Group

The species of this group are characterized by having only an
apical band of spots on the fore wing when fully matured.

Revision of the Species Typhlocyba 1193

Empoa querci Fitch
(PL LXXXVIII, fig. 3)

Empoa querci Fitch, Ann. Rpt. New York State Cab. Nat. Hist., vol. 4, 1851,
p. 63. (Reprint) Rpt. New York State Mus. Nat. Hist., vol. 46, 1893, p. 403.

Typhlocyba querci, Woodworth, Psyche, vol. 5, no. 157-159, May-July, 1889,
pp. 214.

Typhlocyba gillettei var. fitchii McAtee, Proc. U. S. Nat. Mus., vol. 68, art. 18,
1926, p. 25, pis. 1-6.

Typhlocyba querci var. querci, Young, Bull. Univ. Kansas Sci. Bull., vol. 35,
' 1952, p. 103.

Resembling Ejnpoa elmata but distinguished from this species
by lacking brown spot in inner basal cell of fore wing.

Length. — 3.25-3.5 mm.

Color. — Head, pronotum, and scutellum white to bright yellow-
orange in deeply colored specimens; fore wing white to bright yel-
low-orange to shortly anterior to cross veins, with brown spots in
apices of second and third basal and in first apical cells, rarely with
traces of dark color in first basal cell, and slight traces of brown
color at bases of second and fourth apical cells; veins white to yel-
low-orange, hyaline areas anterior to cross veins in inner three basal
cells, apical cells hyaline.

Specimens of this species have been collected on Quercus borealis
and Q. palustrus in Douglas County, Kansas, and from Q. borealis
in Milwaukee, Wisconsin, by the author.

Specimens have been seen from the following localities: Ten-
nessee: Gatlinburg, July 21; Kentucky: Jellico, August 15; North
Carolina: Brevard, June 20; Michigan: Muskeegon, July 21; Wis-
consin: Milwaukee, June 30-July 5; Kansas: Douglas County, June
6, 8, 10, 12, 14; Arkansas: Fayetteville.

Types. — Allotype female of Typhlocyba gillettei var fitchii, Wash-
ington, D. C, here designated neoholotype of querci, and holotype
male of Typhlocyba gillettei var. fitchii, Washington, D. C, July 16,
1885, here designated neoallotype of querci, in the U. S. National
Museum Collection.

Empoa aureotecta Sanders and DeLong

Empoa aureotecta Sanders and DeLong, Ann. Ent. Soc. America, vol. 10, no. 1,

March, 1917, pp. 93-94, pis. 8-9.
Typhlocyba aureotecta, McAtee, Proc. U. S. Nat. Mus., vol. 68, art. 18, 1926

pp. 20, 26, 40, 42.
Typhlocyba gillettei var. russeola McAtee, loc. cit., p. 26, {new synonymy).
Typhlocyba gillettei var. saffrana McAtee, op. cit.
Typhlocyba querci var. russeola, Young, Bull. Univ. Kansas Sci. Bull., vol. 35,

pt. 1, 1952, p. 103.
Typhlocyba querci var. saffrana, Young, op. cit.

Resembles Empoa elmata in color pattern, but differs in having

1194 The University Science Bulletin

fore wing with distinct lunate white marks anterior to brown spots
of apical band.

Length. — 3.25-3.75 mm.

Color. — Head, pronotum, and scutellum yellowish white to orange-
yellow; fore wings with ground color light rose to bright reddish-
orange, extending to shortly basad of cross veins, with brown spots
in apices of three inner basal, and in first and fourth apical cells,
lunate white spots anterior to spots in inner three basal and fourth
apical cells; remaining areas of apical cells fumose hyaline; abdomen
yellow to reddish-orange, without dark markings.

The reported host for this species is Quercus sp.

Specimens have been seen from the following localities: Vir-
ginia: Mountain Lake, July 11; New York: Heart Lake, Essex
County, August 25; Pennsylvania: Northeast, July 4; Ohio: Dela-
ware County, June 26, July 20; Wisconsin: Osceola, July 20; Madi-
son, July 9; Milwaukee, June 28.

Types. — Holotype female, in the collection of D. M. DeLong,
Columbus, Ohio; neoallotype male, Milwaukee, Wisconsin, June 28,
1950, P. J. Christian, and one paraallotype male, Delaware Co.,
Ohio, June 26, 1947, D. J. & J. N. Knull, here designated, in the
Snow Entomological Collections of the University of Kansas; para-
allotype males: two, Mountain Lake, Virginia, July 11, 1938, and
one, July 23, 1940, L. J. and M. J. Milne, in the U. S. National
Museum Collection.

Empoa elmata sp. nov.

Resembling Empoa aureotecta specimens which tend toward
yellow-orange in color, but differs by not having distinct white
lunate spots anterior to apical band of spots on fore wing.

Length. — 3.5 mm.

Color. — Head, pronotum, and scutellum milky white to bright
orange yellow; fore wings whitish-hyaline to opaque milky white,
yellow, or orange yellow, usually with four to five small brown spots
along cross veins in inner three basal and inner two apical cells,
rarely with apical half of fourth and all of third apical cells brown;
apical cells whitish or yellowish hyaline; abdomen with dorsum of
each segment black or brown on basal half, apical half yellow,
lateral sixth entirely yellow on some specimens, venter with basal
three segments black on basal half, remaining portions yellow.

This species has been collected by the author on Uhnus americana
in Milwaukee, Wisconsin, and in Douglas County, Kansas.

Types. — Holotype male and sixteen male paratypes, Douglas

Revision of the Species Typhlocyba 1195

County, Kansas, June 4, 1949, P. J. Christian; allotype female and
one male paratype, Douglas County, Kansas, June 6, 1949, P. J.
Christian; additional paratypes as follows: two males and two fe-
males, Cheboygan County, Michigan, July 18, 1935, H. B. Hunger-
ford; one male, Long Lake, New York, July 28, 1946, L. D. Beamer;
one male, June 8, one male, June 11, one male, June 13, two males,
June 14, 1929, and four males June 3, 1930, Douglas County, Kansas,
P. B. Lawson; one male, June 5, 1948, Wichita, Kansas, P. J. Chris-
tian; one male, Jefferson County, Kansas, June 15, 1950, P. J. Chris-
tian; one male, May 27, six males, May 30, nine males, May 31, one
male, June 1, one male, June 7, five females, June 14, three females,
June 18, two females, June 24, 1949, one male, June 5, one male,
June 6, four males, June 12, and three females June 21, 1950, Doug-
las County, Kansas, P. J. Christian; thirty-seven males, June 27, five
males, June 8, three males, June 29, eight males and one female,
July 1, one female, July 2, five males and two females, July 3, 1950,
Milwaukee, Wisconsin, P. J. Christian; allotype female of Typhlo-
cyba gillettei var. saffrana, Douglas County, Kansas, August, 1923,
W. Robinson, in the Snow Entomological Collections of the Uni-
versity of Kansas; one male paratype June 10, 1947, Louisville,
Kentucky, D. A. Young, in the U. S. National Museum Collection;
one male, Sta. 10, August 11, 1924, a-52, in the collection of Dr. D.
M. DeLong.

Empoa apicata (McAtee)

Typhlocyba gillettei var. apicata McAtee, Proc. U. S. Nat. Mus., vol. 68, art.

18, 1926, pp. 25-26.
Typhlocyba querci var. apicata, Young, Bull. Univ. Kansas Sci. Bull., vol. 35,

pt. 1, 1952, p. 103.

Resembling single-banded specimens of Empoa casta, but easily
distinguished by being smaller in size, and occurring on Ostrya
virginiana rather than on species of Quercus.

Length.— 2.75-3.0 mm.

Color. — Head, pronotum, and scutellum white to light yellow;
fore wings white to deep yellow-orange, with a transverse band of
black spots along cross veins in inner three basal and in first, base
of second, and most of third and fourth apical cells; abdomen with
dorsum of basal three segments black medially, dorsum of pygofer
and anal tube black, venter white to light yellow.

This species has been collected by the author from Ostrya vir-
giniana in Milwaukee, Wisconsin, June 27, 28, July 5, and in Douglas
County, Kansas, May 30, June 1, 7, in association with Ribautiana
parapiscator and R. multispinosa.

1196 The University Science Bulletin

Type. — Holotype male and paratypes in the U. S. National Mu-
seum Collection; paratypes in Illinois State Natural History Survey
Collection; neoallotype female, Douglas County, Kansas, June 7,
1950, P. J. Christian, here designated, in the Snow Entomological
Collections of the University of Kansas.

The type series of McAtee includes a number of specimens of
Empoa casta from Quercus alba, but the holotype is more nearly
like specimens from Ostrya virginiana than from Quercus alba.

The Casta Group

The speries of this group are characterized by usually having only
the apical and median bands dark.

Empoa casta (McAtee) (new combination)

Tuphlocyba gillettei var. casta McAtee, Proc. U. S. Nat. Mus., vol. 68, art. 18,

' 1926, pp. 26-27.
Tuphlocyba querci var. casta, Young, Bull. Univ. Kansas Sci. Bull., vol. 35,

' pt. 1, 1952, p. 103.

Resembling Empoa caryata in color markings, but distinguished
by specimens of the same sex being .25-.5 mm. longer, by having
median band usually noticeably irregular, and occurring on species
of Quercus rather than on Carya.

Length.— 3.25-3.75 mm.

Color. — Head, pronotum, and scutellum usually pale yellowish-
white to light yellow, scutellum sometimes deep yellow to light
yellow-brown; fore wings yellowish-white to whitish-hyaline, with
a narrow light brown median crossband irregular in outline, some-
times indicated by several spots or only by slight traces of brown
at the costal margin, apical band of the same color, variable in the
number of spots from only four in inner three basal and first apical
cells to eight or nine in inner three basal and apical cells; abdomen
with dorsum of each segment black on basal half, venter with basal
half of each segment black in males, females with venter usually
yellow or white. Specimens with only four spots making up the
apical band usually lack the median band also.

This species was found to be very abundant in Milwaukee, Wis-
consin, where the author collected over a thousand specimens from
a wide variety of plants. Many of the specimens taken were seeking
shelter from the wind, but specimens were found to consistently
occur on Quercus macrocarpa and Ulmus americana. Specimens
from these two hosts form a complete, closely intergrading series
extending from typical casta specimens to typical elmata specimens.

In Douglas County, Kansas, specimens were found to be abundant

Revision of the Species Typhlocyba 1197

on Quercus macrocarpa in 1949, and were of the typical casta
pattern, while specimens were rare on Ulmus americana, and were
consistently specimens of the elmata pattern, lacking intergrading
specimens between these two species.

Because of the sharp line of distinction between these forms in
Douglas County, Kansas, the author has chosen to regard them as
distinct species. The intergradation which is found in the specimens
from Milwaukee may be due to a shift of specimens from one host
to another during a part of their development, resulting in speci-
mens which are extremes in variation from the typical forms which
are intermediate between the typical forms.

Specimens from a white oak species in Arizona, and specimens
from Quercus alba from Milwaukee, Wisconsin, are indistinguish-
able from typical specimens of casta found on Quercus macrocarpa,
but show no tendency toward a development of intermediate speci-
mens as are found on Quercus macrocarpa.

A series of specimens from Quercus species? from Franklin, North
Carolina, differs from typical casta specimens in being slightly larger
in size, and in having the median transverse band sharply distinct
with smooth margins and the ground color of opaque light yellow.
This form may eventually prove to be a distinct species.

Specimens have been seen from the following localities: Ontario:
Brockville, August 12; Massachusetts: Woods Hole, July 7; New
York: Canisteo, August 14; Elba, August 23; Ithaca, June 17, 27;
Minetto, August 30; West Nyack, June 11; Sea Cliff; Pennsylvania:
Hummelstown, July 7; Duncannan, June 24; West Virginia: Great
Cacapon, July 4; Washington D. C, June 4; Virginia: Mountain
Lake, July 12, 23; Kentucky: Berea, June 14; Tennessee: Gatlinburg,
June 20, 24, 25; North Carolina: Franklin, August 17; Cherokee, July
19; Raleigh, June 20; Florida: Ocala, April 29; Ohio: Summit County,
June 17; Barberton, August 13; Wooster, July 5, 8; Columbus, Oc-
tober 16; Illinois: Urbana, June 8, 9; Bellwood, June 21; Aroma Park,
July 8; Bell Smith Springs, July 16; Palos Park, June 23; Roseville,
September 1; Wisconsin: Milwaukee, June 26-July 5, 18; Madison,
July 2, 9; Cranmoor, June 21, 27, September 3; Wisconsin Rapids,
June 21; Blue River, July 28; Lake Wingra, July 18; Minnesota: St.
Anthony Park, June 23; Itasca County, July 26, 28, 29; St. Paul,
August 25; Ramsey County, July 20; South Dakota: Custer, August
26; Iowa: Ames, June 16, September 20-22; Muscatine, June 10, 15;
Ledges State Park, June 29; Missouri: Neosho, May 28; Arkansas:
Washington County, June 30; Rodgers, July 10; Louisiana: Madi-

1198 The University Science Bulletin

sonville, June 11; Kansas: Douglas County, June 2-July 20; Wyan-
dotte County, June 23; Leavenworth County, June 28-30; Atchison
County, July 10; Coffey County, June 19; Jefferson County, June 15;
Cherryvale, June 9; Leon, June 20; Colorado: Royal Gorge, July 3;
Garden of the Gods, August 19; Ft. Collins, August 6; Sloss, August
17; Creede, July 22; Arizona: Oak Creek Canyon, August 9; New
Mexico: Sapello, July 25; Ruidoso, October 6.

Empoa caryata sp. nov.

Resembling Empoa casta but differing in having color markings
chestnut brown rather than yellowish-brown, in having dark spot
absent from inner basal cell or reduced to only a slight trace, in
having margins of median band sharply defined and smooth, and in
occurring on species of Carya rather than species of Quercus.

Length. — 3.0-3.5 mm.

Color. — Head and pronotum whitish-yellow to yellow, scutellum
usually of the same color but sometimes chestnut brown; fore wings
with chestnut brown markings forming a narrow median transverse
band one third as broad as long and usually parallel sided, sharply
distinct from ground color, apical band composed of chestnut brown
spots in second and third basal, all of first, base of second, and
all of third apical cells, rarely a trace of dark color in first basal
cell, ground color of fore wing pale white to light yellow; abdomen
white to light yellow, usually without dark markings.

Types. — Holotype male, allotype female, two male and thirty-
seven female paratypes, Douglas County, Kansas, June 11, 1949, P.
J. Christian; additional paratypes: one female, July 5, two males and
one female, July 1, 1939, Clarksville, Tennessee, R. H. Beamer; one
female, August 16, 1946, New Haven, Connecticut, R. H. Beamer;
two males and one female, Tuskahoma, Oklahoma, R. H. Beamer;
ten males and forty-one females, June 11, 1949, Douglas County,
Kansas, R. H. Beamer; one female, Rosedale, Kansas, June 23, 1928,
E. P. Breakey; two males and three females, June 12, thirty-three
females, June 19, one female, June 21, thirteen females, June 24,
two females, June 28, 1949, one male and twenty-two females, June
10, four males and four females, June 12, 1950, Douglas County,
Kansas, P. J. Christian; eight males and fourteen females, June 30,
two males, July 4, 1950, Milwaukee, Wisconsin, P. J. Christian, in
the Snow Entomological Collections of the University of Kansas;
one male and five females, July 9, 1948, Ross and Burks, (on
hickory), N. Chicago, Illinois, in the Collection of the Illinois State
Natural History Survey; one female, June 22, 1918, Mercer County,

Revision of the Species Typhlocyba 1199

Ohio, R. F. Hussey, (on Carya sp. ), in the University of Michigan
Collection; one male and six females, July 2, 1939, Ithaca, New York,
P. A. Readio; one male, Elba, New York, in the Cornell University
Collection.

Empoa platana sp. nov.

Resembling Empoa casta in having scutellum light colored but
differing in having apical band composed of only five brown spots
in inner three basal and inner two apical cells, median band fre-
quently not reaching costal margin, and ground color opaque
milky-white.

Length.— 3.25-3.75 mm.

Color. — Head and pronotum chalky or milky-white to pale yel-
lowish-white, scutellum pale yellowish-brown on disc; fore wing
with ground color chalky white to yellow, with a narrow light brown
median band sometimes extending to costal margin, sometimes re-
stricted to clavus, apical band composed of five light brown spots in
inner three basal cells and in inner two apical cells; cross veins and
apical veins milky white, a border of the same color the width of
vein along veins; apical cells whitish-hyaline with traces of brown
along outer margins of third and fourth apical cells; abdomen
chalky white to light yellow without dark markings.

The host plant for this species is Platanus occidentalis.

Types. — Holotype male, allotype female, Louisville, Kentucky,
June 10, 1947, in the U. S. National Museum Collection; paratypes:
three males and four females, same data as type, in the collection of
Dr. D. A. Young, Jr.; two females and two males, same data as type,
in the Snow Entomological Collections of the University of Kansas;
three females, Columbus, Ohio, October 20, 1920, in the University
of New Hampshire Collection; two females, Lexington, Kentucky,
August 27, 1915, in the U. S. National Museum Collection; one male
and three females, Harrisburg, Pennsylvania, June 13, 1921, in the
collection of Dr. D. M. DeLong.

The Gillettei Group

The species of this group are characterized by usually having the
scutellum dark colored and by having a median and apical band
of the same color. Some of the species in this group show poly-
morphism, having forms with color patterns which are more exten-
sive than the pattern found on specimens of Empoa gillettei.

46—6490

1200 The University Science Bulletin

Empoa gillettei Van Duzee (new combination)

Typhlocyha bifasciata Gillette and Baker, Bull. Colorado Agr. Exp. Sta., no. 31,

Tech. ser. no. 1, 1895, p. 111. (nee. bifasciata Boheman, 1852).
Empoa bifasciata, Van Duzee, Check List of Hemiptera (excepting the Aphid-

idae, Aleurodidae and Coccidae) of America North of Mexico, 1916, p. 77.
Empoa querci var. bifasciata, Sanders and DeLong, Ann. Ent. Soc. America,

vol. 10, no. 1, 1917, p. 93.
Empoa querci var. gillettei Van Duzee, Tech. Bull, California Agr. Exp. Sta.

Ent., vol. 2, 1917, p. 708. (nom. nov. for Typhlocyba bifasciata Gillette and

Baker).
Typhlocyba querci var. gillettei, McAtee, Canadian Ent., vol. 51, no. 8, 1919,

p. 225.
Typhlocyba gillettei var. gillettei, McAtee, Proc. U. S. Nat. Mus., vol. 68, art.

18, 1926, pp. 23-24, 27-28.

Resembling Empoa acericola but differing in having spot in inner
basal cell as large as in other basal cells, median band nearly
parallel margined, ground color deeper yellow, and in occurring on
Alnus sp.

Length. — 3.25-3.5 mm.

Color. — Head white, pronotum white with yellow markings on
anterior half, laterad of disc; scutellum deep orange yellow to light
yellowish brown; fore wings light yellowish white, subhyaline, with
a narrow transverse chestnut-brown median band one third to one
fifth as broad as width of wing, with an apical band of the same
color over cross veins filling apices of inner three basal cells and
most of apical cells, apical two thirds of second apical cell fumose
hyaline and less deeply colored than other apical cells; abdomen
yellow with basal half of dorsum of three basal segments black,
dorsum of male pygofer, and anal tube chestnut brown, venter
yellow or whitish yellow.

A large series of specimens collected on Alnus, Carrizo Creek,
Arizona, June 16, 1950, seen by the author, more nearly resemble
the type of Typhlocyba bifasciata Gillette and Baker, from Colo-
rado, than do specimens of any of the other species in the group.

Types. — Holotype female, type of Typhlocyba bifasciata Gillette
and Baker, in the U. S. National Museum Collection.

Empoa acericola sp. nov.

Resembling Empoa gillettei in color pattern but distinguished by
lacking dark spot in inner basal cell of fore wing.

Length.— 3.25-3.75 mm.

Color. — Head and pronotum milky-white to light yellow; scutel-
lum chestnut-brown; fore wing with a median transverse chestnut-
brown band one third as wide as width of fore wing, of nearly uni-
form width, margins somewhat irregular, an apical band of same
color composed of spots in second and third basal cells, and in all

Revision of the Species Typhlocyba 1201

of first, basal third of second, and apical two thirds of fourth apical
cells; apical two thirds of second and all of third apical cells strongly
fumose-hyaline, areas anterior to bands milky-white to light yellow;
abdomen with dorsum of basal three segments brown on basal half,
dorsum of pygofer and anal tube brown, venter white to yellow.

This species has been collected by the author in Douglas County,
Kansas, and in Milwaukee, Wisconsin, from Acer saccharinum.

Types. — Holotype, allotype, and three female paratypes, June 12,
1950, Douglas County, Kansas, P. J. Christian; additional paratypes:
two males, June 9, one male, June 15, one male, June 19, 1940,
Muscatine, Iowa, and one male, June 12, 1940, Davenport, Iowa,
D. R. Lindsay; two males and one female, June 5, one female, June
28, 1949, Douglas County, Kansas, R. H. Beamer; one male, May 31,
two males and six females, June 6, two females and one male, June
7, one female, June 9, one female, June 15, two females, July 1, one
female, July 15, 1949; seven males, June 3, six males, June 5, twenty-
two males, June 8, three males and seven females, June 10, one male
and seven females, June 21, 1950, Douglas County, Kansas, P. J.
Christian; five males, June 28, five males, June 29, two males and
two females, June 27, 1950, Milwaukee, Wisconsin, P. J. Christian,
in the Snow Entomological Collections of the University of Kansas.

Empoa venusta (McAtee) (new combination)

Typhlocyba gillettei var venusta McAtee, Proc. U. S. Nat. Mus., vol. 68, art. 18,

1926, p. 29.
Typhlocyba querci var. venusta, Young, Bull. Univ. Kansas Sci. Bull., vol. 35,

pt. 1, 1952, p. 103.

Resembling Empoa gillettei, but distinguished by having median
band usually broader at middle than at margins. Dark form dis-
tinguished by having anterior half of fore wing chestnut-brown ex-
cept for a narrow yellow band along costal margin.

Length.— 3.25-3.75 mm.

Color. — Head and pronotum yellowish white to bright yellow;
scutellum chestnut-brown, apex sometimes yellow; fore wing with a
broad, transverse, chestnut-brown median band, its greatest width
half as broad as wing, apical band chestnut-brown, composed of
spots in inner three basal cells and all of first, basal third of second,
all of third and apical two thirds of fourth apical cells, whitish
yellow to yellow on basal fourth of wing and between bands, a
narrow whitish-hyaline band basad of dark spots in basal cells
and in basal third of fourth apical cell, disc of second apical cell
hyaline; dark form specimens differ in having area anterior to
median band also chestnut-brown except for a narrow yellow band

1202 The University Science Bulletin

along costal margin; abdomen yellow with dorsum of basal three
segments chestnut-brown, some specimens with basal half of dorsum
of all segments chestnut-brown.

A large series of specimens of this species has been collected from
Tilea americana, in Milwaukee, Wisconsin, by the author.

Specimens have been seen from the following localities: New
Hampshire: Bretton Woods, August 31; Connecticut: New Haven,
August 16, 18, 20; New York: Albany, June 20; Minetto, August 18,
20, 24, 30; Ontario: Trenton, August 25, 29; Vineland Station, June
16; Ottawa, June 27; Toronto, August 8; Ohio: Sandusky, Cedar
Point, July 11; Barberton, August 13; Milan, September 1; Cleve-
land, September 3; Michigan: Lake Gogebic, August 18; Cheboygan
County, July 30; Minnesota: Itasca County, July 20, 26; Wisconsin:
Amery, August 11, 13; Polk County, July; Wisconsin Rapids, August
11; Milwaukee, June 26-July 7; Soutli Dakota: Brookings, June 17,
27, July 4, August 10, 17, September 15; Big Stone, August 27.

Types. — Holotype male, allotype female, and paratype male in
U. S. National Museum Collection; paratype male in the Snow
Entomological Collections of the University of Kansas.

Empoa scripta (McAtee) (new combination)

Typhlocyba querci var. scripta McAtee, Canadian Ent., vol. 51, no. 8, 1919,

p. 226.
Typhlocyba gillettei var. scripta, McAtee, Proc. U. S. Nat. Mus., vol. 68, art. 18,

1926, p. 28.
TypJiIocyba gillettei subsp. oregonensis Beamer, Canadian Ent., vol. 75, no. 7,

1943, p. 133. (new synonymy).
Typhlocyba querci var. scripta, Young, Bull. Univ. Kansas Sci. Bull., vol. 35,

pt. 1, 1952, p. 103.

Resembling Empoa venusta in color pattern of lightly marked
forms, distinguished in some cases by having median transverse
band of nearly uniform width throughout its length, dark color
pattern easily distinguished by having a longitudinal dark marking
connecting median and apical bands.

Length.— 3.25-3.75 mm.

Color. — Head and pronotum yellowish white to yellow, scutellum
chestnut-brown to dark yellow-brown; fore wing with variable
pattern of chestnut-brown markings, usually with median transverse
band nearly half as broad as fore wing, apical band of spots in
apices of inner three basal cells, in all of first, base of second, and in
apical two thirds of fourth apical cells, ground color yellowish-
white to light yellow; some heavily marked specimens with longi-
tudinal stripe from base of fore wing to cross veins, clavus yellow
to white along basal and apical third of commissural margin, outer

Revision of the Species Typhlocyba 1203

two basal cells yellow to white on basal and apical third; very
heavily marked specimens with only a narrow band of yellow on
basal and apical third of outer basal cell.

Partly teneral specimens are light yellowish brown, but have the
same color pattern. The type series of Typhlocyba gillettei var.
oregonensis is composed entirely of such specimens.

The host for this species is not known definitely, but field notes
made at the time of collection of several short series of specimens,
include only willow and alder. None of the specimens seen from
alder are like specimens of this species so that the more likely host
is some species of willow ( Salix ) .

Specimens have been seen from the following localities: Ten-
nessee: Great Smoky Mountains National Park, June 24, July 20,
September 1; New York: Onteora Mountain, Green County; Keene
Valley, Essex County, August 24; Idaho: Bliss, July 7; Oregon:
Dixie, Grant County, July 8; Haines, Baker County, July 10; British
Columbia: Hope, August 1.

Types. — The holotype female of this species was destroyed by
fire in 1946, in the Nova Scotia Department of Agriculture. A fe-
male specimen, Alum Cave Area, Great Smoky Mountains National
Park, September 1, 1948, Ross and Stannard, agreeing with the
description of the type in every particular, here designated neoholo-
tijpe, in the Illinois State Natural History Survey Collection; a male
specimen, Hope, British Columbia, August 1, 1931, R. H. Beamer,
agreeing with the type description, here designated neoallotype, in
the Snow Entomological Collections of the University of Kansas.

Empoa latifasciata sp. nov.

Resembling Empoa spinosa in size and in color pattern, but easily
distinguished from this species by having hook on anterior margin
of apical half of aedeagal shaft arising from base of median plate
rather than from middle.

Length. — 3.75-4.25 mm.

Color. — Head and pronotum white to bright yellow; scutellum
chestnut-brown; fore wings with a broad transverse median band of
chestnut-brown one half as broad at commissural margin as width
of fore wing, reduced to nearly half this width at costal margin,
posterior margin of band nearly straight but with anterior margin
strongly rounded, apical band of same color covering apices of inner
three basal cells and all of apical cells, apical cells subhyaline with
apical two thirds of second apical cell hyaline, veins in apical band
usually fumose, area anterior to median band and between median

1204 The University Science Bulletin

and apical bands whitish-yellow to orange-yellow with whitish-
hyaline areas anterior to apical band in inner three basal cells.

A large series of specimens has been collected from a species of
Alnus in Sapello, New Mexico.

Types. — Holotype male, allotype female, eight male and twenty-
eight female paratypes, July 24, 1950, Sapello, New Mexico, R. H.
Beamer; three female paratypes, August 1, 1931, Hope, British
Columbia, R. H. Beamer; one female paratype, August 13, 1928,
Harris County Texas, L. D. Beamer, in the Snow Entomological
Collections of the University of Kansas; one female, August 1, 1916,
Maine Agricultural Experiment Station, Orono, Maine; one female,
September 7, 1922, Durham, New Hampshire; two females, Septem-
ber 2, 1928, Fabyans, New Hampshire; 1 female August 19, 1928,
Alton, New Hampshire, in the New Hampshire State University
Collection.

The Vestita Group

This group is characterized by having the pronotum dark colored.
At present it is represented by only one species although three
specimens of one, and single specimens of two other color forms
which may prove to be species of this group have been seen by the
author. Until the hosts for these are known and a sufficient number
of specimens is on hand to be able to tell whether or not these are
distinct species or only extremes of variability of some described
species, the author considers it best for them to remain undescribed.

Empoa vestita (McAtee) (new combination)

Typhlocyba gillettei var. vestita McAtee, Proc. U. S. Nat. Mus., vol. 68, art. 18,

1926, p. 29.
Typhlocyba querci var. vestita McAtee, Young, Bull. Univ. Kansas Sci. Bull.,

vol. 35, pt. 1, 1952, p. 103.

Resembling the most darkly colored specimens of Empoa scripta,
but easily distinguished by having the pronotum and usually the
head dark chocolate-brown or greenish brown.

Length.— 3.25-3.5 mm.

Color. — Head, pronotum, and scutellum greenish- or yellowish-
brown to chocolate-brown; fore wings greenish- to yellowish-brown,
rarely deep chestnut-brown, with areas in apices of inner three basal
cells and areas of apical cells bordering on cross veins deeper brown,
veins on apical third of fore wing lighter than surrounding areas,
costal vein yellow to reddish-brown, deeply colored specimens of
uniform color intensity throughout; face, venter of thorax, and
venter of abdomen yellow; basal half of each abdominal segment
dark brown; male plates yellow to buff.

Revision of the Species Typhlocyba 1205

More than a thousand specimens of this species have been col-
lected from Ulmus fulva in Douglas County, Kansas, where it has
been very abundant. A few specimens were taken from the same
host in Milwaukee, Wisconsin.

Specimens have been seen from the following localities: New
York: Norris, August 15; Pennsylvania: Greensburg; Michigan: Che-
boygan County, July 16; Wisconsin: Amery, August 14; Milwaukee,
June 27-July 4; Illinois: Castle Rock, Grand Detour, July 2; Apple
River Can. State Park, July 11; Kansas: Clay Center, July 2; Douglas
County, June through July.

Types. — Holotype male, allotype female, and paratypes, in the
Snow Entomological Collections of the University of Kansas; para-
types in U. S. National Museum Collection.

The Spinosa Group

The species making up this group is marked like some species of
the Gillettei Group, but is distinguished by having the aedeagal
shaft with hook on anterior margin arising from middle of plate
rather than from basal attachment as in all of the species in preced-
ing groups, and with median pair of apical processes equal in length
to the second pair of apical processes and crossing near the apex
rather than near the middle.

Empoa spinosa ( Beamer ) ( new combination )

(PI. LXXXVIII, fig. 2)

Typhlocyba spinosa Beamer, Canadian Ent., vol. 75, no. 7, 1943, pp. 131-132.

Resembling Empoa latifasciata in external appearance, but with
hook on anterior margin of aedeagal shaft arising from middle of
plate rather than from base.

Length. — 3.75-4.25 mm.

Color. — Head and pronotum light to deep yellow; scutellum
chocolate brown; fore wings with a broad transverse median band
one third to one half as broad as fore wing, apical band chocolate
brown and covering apices of inner three basal cells, basal half of
first, basal fourth of second, all of third, and all of fourth apical
cells; apical half of first and apical three fourths of second apical
cells fumose-hyaline or whitish-hyaline, remaining areas of fore
wing yellowish white to yellow; abdomen with dorsum of basal
three segments entirely chocolate brown, basal half of remaining
segments and dorsal half of pygofer chocolate brown, cross veins
and apical veins white to slightly fumose, whitish-hyaline areas
anterior to apical band in inner three basal cells.

1206 The University Science Bulletin

Genital capsule. — Male pygofer with dorsal angle more pro-
nounced than in Empoa querci, only slightly rounded at apex, ven-
tral angle broadly rounded, not forming a distinct lobe.

Internal male genitalia. — Aedeagus with median pair of apical
processes elongate, crossing at outer fourth, second pair of apical
processes equal to median pair in length, with apices nearly meet-
ing, lateral pair of apical processes exceeding both inner pairs in
length; median plate on dorsal half of anterior margin of aedeagal
shaft produced apically as a large dorsoanteriorly directed hook
arising from middle of anterior margin of plate.

Specimens have been seen from the following localities: Alaska:
Ft. Yukon, July 15; Colorado: Sloss, August 17. The type series was
collected from huckleberry ( Vaccinium sp. ) .

Types. — Holotype male, allotype female, and female paratypes,
in the Snow Entomological Collections of the University of Kansas.

The Albicans Group

This group composed of only a single species at present, is dis-
tinguished from the other species groups by lacking an apical band
of dark colored spots, and by having median plate on anterior
margin of shaft of aedeagus smoothly rounded rather than pro-
duced as a hook.

Empoa albicans Walsh
(PI. LXXXVIII, fig. 1)

Empoa albicans Walsh, The Prairie Farmer, (n. s.) vol. 10, no. 10, September

6, 1862, p. 149.
Typhlocijba albicans, Woodworth, Psyche, vol. 5, no. 157-159, 1889, p. 214.
Typhlocyba ajmba var. pallens McAtee, Proc. U. S. Nat. Mus., vol. 68, art.

18, 1926, p. 22. {new synonymy).
Typhlocyba cymba var. cymba McAtee, Canadian Ent, vol. 50, no. 11, 1918,

pp. 360-361.
Typhlocijba cymba var. grata McAtee, Canadian Ent., vol. 51, no. 8, 1919,

p. 226.
Typhlocyba cymba var. unipnncta McAtee, Proc. U. S. Nat. Mus., vol. 68, art.

18, 1926, p. 23. (nee. Typhlocyba unipuncta Matsumura, 1908).

Resembling Typhlocyba put7nani, T. attenuata, and Edwardsiana
dejecta in size and outward appearance when lacking dark markings
on dorsum, easily distinguished from these by having three pairs of
apical processes on aedeagus.

Length. — 4.0-4.5 mm.

Color. — Head, pronotum, and in some specimens entire dorsum
yellowish-white to light yellow; scutellum dark brown on specimens
with dark markings; fore wings if marked, with dark brown mark-
ings varying from having only a trace of a spot in apex of clavus,
to having entire clavus and most of inner two basal cells dark brown;

Revision of the Species Typhlocyba 1207

abdomen with basal three segments brown on basal half, venter
yellowish white.

Genital capsule. — Male pygofer, in lateral aspect, with ventral
angle forming a slight lobe arising mesad of ventral margin; ventral
margin obliquely curving dorsocaudad and joining posterior margin
near middle.

Internal male genitalia. — Aedeagus with posterior margin of shaft
strongly curving dorsad from base, apical processes directed dorso-
cephalad, median pair nearly parallel to each other, not crossing
apically, second pair short, one third length of median pair, lateral
pair nearly of equal length to median pair; median plate on dorsal
half of anterior margin of shaft of aedeagus evenly rounded, not
produced as a hook.

Four short series of specimens each of which includes three or
four of the described varieties of McAtee have been seen. In addi-
tion to these varieties, intermediate forms also occur and seem
to indicate that these forms are only stages in the maturing of some
adult specimens such as has been seen in other species of the genus.
A single specimen with only the scutellum dark brown has been
seen, which seems to indicate that some specimens may terminate
color formation at various stages before attaining full coloration.
Other specimens entirely lacking dark markings also appear to be
fully matured. These may prove to be only color forms of a single
species with various individuals terminating color formation at
different stages, or they may prove to be specimens of closely re-
lated species. Because of insufficient numbers of specimens and
inability to study this species in the field, the author has not been
able to settle this question.

In the absence of the type specimen of Empoa albicans the iden-
tity of this species has long been held in question. The original
description, reprinted below, is sufficient to indicate that this species
belongs in the Typhlocyba Complex.

"Empoa albicans — New species, whitish. Eyes fuscous. Two or
three of the basal and of the terminal joints of abdomen fuscous at
tip; ovipositor black; elytra subhyaline, at tip a little cloudy; tri-
angular cell peduncled; apex of vein which forms the inner cell not
attaining half the distance to the apex of elytrum; wings hyaline.
Length to tip of wings nearly one-fifth of an inch."

After a consideration of all of the North American species in the
Typhlocyba Complex, only four species could be found which were
whitish, and one sixth of an inch or more in length. Of these four,
only one had dark markings on the dorsal segments of the abdomen,

1208 The University Science Bulletin

Typhlocyba cijmba var. pollens. Since this is the only known species
which fits the description of the type specimen, the author has
placed it in synonymy with Empoa albicans although the type
variety of Typhlocyba cymba has color markings which do not fit
the description of E. albicans.

Specimens have been seen from the following localities: Nova
Scotia: Halifax, August 9; Wolfville; Quebec: Newaygo, July 30,
Ridgland, August 15; Ontario: Toronto, July 9; Vineland Station,
July 1; Maine: Portland, July 9; Mt. Katahdin, August; Mt. Bethel,
July 7; Massachusetts: Cambridge, August 19; New Hampshire:
Wonalancet, July 1; Jackson, September 18; Durham, August 21;
New York: Rock City, July 4; Ithaca, July 26; Gowanda, August 2,
9; Stanford, August 2; Catskill, September 17; Heart Lake, Essex
County, August 23; Onteora Mt. Green County; Michigan: Cran-
berry Lake, August 9; Cheboygan County, July 7, 26; Douglas Lake,
July 13; Agricultural College, June 21; Lake City, July; Wisconsin:
Hazelhurst, July 12.

Types. — Neoholotype female, neoallotype male, Newaygo, Que-
bec, July 30, 1929, Parrish, here designated, in the Snow Entomo-
logical Collections of the University of Kansas.

The following type specimens of varieties of Typhlocyba cymba
McAtee have been destroyed by fire in the Nova Scotia Department
of Agriculture, in 1946: Holotype specimens of varieties cymba,
grata, pollens, and unipuncta; allotype and paratypes of variety
pollens. Paratypes of var. pollens are in the Illinois State Natural
History Survey Collection, and in the U. S. National Museum Col-
lection. Paratypes of var. unipuncta are in the Iowa State College
Collection, and in the U. S. National Museum Collection.

Genus Edwardsiana Jazykov (Zachvatkin)

(Pis. LXXXIX-XCII)

Edwardsiana Jazykov (Zachvatkin), Rev. Russe d'Ent, vol. 23, nos. 3-4, 1929,
pp. 262-265.

Type of the genus, Cicada rosae Linnaeus, by original designa-
tion.

Fore wings. — Inner and outer apical cells short, not attaining
wing apex; second apical cell much broader at apex than at base;
third apical cell petiolate; wing apex smoothly rounded.

Genital capsule. — Male plate in ventral aspect, broad on basal

Revision of the Species Typhlocyba 1209

half, strongly reduced near middle, in lateral aspect gradually
curved dorsad on apical half, apical third slightly enlarged but not
spatulate, a row of short stout setae on outer half submarginally
along dorsal margin, a few scattered setae laterally on apex, and a
single large macroseta on base of male plate near outer basal angle;
pygofer, in lateral aspect, usually broadly rounded, with a pro-
tuberance on dorsocaudal margin sometimes enlarged as a broad
angular arm, ventral angle forming a broadly rounded lobe, rarely
with a short caudally directed hook; a large group of macrosetae
just dorsad of outer basal angle of male plate, numerous setae pos-
terior to these and extending across disc, a patch of macrosetae on
inrolled margin of dorsal angle.

Internal male genitalia. — Style elongate, slender, curving latero-
dorsad, with apex abruptly curved lateroventrad and reduced, ap-
pearing setiform, a row of setae along outer margin, mesal margin
without setae but with a few alveoli on outer third; connective Y-
shaped to triangular, slightly longer than broad; aedeagus without
atrial processes, shaft usually slender, gradually tapering to apex,
rarely with anterior margin inflated, in lateral aspect as a broad thin
plate (rosae), with two pairs of apical processes unbranched, or
with one or both pairs branched; preatrial arm long; aedeagal apo-
deme a slender bar, laterally compressed, nearly two thirds length
of shaft, usually forming an acute angle with the preatrial arm.

Female. — With posterior margin of eighth abdominal sternite
strongly produced medially as an acute lobe as seen in E. rosae.
(PI. LXXXVII, figs. 13a, b).

The head in dorsal aspect narrower than pronotum, longer me-
dially than next the eye, anterior margin of crown broadly rounded;
pronotum short and broad, lateral margins strongly divergent pos-
teriorly, posterior margin smoothly, shallowly convex, pleural por-
tion broader than ocellocular area.

The species are usually pale white or yellow, occasionally with
dark markings.

The genus is holarctic in distribution.

The following key to the North American species of the genus
is based primarily on characteristics of the male genitalia.

Key to the Species of Edwardsiana

1. Aedeagal shaft with two paired processes and one mesal unpaired

process. (Pi. XC, fig. 6) australis p. 1220

Aedeagal shaft with only paired processes 2

2. Aedeagal shaft with one or both pairs of apical processes branch-

ing 3

Aedeagal shaft with apical processes unbranched 10

1210 The University Science Bulletin

3. Aedeagal shaft with both pairs of apical processes branching ... 4
Aedeagal shaft with only mesal pair of processes branching 5

4. With mesal pair of processes branching near base; pygofer only

slightly produced on dorsal angle. ( PI. LXXXIX, fig. 1 ) ,

lethierryi p. 1213
With mesal pair of processes branching near middle; pygofer
strongly produced dorsad as a broad triangular projection.
(PL LXXXIX, fig. 4) bergmani p. 1213

5. With lateral pair of apical processes strongly curved laterocaudad.

(PL XC, fig. 1 ) expanda p. 1215

With lateral pair of apical processes straight, directed cephalad
or dorsad 6

6. Mesal pair of processes fused on basal two thirds, with dorsal

branch arising from middle of fused portion. (PL XCI, fig. 2),

pseudocommissuralis p. 1222
Mesal pair of processes not fused on basal two thirds 7

7. Mesal pair of processes branching near middle. (PL XC, fig. 2),

frustrator p. 1216
Mesal pair of processes branching near base 8

8. Dorsal branch of mesal processes broad, strongly curving dorsad

anteriorly, dorsal margin concave. (PL XC, fig. 3) . . . plebeja p. 1217
Dorsal branch of mesal processes narrow, nearly straight, di-
rected cephalad 9

9. Apices of branches of mesal processes of aedeagus converging

apically, forming nearly a circle in anterior aspect. (PL XC,

fig. 5 ) candidula p. 1219

Apices of branches of mesal processes of aedeagus diverging api-
cally. (PL XC, fig. 4) prunicola p. 1218

10. Anterior margin of aedeagal shaft laterally compressed to a broad

flat plate, lateral apical processes broad, curving strongly dor-
socephalad, diverging laterad. (PL LXXXIX, fig. 2) . . . rosae p. 1211
Anterior margin of aedeagal shaft not compressed, lateral apical
processes slender, directed dorsad, appearing as a continuation
of shaft in lateral aspect 11

11. Pygofer with ventral angle strongly produced as a short acute

caudally directed process 12

Pygofer with ventral angle evenly rounded, or only slightly pro-
duced 13

12. Lateral processes of aedeagus half to one third length of mesal

processes. ( PL XCII, fig. 5) euphrante p. 1228

Lateral processes of aedeagus only slightly shorter than mesal
processes, mesal processes strongly curved dorsad on apical
third. (PL XCII, fig. 2) delongi p. 1226

13. Mesal processes of aedeagus sharply directed ventrad, convex on

anterior margin. (PL XCI, fig. 3) dejecta p. 1222

Mesal processes of aedeagus not directed ventrad, straight, or
slightly concave on dorsal margin 14

14. With dorsal surface of mesal processes of aedeagus concave,

strongly curving dorsad near outer half or third 15

With dorsal surface of mesal processes of aedeagus straight, or
slightly convex 16

Revision of the Species Typhlocyba 1211

15. Lateral processes of aedeagus one third as long as mesal processes;

fore wings entirely white. (PL XCI, fig. 4) dorsti p. 1225

Lateral processes of aedeagus nearly as long as mesal processes;
fore wings with commissural margins black. (PL XCII, fig- 1 ),

nigripennis p. 1225

16. With mesal processes of aedeagus distinctly shorter than lateral

processes 17

With mesal processes of aedeagus equal to or slightly longer than
lateral processes; fore wings with commissural margins and part
of clavus usually black. (PL XCII, fig. 3) commissuralis p. 1223

17. With mesal processes of aedeagus less than half as long as lateral

processes, fused on basal third. (PL XCI, fig. 1) projecta p. 1221

With mesal processes of aedeagus two thirds as long as lateral

processes, not fused on basal third. (PL XCII, fig. 4) ... . ariste p. 1227

Edivardsiana rosae ( Linnaeus )
(PL LXXXIX, fig. 2)

Cicada rosae Linnaeus, Systema Naturae, ed. 10, 1758, p. 439. (Engelmann

Reprint, ed. 10, p. 439, 1894).
Cicada {Tettigonia) rosae, Geoffroy, Histoire d'lnsectes, 1762, p. 428.
Typhlocyba rosae, Herrich-Schafter, Fauna Insectorum Germanicae initia; oder

Deutschlands Insecten gesammelt und herausgegeben von D. G. W. F.

Panzer. Fortgesetzt von G. A. W. Herrich-Schaffer, vol. 124, 1834, p. 10.
Cicadula rosae, Zetterstedt, Insecta Lapponica, 1840, p. 300.
Tettigonia rosae, Harris, Insects Injurious to Vegetation, 1842, p. 199.
Typhlocyba pteridis Dahlbom, Handlingar. Kongliga Svenska Vetenskaps

Akademien, 1850, pp. 179-181.
Anomia rosae, Fieber, Katalogue der europaischen Cicadinen, 1872, p. 15.
Typhlocyba lactea Douglas, Ent. Monthly Mag., ser. 1, vol. 12, 1875, p. 77.
Empoa rosae, Weed, American Garden, July 1889, p. 257.
Edivardsiana rosae, Jazykov ( Zachvatkin ) , Revue Russe, Ent., vol. 23, nos. 3-4,

1929, p. 265.

Resembling E. candidula in external appearance, but easily dis-
tinguished from this species and other North American species of
the genus by having the anterior margin of the aedeagal shaft pro-
duced into a flat plate, appearing greatly enlarged in lateral aspect.

Length. — 3.5-3.75 mm.

Color. — White to pale yellow, without dark markings.

Genital capsule. — Male pygofer, in lateral aspect, with posterior
margin nearly vertical, slightly slanting ventrocaudad; dorsal angle
produced, forming an obtuse angle; ventral angle broadly rounded
projecting slightly beyond dorsal angle.

Internal male genitalia. — Aedeagus, shaft with anterior margin
produced in a flat plate on apical two thirds, appearing greatly
swollen near middle in lateral aspect, with both pairs of unbranched
apical processes directed laterocephalad, lateral pair greatly broad-
ened near middle, gradually tapering from middle to acute apex,
median pair less broad and with apices curving mesad.

A single abnormal male with processes on each side of aedeagal
shaft fused for most of their length, resembling var. manca of Ribaut

1212 The University Science Bulletin

(1936), but differing in having apices of inner pair of processes
forming three short branches, has been seen from Catskill, New
York, taken with normal males of this species, and has been drawn
for comparison with normal specimens. (PI. LXXXIX, figs. 2f, g).

The approved common name for this species is "The Rose Leaf-
hopper" (Muesbeck, 1950, p. 138).

Specimens taken on Malus, Rosa, and Ruhus have been seen.
Specimens seen are from the following localities: Nova Scotia;
Ontario: Trenton, June 11; Massachusetts: Boston, July 27; Lexing-
ton, September 16; New Hampshire: Durham, August 20, 29; Sep-
tember 3, 7, 11, 13, 14, 22; New York: Ithaca, July; Catskill, October
19; Pennsylvania: Northeast, June 7, 8, July 4, October 15; Hazel-
ton, June 12; Ohio: Cleveland, September 1, Columbus, May 31,
Wooster, August 14; Minnesota: Ramsay County, August 22; Wis-
consin: Green Bay, June 10; Milwaukee, June 26-July 5; Bayfield,
September 10; Racine, September 7; Illinois: Urbana, October 1,
November 4, 10; Iowa: Davenport, June 8; Colorado: Ft. Collins,
September 23; Utah: Salt Lake, June 4; Logan, June 11, 26; Sep-
tember 20, 22, October 17; Magna, June 7; Millville, August 2, 3,
September 13; Providence, July 18, September 19, 22, October 4;
Granite, July 16; Riverside, August 12; Provo, September 17; Hooper,
October 14, 16; Fountain, October 22; Richfield, June 9, August 7;
Farmington, September 27; Idaho: Sandpoint, July 3; Shoshone
Basin, July 27; Moscow, October 16; Jerome, June 15; British
Columbia: Oliver, August 6; Vernon, August 5, 23; Vancouver,
August 4; Victoria, October 29, 30; Washington: Buckley, July 6;
Republic, August 5; Wenatchee, June 26; Pullman, July 2, 3; Puyal-
lup, June 29, July 5; Sumner, October 20; Northwest of Yakima,
June 19; Oregon: Yoncalla, July 12; Hood River, August 20;
Gresham, September 12; Tillanook, October 15; Gilliam County,
30 Mile Creek, June 24; Freewater, August 23; Corvallis, October;
LaGrenda, September 11; Jacksonville; Ashland; Waldport, October
22; Rogue River, September; Grant's Pass, September; West of
Junction City, October; North of Cobury, October; South of Peoria,
October; North of Kirby, September; North of Gold Hill, September;
Lancaster, October; Azalea, September, October; Brentwood, Au-
gust 25; Astoria, August 1; Portland, August, October; Salem, Octo-
ber 21; California: San Jacinto Mountains, July 21; Mountain View,
September 21, 24; San Francisco, July 25; Quincy, July 23; Sweden:
Rystad, Frosta, June 20; Germany: Bornkagen, Eichsfeld, October
8; Poland: Warsaw, June.

Determination of this species is based on figures by Ribaut (1936).

Revision of the Species Typhlocyba 1213

Edwardsiana lethierryi (Edwards)
(PL LXXXIX, fig. 1)

Typhlocyba lethierryi Edwards, Ent. Monthly Mag., vol. 17, 1881, p. 224.
Anomia lethierryi, Edwards, Ent. Monthly Mag., vol. 64, 1928, p. 82.
Edwardsiana lethierryi, China, Ent. Monthly Mag., vol. 86, 1950, p. 248.

Resembling E. hippocastana (Edwards) (1888), and E. bergmani
Tullgren, but differs from the former in not having the dorsal branch
of mesal pair of processes on aedeagus rebranching, and differs from
the latter by having the mesal pair of processes branching nearly
from the base.

Length. — 3.5-3.75 mm.

Color. — Dorsum yellowish-white to yellow, without dark mark-
ings.

Genital capsule. — Male py gofer, in lateral aspect, with posterior
margin strongly convex, dorsal protuberance small, ventral angle a
large smoothly rounded lobe.

Internal male genitalia. — Aedeagus with shaft slender, elongate,
curving dorsocephalad, with two pairs of branching apical processes,
lateral pair directed dorsolaterad to fork, dorsal branch shorter than
ventral branch and directed dorsad, ventral branch directed ventro-
laterad; median pair of processes branching near base, dorsolateral
branch one third length of ventral branch.

A large series of male specimens has been collected by the author
from Acer platanoides, in Milwaukee, Wisconsin.

Specimens have been seen from the following localities: Nova
Scotia: Truro, August 17; Massachusetts: Cambridge, October 13,
20, 24; New Hampshire: Durham, September 9; New York: Salem,
June 26, New York City, June 29, 1900; Quebec: St. Anne's, Sep-
tember 2; Wisconsin: Milwaukee, June 26, 27, 29; England: Cam-
bridge, October 17.

Determination of this species is based on figures by Ribaut ( 1936).

Edwardsiana bergmani var. bergmani (Tullgren)

(PL LXXXIX, fig. 4)

Typhlocyba bergmani Tullgren, Ent. Tidskr., vol. 37, 1916, pp. 65-69.
Anomia bergmani, Edwards, Ent. Monthly Mag., vol. 64, 1928, p. 82.
Edwardsiana bergmani, China, Ent. Monthly Mag., vol. 86, 1950 p. 248.

Resembling E. lethierryi externally and in shape of aedeagus, but
distinguished by having mesal pair of apical processes of the aedea-
gus branching near middle rather than near base, and by having a
broad dorsal process on pygofer.

Length. — 3.75 mm.

1214 The University Science Bulletin

Color. — Dorsum light yellow without dark markings.

Genital capsule. — Male py gofer in lateral aspect, with posterior
margin rounded on lower two thirds, deeply excavated on dorsal
third, dorsal angle directed dorsad as a broad triangular, apically
rounded process, ventral angle forming a slight lobe.

Internal male genitalia. — Aedeagus, with both pairs of apical
shaft processes branching, lateral pair with branches of nearly equal
length, directed dorsad, apices slightly curving toward each other;
mesal pair of processes branching near middle, both branches
slightly concave on dorsal surface, directed cephalad, ventral
branches with apices converging mesad; aedeagal shaft strongly
convex on posterior margin, recurved near apex, anterior margin
more strongly recurved than posterior, width nearly uniform from
base to apex in lateral aspect.

Only a single specimen of this variety from Arvidajaur, Sweden,
September 20, in the collection of Dr. Frej Ossiannilsson of Uppsala,
Sweden, has been seen. Illustrations for this variety have been
made from this specimen to show how it differs from the following
variety.

Edwardsiana bergmani var. ariadne ( McAtee )
(PI. LXXXIX, fig. 3)

Ti/phlocuba ariadne McAtee, Proc. U. S. Nat. Mus., vol. 68, art. 18, 1926,
pp. 14-15.

Length. — 4.0-4.25 mm. (Alaska and Montana specimens).
3.75 mm. ( Oregon specimens ) .

3.50 mm. ( Michigan, Pennsylvania and Maine speci-
mens ) .

Color. — Light yellow to light yellow-orange, disc of pronotum and
fore wings deeper yellow.

Genital capsule. — Male pygofer, in lateral aspect, with posterior
margin rounded on lower two thirds, dorsal angle as in type variety,
ventral lobe more pronounced on specimens from Michigan, Penn-
sylvania, and Maine.

Internal male genitalia. — Aedeagus similar to that of type variety
but with shaft strongly convex on both anterior and posterior mar-
gins, broader on apical two thirds and apex not recurved, length of
apical processes proportionately longer on specimens from Alaska,
Montana, and Oregon, lateral processes directed strongly laterad,
length of processes on specimens from Michigan, Pennsylvania, and
Maine nearly of the same proportion as those of the specimen of the

Revision of the Species Typhlocyba 1215

type variety but with shaft distinctly characteristic of that of the
holotype of variety ariadne, from Vancouver, British Columbia.

The type of variability shown by this species is like that found
between Edwardsiana australis Froggatt and E. crataegi (Ed-
wards), still regarded as distinct species. Because of the scarcity
of male specimens, the limited number of localities from which
specimens have been collected, and the wide distribution of the
species, the author has considered it best to regard ariadne as a
variety of bergmani until conclusive evidence can be seen for re-
garding it as a distinct species.

Specimens have been seen from the following localities: Alaska:
Ft. Yukon, July 15; Matanuska, July 12, 19, 21, 27, 28, August 17,
September 24, 27, 28, October 2, 5, 6; Northeast of Anchorage,
August 4; Sleese Highway, September 4; Anchorage, July 20; North
of Paxton's Lodge 27-52 miles, July 28; British Columbia: Hope,
August 1; Vancouver; Kalso, July 13, 17, June 22; Oregon: Mount
Hood, July 3; Bonnville, July 4; Tillamook, October 15; Washington
Shelton, July 24; Sumner, August 20, September 18; Kalama, July 21
Conway, July 28; Quinault, July 26; Montana: Haugan, August 9
Michigan: Au Train, August 21; Pennsylvania: Hartstown Bog,
September 13; Maine: Dixfield, October 20.

Types. — Holotype male, allotype female, and paratypes in the
U. S. National Museum Collection; paratypes in the collection of
W. Downes.

Edwardsiana expanda (DeLong and Johnson) (new combination)

(PI. XC, fig. 1)

Typhlocyba expanda DeLong and Johnson, Ent. News, vol. 47, no. 4, April,
1936, p. 104.

Resembling E. bergmani var. ariadne in external appearance, but
distinguished by having aedeagus with lateral pair of apical proc-
, esses unbranched and strongly recurved laterodorsad.

Length. — 4.0-4.5 mm.

Color. — White to light yellow without dark markings.

Genital capsule. — Male pygofer in lateral aspect very similar to
that of E. lethierryi, but with ventral lobe more pronounced.

Internal male genitalia. — Aedeagus with lateral pair of apical
processes unbranched, elongate, longer than mesal pair, closely ap-
pressed to shaft on basal fourth and directed dorsad, apical three
fourths sharply recurved laterodorsad; mesal pair of processes di-
rected laterocephalad branching near middle, dorsal branch slightly

47—6490

1216 The University Science Bulletin

longer than ventral branch, branches diverging from base and
slightly curving toward each other at apices in lateral aspect.

Two abnormal specimens of this species which have the lateral
processes shorter than usual and not recurved, have been seen, and
superficially resemble the illustration of Eclwardsiana kemneri
Ossiannilsson, but are distinguished by having the mesal pair of
processes like those of typical specimens of E. expanda. (see Plate
XC,fig. Id).

Specimens have been seen from the following localities: Alaska:
Ft. Yukon, July 15; North of Paxton's Lodge 27-52 miles, July 28;
Northwest Territories: Good Hope, August 23; Oregon: Hood,
August 22; California: Siskiyou Mountains, June 24; Colorado:
Cameron Pass, June 24; Estes Park, August 21, 25.

Types. — Holotype male, allotype female, and female paratype, in
the collection of Dr. D. M. DeLong, Columbus, Ohio.

Eclwardsiana frnstrator (Edwards)

(PI. XC, fig. 2)

Typhlocyba frustrator Edwards, Ent. Monthly Mag., vol. 44, 1908, p. 84.
Anomia frnstrator, Edwards, Ent. Monthly Mag., vol. 64, April, 1928, p. 82.
Typhlocyba solearis Ribaut, Bull. Soc. Hist. Nat. Toulouse, vol. 61, pt. 1, 1931,

p. 339.
Edwardsiana frustrator, China, Ent. Monthly Mag., vol. 86, 1950, p. 248.

Resembling E. prunicola in the shape of the aedeagus, but dis-
tinguished from it by having mesal pair of apical processes with
branches as long as unbranched portion, and aedeagal shaft more
slender and nearly straight.

Length. — 3.5 mm.

Color. — Pale yellowish white to light yellow without dark mark-
ings.

Genital capsule. — Male pygofer, in lateral aspect, with dorsal
angle smoothly rounded, ventral angle slightly angular and rounded
apically.

Internal male genitalia. — Aedeagus with shaft slender, elongate,
nearly straight, directed dorsad, lateral pair of apical processes
unbranched, slightly concave on dorsal surface, directed cephalad
for half the length of mesal pair, mesal pair of apical processes
branching near middle, directed cephalad, with dorsal branch con-
vex on dorsal surface and with ventral branch slightly concave on
dorsal surface, branches less divergent than in E. expanda.

This species has not previously been recorded from North America
and has recently been introduced from Europe, where it has been
found on "nut bushes" and on "sycamore" according to China

Revision of the Species Typhlocyba 1217

(1943). Specimens seen have been taken on Corylus. Specimens
have been seen from the following localities: Washington: Shelton,
July 24, 1949; Oregon: McMinnville, August 22, 1946; Salem, Sep-
tember 29, 1947; Portland, October, 1948; Wheatlend, October 15,
1949; Orleans, October, 1949; West of Junction City, October, 1949;
North of Coburg, October, 1949; Clackomas, October, 1950;
Sweden: Linkoping, July 7, 1935 on Acer platanus.

Determination of this species is made on the basis of figures of
the male genitalia of Typhlocyba solearis by Ribaut (1931b, 1936),
reprinted by China ( 1943 ) as representative of the type of T. frus-
trator Edwards.

Edwardsiana plebeja (Edwards)
(PI. XC, fig. 3)

Typhlocyba plebeja Edwards, Ent. Monthly Mag., vol. 50, 1914, pp. 168, 172.
Anomia plebeja, Edwards, Ent. Monthly Mag., vol. 64, April, 1928, p. 82.
Typhlocyba divergens Ribaut, Bull. Soc. Hist. Nat. Toulouse, vol. 61, pt. 1,

1931, pp. 339-341.
Edwardsiana plebeja, China, Ent. Monthly Mag., vol. 86, 1950, p. 248.

Resembling E. pranicola in external appearance and in male geni-
talia, but differing in having dorsal branch of mesal pair of aedea-
gal processes curving strongly dorsad, and by having margins of
pygofer forming nearly a semicircle in lateral aspect.

Length.— 3.25-3.75 mm.

Color. — Yellowish-white to light yellow without dark markings.

Genital capsule. — Male pygofer, in lateral aspect, with posterior
margin outlining a semicircle from base of anal tube to middle of
ventral lobe, dorsal and ventral angles reduced.

Internal male genitalia. — Aedeagus with shaft evenly convex on
posterior margin from base of preatrial shaft to apex, gradually
reduced toward apex, lateral pair of apical processes unbranched,
short, one third as long as mesal pair and directed dorsolaterad,
mesal pair of apical processes branching at basal fourth, dorsal
margins of both branches concave, dorsal branch broad in lateral
aspect and strongly curving dorsad, ventral branch shorter and di-
rected ventrocephalad.

A large series of specimens including nymphs labeled Ulmus
americana, and several specimens labeled "Weeping Elm" have been
seen.

Specimens have been seen from the following localities: Massa-
chusetts: Wood's Hole, July 9, 1900; Cambridge; New York: Sep-
tember 1, 1919 (a series of 18 specimens made paratypes of T. com-
missuralis var. munda McAtee); Washington D. C: May 24, 26;

1218 The University Science Bulletin

Virginia: Arlington, October 2. Mrs. J. N. Knull informs me she
has specimens of this species taken in Ohio.

This species has not previously been recorded as occurring in
North America, although the first specimens were collected in 1900.

Determination of specimens is based on the figures of the male
genitalia of Typhlocijba divergens Ribaut, (1931b, 1936), reprinted
by China (1943) as representative of the type of T. plebeja Ed-
wards.

Edwardsiana prunicola ( Edwards )

(PI. XC, fig. 4)

Typhlocijba prunicola Edwards, Ent. Monthly Mag., vol. 50, 1914, pp. 168, 172.
Anomia prunicola, Edwards, Ent. Monthly Mag., vol. 64, April, 1928, p. 82.
Typhlocijba barbata Ribaut, Bull. Soc. Hist. Nat. Toulouse, vol. 61, pt. 1, 1931,

pp. 338-339.
Typhlocijba pruni DeLong and Davidson, Ohio J. Sci., vol. 34, no. 3, 1934,

pp. 161-162, figs. 1-3.
Typhlocijba pruniella DeLong, Ohio J. Sci., vol. 44, no. 6, 1944, p. 272. (n. n.

for pruni DeLong and Davidson, nee. pruni Edwards).
Edwardsiana prunicola, China, Ent. Monthly Mag., vol. 86, 1950, p. 248.

Resembling E. plebeja in outward appearance and in male geni-
talia, but distinguished by having median pair of apical processes
of aedeagus branching near base, dorsal branch slender and longer
than ventral branch, nearly straight and directed cephalad, and
by having dorsal and ventral angles of pygofer produced.

Length.— 3.75-4.0 mm.

Color. — Uniform light yellowish white to yellow without dark
markings.

Genital capsule. — Male pygofer, in lateral aspect, with dorsal
angle forming a rounded protuberance, ventral angle slightly pro-
duced forming nearly a right angle.

Internal male genitalia. — Aedeagus with lateral pair of apical
processes unbranched and curving dorsocephalad and diverging
laterally, mesal pair of apical processes branching near basal third,
dorsal branch nearly straight, directed cephalad forming nearly a
right angle with the axis of shaft, ventral branch directed ventrad
toward base of aedeagal apodeme in lateral aspect, apices of both
pairs of processes diverging laterad.

Specimens have been seen from the following localities: New
York: Albany, August 17, 1900; Utah: Provo, September 17; Logan,
June 26; Bountiful, June 20, July 2; Idaho: Boise, July 11; Parma,
September 3; Washington: Puyallup, July 5; Yakima, August; Ore-
gon: The Dalles; Milton, June 8; Dufur; Ashland; South of Peoria;
California: Berkeley, May 15; San Jose, April 22; Sweden: August
14, "Primus"; Rumania: Bucharest; Bohemia.

Revision of the Species Typhlocyba 1219

Determination of specimens is based on the figures of the male
genitalia printed by China (1943). Specimens have been seen
which have the pygofer rounded on ventral angle in addition to
specimens similar to each of the drawings of the pygofer shown by
China. The shape of the aedeagus is identical in specimens having
variation in the shape of posterior margin of pygofer which seems to
indicate that these forms are only degrees of variation within the
species and insufficient as evidence for making prunicola and bar-
bata distinct species.

Edwardsiana candidula (Kirschbaum)
(PI. XC, fig. 5)

Tijphlocyba candidula Kirschbaum, Jharb., Ver. Nat. Nassu., vol. 21-22, 1868,

p. 185.
Anomia candidula, Edwards, Ent. Monthly Mag., vol. 64, April, 1928, p. 82.
Edwardsiana candidula, China, Ent. Monthly Mag., vol. 86, 1950, p. 248.

Resembling E. expanda in outward appearance and in shape of
male genitalia, but distinguished by having lateral pair of processes
on aedeagus directed dorsolaterad, not sharply recurved, apices of
both branches of mesal pair of processes curving mesad.

Length. — 3.5-3.75 mm.

Color. — Uniform milky-white without dark markings.

Genital capsule. — Male pygofer, in lateral aspect, with dorsal
and ventral angles reduced, forming a single caudally directed,
apically rounded lobe.

Internal male genitalia. — Aedeagus with shaft strongly convex
on posterior margin, smoothly curving and evenly tapered to apex,
lateral pair of apical processes unbranched, slender, directed dorso-
laterad and appearing as a continuation of the shaft in lateral aspect,
length equal to that of the mesal processes, mesal pair of processes
of the aedeagus branching near base, branches slender, slightly di-
verging and curving laterocephalad outlining nearly a complete
circle in cephalic aspect.

A large series of specimens of this species has been collected by
the author from Populus alba in Milwaukee, Wisconsin. Specimens
have been seen from the following localities: Wisconsin: Mil-
waukee, June 26-July 5; Minnesota: St. Paul, July 8, 11; Oregon:
North of Coburg, October; Germany: Buchen, September 30, 1934,
determined by W. Wagner.

Determination of specimens of this species has been based on
figures of the male genitalia according to Ribaut ( 1936 ) .

1220 The University Science Bulletin

Edwardsiana australis (Froggatt) (new combination)

(PI. XC, fig. 6)

Empoasca australis Froggatt, Agr. Gaz. New South Wales, vol. 29, 1918,

pp. 568-571.
Typhlocyba australis, Myers, Proc. Linn. Soc. New South Wales, vol. 46, 1921,

pp. 473-474.
Typhlocyba froggatti (Baker), Philippine J. Sei., vol. 27, 1925, p. 537. (nom.

nov. for australis Froggatt, nee. australis Walsh 1862.)
Typhloctjba Xanthippe McAtee, Proc. U. S. Nat. Mus., vol. 68, art. 18, 1926,

p. 14.
Empoa ( Typhloci/ba ) malini DeLong, J. Econ. Ent., vol. 19, no. 3, 1926,

pp. 469-470.
Ttiphlocyba oxyacanthae Ribaut, Bull. Soc. Hist. Nat. Toulouse, vol. 61, pt. 1,

" 1931, pp. 334-335.
Edwardsiana froggatti, China, Ent. Monthly Mag., vol. 86, no. 1035, (4th ser.

vol. XI, no. 128) 1950, pp. 243-248.

This species is not Erythroneura australis Walsh (1862), the
synonymy of which is given on p. 1142.

Resembling E. crataegi in external appearance, and in male geni-
talia, but differing in having lateral pair of apical processes on
aedeagus shorter, distinguished from other species by having a
median unpaired process as well as two pairs of unpaired processes.

Length. — 3.5-3.75 mm.

Color. — Dorsum yellow to orange-yellow; fore wings with fumose
areas along commissural suture covering outer half of clavus, apices
of inner three basal cells, and all of hyaline apical cells.

Genital capsule. — Male pygofer, in lateral aspect, with posterior
margin nearly straight, obliquely slanting ventrocaudad, dorsal
posterior margin excavated and with a short projection arising on
dorsal margin.

Internal male genitalia. — Aedeagus with shaft broad on basal
half, apical half slender, with two pairs of unbranched apical proc-
esses and a single unpaired median process; lateral pair of apical
processes shorter than mesal pair, curving laterodorsad and caudad;
mesal pair of processes strongly concave on dorsal surface and curv-
ing dorsolaterad anteriorly; unpaired process arising between mesal
pair of processes and strongly convex on dorsal margin curving
dorsocephalad.

Several short series of specimens of this species were collected
by the author in Milwaukee, Wisconsin, from: Crataegus sp., Mains
sp., Aesculus hippocastanum, and Primus sp.

Specimens have been seen from the following localities: Massa-
chusetts: Lexington, June 19; Cambridge; Boston, June 23; Con-
necticut: New Haven, August 18, 20; Mystic, August 19; New York:
New York City, June 22; Ontario: Vineland Station, September 2,

Revision of the Species Typhlocyba 1221

8, 13; Ohio: Wooster, July 5; Columbus, July; Minnesota: St. Paul,
August 20; Wisconsin: Milwaukee, June 26-July 5; Utah: Castle
Dale, August 27; British Columbia: Vancouver, August 4; Washing-
ton: Puyallup, July 5; Oregon: Portland, October; Neskowin, Oc-
tober 3; Clachomas, October 13; California: Sequel, June 7; Chile:
Santiago Quinta Normal, December, 1946; El Pino Angol, February
27, March 24, 1932, on apple.

Although Mrs. J. N. Knull has informed me (in correspondence)
that she has seen a specimen of E. crataegi among specimens of E.
froggatti from Wooster, Ohio, the author has not seen any speci-
mens of crataegi in the samples of this species examined. It is
quite possible that this species is only a variety of crataegi, and if
this is the case additional specimens of crataegi should be found in
North America.

Determination of specimens of this species are based on figures
of the male genitalia printed by W. E. China (1943).

Edwardsiana projecta sp. nov.
(PI. XCI, fig. 1)

Resembling E. candidula in outward appearance, but easily dis-
tinguished by having mesal pair of processes on aedeagus one fourth
the length of lateral processes, and fused on basal half, appearing
as single branching median process.

Length. — 3.5-4.0 mm.

Color. — Uniform milky-white without dark markings.

Genital capsule. — Male pygofer, in lateral aspect, with posterior
margin nearly straight, slanting obliquely ventrocaudad, dorsal
angle more prominent than in preceding species, ventral angle
forming a broad, caudally protruding lobe.

Internal male genitalia. — Aedeagus very similar to that of E.
fraterada (Edwards) but with mesal pair of apical processes un-
branched; shaft proportionately short and stout as compared to
other species, scarcely longer than aedeagal apodeme, apical half
straight and only slightly reduced in diameter toward apex; lateral
processes slender, widely diverging laterodorsad, posterior margin
convex in lateral aspect; mesal processes straight, directed cephalo-
dorsad, one fourth the length of lateral processes, fused on basal
third and appearing as a single branched process, apical two thirds
as long as broad.

Types. — Holotype male and three male paratypes, LaVeta Pass,
Colorado, July 28, 1937, L. D. Tuthill, in the Iowa State College
Collection.

1222 The University Science Bulletin

Edwardsiana pseudocommissuralis sp. nov.
(PI. XCI, fig. 2)

Resembling E. commissuralis in color markings, but with mesal
pair of apical processes on aedeagus branching.

Length.— 3.75-4.0 mm.

Color. — Head and pronotum yellow to yellow-orange; scutellum
brown medially, outer angles yellow; fore wings yellow to yellow-
orange, with black longitudinal band along commissural margin on
outer half of clavus extending to cross veins, apices of basal and
all of apical cells fumose-hyaline.

Genital capsule. — Male pygofer, in lateral aspect, with posterior
margin strongly rounded, similar to pygofer in E. plebeja.

Internal male genitalia. — Aedeagus with shaft slender, slightly
enlarged on basal half, posterior margin slightly angled near middle,
with two pairs of apical processes; lateral pair of apical processes
unbranched, exceeding mesal pair in length, directed dorsolaterad,
sinuate on apical two thirds; mesal pair of apical processes fused on
basal two thirds, with a pair of short straight branches arising at
basal third, directed dorsocephalad parallel to each other, apical
two thirds strongly curving laterodorsad anteriorly, apical third
divided into two branches which diverge laterad.

Types. — Holotype male, Trinity Bay, Quebec, August 20, 1929,
W. J. Brown, in the Canadian National Collection; allotype female
and female paratype, Crawford Notch, New Hampshire, August 21,
1934, P. W. Oman, in the U. S. National Museum Collection.

Edwardsiana dejecta sp. nov.
(PI. XCI, fig. 3)

Resembling Typhloajba putmani in external appearance, but
easily distinguished by the absence of atrial processes on the aedea-
gus, and by having mesal pair of apical processes unbranched and
directed sharply ventrad from base.

Length. — 4.0-4.5 mm.

Color. — Uniformly light yellowish-white to yellow without dark
markings.

Genital capmle. — Male pygofer, in lateral aspect, with posterior
margin nearly straight, dorsal angle slightly produced, ventral angle
slightly angular.

Internal male genitalia. — Aedeagus with shaft elongate, slightly
enlarged on basal two thirds, reduced on outer third and slightly
sinuate, with two pairs of unbranched apical processes on aedeagus,

Revision of the Species Typhlocyba 1223

lateral pair half as long as mesal pair, nearly straight, diverging
laterodorsad anteriorly, mesal pair of apical processes sharply
directed ventrad from base, broadly joined basally by a transverse
plate, only slightly diverging and curving toward base of shaft.

A large series of specimens has been collected by the author
from Cornus stolonifera, in Milwaukee, Wisconsin, in association
with Typhlocyba putmani, a species almost identical with it in ex-
ternal appearance.

Types. — Holotype male and fifty-nine paratype males, Milwaukee,
Wisconsin, July 1, 1950; allotype female and twenty-three female
paratypes, July 3, 1950, Milwaukee, Wisconsin; one male paratype
June 26, 1950, Milwaukee, Wisconsin; seven male paratypes, July 4,
1950, Milwaukee, Wisconsin, P. J. Christian; types in the Snow En-
tomological Collections of the University of Kansas; one male para-
type, July 13, 1940, and two female paratypes, July 16, 1940, Vine-
land Station, Ontario, W. L. Putman, in the Canadian National Col-
lection.

The Commissuralis Complex

The following six species compose a group which on the basis of
male genitalia appear to be more closely related to each other than
to other species in the genus.

Edwardsiana commissuralis (Stal) (new combination)

(PL XCI, fig. 3)

Tijphloct/ba commissuralis Stal, Stett. Ent. Zeit., vol. 19, 1858, p. 196. (Re-
print by Gillette, 1898, Proc. U. S. Nat. Mus., vol. 20, pp. 769~-770. )

Kybos commissuralis, Fieber, Verh. Zool.-bot. Ges. Wien., vol. 16, 1866, p. 508.

Empoasca commissuralis, Ashmead, Harriman Alaskan Exped., vol. 8, 1904,
p. 135.

Empoa commissuralis, Van Duzee, Trans. San Diego Soc. Nat. Hist., vol. 2,
no. 1, 1914, p. 57.

Typhlocvba commissuralis var. munda McAtee, Proc. U. S. Nat. Mus., vol. 68,
art. 18, 1926, p. 12.

Resembling E. pseudocommissuralis in color markings, but with
apical processes of aedeagus unbranched, lateral pair as long as or
slightly shorter than median pair, pygofer with posterior margin
oblique and with ventral angle smoothly rounded.

Length. — 3.0-4.5 mm.

Color. — Head, pronotum, and lighter portions of fore wing light
yellowish-white to orange-yellow; scutellum and commissural
margin of fore wings usually dark black, width of black on wing
varies from a narrow line to two-thirds the width of clavus, apical
cells fumose, heavily pigmented specimens with dorsal surface of
head near base, pronotum, apical half of clavus, and basal fourth

1224 The University Science Bulletin

of costal margin dark black, lighter portions deep yellow and apical
cells strongly fumose; abdomen yellow, margin of each segment of
dorsum black, pygofer with heavily sclerotized portions black.

Genital capsule. — Male pygofer, in lateral aspect, with posterior
margin evenly rounded from base of anal tube to ventral angle,
slightly produced as a broad ventrally directed lobe.

Internal male genitalia. — Aedeagus slender, gradually tapering to
apex, curving cephalad from base, with two pairs of unbranched
apical processes, processes slightly sinuate, the lateral pair directed
dorsocephalad diverging laterad and continuing in the same axis
as the shaft in lateral aspect, mesal pair of processes directed ven-
trocephalad and diverging laterad.

The host plants known for this species are Alnus viridis and A.
rhombifolia. A series of specimens is also on hand collected on
"flowering dogwood".

Specimens have been seen from the following localities: Alaska:
Fairbanks, July 24, 28; Valdez, August 24; Matanuska, July 21;
Northwest Territories: Aklavik, July 18; British Columbia: Van-
couver, August 4, 8, September 3; Saanich District, September 12;
Gorgon Head, September 12; Victoria, November 11; Washington:
Buckley, July 6; Sumner, August 20; Puyallup, July 6; Shelton, July
24; Oregon: Rogue River, August 16; Lee's Camp, September 23;
The Dalles, October 8; Scapoose, October 16; Mount Hood, July 3,
8; McMinnville, August 15, 19, 21; Tillamook, October 15; Pioneer,
October 15; Booneville, July 4; Bend, July 2, 3; Dixie, July 8; North
Powder, July 13; Pendleton, July 14; California: North of Cassel 5
miles, July 15; Muir Woods, September 5; Tahoe South Alpine
Creek, July 15; Cucamonga, December 12; Eureka, July 23; Bray,
June 30; Alameda County, July 19; Lemon Cove, July 24; Straw-
berry, August 8; Stinson Beach, August 15; Donner Lake, June 6;
Colfax, June 23; Wyoming: Laramie, August 16; Idaho: Alturas
Lake, July 19; Utah: Emery, August 16; Colorado: Pagosa Springs;
Sloss, August 17; Durango, July 17.

Types. — Holotype male, Sitka, Alaska, (present location of type
unknown ) .

Variety munda, holotype, allotype and paratypes, in the U. S.
National Museum Collection. One paratype male, Vancouver,
British Columbia, is a specimen of E. bergmani var. ariadne, and
a series of paratypes from New York are specimens of E. plebeja.
Some of the specimens of this variety are only teneral specimens,
but others appear to be fully matured yet lack the dark markings
of the type variety.

Revision of the Species Typhlocyba 1225

Specimens bearing a black commissural line, and occurring in
Northeastern United States and Southeastern Canada, will probably
be specimens of E. pseudocommissuralis. Specimens of Ossiannils-
sonola phryne may also be confused with it, but differ by having
black parenthesis-shaped markings on the scutellum and pronotum,
and a black spot in each of inner three basal cells.

Edwardsiana dorsti ( Ossiannilsson ) (new combination)

(PI. XCI, fig. 4)

Typhlocyba dorsti Ossiannilsson, Ent. Tidskr., vol. 57, no. 1, 1936, pp. 10-11,

figs. i-3.
Typhlocyba enascora DeLong and Johnson, Ent. News, vol. 47, no. 4, April,

1936, pp. 103-104, figs. 1-6. {new synonymy).

Resembling E. euphrante in outward appearance and in propor-
tion of apical processes of aedeagus, but differs in having posterior
margin of pygofer smoothly rounded and without hook on ventral
angle, and in having median pair of apical processes parallel to each
other and strongly curving dorsocephalad on apical half.

Length. — 3.5 mm.

Color. — Light yellowish-white to light yellow without dark mark-
ings.

Genital capsule. — Male pygofer, in lateral aspect, with posterior
margin forming a broadly rounded lobe, ventral angle slightly
pointed.

Internal male genitalia. — Aedeagus with shaft slender, gradually
reduced toward apex, with two pairs of unbranched apical processes,
lateral pair straight and directed dorsolaterad, less than half as long
as mesal pair, mesal pair of apical processes strongly curving dorso-
cephalad on apical half, fused at base and continuing nearly parallel
to each other to apex.

Specimens from the following localities have been seen: Idaho:
Idaho Falls, July 27; Oregon: Dufur; Nevada: Ormsby, July.

Types. — Holotype male, in the collection of Dr. Frej Ossiannils-
son, Uppsala, Sweden. The aedeagus of the holotype has been lost
since its description, but the original drawings are sufficiently clear
to recognize it as the same species as the holotype of Typhlocyba
enascora. The description of dorsti was mailed on February 2, 1936,
while that of enascora was mailed April 12, 1936.

Edwardsiana nigripennis sp. nov.

(PI. XCII, fig. 1)

Resembling E. commissuralis externally by having commissural
vein black on basal two thirds and in its large size, but distinguished

1226 The University Science Bulletin

by having median pair of apical processes strongly curving dorso-
cephalad and in having all of aedeagus but the apical third of shaft
black rather than brown.

Length. — 4.0 mm.

Color. — Light yellowish-white to yellow with commissural vein
brown on basal two thirds.

Genital capsule. — Male py gofer, in lateral aspect, with posterior
margin nearly straight, obliquely slanting ventrocaudad, dorsal
angle slightly produced to a point, ventral angle slightly produced
caudally to a point.

Internal male genitalia. — Aedeagus with shaft slender, with two
pairs of unbranched apical processes of nearly equal length, lateral
pair of processes slender, directed dorsolaterad in the same axis as
the shaft, median pair of processes strongly curving dorsolaterad
from base, preatrial arm and basal two thirds of shaft black, apical
third dark brown as in other species.

Types. — Holotype male, allotype female, four male and three fe-
male paratypes, Stinson Beach, California, August 15, 1938, R. H.
Beamer, in the Snow Entomological Collections of the University
of Kansas.

Edwardsiana delongi sp. nov.
(PL XCII, fig. 2)

Resembling E. nigripennis in shape of the aedeagus, and E. eu-
phrante in the shape of the pygofer, distinguished from the first by
having hook on ventral angle of pygofer, and from the second by
having apical processes of aedeagus of nearly equal length.

Length.— 3.25-3.5 mm.

Color. — Light yellowish-white to yellow without dark markings.

Genital capsule. — Male pygofer, in lateral aspect, with posterior
margin nearly straight and slanting ventrocaudad, dorsal angle
slightly produced, ventral angle with a short caudally directed hook.

Internal male genitalia, — Aedeagus with two pairs of unbranched
apical processes, lateral pair directed strongly laterad on basal half
and curving dorsocephalad on apical half, median pair of processes
strongly curving dorsocephalad and only slightly distant from each
other throughout their length.

Types. — Holotype male, allotype female, and one male paratype,
Bray, California, June 30, 1935, R. H. Beamer, and one male para-
type, Dixie, Oregon, July 8, 1939, R. H. Beamer, in the Snow Ento-
mological Collections of the University of Kansas; twenty-four fe-

Revision of the Species Typhlocyba 1227

male and sixteen male paratypes, Bray, California, June 30, 1935,
P. W. Oman, and eight female and four male paratypes, Kalama,
Washington, July 4, 1935, P. W. Oman, in the U. S. National Mu-
seum Collection; one male, Grant's Pass, Oregon, September, 1949,
S. M. Dietz, in the Oregon State Department of Agriculture Collec-
tion, Salem, Oregon; two males and one female paratype, south of
San Francisco, California, October 2, 1915, O. E. Essig, on Wild
Blackberry, in the University of California Collection; some para-
types from Washington, Oregon, and S. of San Francisco, differ in
having the ventral hook of the pygofer reduced.

Edwardsiana ariste (McAtee) (new combination)

(PI. XCII, fig. 4)

Typhlocyba ariste McAtee, Proc. U. S. Nat. Mus., vol. 68, art. 18, 1926, p. 13.

Resembling E. australis (Froggatt) in size and color markings,
and E. commissuralis in form of male genitalia, differing by having
only two pairs of apical processes on aedeagus and in having the
lateral pair exceeding the median pair by one fourth.

Length. — 3.5-3.75 mm.

Color. — Yellowish-white to slightly greenish-yellow or yellow,
with fumose spots in apices of basal and all of apical cells, slightly
fumose along commissural margin.

Genital capsule. — Male pygofer, in lateral aspect, with posterior
margin convex, with ventral angle slightly produced caudad as a
small apically rounded lobe.

Internal male genitalia. — Aedeagus slender on distal half, with
two pairs of unbranched apical processes, lateral pair of processes
one-fourth longer than median pair, slender and widely diverging
laterodorsad appearing as a continuation of apex of shaft in lateral
aspect, median pair of processes slender and straight, diverging
laterocephalad.

This species occurs on Ribes. Specimens have been seen from
the following localities: New Mexico: Pecos, September 1; Colo-
rado: Creede, July 2, 3, 16; Estes Park, July 18, August 25; Ft.
Collins, July 19.

Types. — Holotype male, in the U. S. National Museum Collec-
tion; female, Creede, Colorado, July 16, 1938, L. D. Tuthill, col-
lected with males of this species, here designated neoallotype, in
the Iowa State College Collection.

1228 The University Science Bulletin

Edwardsiana euphrante (McAtee) (new combination)

(Pi. XCII, fig. 5)

TyphJocyba euphrante McAtee, Proc. U. S. Nat. Mus., vol. 68, art. 18, 1926,
pp. 12-13.

Resembling E. ariste in external appearance and in form of the
male genitalia, but distinguished by having lateral pair of apical
processes of aedeagus one third length of median pair and by having
ventral hook on pygofer produced to an acute apex.

Length. — 3.5 mm.

Color. — Yellowish-white to dark yellow, fore wings sometimes
light greenish-yellow, apical and apices of basal cells slightly fumose.

Genital capsule. — Male pygofer, in lateral aspect, with posterior
margin slightly convex, slanting ventrocaudad, dorsal angle not pro-
duced, ventral angle with a short, acute caudally directed hook.

Internal male genitalia. — Aedeagus with shaft broad on basal
two thirds and reduced on apical third, with two pairs of un-
branched apical processes, lateral pair straight, one third length of
median pair, directed laterodorsad, median pair straight, directed
laterocephalad.

Specimens labeled "wild currant" have been seen. No other host
plant records are known.

Specimens have been seen from the following localities: British
Columbia: Vancouver, August 8; Hope, August 1; Washington:
South of Cheney, July 9; Kalama, July 4; Oregon: South of Bend,
July 2; Bend, July 2; Galena Summit, June 3; California: Clear Lake,
July 21; Idaho: Craters of the Moon, June 29; Utah: Fish Lake,
August 16; Soldier, August 13, (type series); Soldier Summit, Au-
gust 13; Colorado: Kremmling, July 17. A single male specimen
from 2 miles west of St. Louis, Missouri, is very similar to speci-
mens of this species, but lacks the ventral hook on the pygofer.

Types. — Holotype male, allotype female, and two male paratypes,
in the U. S. National Museum.

Discussion of the Commissuralis Complex

Although the preceding six species have been described as
sharply distinct species, intermediate forms have been found to
occur. Two large series of specimens have been seen which are
morphologically intermediate between E. ariste and E. euphrante.
E. ariste is characterized as having the lateral pair of apical proc^
esses of the aedeagus one-fourth longer than the median pair, and
having the posterior margin of the pygofer strongly convex with a
short rounded apical lobe. E. euphrante is characterized as having

Revision of the Species Typhlocyba 1229

the lateral pair of apical processes of the aedeagus one third as
long as the median pair, and having the posterior margin of the
pygofer strongly convex with the ventral angle strongly produced
caudally as a short acute hook.

Two large series of specimens have been seen which are inter-
mediate between these two species. One of these series taken on
Ribes inebrians, near Flagstaff, Arizona, is close to E. ariste, and
has both pairs of apical processes of the aedeagus of nearly equal
length, the posterior margin of the pygofer is convex in the ma-
jority of specimens with the ventral angle only slightly lobed and
apically rounded. Some specimens which appear to be subnormal
have the posterior margin of the pygofer straight and nearly ver-
tical. The second series of specimens is intermediate between the
preceding series and E. euphrante, having the lateral pair of apical
processes of the aedeagus only one half as long as the median pair,
and having the ventral lobe of the pygofer only slightly lobed and
apically rounded. This series was taken on Ribes sp. near Hot
Sulfur Springs, Colorado.

E. dorsti is another species which does not differ from the pre-
ceding two species and their intermediates in external appearance,
but differs only in that the pygofer is broadly rounded on the ventral
lobe, and that the apical processes of the aedeagus which are of the
same proportionate length as those found on E. euphrante have
the median pair strongly curving dorsocephalad on the apical third.
The author has seen only four specimens of this species so that the
interspecific variability is not known.

Some specimens of a fourth species, E. commisstiralis are also
very similar to the preceding species in size, and differ from the
series of specimens from Flagstaff, Arizona, only in the presence of
the commissural black stripe on the fore wings, and in the shape of
the ventral lobe of the pygofer which is broadly rounded apically
and does not form a small ventral lobe. The great difference in the
size of specimens from different localities ( 3.0-4.5 mm. ) and in the
relative amounts of black color composing the wing markings indi-
cates the degree of external variability which one of these species
may show, although on the basis of the shape of the male genitalia
this constitutes only one species.

The larger specimens of E. commissuralis are similar to E. nigri-
pennis in size and the narrow dark line along the commissural
margin in nigripennis strengthens the similarity although the geni-
talia are easily recognized as those of distinctly different species.

The sixth species in this complex, E. delongi, strongly resembles

1230 The University Science Bulletin

E. euphrante in the shape of the posterior margin of the pygofer, but
its greater length, and the size, relative lengths, and shape of the
aedeagal processes set it off from euphrante as a distinct species.

A number of abnormal specimens, some taken singly, others in
association with normal specimens have been seen. Some of these
showed evidences of the presence of internal parasites while in others
no evidences of the parasitism could be seen. Many of these speci-
mens were intermediate in structure of the aedeagus and pygofer
so that the species to which they belonged could not be determined.
Although some of the specimens in the two large series of inter-
mediate forms mentioned in relation to E. ariste and E. euphrante
were apparently abnormal in structure, sufficient numbers of ap-
parently normal specimens were on hand to indicate that something
other than parasitism must account for so many specimens having
the same degree of modification.

Whether this complex of species represents the extremes of varia-
tion of one or two species, or whether each of these is a distinct
species cannot be conclusively decided from the evidence that is on
hand. Only by the collection of numerous samples of specimens
throughout the range of these species will it be possible to determine
the relationships between these species.

GLOSSARY OF TERMS

aedeagus. — The sclerotized intromittent organ of the male, com-
posed of base, dorsal apodeme, preatrial arm, shaft, and atrial

processes, though some of these may be absent.
aedeagal apodeme. — A dorsal arm arising from the base of the

aedeagus, for muscle attachment.
alveolus. — A socket in the cuticula for a seta, vestigial on inner

margin of styles.
apical processes. — Processes arising from the shaft of the aedeagus

at, or distad of outer third.
atrial processes. — Paired processes arising from the base of the

aedeagus ventrad or laterad of the atrium, rarely fused with shaft

and thus becoming shaft processes.
atrium. — A large opening in the base of aedeagus through which

the gonoduct passes to the base of the shaft.
apical cells. — Areas of the wings distad of cross veins, marked off

by apical veins, numbered outward from mesal or caudal margin

of the wing.
basal cells. — Areas of the wings basad of cross veins, marked off by

longitudinal veins, numbered from the mesal, or caudal margin,

outward.

Revision of the Species Typhlocyba 1231

base of aedeagus. — Part of aedeagus to which shaft, atrial processes,
aedeagal apodeme, and preatrial arm are attached, sometimes
massive, sometimes reduced and indistinct.

brachial cell. — First basal cell of fore wing, lying along claval
suture.

connective. — A ventrally located plate with styles attached later-
ally, and aedeagus attached posteriorly.

dorsal angle. — Portion of pygofer bordering on dorsal half of caudal
margin, and caudal half of dorsal margin.

genital capsule. — Ninth abdominal segment, composed of pygofer
and plates.

gonopore. — Distal opening of gonoduct.

internal male genitalia. — Composed of aedeagus, connective, and
styles.

macrosetae. — Large setae with diameter several times that of micro-
setae, with large lumen, arising from alveolus.

microsetae. — Filiform setae, short or greatly elongate, lumen small
or indistinct, arising from alveolus.

plate. — An elongate lobe arising from the venter of the genital cap-
sule of the male, functioning as a clasping organ.

preatrial arm. — Ventral arm of aedeagus extending cephalad from
ventral margin of atrium.

process. — A cuticular projection sometimes elongate, slender, heavily
sclerotized ( on aedeagus ) ; sometimes a modified lobe of cuticula
terminating as a hook or spine (on pygofer, style, or plate).

pygofer. — Side of genital capsule.

pygofer hooks. — Projections from dorsal, posterior, or ventral mar-
gins of pygofer, strongly sclerotized or not, apex acutely pointed.

shaft. — Usually a sclerotized tubular portion of aedeagus through
which the gonoduct passes; in the genus Ossiannilssonola reduced
to a membranous structure between atrial processes.

shaft processes. — Processes arising from shaft of aedeagus, usually
paired.

spine. — A heavily sclerotized, acute, cuticular appendage, not ar-
ticulate.

style. — One of the paired clasping organs comprising the inner male
genitalia.

Typhlocyba Complex. — Those genera of leaf hoppers which have
two apical cells open in the hind wing, and have the fore wing
with first apical cell not reaching the apex of the wing, and third
apical cell usually triangular and stalked.

48—6490

1232 The University Science Bulletin

ventral angle. — Portion of pygofer bordering on ventral half of
caudal margin, and caudal half of ventral margin.

ventral lobe. — Lobe of pygofer just dorsad of outer basal angle of
male plate.

LITERATURE CITED
Andison, Harry.

1950. The Rramble Leafhopper, Typhlocyha tenerrima H.-S. (Homoptera:
Cicadellidae ) , A Destructive European Insect New to the Pacific
Northwest. Canadian Ent, vol. 82, no. 3, 1950, pp. 68-70.
Baker, Charles Fuller.

1925. Nomenclatorial Notes on the Jassoidea, IV. Philippine J. Sci., vol.
27, no. 6, 1925, p. 537.

Beamer, Raymond Hill.

1943. Some New Species of Typhlocyha ( Homoptera, Cicadellidae ) Ca-
nadian Ent., vol. 75, no. 7, 1943, pp. 131-133.
China, W. E.

1943. New and Little-known Species of British Typhlocybidae (Homop-
tera) with Keys to the Genera Typhlocyha, Erythroneura, Dikra-
neura, Notus, Empoasca and Alehra. Trans. Soc. Brit. Ent., vol.
8, pt. 4, 1943, pp. 111-153, figs. 1-14.

1950. A Check List of the British Hemiptera-Homoptera Auchenorhyncha.
Ent. Mon. Mag., vol. 86, no. 1035, 1950, pp. 243-248. 4th ser.,
vol. 11, no. 128.

DeLong, Dwight Moore.

1926. A New and Important Species of Leafhopper Injuring Apple in
Ohio. J. Econ. Ent., vol. 19, no. 3, 1926, pp. 469-470, fig. 23.

1944. Nomenclatorial Notes on Cicadellidae. Ohio J. Sci., vol. 44, no. 6,

1944, p. 272.

DeLong, D. M., and Davidson, Ralph H.

1934. A New Species of Typhlocyha (Homoptera Cicadellidae) Injurious
to Prune in the Pacific Northwest. Ohio J. Sci., vol. 34, no. 3,
1934, pp. 161-162, figs. 1-3.

DeLong, D. M., and Johnson, Dorothy M.

1936. Six New Species of Typhlocyha from the United States. ( Homop-
tera: Cicadellidae). Ent. News, vol. 47, no. 4, 1936. pp. 101-
104, figs. 1-6.

Dlablola, Jiri.

1950. A Revision of Leaf-hoppers in Melichar's Collection. Acta Musei

Moraviae, vol. 35, 1950, pp. 1-16.
1946. Description De Deux Nouvelles Especes Et Plusieurs Remarques

Sur Les Especes Peu Connues D'Europe Centrale (Homopt., Au-

chenorry.) Acta Ent. Mus. Prague, vol. 24, 1946, no. 314, pp. 97-

106.

Edwards, James.

1888. Descriptions of Four New Species of Typhlocyha. Ent. Monthly
Mag., vol. 25, 1888-1889, pp. 157-158, figs. a-1.

Revision of the Species Typhlocyba 1233

1928. On the Genus Anomia Fieber, with Descriptions of Two New
Species. Ent. Monthly Mag., vol. 64, 1928, pp. 79-85, figs. a-f.
3rd ser., vol. 14.

Fabricius, Johann Christian.

1794. Entomologia systematica emendata et auct. Secundum classes,
ordines, genera, species adjectis synonimis, locis, observationibus,
descriptionibus. vol. 4, 1794, pp. 1-472.

Froggatt, Walter Wilson.

1918. The Apple-leaf Jassid. (Empoasca australis). Agr. Gaz. New
South Wales, vol. 24, pp. 341-344, 3 figs.

Gillette, C. P., and Baker, C. F.

1895. A Preliminary list of the Hemiptera of Colorado. Bull. Colorado
Agr. Exp. Sta., no. 31, Tech. ser. no. 1, pp. 1-137, figs.
Jacobi, Arnold.

1941. Die Zikadenfauna der Kleinen Sundainseln. Nach der Expedi-
tionsausbeute von R. Rensch. Zool. Jahr. Syst. vol. 74, 1941, pp.
277-321, 1 pi.

Knull, Dorothy M. Johnson.

1944. Descriptions of Six Typhlocybas from the United States (Homop-
tera: Cicadellidae). Ohio J. Sci., vol. 44, no. 6, 1944, pp. 269-272,
1 pi.

1945. Eleven New Leafhoppers with Notes on Others. (Homoptera:
Cicadellidae). Ohio J. Sci., vol. 45, no. 3, 1945, pp. 103-110,
pis. 1-2.

LlNNAVUORI, R.

1949. Hemipterologisches aus Finnland. Ann. Ent. Fennici. vol. 15,
no. 4, 1949, pp. 145-146, figs.

Matsumura, Shonen.

1908. Neue Cicadinen aus Europa und Mittelmeergebiet. J. Coll. Sci.
Tokyo, vol. 23, no. 6, pp. 1-46, figs. 1-8.

1931. A Revision of the Palaearctic and Oriental Typhlocybid-genera with
Descriptions of New Species and New Genera. Ins. Matsumurana,
vol. 6, no. 2, 1931, pp. 55-91, pis. 2-3, text figs. 1-6.

1932. A Revision of the Palaearctic and Oriental Typhlocybid-genera with
Descriptions of New Species and New Genera. Ins. Matsumurana,
vol. 6, no. 3, 1932, pp. 93-120.

Medler, John T.

1942. The Leafhoppers of Minnesota (Homoptera: Cicadellidae). Min-
nesota Agr. Exp. Sta. Tech. Bull. no. 155, 1942, pp. 1-196, pis. 9.

McAtee, Waldo Lee.

1926. Revision of the American Leaf Hoppers of the Jassid Genus Typh-
locyba. Proc. U. S. Nat. Mus., vol. 68, art. 18, 1926, pp. 1-47,
pis. 1-6.

MUESBECK, C. F. W.

1950. Common Names of Insects Approved by the American Association
of Economic Entomologists. J. Econ. Ent., vol. 43, no. 1, 1950,
pp. 117-138.

1234 The University Science Bulletin

Myers, John G.

1921. The Australian Apple Leafhopper (Typhlocyha australia Frogg.).
Proc. Linn. Soc. New South Wales, vol. 46, 1921, pp. 473-474,
figs. 1-4.

Oman, Paul W.

1949a. A Leafhopper Injurious to Cultivated Prune in the Western United

States. J. Econ. Ent., vol. 41, no. 6, 1949, p. 983.
1949b. The Nearctic Leafhoppers — A Generic Classification and Check
List. Mem. Ent. Soc. Washington, no. 3, 1949, pp. 1-253, pis. 44.

Osborn, Herbert.

1928. The Leafhoppers of Ohio. Ohio State Univ. Bull., vol. 32, no. 27,
1928, pp. 199-374, figs. 1-111. Ohio Biological Survey Bulletin 14
(vol. 3, no. 4).

Ossiannilsson, Frej.

1935. Bidrag till kannedomen om Sveriges Homoptera Cicadina. II Ent.
Tidskr., vol. 56, nos. 3-4, 1935, pp. 129-137, figs. 1-19.

1936. A New North American Species of Typhlocyha (Homoptera, Cica-
dellidae), Typhlocyha dorsti n. sp. Ent. Tidskr., vol. 57, no. 1,
1936, pp. 10-11, figs. 1-3.

Ribaut, Henri.

1931a. Les especes francaises du groupe Typhlocyha ulmi (L.) (Homop-

tera-Typhlocybidae ) . Bull. Soc. Hist. Nat. Toulouse, vol. 59, pt. 1,

1931, pp. 280-291, figs. 1-29.
1931b. Especies nouvelles du groupe Typhlocyha rosae (L. ) (Homoptera-

Typhlocybidae ) . Bull. Soc. Hist. Nat. Toulouse, vol. 59, pt. 1,

1931, pp. 333-342, figs. 1-38.
1936. Homopteres Auchenorhynques I. ( Typhlocybidae ) . Faune de

France, vol. 31, 1936, pp. 1-231, figs. 1-629.

Ross, Herbert H., and DeLong, D. M.

1949. New Eastern Species and a Newly Reported Introduction of Typh-
locyha (Homoptera, Cicadellidae). Ohio J. Sci., vol. 49, no. 3,
1949, pp. 115-118, pis. 2.

Slosson, Annie Trumbull.

1906. Additional List of Insects Taken in Alpine Region of Mt. Wash-
ington. Ent. News, vol. 17, no. 9, 1906, pp. 323-326.

Wagner, W.

1935. Die Zikaden der Nordmark und Nordwest-Deutschlands. Verh.
ver. Naturw. Heimatforsch., vol. 24, no. 1, 1935, pp. 1-44, figs. 1-5.

Walsh, Benjamin Dann.

1862. Fire Blight. Two New Foes of the Apple and Pear. Prairie Farmer
(n. s.), vol. 10, no. 10, 1862, pp. 147-149, figs. 1-7.

WlGGLESWORTH, V. B.

1947. The Principles of Insect Physiology, edn. 3, 1947, pp. 1-434, figs.
316.
Woodworth, Charles William.

1889. North American Typhlocybini. Psyche, vol. 5, no. 157-159, May-
July, 1889, pp. 211-214.

Revision of the Species Typhlocyba 1235

Young, David A., Jr.

1952. A Reclassification of Western Hemisphere Typhlocybinae (Homop-
tera, Cicadellidae). Bull. Univ. Kansas Sci. Bull., vol. 35, pt. 1,
1952, pp. 1-217.

Zakhvatkin, Alexis A. (Jazykov).

1929. Description d'un nouvelle espece du genre Edwardsiana Jaz. 1929
(Homoptera, Eupterygidae ) des environs de Moscou. Rev. Russe
d'Ent., vol. 23, nos. 3-4, 1929, pp. 262-265, figs. 1-5.
1947. Homoptera — Cicadina from north-western Persia. I. [In Russian
with English Summary.] Rev. Ent. URSS, vol. 28, nos. 3-4, ( 1945),
1947, pp. 106-115, 22 figs.

1236

The University Science Bulletin

INDEX TO GENERA AND SPECIES

PAGE

acericola, Ernpoa 1200

alabamaensis, Typhlocyba 1177

albicans, Empoa 1206

andromache, Typhlocyba 1176

Anomia 1160

antigone, Ossiannilssonola 1139

apicata, Empoa 1195

appendiculata, Ossiannilssonola, 1150

ariadne, Edwardsiana 1214

ariste, Edwardsiana 1227

arsinoe, Typhlocyba 1166

alhene, Typhlocyba 1166

attenuata, Typhlocyba 1172

aureotecta, Empoa 1193

australis, Edwardsiana 1220

australis, Ossiannilssonola 1142

bangsoni, Ossiannilssonola .... 1145

barbata, Typhlocyba 1218

beamed, Henribautia 1118

berenice, Ossiannilssonola 1136

bergmani, Edwardsiana 1213

bifasciata, Typhlocyba, Bohe-

man 1145

bifasciata, Typhlocyba, Gillette

& Baker 1200

callosa, Typhlocyba 1133

candidula, Edwardsiana 1219

caryata, Empoa 1198

Cassiopeia, Typhlocyba 1181

casta, Empoa 1196

clara, Typhlocyba 1174

clymene, Ossiannilssonola 1140

commissuralis, Edwardsiana . . . 1223

crassa, Typhlocyba 1182

crataegi, Edwardsiana 1220

cruciata, Ribantiana 1123

cymba, Typhlocyba 1206

danae, Ossiannilssonola 1149

debilis, Typhlocyba 1123

dejecta, Edwardsiana 1222

delongi, Edwardsiana 1226

discincta, Typhlocyba 1139

diver gens, Edwardsiana 1217

dorsti, Edwardsiana 1225

duplicata, Typhlocyba, Jacobi. . 1107
duplicata, Ossiannilssonola, ( Mc-

Atee) 1146

PAGE

Edwardsiana 1208

elmata, Empoa 1194

Empoa 1187

enascora, Typhlocyba 1225

escana, Typhlocyba 1174

cuphrante, Edwardsiana 1228

cunjdice, Typhlocyba 1149

expanda, Edwardsiana 1215

fitchii, Typhlocyba 1193

flavomarginata, Ossiannilssonola, 1157

foliosa, Ribantiana 1126

fratercula, Edwardsiana 1208

froggatti, Typhlocyba 1220

frustrator, Edwardsiana 1216

gillettei, Empoa 1200

grata, Typhlocyba 1206

Henribautia 1115

hermione, Ossiannilssonola .... 1137

hinei, Ossiannilssonola 1144

hippocastani, Edwardsiana .... 1108

hockingensis, Typhlocyba 1169

hubbardi, Henribautia 1117

inflata, Typhlocyba 1186

jacobii, Typhlocyba 1107

kemneri, Edwardsiana 1216

knulli, Ossiannilssonola 1154

lactea, Typhlocyba 1211

lancifer, Typhlocyba 1184

latifasciata, Empoa 1203

lethierryi, Edwardsiana 1213

luculla, Ribantiana 1127

malini, Typhlocyba 1220

manca, Edwardsiana 1211

Mcateeana 1130

mcateei, Ossiannilssonola 1148

media, Typhlocya 1157

medleri, Typhlocyba 1169

melite, Typhlocyba 1176

misella, Typhlocyba 1122

modesta, Typhlocijba 1167

multispinosa, Ribantiana 1125

munda, Typhlocijba 1223

nicarete, Ossiannilssonola 1143

nigricephala, Henribautia 1116

nigripennis, Edwardsiana 1225

niobe, Typhlocyba 1183

Revision of the Species Typhlocyba

1237

PAGE

nitidula, Typhlocyba 1145

ocellata, Eupteryx 1121

oneka, Typhlocyba 1165

oregonensis, Typhlocyba 1202

Ossiannilssonola 1132

oxyacanthae, Typhlocyba 1220

pollens, Typhlocyba 1206

parapiscator, Ribautiana 1123

persephone, Typhlocyba 1184

phryne, Ossiannilssonola 1152

piscator, Ribautiana 1124

platana, Etnpoa 1199

plcbeja, Edioardsiana 1217

pomaria, Typhlocyba 1170

projecta, Edwardsiana 1221

pruni, Typhlocyba 1218

pnmicola, Edioardsiana 1218

pruniella, Typhlocyba 1218

pseudocom missuralis, Edwardsi-
ana 1222

pteridis, Typhlocyba 1211

putmani, Typhlocyba 1180

quadrata, Ossiannilssonola .... 1155

querci, Empoa 1193

que reus, Typhlocyba 1163

Ribautiana 1119

rosae, Edioardsiana 1211

rossi, Ossiannilssonola 1153

rubi, Typhlocyba 1122

rubriocellata, Typhlocyba 1174

russeola, Empoa 1193

PAGE

saffrana, Typhlocyba 1193

scalaris, Typhlocyba 1123

sciotoensis, Ribautiana 1128

scorta, Tij)>hlocyba 1157

scripta, Empoa 1202

sellata, Typhlocyba 1150

serrula, Ossiannilssonola 1147

sexnotata, Mcatecana 1131

shawneeana, Typhlocyba 1178

solearis, Edwardsiana 1216

sollisa, Typhlocyba 1182

spinosa, Empoa 1205

subpulchra, Ossiannilssonola . . . 1152

surcula, Typhlocyba 1175

surda, Ribautiana 1129

tenerrima, Ribautiana 1122

tortosa, Typhlocyba 1185

transviridis, Typhlocyba 1179

troza, Ossiannilssonola 1156

tunicarubra, Ossiannilssonola . . 1141

Typhlocyba 1160

idmi, Ribautiana 1121

unca, Ribautiana 1129

unipuncta, Typhlocyba, Matsu-

mura 1206

unipuncta, Typhlocyba, McAtee, 1206

venusta, Empoa 1201

vesta, Typhlocyba 1157

vestita, Empoa 1204

volans, Ossiannilssonola 1138

Xanthippe, Typhlocyba 1220

1238 The University Science Bulletin

PLATE LXXIII

Fig. 1. Henribautia nigricephala (Beamer)

la. Left side of pygofer, lateral aspect.

lb. Aedeagus, left lateral aspect.

lc. Aedeagus, posterior aspect.

Id. Right style, ventral aspect.

le. Connective, ventral aspect.

If. Left plate, ventral aspect.

lg. Left plate, left lateral aspect.

lh. Left fore and hind wing.

li. Head of male and female, dorsal aspect.
Fig. 2. Henribautia beameri sp. nov.

2a. Left side of pygofer, lateral aspect.

2b. Aedeagus, left lateral aspect.

2c. Aedeagus, posterior aspect.
Fig. 3. Henribautia hubbardi (McAtee)

3a. Left side of pygofer, lateral aspect.

3b. Aedeagus, left lateral aspect.

3c. Aedeagus, posterior aspect.
Fig. 4. Mcateeana sexnotata ( Van Duzee )

4a. Left side of pygofer and left plate, lateral aspect.

4b. Aedeagus, left lateral aspect.

4c. Aedeagus, posterior aspect.

4d. Aedeagus, dorsal aspect.

4e. Right style, ventral aspect.

4f. Connective, ventral aspect.

4g. Left plate, ventral aspect.

4h. Left fore and hind wing.

4i. Head of male and female, dorsal aspect.

Revision of the Species Typhlocyba
PLATE LXXIII

1239

4o. M. sexnotola

1240 The University Science Bulletin

PLATE LXXIV

Fig. 1. Ribantiana uhni (Linnaeus)

la. Left side of pygofer and left plate, lateral aspect.

lb. Aedeagus, left lateral aspect.

lc. Aedeagus, posterior aspect.

Id. Right style, ventral aspect.

le. Connective, ventral aspect.

If. Left plate, ventral aspect.

lg. Left fore and hind wing.

lh. Head of male and female, dorsal aspect.
Fig. 2. Ribantiana tenerrima (Herrich-Schaffer)

2a. Pygofer, left side, lateral aspect.

2b. Aedeagus, left lateral aspect.

2c. Aedeagus, posterior aspect.
Fig. 3. Ribautiana parapiscator sp. nov.

3a. Left side of pygofer, lateral aspect.

3b. Aedeagus, left lateral aspect.

3c. Aedeagus, posterior aspect.
Fig. 4. Ribautiana unca (McAtee)

4a. Left side of pygofer, lateral aspect.

4b. Aedeagus, left lateral aspect.

4c. Aedeagus, posterior aspect.

4d. Tip of aedeagus, left lateral aspect. Oregon specimen.

4e. Tip of aedeagus, left lateral aspect. California specimen.

4f. Tip of aedeagus, left lateral aspect. Missouri specimen.

4g. Tip of aedeagus, left lateral aspect. Colorado specimen.

Revision of the Species Typhlocyba

1241

PLATE LXXIV

4o R. unca

1242 The University Science Bulletin

PLATE LXXV

Fig. 1. Ribautiana luculla (Medler)

la. Left side of pygofer, lateral aspect.

lb. Aedeagus, left lateral aspect.

lc. Aedeagus, posterior aspect.
Fig. 2. Ribautiana foliosa ( Knull )

2a. Left side of pygofer, lateral aspect.

2b. Aedeagus, left lateral aspect.

2c. Aedeagus, posterior aspect.
Fig. 3. Ribautiana piscator (McAfee)

3a. Left side of pygofer, lateral aspect.

3b. Aedeagus, left lateral aspect.

3c. Aedeagus, posterior aspect.
Fig. 4. Ribautiana multispinosa sp. nov.

4a. Left side of pygofer, lateral aspect.

4b. Aedeagus, left lateral aspect.

4c. Aedeagus, posterior aspect.
Fig. 5. Ribautiana sciotoensis (Knull)

5a. Aedeagus, left lateral aspect.

5b. Aedeagus, posterior aspect.

Revision of the Species Typhlocyba

1243

PLATE LXXV

5a R. selotoensis

1244 The University Science Bulletin

PLATE LXXVI

Fig. 1. Ossiannilssonola berenice (McAtee)

la. Left side of pygofer and left plate, lateral aspect.

lb. Aedeagus, left lateral aspect.

lc. Aedeagus, posterior aspect.

Id. Right style, ventral aspect.

le. Connective, ventral aspect.

If. Left plate, ventral aspect.

lg. Left fore and hind wing.

lh. Head of male and female, dorsal aspect.

li. Left side of pygofer, dorsal angle, posterior aspect.
Fig. 2. Ossiannilssonola hermione ( McAtee )

2a. Left side of pygofer, lateral aspect.

2b. Aedeagus, left lateral aspect.

2c. Aedeagus, posterior aspect.

2d. Left side of pygofer, dorsal angle, posterior aspect.
Fig. 3. Ossiannilssonola volans (McAtee)

3a. Left side of pygofer, lateral aspect.

3b. Aedeagus, left lateral aspect.

3c. Aedeagus, posterior aspect.

3d. Left side of pygofer, dorsal angle, posterior aspect.

Revision of the Species Typhlocyba

1245

PLATE LXXVI

1246 The University Science Bulletin

PLATE LXXVII

Fig. 1. Ossiannilssonola antigone (McAtee)

la. Left side of pygofer, lateral aspect.

lb. Aedeagus, left lateral aspect.

lc. Aedeagus, posterior aspect.

Id. Left side of pygofer, dorsal angle, posterior aspect.
Fig. 2. Ossiannilssonola clymene (McAtee)

2a. Left side of pygofer, lateral aspect.

2b. Aedeagus, left lateral aspect.

2c. Aedeagus, posterior aspect.

2d. Left side of pygofer, dorsal angle, posterior aspect.
Fig. 3. Ossiannilssonola tunicarubra (Gillette)

3a. Left side of pygofer, lateral aspect.

3b. Aedeagus, left lateral aspect.

3c. Aedeagus, posterior aspect.

3d. Left side of pygofer, dorsal angle, posterior aspect.
Fig. 4. Ossiannilssonola australis (Walsh)

4a. Left side of pygofer, lateral aspect.

4b. Aedeagus, left lateral aspect.

4c. Aedeagus, posterior aspect.

4d. Left side of pygofer, dorsal angle, posterior aspect.
Fig. 5. Ossiannilssonola hinei ( Knull )

5a. Left side of pygofer, lateral aspect.

5b. Aedeagus, left lateral aspect.

5c. Aedeagus, posterior aspect.

5d. Left side of pygofer, dorsal angle, posterior aspect.

Revision of the Species Typhlocyba

1247

PLATE LXXVII

5«. 0. hinei

49-6490

1248 The University Science Bulletin

PLATE LXXVIII

Fig. 1. Ossiannilssonola bangsoni sp. nov.

la. Left side of pygofer, lateral aspect.

lb. Aedeagus, left lateral aspect.

lc. Aedeagus, posterior aspect.

Id. Left side of pygofer, dorsal angle, posterior aspect.

le. Apex of right style, ventral aspect.
Fig. 2. Ossiannilssonola duplicata ( McAtee )

2a. Left side of pygofer, lateral aspect.

2b. Aedeagus, left lateral aspect.

2c. Aedeagus, posterior aspect.

2d. Left side of pygofer, dorsal angle, posterior aspect.
Fig. 3. Ossiannilssonola serrula (Ross and DeLong)

3a. Left side of pygofer, lateral aspect.

3b. Aedeagus, left lateral aspect.

3c. Aedeagus, posterior aspect.

3d. Left side of pygofer, dorsal angle, posterior aspect.
Fig. 4. Ossiannilssonola mcateei sp. nov.

4a. Left side of pygofer, lateral aspect.

4b. Aedeagus, left lateral aspect.

4c. Aedeagus, posterior aspect.

4d. Left side of pygofer, dorsal angle, posterior aspect.
Fig. 5. Ossiannilssonola danae (McAtee)

5a. Left side of pygofer, lateral aspect.

5b. Aedeagus, left lateral aspect.

5c. Aedeagus, posterior aspect.

5d. Left side of pygofer, dorsal angle, posterior aspect.

5e. Apex of right style, ventral aspect.

Revision of the Species Typhlocyba

1249

PLATE LXXVIII

5o 0. doi.ue

1250 The University Science Bulletin

PLATE LXXIX

Fig. 1. Ossiannilssonola appendiculata (Malloch)

la. Left side of pygofer and left plate, lateral aspect.

lb. Aedeagus, left lateral aspect.

lc. Aedeagus, posterior aspect.

Id. Left side of pygofer, dorsal angle, posterior aspect.

le. Right style, ventral aspect.

If. Left plate, ventral aspect.
Fig. 2. Ossiannilssonola phryne (McAtee)

2a. Left side of pygofer and left plate, lateral aspect.

2b. Aedeagus, left lateral aspect.

2c. Aedeagus, posterior aspect.

2d. Left side of pygofer, dorsal angle, posterior aspect
Fig. 3. Ossiannilssonola rossi sp. nov.

3a. Left side of pygofer, lateral aspect.

3b. Aedeagus, left lateral aspect.

3c. Aedeagus, posterior aspect.

3d. Left side of pygofer, dorsal angle, posterior aspect.
Fig. 4. Ossiannilssonola knulli sp. nov.

4a. Left side of pygofer, lateral aspect.

4b. Aedeagus, left lateral aspect.

4c. Aedeagus, posterior aspect.

4d. Left side of pygofer, dorsal angle, posterior aspect.

4e. Apex of right style, ventral aspect.

Revision of the Species Typhlocyba

1251

PLATE LXXIX

1252 The University Science Bulletin

PLATE LXXX

Fig. 1. Ossiannilssonola troza (Ross and DeLong)

la. Left side of pygofer, lateral aspect.

lb. Aedeagus, left lateral aspect.

lc. Aedeagus, posterior aspect.

Id. Left side of pygofer, dorsal angle, posterior aspect.
Fie. 2. Ossiannilssonola flavomarginata (Gillette and Baker)

2a. Left side of pygofer and left plate, lateral aspect.

2b. Aedeagus, left lateral aspect.

2c. Aedeagus, posterior aspect.

2d. Left side of pygofer, dorsal angle, posterior aspect.

2e. Right style, ventral aspect.
Fig. 3. Ossiannilssonola quadrata (DeLong and Johnson)

3a. Left side of pygofer, lateral aspect.

3b. Aedeagus, left lateral aspect.

3c. Aedeagus, posterior aspect.

3d. Left side of pygofer, dorsal angle, posterior aspect.

3e. Apex of right style, ventral aspect.

Revision of the Species Typhlocyba

1253

PLATE LXXX

1254 The University Science Bulletin

PLATE LXXXI

Fig. 1. Typhlocyba quercus (Fabricius)

la. Left side of pygofer and left plate, lateral aspect.

lb. Aedeagus, left lateral aspect.

lc. Aedeagus, posterior aspect.

Id. Right style, ventral aspect.

le. Connective, ventral aspect.

If. Left fore and hind wing.

lg. Head of male and female, dorsal aspect.
Fig. 2. Typhlocyba oneka Knull

2a. Left side of pygofer, lateral aspect.

2b. Aedeagus, left lateral aspect.

2c. Aedeagus, posterior aspect.

2d. Right style, ventral aspect.

2e. Connective, ventral aspect.
Fig. 3. Typhlocyba athene McAtee

3a. Left side of pygofer, lateral aspect.

3b. Aedeagus, left lateral aspect.

3c. Aedeagus, posterior aspect.
Fig. 4. Typhlocyba arsinoe McAtee

4a. Left side of pygofer, lateral aspect.

4b. Aedeagus, left lateral aspect.

4c. Aedeagus, posterior aspect.

Revision of the Species Typhlocyba

1255

PLATE LXXXI

1256 The University Science Bulletin

PLATE LXXXII

Fig. 1. Typhlocyba modesta Malloch

la. Left side of pygofer, lateral aspect.

lb. Aedeagus, left lateral aspect.

lc. Aedeagus, posterior aspect.

Id. Right style, ventral aspect.

le. Connective, ventral aspect.
Fig. 2. Typhlocyba medleri sp. nov.

2a. Left side of pygofer, lateral aspect.

2b. Aedeagus, left lateral aspect.

2c. Aedeagus, posterior aspect.

2d. Right style, ventral aspect.

2e. Connective, ventral aspect.
Fig. 3. Typhlocyba hockingensis Knull

3a. Left side of pygofer, lateral aspect.

3b. Aedeagus, left lateral aspect.

3c. Aedeagus, posterior aspect.

3d. Aedeagus, dorsal aspect.

3e. Right style, ventral aspect.

3f. Connective, ventral aspect.
Fig. 4. Typhlocyba pomaria McAtee

4a. Left side of pygofer, lateral aspect.

4b. Aedeagus, left lateral aspect.

4c. Aedeagus, posterior aspect.

4d. Right style, ventral aspect.

4e. Connective, ventral aspect.

Revision of the Species Typhlocyba

1257

PLATE LXXXII

1258 The University Science Bulletin

PLATE LXXXIII

Fig. 1. Typhlocyba attenuata sp. nov.

la. Left side of pygofer, lateral aspect.

lb. Aedeagus, left lateral aspect.

lc. Aedeagus, posterior aspect.

Id. Right style, ventral aspect.

le. Connective, ventral aspect.
Fig. 2. Typhlocyba rtibriocellata Malloch

2a. Left side of pygofer, lateral aspect.

2b. Aedeagus, left lateral aspect.

2c. Aedeagus, posterior aspect.

2d. Right style, ventral aspect.

2e. Connective, ventral aspect.
Fig. 3. Typhlocyba surcula DeLong and Johnson

3a. Left side of pygofer, lateral aspect.

3b. Aedeagus, left lateral aspect.

3c. Aedeagus, posterior aspect.

3d. Right style, ventral aspect.

3e. Connective, ventral aspect.
Fig. 4. Typhlocyba andromache McAtee

4a. Left side of pygofer, lateral aspect.

4b. Aedeagus, left lateral aspect.

4c. Aedeagus, posterior aspect.

4d. Right style, ventral aspect.

4e. Connective, ventral aspect.

Revision of the Species Typhlocyba

1259

PLATE LXXXIII

1260 The University Science Bulletin

PLATE LXXXIV

Fig. 1. Typhlocyba melite McAtee

la. Left side of pygofer, lateral aspect.

lb. Aedeagus, left lateral aspect.

lc. Aedeagus, posterior aspect.

Id. Right style, ventral aspect.

le. Connective, ventral aspect.
Fig. 2. Typhlocyba alabamaensis sp. nov.

2a. Left side of pygofer, lateral aspect.

2b. Aedeagus, left lateral aspect.

2c. Aedeagus, posterior aspect.
Fig. 3. Typhlocyba shawneeana Knull

3a. Left side of pygofer, lateral aspect.

3b. Aedeagus, left lateral aspect.

3c. Aedeagus, posterior aspect.

3d. Right style, ventral aspect.
Fig. 4. Typhlocyba transviridis sp. nov.

4a. Left side of pygofer, lateral aspect.

4b. Aedeagus, left lateral aspect.

4c. Aedeagus, posterior aspect.

4d. Right style, ventral aspect.

Revision of the Species Typhlocyba

1261

PLATE LXXXIV

3o. Tshowneeono

1262 The University Science Bulletin

PLATE LXXXV

Fig. 1. Typhlocyba putmani Knull

la. Left side of pygofer, lateral aspect.

lb. Aedeagus, left lateral aspect.

lc. Aedeagus, posterior aspect.

Id. Right style, ventral aspect.

le. Connective, ventral aspect.
Fig. 2. Typhlocyba Cassiopeia Knull

2a. Left side of pygofer, lateral aspect.

2b. Aedeagus, left lateral aspect.

2c. Aedeagus, posterior aspect.

2d. Right style, ventral aspect.

2e. Connective, ventral aspect.
Fig. 3. Typhlocyba crassa DeLong and Johnson

3a. Left side of pygofer, lateral aspect.

3b. Aedeagus, left lateral aspect.

3c. Aedeagus, posterior aspect.

3d. Right style, ventral aspect.

3e. Connective, ventral aspect.

Revision of the Species Typhlocyba
PLATE LXXXV

1263

50—6490

1264 The University Science Bulletin

PLATE LXXXVI

Fig. 1. Typhlocyba sollisa Ross and DeLong

la. Left side of pygofer, lateral aspect.

lb. Aedeagus, left lateral aspect.

lc. Aedeagus, posterior aspect.
Fig. 2. Typhlocyba tortosa Ross and DeLong

2a. Left side of pygofer, lateral aspect.

2b. Aedeagus, left lateral aspect.

2c. Aedeagus, posterior aspect.

2d. Right style, ventral aspect.
Fig. 3. Typhlocyba persephone McAtee

3a. Left side of pygofer, lateral aspect.

3b. Aedeagus, left lateral aspect.

3c. Aedeagus, posterior aspect.

3d. Right style, ventral aspect.

3e. Right style, apex, lateral aspect.

3f. Right style, apex, posterior aspect.

3g. Connective, ventral aspect.
Fig. 4. Typhlocyba niobe McAtee

4a. Left side of pygofer, lateral aspect.

4b. Aedeagus, left lateral aspect.

4c. Aedeagus, posterior aspect.

Revision of the Species Typhlocyba

1265

PLATE LXXXVI

1266 The University Science Bulletin

PLATE LXXXVII

Fig. 1. Typhlocyba infiata sp. nov.

la. Left side of pygofer, lateral aspect.

lb. Aedeagus, left lateral aspect.

lc. Aedeagus, posterior aspect.

Id. Right style, ventral aspect.

le. Connective, ventral aspect.
Figs. 2-13: Eighth abdominal sternite, female, a. Ventral, b. Lateral

aspect.
Fig. 2. Henribautia nigricephala (Beamer)
Fig. 3. Ribautiana piscator (McAtee)
Fig. 4. Ribautiana ulmi (Linnaeus)
Fig. 5. Mcateeana sexnotata ( Van Duzee )
Fig. 6. Ossiannilssonola berenice (McAtee)
Fig. 7. Ossiannilssonola flavomarginata (Gillette and Baker)
Fig. 8. Typhlocyba quercus (Fabricius)
Fig. 9. Typhlocyba attenuata sp. nov.
Fig. 10. Typhlocyba persephone McAtee
Fig. 11. Typhlocijba modesta Gibson
Fig. 12. Typhlocyba pomaria McAtee
Fig. 13. Edwardsiana rosae ( Linnaeus )

Revision of the Species Typhlocyba

1267

PLATE LXXXVII

80. T. quercuS

1268 The University Science Bulletin

PLATE LXXXVIII

Fig. 1. Empoa albicans Walsh

la. Left side of pygofer, lateral aspect.

lb. Aedeagus, left lateral aspect.

lc. Aedeagus, posterior aspect.

Id. Right style, ventral aspect.

le. Connective, ventral aspect.
Fig. 2. Empoa spinosa (Beamer)

2a. Left side of pygofer, lateral aspect.

2b. Aedeagus, left lateral aspect.

2c. Aedeagus, posterior aspect.

2d. Right style, ventral aspect.

2e. Connective, ventral aspect.
Fig. 3. Empoa querci Fitch

3a. Left side of pygofer, lateral aspect.

3b. Aedeagus, left lateral aspect.

3c. Aedeagus, posterior aspect.

3d. Right style, ventral aspect.

3e. Connective, ventral aspect.

Revision of the Species Typhlocyba
PLATE LXXXVIII

1269

1270 The University Science Bulletin

PLATE LXXXIX

Fig. 1. Edwardsiana lethierryi (Edwards)

la. Left side of pygofer, lateral aspect.

lb. Aedeagus, left lateral aspect.

lc. Aedeagus, anterior aspect.
Fig. 2. Edwardsiana rosae (Linnaeus)

2a. Left side of pygofer, lateral aspect.

2b. Aedeagus, left lateral aspect.

2c. Aedeagus, anterior aspect.

2d. Right style, ventral aspect.

2e. Connective, ventral aspect.

2f. Apex of aedeagus, left lateral aspect, rosae variation.

2g. Apex of aedeagus, anterior aspect, rosae variation.
Fig. 3. Edwardsiana bergmani var. ariadne ( McAtee )

3a. Left side of pygofer, lateral aspect, (Maine specimen).

3b. Aedeagus, left lateral aspect, ( Maine specimen ) .

3c. Aedeagus, anterior aspect, ( Maine specimen ) .

3d. Left side of pygofer, lateral aspect, ( Montana specimen ) .

3e. Aedeagus, left lateral aspect, (Montana specimen).

3f. Aedeagus, anterior aspect, (Montana specimen).
Fig. 4. Edwardsiana bergmani var. bergmani (Tullgren)

4a. Aedeagus, left lateral aspect, (Swedish specimen).

4b. Aedeagus, anterior aspect, (Swedish specimen).

Revision of the Species Typhlocyba

1271

PLATE LXXXIX

4a. vor. bergmani

1272 The University Science Bulletin

PLATE XC

Fig. 1. Edwardsiana expanda (DeLong and Johnson)

la. Left side of pygofer, lateral aspect.

lb. Aedeagus, left lateral aspect.

lc. Aedeagus, anterior aspect.

Id. Apex of aedeagus, left lateral aspect (abnormal specimen).
Fig. 2. Edwardsiana frustrator (Edwards)

2a. Left side of pygofer, lateral aspect.

2b. Aedeagus, left lateral aspect.

2c. Aedeagus, anterior aspect.
Fig. 3. Edwardsiana plebeja (Edwards)

3a. Left side of pygofer, lateral aspect.

3b. Aedeagus, left lateral aspect.

3c. Aedeagus, anterior aspect.
Fig. 4. Edwardsiana prunicola (Edwards)

4a. Left side of pygofer, lateral aspect.

4b. Aedeagus, left lateral aspect.

4c. Aedeagus, anterior aspect.
Fig. 5. Edwardsiana candidula (Kirschbaum)

5a. Left side of pygofer, lateral aspect.

5b. Aedeagus, left lateral aspect.

5c. Aedeagus, anterior aspect.
Fig. 6. Edwardsiana australis (Froggatt)

6a. Left side of pygofer, lateral aspect.

6b. Aedeagus, left lateral aspect.

6c. Aedeagus, anterior aspect.

Revision of the Species Typhlocyba

1273

PLATE XC

1274 The University Science Bulletin

PLATE XCI

Fig. 1. Edwardsiana projecta sp. nov.

la. Left side of pygofer, lateral aspect.

lb. Aedeagus, left lateral aspect.

lc. Aedeagus, anterior aspect.

Id. Aedeagus, apex, posterodorsal aspect.
Fig. 2. Edwardsiana pseudocommissuralis sp. nov.

2a. Left side of pygofer, lateral aspect.

2b. Aedeagus, left lateral aspect.

2c. Aedeagus, anterior aspect.
Fig. 3. Edwardsiana dejecta sp. nov.

3a. Left side of pygofer, lateral aspect.

3b. Aedeagus, left lateral aspect.

3c. Aedeagus, anterior aspect.
Fig. 4. Edwardsiana dorsti ( Ossiannilsson )

4a. Left side of pygofer, lateral aspect.

4b. Aedeagus, left lateral aspect.

4c. Aedeagus, anterior aspect.

Revision of the Species Typhlocyba

1275

PLATE XCI

f. dorst

1276 The University Science Bulletin

PLATE XCII

Fig. 1. Edwardsiana nigripennis sp. nov.

la. Left side of pygofer, lateral aspect.

lb. Aedeagus, left lateral aspect.

lc. Aedeagus, anterior aspect.
Fig. 2. Edwardsiana delongi sp. nov.

2a. Left side of pygofer, lateral aspect.

2b. Aedeagus, left lateral aspect.

2c. Aedeagus, anterior aspect.
Fig. 3. Edwardsiana commissuralis ( Stal )

3a. Left side of pygofer, lateral aspect.

3b. Aedeagus, left lateral aspect.

3c. Aedeagus, anterior aspect.
Fig. 4. Edwardsiana ariste (McAtee)

4a. Left side of pygofer, lateral aspect.

4b. Aedeagus, left lateral aspect.

4c. Aedeagus, anterior aspect.
Fig. 5. Edwardsiana euphrante (McAtee)

5a. Left side of pygofer, lateral aspect.

5b. Aedeagus, left lateral aspect.

5c. Aedeagus, anterior aspect.

Revision of the Species Typhlocyba

1277

PLATE XCII

5a. F. tuphronte

THE UNIVERSITY OP KANSAS

SCIENCE BULLETIN

Vol. XXXV, Pt. II] September 10, 1953 [No. 10

The Ambrysus * of Mexico ( Hemiptera, Naucoridae)

Ira La Rivers 2

FAMILY NAUCORIDAE (Fallen) 1814

Subfamily AMBRYSINAE Usinger 1941

Genus Ambrysus Stal 1862

(Logotype signoreti Stal)

Ambrysus Stal 1862, Stet. Ent. Zeit. 23:459; 1865, Hemip. Afr., 3:174; 1876,

Enum. Hemip., 5:141, 143.
Ambrysus, Uhler, 1872, Hayden's Surv. Terr., Rept. for 1871:423; 1876, Bull.

U.S.G.S. 1:337; 1877, Wheeler's Rept. Chief Eng. for 1877:1331; 1884, in

Kings. Nat. Hist., 2:249, 260; 1886, Brook. Ent. Soc.; 1894, Proc. Calif.

Acad. Sci., ser. 2, 4:291.
Ambrysus, Berg, 1879, Hemip. Argent, enum. spec. nov. desc, 8vo.
Ambrysus, Gillette and Baker, 1895, Hemip. Colo. :63.
Ambrysus, Montandon, 1897, Bull. Paris Mus. d'Hist. Nat. 3:124; 1897, Verh.

zool.-bot. Ges. Wien 47:6; 1897, Bull. Soc. Sci. Buc.-Roum. 7(3-4) :282;

1909, ibid, 17(5-6) :316, 18(1 ):43; 1910, ibid, 18(5-6) :180, 19(3) :438.
Ambrysus, Champion, 1900, Biol. Centr.-Amer., Heterop. 2:355-358.
Ambrysus, Kirkaldy, 1906, Trans. Amer. Ent. Soc, 32:151.
Ambrysus, Kirkaldy and Bueno, 1908, Proc. Ent. Soc. Wash., 10:185.
Ambrysus, Snow, 1906, Trans. Kans. Acad. Sci., 20(1):180.
Ambrysus, Van Duzee, 1916, Proc. N. Y. Ent. Soc; 1917, U. C. Ent. Bull. 2:458.
Ambrysus, Hungerford, 1919, Univ. Kans., Sci. Bull., 11:198.
Ambrysus, Usinger, 1941, Ann. Ent. Soc. Amer. 34(1 ):11, 15; 1946, Univ.

Kans., Sci. Bull., 31(1):185.
Ambrysus, La Rivers, 1949, Bull. S. Calif. Acad. Sci., 47(3) :103; 1949, Ann.

Ent. Soc. Amer., 41(3) :371; 1950, Pan-Pac Ent., 26(1 ):19.

Because of the superficial, external similarity between many of
the species, it is convenient to characterize the genus in some de-
tail, making it possible to eliminate much repetitive material in the
individual species descriptions:

1. Dr. Usinger in Univ. of Kansas Science Bull. XXXI, pp. 182-210, 1946, described
some new species of Ambrysus from the Frances Huntington Snow Collections and this paper
is the culmination and with its illustrated key is a companion work to be used with "A
Revision of the Genus Ambrysus in the United States" by Ira La Rivers in the Univ. of
California Publications in Entomology VIII, No. 7, pp. 277-338, 1951, and is based to a
considerable extent upon Univ. of Kansas material. — H. B. Hungerford.

2. Department of Biology, University of Nevada, Reno, Nevada.

(1279)
51—6490

1280 The University Science Bulletin

General appearance: small and rotund to large and ovate — size
6-16 mm. long, 3.5-8.0 mm. wide. Dorsum generally distinctly bi-
colored, the head and prothorax slightly lighter in color than the
remainder, emboliar edges usually lighter than hemelytra, the latter
with or without mottling. Venter generally lighter than abdomen,
darkening somewhat in anterior half of body, with or without
median, anterior mottling.

Head: glistening and impunctate to roughened and strongly
punctate, greenish, yellowish, brown or combinations of these colors,
varying from smoothly curved between the eyes to distinctly pro-
tuberant before eyes. Eyes generally much darker in color than
head, brown to black, often drying in pinned specimens to a pearly
gray; outer and posterior margins either forming a continuous
curvature, or producing an angulation at point of contact. Labrum
same color as front of head, or darker; mouthparts generally darken-
ing at tip.

Pronotum: varying from nearly impunctate and glistening to
rough and strongly punctate, and in color from greenish through
yellow to deep black-brown. Combined length of pronotum and
head varying from more than two thirds greatest pronotal width
to distinctly less than two thirds such width A thin, transverse,
dark posterior pronotal line prominently present, the remainder of
the posterior pronotal border behind the line predominantly whitish
in color. Lateral pronotal edges varying from evenly curving cau-
dad to nearly straight; also varying from smooth to serrate. Pos-
terior pronotal angles may be lacking ( i. e., well rounded ) or
sharply prominent. Venter generally darker medially, with yellow
pile prominent about centrum and posterior edge.

Scutellum: always strongly shagreened with shallow but dense
punctation, each puncture whitish at bottom. Varying in color
from unicolorously dark-brown to various mottlings of light brown
and yellow, in some cases, predominantly yellow.

Hemelytra: nearly always with some glisten, punctate, each
puncture the seat of a white spot, varying in color from unicolorous
dark-brown to a well-developed mottled pattern of brown and light,
bright yellow, the embolium almost always distinctly and conspic-
uously lighter than the rest of hemelytra. Hemelytra may strongly
or weakly expose the lateral connexival edges, and may or may not
attain abdominal tip.

Venter: the prothoracic venter has been discussed above. Re-
mainder of venter is generally distinctly bicolored, the abdomen
golden to brown due to the short, golden hydrofuge pelt, and lighter

La Rivers: The Ambrysus of Mexico 1281

in color than the velvety-appearing meso- and metathoracic venters,
which latter may have rich brown mottling centrally. Connexival
segments may exhibit various combinations of spination or nonspi-
nation on posterolateral angles; if spined, the first, and often the first
and second angles are generally nonspinose.

Legs: Prolegs — coxa elongate, globular, green, yellow or brown,
smooth, glistening, darkening at distal tips, flattened to receive heel
of femur; trochanter well developed, smooth, green, yellow or
brown, with tufts of yellow hairs on anterior edge; femur smooth,
glistening, green, yellow or brown, widest near proximal end, nar-
rowing rapidly to distal end, i. e., typically incrassate, and with
characteristic short, very dense mat of hairs along front border
which serves as a resting groove for the tibia when closed against
femur; tibia very long, slender, smooth, yellow to brown, occasion-
ally greenish, curved most strongly in distal part where, with the
single tarsal segment, it forms a continuous, curved, grasping instru-
ment— combined tibia-tarsus-claw, when closed, either slightly ex-
ceeding or slightly shorter than, adjacent edge of femur.

Mesolegs — coxa long, greenish, yellow or brownish, somewhat
angularly globular, beset with short, dense golden pile, slightly
curved from posterior end and weakly laterad, to anterior end, the
outer face flat for reception of basal part of femur; trochanter large,
similar in color to coxa, smooth distally, pilose proximally; femur
very long, narrow, smooth, similar in color to coxa, glistening, com-
pressed dorsoventrally, some golden pilosity on outer length; tibia
greenish, yellow or light-brown, smooth, glistening, narrow, strongly
armed with longitudinal rows of strong, reddish-brown spines, ar-
ranged in series along the four rounded "angles" formed by the slight
dorsoventral compression of tibia — distal end ventrally with from
one and one-half to a maximum of eight complete or partial trans-
verse rows of spines set across tibial width, the last row at extreme
distal tip; tarsus slender, smooth, glistening, green to yellow, pilose
beneath, three-segmented, the first segment minute and difficult to
see, even in ventral view, terminating in two claws of about same
color as tarsus, darkening at tips, generally only weakly or moder-
ately curved.

Metalegs — coxa swollen, globular, green, yellow or light brown,
well haired with short, dense golden pile, flattened ventrolaterally
for reception of basal part of femur; trochanter well developed,
similar in color to coxa, pilose proximally, smooth and glistening
distally; femur long, narrow, smooth, glistening, green, yellow or
brown, dorsoventrally compressed, weakly spinulose along outer

1282 The University Science Bulletin

margin, and bearing a thin mat of short, yellow pile on inner margin;
tibia long, narrow, glistening, yellow or green, thickly beset with
brownish-red spines as in mesotibia, but spines longer, more promi-
nent, and more evidently unequal in size, long and short spines
alternating, arranged along four "corners" formed by the slight
dorseventral flattening — distal end ventrally with from one and
one-half to a maximum of eight partial or complete transverse
rows of spines set across width of tibia, the last row at extreme
tibial apex — inner margin cushioned with a solid, dense mat of long,
silky golden swimming hairs; tarsus slim, smooth, long, narrow,
green to yellow and brown, three segmented, the first segmented diffi-
cult to see, spinulose and pilose beneath, with two claws markedly
darkening at tips and weakly to moderately curved.

Key to the Mexican Species of Ambrysus

1. Presternum fused to propleura, both on the same plane 2*

Presternum free from propleura, most definitely so medially,

where presternum disappears posteriorly beneath fore edge

of propleura 3

2(1). Male genital process short, somewhat platelike, roundly
capitate, distinctly less than twice as long as wide; female
subgenital plate-apex widely concave, the concavity armed
with two small teeth or projections near middle. (Pi. 93,
fig. 1 ) pijgmaeus

Male genital process moderately long, parallel-sided to

slightly narrowing towards tip, distinctly more than twice as
long as wide; female subgenital plate not so constructed.

( Pi. 93, fig. 2 ) circumcinctus caliginosus

3(1). Female with two elevated almost peglike angles arising
from caudal edge of sternite V at lateral anterior edges of
subgenital plate; connexiva spineless; prosternal ridge sub-
equal in length to union of propleura along median fine; fe-
male subgenital plate tipped with a low, blunt angle; male
genital process short, foot-shaped, strongly turned exter-
nally. ( Pi. 93, figs. 3 and 4 ) melanopterus

Female lacking the two elevated angles on caudal edge of

sternite V; species without the remaining combination of

characters 4

4(3). Lateral edges of abdominal segments III-IV strongly and
distinctly serrate in contrast to smooth edges of segments
I-II; small, rotund species (species generally with some de-
velopment of the laterocaudal angle on the female Vth
sternite, from a thin, sharp spine to a broad, low but definite
sinuosity) 5

Lateral edges of all abdominal segments smooth or, if ser-
rate, the serration is gradually developed anteriorly to pos-
teriorly 10

0 Numbers refer to figures, pis. XCIII and XCIV.

La Rivers : The Ambrysus of Mexico 1283

5(4). Males completely lacking a genital process or any indication
of such a process, the point usually occupied by it on caudal
edge of tergite V being smoothly rounded 6

Males always with some indication of the genital process,

from a definite angle or short, stubby process to a fully de-
veloped hook 8

6(5). Female subgenital plate quadrisinuate at apex, the lateral
shoulder-sinuosities low, the central sinuosities prominent,
appearing as a medially cleft process; female V sternite
bispinated at left posterolateral edge, the usual connexival
spine accompanied by an angulate projection arising inward
from the laterocaudal margin of the sternite (— latero-
caudal spine or angle ) . (Pi. 93, fig. 5 ) parviceps

Female subgenital plate simply and strongly concave at apex;

Vth female sternite unispinate at left posterolateral edge
due to loss of outer, true connexival spine, its place occupied
by only a smooth curve ( pudicus ) 7

7(6). Depth of concavity at apex of female subgenital plate less
than 50% of width of concavity; smaller, measuring 7.0-
7.5 mm. in length. ( Pi. 93, fig. 6 ) pudicus pudicus

Depth of concavity at apex of female subgenital plate more

than 50% of width of concavity; larger, measuring 7.5-
9.0 mm. in length. (Pi. 93, fig. 7) pudicus barberi

8(5). Posterolateral connexival spine of female abdominal seg-
ment V absent, laterocaudal process spinelike, very prom-
inent; female subgenital plate quadriangulate at apex, the
lateral shoulders subdued, the central angles prominent
( /. e., extending more caudad ) . (PI. 93, fig. 8 ) abortus

Posterolateral connexival spine of female abdominal seg-
ment V present, laterocaudal process subdued, either a low,
blunt angle or a rounded sinuosity; female subgenital plate
smoothly and weakly concave at apex in A. hungerfordi
hungerfordi, most angulate in A. h. angularis,

( hungerfordi ) 9
9(8). Female subgenital plate smoothly and evenly rounded at
apex, tip concave; laterocaudal angle low, rounded, much
closer to posterolateral spine than to base of subgenital
plate. ( PI. 93, figs. 9 and 10 ) hungerfordi hungerfordi

Female subgenital plate slightly more angulate at apex,

the sinuosities on each side of the median terminal concavity
being flat-topped; laterocaudal angle low but rather sharp,
about midway between connexival spine and base of sub-
genital plate. (Pi. 93, figs. 11 and 12) hungerfordi triunfo

Female subgenital plate with terminal outline more quad-
risinuate, nearly angulate, but rather than sinuosities on each
side of median terminal concavity being flat-topped or
rounded, they are blunt-angulate; laterocaudal angle absent,
unmarked by even a sinuosity along the border. (Pi. 93,
figs. 13 and 14) hungerfordi angularis

1284 The University Science Bulletin

10(4). Posterior slope behind median prosternal ridge with two
tuberculations on each side of median line directly behind
caudal end of median ridge, producing a longitudinal median
trough- or indentation-effect; female possessing a strong,
broad and rather blunt laterocaudal angle; male genital
process large and strongly boot-shaped, the sole of the boot
straight; tip of female subgenital plate broadly, weakly and
simply concave; large species. (Pi. 93, figs. 15 and 16),

puncticolhs

Posterior slope behind median prosternal ridge flat; at most,

with a few weak transverse rugosities which never produce a
median longitudinal furrow; females lacking the latero-
caudal angle; species without the remaining combination of
characters 11

11(10). Combined length of pronotum and head more than two-
thirds greatest pronotal width; head and pronotum highly
polished, smooth and glistening, without evidences of puncta-
tion. (PI. 93, fig. 17) pulchellus

Combined length of pronotum and head less than two thirds

greatest pronotal width; head and pronotum always with
some discernible punctation, even though shiny 12

12(11). Female subgenital plate the most distinctive in the genus
for this area, semitubular in shape, narrowing sharply from
base to apex, and protruding caudad of abdominal tip;
valvulae, when projecting beyond subgenital tip, seen to be
a rather sharp, smooth, double-pointed ovipositor. (Pi. 93,
fig. 18 ) vanduzeei

Female subgenital plate not so constructed and never pro-
jecting beyond abdominal tip; valvulae, when projecting,
are hirsute 13

13(12). Female subgenital plate dominantly concave at apex, i.e.,
regardless of degree of development of lateral terminal
angles, whether rounded or spinose, a well-marked concavity
occupies the apex and is at least half as wide as the sub-
genital plate at that point 14

Female subgenital plate variously shaped, multisinuate at

apex, but never with a dominant concavity 15

14 ( 13). Female subgenital plate simply and broadly concave at apex,
lateral angles simple and sharp, but not spinose; male genital
process broadened and somewhat spatulate. (Pi. 93, fig.
19 ) huenoi

Female subgenital plate complexly and broadly concave at

apex, lateral angles set inward from edge and prolonged
caudad into strong, narrow, but not particularly sharp,
spines; male genital process narrow, nonspatulate. (Pi. 93,
fig. 20) mormon australis

15(13). Lateral angles of female subgenital plate-apex rounded, gen-
erally shorter (i. e., more cephalad in position) than median
angles or sinuosities 16

La Rivers: The Ambrysus of Mexico 1285

Lateral angles of female subgenital plate-apex pointed, angu-

late or spinose, shorter or longer than median angles or
sinuosities 24

16(15). Apex of female subgenital plate asymmetrical, from ventral
view, with right side extending farther caudad than left side,
all angles rounded, the apex trisinuate ( male genital process
short and moderately broad). ( Pi. 94, fig. 21 ) scalenus

Apex of female subgenital plate symmetrical, one side of

plate not longer than the other side 17

17(16). Male genital process short, narrow, moderately curved; fe-
male subgenital plate weakly quadrisinuate at apex, the lat-
eral angles low (i. e., more cephalad in position), the median
angles extending more caudad and appearing almost as a
single bluntly rounded tip. ( Pi. 94, fig. 22 ) convexus

Without the above combination of characters 18

18(17). Embolia remarkably inflated, almost angularly so — width of
embolia 43 ± 1% of emboliar length ( male genital process
very short and broad, female subgenital plate with lateral
apical angles low, median angles projecting more caudad,
not prominent ) . (Pi. 94, figs. 23, 24 ) inflatus

Embolia never so inflated (species without remaining com-
bination of characters) 19

19(18). Male genital process short but very broad at apex, decidedly
boot-shaped, with a well-developed toe; female subgenital
plate-apex with lateral angles lower than median angles;
large, comparatively narrow species with a proportionately
large head, noticeable, however, only during direct compari-
sons between species. (Pi. 94, fig. 25) magniceps

Without the above combination of characters 20

20(19). Lateral and median angles of female subgenital plate-apex
on the same plane, /. e., a line drawn through their four tips
being approximately a straight line 21

Lateral angles of female subgenital plate-apex shorter than

median angles or sinuosities, i. e., the median angles project
more caudad than the laterals 22

21(20). Male genital process weakly curved, proportionately broad,
width of process averaging 48% of process-length (using
greatest width in apical half of process, and curving length
of process from tip to base along inner margin). (PI. 94,
fig. 26 ) portheo

Male genital process moderately curved, proportionately

slender, width of process averaging 28% of process-length.

( PI. 94, fig. 27 ) signoreti

22(20). Male genital process simply and weakly capitate, the inner
terminal corner but slightly more prominent than outer ter-
minal corner. ( Pi. 94, fig. 28 ) hydor

Male genital process noncapitate, inner terminal corner

rounded or angulate, in the former case, strongly produced
over outer terminal angle ( lunatus ) 23

1286 The University Science Bulletin

23(22). Male genital process rather goosehead or doghead-shaped,
the inner terminal corner bluntly prolonged; female sub-
genital plate-apex with median angles only moderately pro-
duced caudad of lateral angles. ( Pi. 94, fig. 29 ) , lunatus lunatus

Male genital process parallel-sided to apex, where the inner

terminal corner is sharply angulate, no wider at apex than
over stem; female subgenital plate-apex with median angles
rather markedly produced caudad of lateral angles. (Pi.
94, fig. 30 ) lunatus menoides

24(15). Female subgenital plate asymmetrical on left side, where a
prominent flap occupies the border beyond (laterad of) the
left lateral angle; right border scarcely produced. (Pi. 94,
fig- 31 ) dilatus

Female subgenital plate not markedly or noticeably asym-
metrical from one lateral border to the other 25

25(24). Lateral apical angles of female subgenital plate long and
sharp, spinosely produced, extending considerably caudad
of low, rounded, median angles; male genital process large,
long, and slimly goosehead-shaped, the inner terminal corner
strongly produced, the narrowly rounded end nearly over-
lapping adjacent edge of the process-bearing stemite. (PI.
94, fig. 32 ) cosmius

Lateral apical angles of female subgenital plate shorter than,

or even with, the median angles; species without such a male
genital process 26

26(25). Lateral apical angles of female subgenital plate lower than
median angles or sinuosities ( i. e., more cephalad in position
than the medians ) . ( PI. 94, fig. 33 ) guttatipennis

Lateral apical angles of female subgenital plate approxi-
mately even with the median sinuosities 27

27(26). Lateral apical angles of female subgenital plate long, com-
paratively narrow, sharp and spinosely produced, even with
median, low-rounded angles or sinuosities; median angles set
close together; male genital process progressively narrowing
to tip, inner terminal corner enormously produced into a
straight-edged long process, somewhat like a greatly exag-
gerated, thin foot. ( Pi. 94, fig. 34 ) fuscus

Lateral apical angles of female subgenital plate short, blunt,

often hardly angulate, even with median, low-rounded angles
( i. e., level with the medians ) ; median angles wide-spaced,
as close, or closer to, lateral angles (as, than) to each other;
male genital process not so shaped 28

28(27). Male genital process weakly curved, proportionately broad,
width of process averaging 48% of length of process (using
greatest width in apical half of process, and curving length
of process from tip to base along inner margin ) portheo

Male genital process moderately curved, proportionately

more slender, width of process averaging 28% of process-
length signoreti

La Rivers: The Ambrysus of Mexico
PLATE XCIII

1287

I PYGMAEUS 2 CAUGfNOSUS 3 MELANOPTERUS 4 MELANOPTERUS

5 PARVICEPS 6 PUDICUS 7 BARBERI 8 ABORTUS

9 HUNGERFORDI 10 HUNGERFORDI II TRIUNFO 12 TR1UNF0

13 ANGULARIS 14 ANGULARIS 15 PUNCTICOLLIS 16 PUNCTICOLLIS

17 PULCHELLUS 18 VANDU2EEI 19 BUENOI 20 AUSTRAUS

MEXICAN AMBRYSUS

1288

The University Science Bulletin
PLATE XCIV

21 SCALENUS

2 2 CONVEXUS 23 INFLATUS

24 INFLATUS

25 MAGNICEPS

26 PORTHEO

27 SIGNORETI 28 HYOOR 29 LUNATUS

30 MENOIOES

31 DILATUS

32 COSMIUS

d.--;

34 FUSCUi. 35 PARVICEPS

MEXICAN AMBRYSUS

La Rivers : The Ambrysus of Mexico 1289

DESCRIPTIONS OF SPECIES

Ambrysus pygmaeus sp. nov.

General appearance: small for the genus, slim, streamlined — size
7.0-7.5 mm. long, 3.75-4.0 mm. wide. Dorsum unicolorously deep
brownish; entire surface densely, shallowly, roughly and coarsely
punctate, shiny. Venter unicolorous, slightly darker anteriorly.

Head: densely and coarsely punctate, strongly but smoothly pro-
tuberant between eyes. Eyes blackish, bulging markedly above
head surface, the most prominent in this respect of the entire genus,
resulting in a "popeyed" appearance. Head ratios are:

(1) total length to width (including eyes), 57:: 79 (72%)

(2) anterior distance between eyes to posterior distance, 37:: 52 (71%)

(3) anterior distance between eyes to inner eye length, 37:: 30 (77%)

(4) posterior distance between eyes to greatest length of head posterior
to this line, 52:: 22 (42%).

Pronotum: roughly, coarsely punctulate, a slight transverse rugos-
ity evident in anterior central part just posterior to greatest penetra-
tion of head into pronotum; lateral edges microscopically serrate,
moderately curved from front to hind angles — percent of curvature,
expressed in terms of straight-line distance between anterior and
posterior lateral angles and greatest vertical distance between this
baseline and line of curvature, is 12-13% (av. 80:: 10). Dorsal pro-
notal ratios are:

( 1 ) width between anterior angles to width between posterior angles, or,
in this case, widest part of pronotum, which, unlike most members of
the genus, is not synonymous with posterolateral angles, but is slightly
anterior to them, is 81:: 143 (57%)

(2) median length to greatest width, 46:: 143 (32%)

(3) distance between anterior and posterior angles on same side to perpen-
dicular distance between anterior angle and baseline of pronotum,
67::75 (89%).

Scutellum: unicolorous reddish-brown, rough-punctate. In nor-
mal position, i. e., approximately on a plane surface with remainder
of body, ratio of three sides, anterior and two laterals, is 96::68::68.

Hernelytra: uniform, deep, reddish-brown, densely punctate.
Embolium long, narrow (length to width 81:: 20 = 25%), peculiarly
shaped in that the anterior end is nearly as wide as the posterior
end, whereas in all other Ambrysi known to me, the anterior end is
characteristically and conspicuously narrower than the posterior.
Hernelytra rather strongly exposing lateral connexiva, and not quite
attaining abdominal tip.

Venter: deep reddish-brown. Connexival segments nonspinose,
edges black, nonserrate, curvature weak but present.

1290 The University Science Bulletin

Legs: Prolegs — brownish to yellow; ratio of length to greatest
width of ventral surface of femur 75:: 51 (68%): combined tibio-
tarsus, when closed, slightly exceeding adjacent (proximal) end of
femur, but not conspicuously so.

Mesolegs — yellow to brown; femur stout for the genus, ratio of
length to median width of ventral surface 71::14 (20%) — 1.80 mm.
long; tibia comparatively stout for the genus — distal end ventrally
with two prominent transverse rows of spines set in solid rows
across tibial apex, the last row at extreme tip — ratio of length to
median width 91: :12 ( 13%)— length 1.60 mm.

Metalegs — yellow to brown; femur stout for the genus, ratio of
length to median width of ventral surface 86::15 (17%) — length
2.0 mm. Tibia stout for the genus, distal end ventrally with two
prominent transverse rows of spines set in solid rows across tibial
width, the last row at extreme tibial apex — ratio of length to median
width of ventral surface 86::7.5 (9%) — length 1.5 mm.

Distribution: see types.

Type locality data: Mexico — Mexico (Temescaltepec), 5(vi)33,
H. E. Hinton — R. L. Usinger.

Location of types: holotypic male and allotype in the collections
of the California Academy of Sciences, San Francisco; four para-
types in the Snow Museum, University of Kansas, Lawrence; six
paratypes in the collection of Robert L. Usinger, Berkeley, Cali-
fornia; two paratypes in the collection of the writer, Reno, Nevada.

This small species is perhaps the most distinctive member of the
oblongulus group of Ambrysus and a more exact knowledge of its
position in the group waits on further collecting, particularly in
southern Mexico, south to Costa Rica.

Ambrysus circumduct us caliginosus Usinger
Ambrysus caliginosus Usinger 1946, Univ. Kans., Sci. Bull. 31(1):190.

General appearance: moderately small, slim, streamlined — size
9.0-9.5 mm. long, 5.0-5.25 mm. wide. Dorsum weakly bicolored,
lighter anteriorly; entire surface coarsely punctate. Venter uni-
formly deep reddish-brown, with some vaguely lighter areas an-
teriorly.

Head: yellowish-brown, coarsely punctate, distinctly but smoothly
protuberant before eyes. Eyes rather prominently protuberant
above head surface. Head widely and deeply set into anterior
pronotal border. Head ratios are:

( 1 ) total length to width, 60: :85 (88%)

(2) anterior distance between eyes to posterior distance, 39:: 57 (68%)

La Rivers : The Ambrysus of Mexico 1291

(3) anterior distance between eyes to inner eye margin, 39:: 31 (74%)

(4) posterior distance between eyes to greatest length of head posterior to
this line 57:: 23 (40%).

Pronotum: brownish, coarsely punctate, extreme edge blackish;
lateral edges distinctly, minutely and closely serrate, moderately
curved — percent of curvature 12-13% (av. 95:: 12). Dorsal pronotal
ratios are:

( 1 ) width between anterior angles to greatest width of pronotum ( see A.
pygmaeus for qualifications applicable here), 44:: 89 (49%)

(2) median length to greatest width 31:: 89 (35%)

(3) distance between anterior angles and point of widest spread of pro-
notum on the same side to perpendicular distance between anterior
angle and baseline of pronotum 41:: 48 (85%).

Scutelhtm: deep, uniform reddish-brown, minutely punctulate;
ratio of three sides 123: : 88: : 88.

Hemelytra: uniform punctate; blackish in color with seven dis-
tinct lighter areas as follows: (a) two at bases of elytra; (b) em-
bolia; (c) one medially just behind scutellum; (d) two in posterior
third of hemelytra. Embolium long, narrow (length-to-width
118: :27 = 23%), shaped somewhat as in A. pygmaeus but anterior
third not as wide, proportionately. Hemelytra strongly exposing
connexiva, and just barely attaining abdominal tip to falling just
short of tip.

Venter: deep reddish-brown. Connexiva spineless, characterized
by a slight but definite increase in lateral projection of postero-
lateral angles from anterior to posterior. Connexiva distinctly but
minutely serrate, particularly on posterior segments.

Legs: Prolegs — yellow-brown; ratio of length to greatest width of
ventral femoral surface 92:: 54 (58%); combined tibiotarsus, when
closed, slightly, but definitely, overlapping adjacent (proximal) end
of femur.

Mesolegs — yellow-brown; femur somewhat stout for the genus,
ratio of length to median width of ventral surface 95:: 12 (13%) —
length 2.40 mm.; tibia somewhat stout, distal end ventrally with
two prominent transverse rows of spines set in solid rows across
width of tibia, the last row at extreme tip — ratio of length to median
width 78: :8 ( 10%)— length 2.10 mm.

Metalegs — yellow-brown; femur comparatively stout, ratio of
length to median width 110: :18 (16%)— length 2.85 mm. Tibia
somewhat stout, distal end ventrally with two prominent transverse
rows of spines set solidly across tibial apex, the last row at extreme
tibial end — ratio of length to median width of ventral surface 116: :9
(8%)— length 3.0 mm.

1292 The University Science Bulletin

Recorded distribution: central Mexico (Usinger 1946).

Type locality data: Mexico — "Mexico ( Tejupilco ) , June 18-21,
1933, H. E. Hinton and R. L. Usinger" (Usinger 1946).

Location of types: "Holotype, male, and allotype, female (Cali-
fornia Academy of Sciences)," (Usinger 1946), San Francisco.

Specimens examined: the types and several paratypes from the
type locality, the latter in the Usinger collection, Berkeley, Cali-
fornia.

I am unable to separate caliginosus from circumcinctus on purely
structural characters, and although they seem relatively distinct
when examined side by side, the basis for such distinction, size and
color, is not reliable in Ambrysus; the smaller specimens of cali-
ginosus grade into the larger specimens of circumcinctus in both
respects, and the male and female accessory genitalia, so valuable
in species delineation throughout the remainder of the genus, are
not even subspecifically distinct. At the present, we can only
regard caliginosus as the southern unit of circumcinctus, which
latter was originally described by Montandon from a population
centering on Kerrville, Texas, not far from the Mexican border
(1910). In all probability, A. circumcinctus circumcinctus occurs
in northern Mexico as well, but is so far unreported. A. c. circum-
cinctus averages smaller and lighter than A. c. caliginosus, a fact
which does not disturb any of our theories concerning the changes
undergone by a homogeneous population as it proceeds from the
tropics into colder regions to the north.

Ambrysus melanopterus Stal

Ambrysus melanopterus Stal 1862, Stet. Ent. Zeit. 23:459; 1876, Enum. Hemipt.

5:143.
Ambrysus melanopterus, Uhler, 1876, Bull. U.S.C.S. 1:337.
Ambrysus melanopterus, Montandon, 1897, Verh. zool.-bot. Ges. Wien, 57:12,

19; 1909, Bull. Soc. Sci. Buc.-Roum. 17(5-6) :316.
Ambrysus melanopterus, Champion, 1900, Biol. Centr.-Amer., Heter. 2:357.
Ambrysus melanopterus, Snow, 1906, Trans. Kans. Acad. Sci. 20(1):180.
Ambrysus melanopterus, Kirkaldy and Bueno, 1908, Proc. Ent. Soc. Wash. 10:

186.
Ambrysus melanopterus, Van Duzee, 1917, Univ. Calif. Publ. Ent. 2:458.
Ambrysus melanopterus, Hungerford, 1919, Univ. Kans., Sci. Bull. 11:201.
Ambrysus melanopterus, La Rivers, 1949, Bull. S. Calif. Acad. Sci. 47(3) :108;

1951; Univ. Calif. Publ. Ent. 8(7) :295-297.

General appearance: slightly more than medium-sized for the
genus, moderately slim, and streamlined — size 10-12 mm. long, 5.5-
6.5 mm. wide. Dorsum strongly mottled anteriorly, unicolorous
brownish posteriorly, entire surface minutely rugose-punctulate.
Venter lighter posteriorly.

Head: weakly punctate, longitudinally streaked with alternate

La Rivers : The Ambrysus of Mexico 1293

black-brown and yellowish stripes, smoothly but distinctly pro-
tuberant between eyes. Eyes very slightly protuberant above head
surface, outer and posterior margins meeting at posterolateral
corner of eye in a blunt, but definite, angle, and not blending uni-
formly, smoothly and evenly into one continuous curvature as in
the preceding species; basal or posterior eye margin conspicuously
rimmed with a chitinous bar or raised edge which projects slightly
laterad as posterolateral eye angle to form a weak but discernible
angle, in marked contrast to other members of the genus in this
area. Head ratios are:

(1) total length to width 68:: 115 (59%)

(2) anterior distance between eyes to posterior distance 42:: 62 (68%)

(3) anterior distance between eyes to inner eye length 42: :42

(4) posterior distance between eyes to greatest length of head posterior
to this line 62:: 22 (38%).

Pronotum: shallowly punctate laterally, weakly but distinctly
transversely rugulose behind deepest head penetration; lateral edges
narrowly pale-yellow, inwardly bordered by a wider area of rich
deep brown or black, this bordered in turn inwardly by a pale area
well supplied with small, conspicuous, uniform, brownish-black
spots which may coalesce at one or two places to form short streaks
— the centrum is characterized by a "V"-shaped yellowish area
occupied centrally by a dark-brown spot; edges smooth, non-
serrate, moderately curved — percent of curvatures 10-12% ( av.
128:: 14); posterolateral angles distinct but rounded. Dorsal pro-
notal ratios are:

(1) width between anterior angles to width between posterior angles 56::
115 (49%)

(2) median length to greatest width 44:: 15 (38%)

(3) distance between anterior and posterior angles on same side to per-
pendicular distance between anterior angle and baseline of pronotum,
64::60 (94%).

Scutellum: black, minutely punctulate, ratio of three sides
82::61::62.

Hemehjtra: deep brown to black, minutely punctulate; embolium
long, narrow (length-to-width 138: : 30 = 22%), markedly longitudi-
nally bicolored, outer third light. Hemelytra moderately exposing
connexiva, and not quite, to just, attaining abdominal tip.

Venter: bicolored, abdomen appearing lighter than remainder.
Connexiva spineless at posterolateral angles, edges smooth, non-
serrate; curvature discernible but weak.

Legs: Prolegs — yellow to brown; ratio of length to greatest width
of ventral surface of femur 120:: 72 (62%); combined tibiotarsus,

1294 The University Science Bulletin

when closed, quite markedly overlapping adjacent (proximal) end
of femur.

Mesolegs — yellow to brown; femur long, narrow, ratio of length
to median width of ventral surface 113:: 18 (16%) — length 3.0 mm.;
tibia narrow, distal end ventrally with two prominent, transverse
rows of spines set in solid rows across width of tibia, the last row at
extreme tip — ratio of length to median width 99: :8 (9%) — length
2.5 mm.

Metalegs — yellow to brown; femur narrow, ratio of length to
median width of ventral surface 132: :20 (15%) — length 3.1 mm.;
tibia narrow, distal end ventrally with two prominent transverse
rows of spines set solidly across tibial width, the last row at extreme
tip — ratio of length to median width of ventral surface, 150:: 10
(7%)— length 3.7 mm.

Recorded distribution: southern Mexico to southwestern United
States.

Type locality data: "Mexico."

Location of type: Royal Stockholm Museum, Sweden.

Specimens examined: Guatemala: — Panzos, (iv)20, Jordan
(USNM). Mexico: — Chiapas (Mt. Obando) ( quiet pool of small,
swift stream), 15 (iv)40, el. 3,000 ft., H. M. Smith (UK); Guerrero
(S. of Chilpancingo between Cajones and Rincon), 1 (vii)32, H. M.
Smith (UK); Mexico (Tejupilco), 3(vii)33, H. E. Hinton— R. L.
Usinger (RLU); Tres Marias Islands (Maria Madre Island, Arroyo
Hondo), 17(v)25, G. D. Hanna (RLU). United States :— Ari-
zona: Santa Cruz County (Patagonia), 7(ix)38, C. L. Hubbs &
family (UM).

Judged solely on the basis of the material examined, this dis-
tinctive species does not seem to be particularly common anywhere,
but its center of distribution is patently Mexico.

Ambrysus parviceps Montandon

Ambnjsus parviceps Montandon 1897, Verh. zool.-bot. Ges. Wien, 47:17; 1909,

Bull. Soc. Sci. Buc.-Roum. 17(5-6) :320.
Ambrysus parviceps, Champion, 1900, Biol. Centr.-Amer., Heter., 2:356.
Ambrysus parviceps, Usinger, 1946, Univ. Kans., Sci. Bull., 31(1) :188.
Ambrysus infuscatus Usinger 1946, Univ. Kans., Sci. Bull., 31(1):188.

General appearance: small, rotund species — size 8.0-8.5 mm. long,
5.5-6.0 mm. wide. Dorsum strongly mottled, lighter anteriorly.
Venter light golden-yellow, well furred, dark blotched anteriorly.

Head: groundcolor light yellow, with longitudinal darker, ob-
scure blacker markings superimposed; shiny, impunctate; front
widely and smoothly protuberant between eyes. Eyes with postero-

La Rivers: The Ambrysus of Mexico 1295

lateral angle discernible, a chitinous bar bordering posterior eye
margin; from behind, eyes only slightly protuberant above head
surface. Head ratios are:

( 1 ) total length to width 57 : : 80 ( 71% )

(2) anterior distance between eyes to posterior distance 36:: 46 (78%)

(3) anterior distance between eyes to inner eye margin 36:: 37

(4) posterior distance between eyes to greatest length of head posterior to
this line 46:: 14 (30%)

(5) anterior distance between eyes to greatest length of head anterior to
this line 36:: 8 (17%).

Pronotum: highly polished, impunctate or nearly so; groundcolor
light yellow with varied mottling of suffuse brownish; edges smooth,
unserrate, densely set with long, yellowish hairs; percent of curva-
ture of pronotal edge 20% (av. 50:: 10); posterolateral angles usually
present but not conspicuous and not obviously at widest point of
pronotum. Dorsal pronotal ratios are:

(1) width between anterior angles to width between posterior angles, 42::
100 (42%)

(2) median length to greatest width 28:: 100 (28%)

(3) distance between anterior angle and posterior angle on same side to
perpendicular distance between anterior angle and baseline of pronotum
45::45.

Scutellum: reddish-brown, lighter along lateral margins; shiny,
shallowly punctate; ratio of three sides 67: :49: :47.

Hemehjtra: color of scutellum, with same shagreenation of white-
spotted punctation. Embolium long, of normal width (length-to-
width 104: :33 = 32%), light yellow in anterior two thirds; in un-
rubbed specimens, a conspicuous mat of long, yellow pilosity
borders lateral edge. Hemelytra moderately exposing connexiva,
and just attaining abdominal tip.

Venter: Connexiva spinose, the posterolateral angles sharp on all
segments except I; edges of segments I-II absolutely smooth, un-
serrated, edges of remaining segments serrate, becoming progres-
sively stronger caudally.

Legs: Prolegs — whitish to yellow; ratio of length to greatest
width of ventral surface of femur 66:: 45 (68%); combined tibia-
tarsus, when closed, not overlapping adjacent (proximal) end of
femur.

Mesolegs — whitish to yellow; femur long, narrow, ratio of length
to median width of ventral surface 81:: 16 (20%) — length 2.1 mm.;
tibia narrow, distal end ventrally with one complete, transverse row
of spines set in a solid row across extreme tip, instead of the usual
two rows — in this case, the second row is reduced to two or three

52—6490

1296 The University Science Bulletin

spines on each side, leaving the middle section bare — ratio of length
to median width 97: :15 ( 15%)— length 1.5 mm.

Metalegs — whitish to yellow; femur long, ratio of length to me-
dian width 115: :19 (17%)— length 2.60 mm.; tibia long, distal end
ventrally with one complete prominent row of spines set in a solid
row across the extreme tip instead of the usual two rows — in this
case, the second row is reduced to two or three spines on each side,
leaving the middle section bare — ratio of length to median width of
ventral surface 115: :8 (7%)— length 2.50 mm.

Recorded distribution: central Mexico.

Type locality data: "Mexique."

Location of types: "K. K. Hofmuseum, Vienna, Austria."

Specimens examined: Mexico: — Chihuahua (Rio San Pedro be-
tween Chihuahua and Naica), 22(vi)34, Smith and Dunkle (UK);
(Corriente), 24(viii)37, H. D. Thomas (UK); Guerrero (438 kilom.
S. of Mexico [City]), l(xi)36, H. D. Thomas (UK); (Acapulco),
12(vii)37, H. D. Thomas (UK); (Acapulco), 28(viii)38, L. I.
Lipovsky (UK); Michoacan (Uruapan) (ElSabino) 28(vii)36,
H. D. Thomas (UK); Mexico (Tejupilco), 15(vi)33, H. E. Hinton
— R. L. Usinger (RLU).

The types of A. infuscatus in the California Academy of Sciences,
San Francisco, have been examined as well as paratypes of the
same name in the Usinger collection, Berkeley, California. Dr.
Usinger was able to compare a specimen of A. infuscatus with the
type of A. parviceps in Vienna, and noted the synonymy.

Ambry sus pudicus pudicus Stal

Ambnjsus pudicus Stal 1862, Stet. Ent. Zeit, 23:460.

Ambrysus pudicus, Uhler, 1894, Proc. Calif. Acad. Sci., ser. 2, 4:291.

Ambnjsus pudicus, Montandon, 1897, Verh. zool.-bot. Ges. Wien, 47:12, 17;

1909, Bull. Soc. Sci. Buc.-Roum. 17 (5-6): 320.
Ambrysus pudicus, Champion, 1900, Biol. Centr.-Amer., Heteropt. 2:356.
Ambnjsus pudicus, Van Duzee, 1917, Univ. Calif. Publ. Ent. 2:458.
Ambrysus pudicus, Usinger, 1946, Univ. Kans., Sci. Bull., 31(1):187.
Ambrysus pudicus, La Rivers, 1949, Bull. S. Calif. Acad. Sci. 47(3) :108.

General appearance: small, rotund — size 7.0-7.5 mm. long, 4.5
mm. wide. Dorsum strongly bicolored, light and glistening an-
teriorly. Venter light yellowish, well furred with hydrofuge pile.

Head: groundcolor light yellow; shiny, impunctate, with longi-
tudinal brown streaking on centrum; front widely but smoothly
protuberant between eyes. Eyes somewhat triangular, the straight
inner and outer margins forming two sides of the triangle, the
smoothly curved posterior margin connecting the two; chitinous bar
bordering posterior eye margin quite conspicuous; viewed behind,

La Rivers: The Ambrysus of Mexico 1297

eyes only slightly protuberant above head surface. Head ratios
are:

(1) total length to width (including eyes) 53::78 (68%)

(2) anterior distance between eyes to posterior distance 33:: 48 (69%)

(3) anterior distance between eyes to inner eye margin 33:: 32

(4) posterior distance between eyes to greatest length of head posterior to
this line 48:: 18 (38%)

(5) anterior distance between eyes to greatest length of head anterior to
this line 33::4.5 (14%).

Pronotum: highly polished, impunctate; groundcolor light yel-
lowish, with a varied mottling of suffuse brownish areas and dots;
lateral edges absolutely smooth, sparsely pilose, and smoothly
curved from anterior to posterior angles, percent of curvature 12-
13% (av. 85:: 11); posterolateral angles rather prominent for this
section of the genus, representing approximately the widest point
of pronotum, and distinctly posterior to the thin, transverse pos-
terior pronotal line. Dorsal pronotal ratios are:

(1) width between anterior angles to width between posterior angles 78::
165 (47%)

(2) median length to greatest width 54:: 165 (33%)

(3) distance between anterior angle and posterior angle on same side to
perpendicular distance between anterior angle and baseline of pronotum
77::79 (97%).

Sciitellwn: glistening but minutely punctate; blackish, whitish
along edges; ratio of three sides 115: :74: :70.

Hemelytra: brownish-black, somewhat lighter along scutellar
margins, generally with two well-defined light spots on border be-
tween membrane and corium, and lighter on embolium; shining but
coarsely punctate. Embolium long, of normal width (length-to-
width 94: :30 = 32%), light yellow in anterior two thirds; edge vari-
ably equipped with long, golden hairs. Hemelytra narrowly ex-
posing connexiva, which are densely pilose, with an occasional
longer yellow hair interspersed; hemelytra generally slightly ex-
ceeding abdominal apex.

Venter: bicolored, abdomen lighter yellow than sterna; connexiva
sharply, finely spinose except segment I, the posterolateral angles
produced; edges of abdominal segments I-II smooth, glabrous, in
contrast to marked pilosity and serration of remaining segments.

Legs: Prolegs — whitish to yellow; ratio of length to greatest
width of ventral femoral surface 72:: 46 (64%); combined tibia-
tarsus, when closed, markedly overlapping adjacent (proximal) end
of femur.

Mesolegs — whitish to yellow; femur long, narrow, ratio of length

1298 The University Science Bulletin

to median width of ventral surface 68::12 (18%) — length 1.60 mm.;
tibia narrow, distal end ventrally with two prominent transverse
rows of spines set in solid rows across tibial width, the last row at
extreme tip — ratio of length to median width 84:: 10 (12%) — length
1.35 mm.

Metalegs — yellow to white; femur long, narrow, ratio of length
to median width of ventral surface 84:: 15 (18%) — length 2.10 mm.;
tibia long, narrow, distal end ventrally with two prominent trans-
verse rows of spines set in solid rows across tibial width, the last
row at extreme tibial apex — ratio of length to median width of ven-
tral surface 86: :6 (7%)— length 2.30 mm.

Recorded distribution: California, Wyoming and Mexico.

Type locality data: "Mexico."

Location of types: Royal Stockholm Museum, Sweden.

Specimens examined: Mexico: — Guerrero (Acapidco), 28 ( viii )
38, L. I. Lipovsky (UK); (Rio Papagato, 387 km. S. of Mexico
[City]), 31(x)36, H. D. Thomas (UK); (438 km. S. of Mexico
[City]), l(xi)36, H. D. Thomas (UK); Mexico (Tejupilco), 20(vi)
33, H. E. Hinton and R. L. Usinger (RLU). Guatemala: —
Sanarate, Kellerman (MCZ).

There is little doubt in my mind that the United States records of
this species in the literature are in error. In all probability several
species have been confused as A. pudicus in the United States. The
California records are probably based on A. californicus, while
small specimens of A. mormon, in light of our present knowledge,
could be the only material upon which Wyoming records of
A. pudicus could be based. A. woodburyi could also have been
easily confused with A. pudicus in the early, primitive stages of
Ambrysus taxonomy.

Ambrysus pudicus barbed Usinger

Ambrysus barberi Usinger 1946, Univ. Kans., Sci. Bull. 31(1):189.

General appearance: a medium-sized subspecies, nearly immacu-
late— size 7.5-9.0 mm. long and 4.75-6.0 mm. wide.
Head: ratios are:

( 1 ) total length to width 61 : :89 (69%)

(2) anterior distance between eyes to posterior distance 40:: 55 (73%)

(3) posterior distance between eyes to greatest length of head posterior to
this line 55:: 19 (35%).

Pronotum: ratios (dorsal) are:

( 1 ) width between anterior angles to width between posterior angles
52::106 (49%)

La Rivers: The Ambrysus of Mexico 1299

(2) median length to distance between posterior angles 35:: 106 (33%)

(3) width between anterior angles to distance between anterior angle and
posterior baseline of pronotum 52:: 50 (96%).

Scutellum: ratio of three sides 69: :50: :49.

Hemelytra: essentially as in A. pudicus pudicus.

Venter: Connexival posterolateral angles distinctly but weakly
spinose except those of segment I, and increasing slightly but pro-
gressively posteriorly in size. Connexival margins very strongly
serrate on all segments except I-II which are always perfectly
smooth.

Legs: as in A. pudicus pudicus, but larger.

Recorded distribution: northeastern Mexico (Usinger 1946).

Type locality data: "Victoria, Tampico, Mexico, December 10,
1909, F. E. Bishopp, collector" (Usinger 1946).

Location of types: "Holotype, male and allotype, female (U. S.
National Museum)" (Usinger 1946).

Specimens examined: Mexico: — Tamaulipas (Victoria), 10(xii)
09, F. C. Bishopp (USNM) ( = allotype), (5 mi. S. of Victoria),
5(xi)36, H. D. Thomas (UK); Mexico (Tejupilco), 15(vi)33, H. E.
Hinton and R. L. Usinger (RLU). (The allocation of the type
locality, Victoria, to "Tampico" in the original description should
be changed to Victoria in Tamaulipas, as indicated above. ) United
States:— Texas (Hidalgo County— Mc Allen), 20(xi)32, L. D. Tut-
hill (UK).

The larger A. pudicus barberi, with our present knowledge, must
be considered the northern component of A. pudicus, the smaller
typical subspecies seemingly a central Mexico-to-Central American
entity.

Ambrysus abortus sp. nov.

General appearance: small, rotund species — size 7.5 mm. long,
5.25 mm. wide. Dorsum generally lighter anteriorly, darker and
mottled posteriorly, glistening. Venter generally light, with con-
spicuous dark areas anteriorly.

Head: groundcolor light yellow, shining, glistening, impunctate,
with two rather faded, median, longitudinal rows of brown dots on
centrum; front widely but smoothly protuberant before eyes. Eyes
with posterior bordering chitinous bar somewhat inflated at postero-
lateral angles and extending slightly laterad; eyes but slightly sin-
uous above general head surface. Head ratios are:

( 1 ) total length to width 53: :82 ( 65%)

(2) anterior distance between eyes to posterior distance 34:: 48 (71%)

(3) anterior distance between eyes to inner eye margin 34:: 35

1300 The University Science Bulletin

(4) posterior distance between eyes to greatest length of head posterior
to this line 48:: 15 (31%)

(5) anterior distance between eyes to greatest length of head anterior to
this line 34:: 3.5 (10%).

Pronotum: shiny but not polished, irregularly punctate; ground-
color light yellow with a varied mottling of suffuse brown spots;
lateral edges smooth, unserrate, conspicuously set with long but
not dense yellowish hairs; percent of curvature 14% (av. 85:: 12);
posterolateral angles quite prominent and definite, constituting
widest part of pronotum, and are definitely situated posterior to the
thin, transverse, posterior pronotal line. Dorsal ratios are:

( 1) width between anterior angles to width between posterior angles 42:: 94
(45%).

(2) median length to greatest width 26:: 94 (28%)

(3) distance between anterior angle and posterior angle on same side to
perpendicular distance between anterior angle and baseline of pro-
notum 43:: 42.

Scutellum: blackish with a reddish tinge; lateral edges narrowly
light yellow, surface shiny, but not polished, densely, shallowly and
roughly punctate. Ratio of three sides 61: :41: :41.

Hemelytra: suffusedly mottled with light and dark; punctured as
is scutellum. Embolium short and broad for the genus, length-to-
width 91:: 33 (36%), bicolored, anterior three fourths light yellow;
in unrubbed specimens a thin line of long, sparse yellow hairs along
outer edge which, in older individuals, may be entirely rubbed off.
Hemelytra moderately exposing connexiva, and just attaining ab-
dominal tip.

Venter: abdominal venter light yellow with a dense coat of short
hydrofuge hairs, sterna darker. Connexival posterolateral spines
distinguishable on all segments except I, the spines very definitely
becoming progressively larger caudad; spination is more in the
nature of an acute elongation of the angles than true spination; all
lateral connexiva except I-II strongly dentate.

Legs: Prolegs — whitish; ratio of length to greatest width of ven-
tral femoral surface 64:: 41 (64%); combined tibia-tarsus, when
closed, just attaining adjacent (proximal) end of femur.

Mesolegs — whitish; ratio of length to median width of ventral
femoral surface 70::15 (21%) — length 2.0 mm.; distal tibial end
ventrally with two transverse rows of spines set across tip, the distal
row completely solid across apex, the second or proximal row not
quite extending the full tibial width, leaving a gap near the outer
or anterior end — however, the second row is strong enough to be
considered a full row, in contrast to the condition described for

La Rivers: The Ambrysus of Mexico 1301

A. parviceps and A. hungerfordi where this row fades out medially
and the spinal remnants are considered as only secondary armature
— ratio of length to median width 58: :9 (16%) — length 1.25 mm.

Metalegs — whitish; ratio of length to median width of ventral
femoral surface 95:: 16 (17%) — length 2.25 mm.; distal tibial end
ventrally with two prominent transverse rows of spines set across
tip, the distal row completely solid across apex, the second or
proximal row not extending the full tibial width, leaving a gap near
the outer or anterior end — ratio of length to median width 95: :8
(8%)— length 2.25 mm.

Distribution: see types.

Type locality data: Mexico: — Mexico (Tejupilco), 15(vi)33,
H. E. Hinton and R. L. Usinger.

Location of types: Holotypic male and allotype in the collections
of the California Academy of Sciences, San Francisco; four para-
types in the Snow Museum, University of Kansas, Lawrence [three
paratypes from the type locality, one paratype from Corriente,
24(viii)37, H. D. Thomas (Mexico)]; fourteen paratypes (type
locality) in the collection of Robert L. Usinger, Berkeley, California;
two paratypes (type locality) in the collection of the writer, Reno,
Nevada.

Ambrysus hungerfordi hungerfordi Usinger
Ambrysus hungerfordi Usinger 1946, Univ. Kans., Sci. Bull. 31(1) :192.

General appearance: small, rotund species — size 7.0-7.75 mm.
long, 4.75-5.00 mm. wide. Dorsum bicolored, lighter anteriorly,
glistening but not polished. Venter strongly contrastingly colored
with blackish-browns, yellows, and, in some specimens, greenish.

Head: groundcolor light yellow, often with a greenish cast,
shiny, impunctate; front widely but smoothly protuberant between
eyes. Eyes somewhat angular at posterolateral angle in some
specimens, caused by an angularity of the caudal-bordering chi-
tinous bar; viewed from behind, eyes insignificantly protuberant
above head surface. Head ratios are :

(1) total length to width 52::80 (71%)

(2) anterior distance between eyes to posterior distance 31:: 45 (69%)

(3) anterior distance between eyes to inner eye margin 31:: 35 (88%)

(4) posterior distance between eyes to greatest length of head posterior
to this line, 45:: 13 (29%)

(5) anterior distance between eyes to greatest length of head anterior to
this line 31::4 (13%).

Fronotum: shiny, but not polished, roughly and shallowly punc-
tate; groundcolor light yellowish-brown, often with a greenish cast,

1302 The University Science Bulletin

with a varied mottling of suffuse brownish areas; edges smooth,
unserrate, conspicuously set with long, but not dense, yellowish
hairs; percent of curvature of pronotal edge 15% (86:: 13); postero-
lateral angles usually present, but not conspicuous, and approxi-
mately at widest point of pronotum. Dorsal ratios are:

( 1 ) width between anterior angles to width between posterior angles 41 : :87
(47%)

(2) median length to greatest width 29:. -87 (33%)

(3) distance between anterior angle and posterior angle on same side to
perpendicular distance between anterior angle and baseline of pro-
notum 39:: 40.

Scutellum: deep reddish-brown, shiny but not polished, roughly
punctate; ratio of three sides 117: :74: :73.

Hemelytra: darker than scutellum, weakly variegate-blackish and
brownish, often with a greenish tinge, punctation as in scutellum.
Embolium tending to be somewhat short and wide for the genus,
length-to- width 83:: 30 (36%), light on the anterior two thirds; a
thin but conspicuous line of long, sparse yellow hairs along outer
edge. Hemelytra moderately exposing lateral connexival margins,
and just attaining abdominal apex.

Venter: Connexival posterolateral angles sharply, rather finely
spinose, the spination quite evident on all segments except I; how-
ever, segment I, instead of completely lacking a spine, as is usual,
has a minute, microscopically sharp posterolateral angle, an un-
usual condition for this section of the genus; all connexiva except
I-II are strongly and markedly dentate or serrate, and the posterior
half of segment II may show incipient serration under very high
magnification.

Legs: Prolegs — yellow, yellow-brown, or greenish; ratio of length
to greatest femoral width (ventral surface), 102:: 65 (64%); com-
bined tibia-tarsus, when closed, not distinctly overlapping adjacent
( proximal ) end of femur.

Mesolegs — yellow, yellow-brown, or greenish; ratio of femoral
length to median width of ventral surface 71:: 13 (18%) — length
1.85 mm.; tibial distal end ventrally with one complete, prominent,
transverse row of spines set in a solid row across extreme tip ( as in
A. parviceps), instead of the usual two — in this case, the second
row is reduced to two-three spines on each side, leaving the middle
section free — ratio of length to median width 82:: 14 (17%) — length
1.40 mm.

Metalegs — yellow, yellow-brown to greenish; ratio of femoral
length to median width of ventral surface 93:: 15 (16%) — length

La Rivers: The Ambrysus of Mexico 1303

2.20 mm.; tibial transverse apical spination same as for mesolegs —
ratio of length to median width 92: :9 ( 10%)— length 2.20 mm.

Recorded distribution: Several localities in Mexico (Usinger
1946), one of which (Tejupilco in Mexico), refers rather to A.
abortus.

Type locality data: "San Antonio, Mexico, July 15, 1927, R. H.
Beamer" ( Usinger 1946 ) .

Location of types: "Holotype, male, and allotype, female (Snow
Museum, University of Kansas ) . . . A series of paratypes col-
lected at the same place by both R. H. Beamer and P. A. Readio"
(Usinger 1946).

Specimens examined: Mexico: — Michoacan (San Antonio), 15
(vii)27, P. A. Readio (UK) (paratypes); Sonora (Alamos) (Buro-
paco District), 23 (x)34, H. S. Gentry (RLU), (Navajoa), 5(iii)30,
D. Wright (RLU); (San Bernardo) (Rio Mayo Arroyo), 2(iii)35
(RLU).

Future collecting will probably show that this subspecies occurs
in extreme southwestern United States, as well.

Ambrysus hungerfordi triunfo subsp. nov.

Identical to the typical subspecies in all essential respects, differ-
ing chiefly in the structure of the tip of the female subgenital plate
( see illustration ) .

Distribution: see types.

Type locality data: Mexico: — Baja California (Between Sail Jose
del Cabo and Triunfo) (Albatross Expedition), 1911 (RLU).

Location of types: Holotype male and allotype in collections of
California Academy of Sciences, San Francisco; paratypes in the
collection of Robert L. Usinger, Berkeley, California.

Ambrysus hungerfordi angidaris subsp. nov.

Identical to the typical subspecies in all essential respects, differ-
ing chiefly in the structure of the tip of the female subgenital plate
( see illustration ) .

Distribution: see types.

Type locality data: Mexico: — Guerrero (Rio Agua 437 kilom. S.
of Mexico [City]), 31(x)36, H. D. Thomas (UK).

Location of types: Holotype male and allotype in the Snow Mu-
seum, University of Kansas, Lawrence.

When the variable population of A. hungerfordi has been ade-
quately sampled over its range, the two above-described subspecies
will very likely fade into the general background of the species; at

1304 The University Science Bulletin

the moment they are useful in depicting what are probably the
limits of variation in the species, and will stand as distinct until
more collecting brings to light such intermediate forms as exist in
the populations.

Ambrysus puncticollis Stal

Amhrysus puncticollis Stal 1876, Enum. Hemipt. 5:143.

Ambrysus puncticollis, Montandon, 1897, Verh. zool.-bot. Ges. Wien, 48:13,

20; 1909, Bull. Soc. Sci. Buc.-Roum. 17 (5-6): 322.
Ambrysus puncticollis, Van Duzee, 1917, Univ. Calif. Publ. Ent. 2:459.
Ambrysus puncticollis, Hungerford, 1919, Univ. Kans., Sci. Bull. 11:199, 203.
Ambrysus puncticollis, La Rivers, 1949, Bull. S. Calif. Acad. Sci. 47(3): 108.

General appearance: large for the genus, comparatively long and
narrow — size 13.0-15.5 mm. long, 7.0-9.0 mm. wide. Dorsum weakly
bicolored, lighter and glistening anteriorly. Venter yellowish to
golden, lighter posteriorly.

Head: smooth, polished, impunctate; groundcolor yellow-brown,
two longitudinal median rows of brown dots occupying centrum;
front conspicuously protuberant between eyes. Eyes, viewed be-
hind, slightly but definitely protuberant above head surface. Head
ratios are:

(1) total length to width 42:: 66 (64%)

(2) anterior distance between eyes to posterior distance 29:: 42 (69%)

(3) posterior distance between eyes to greatest length of head posterior to
this line 42:: 16 (38%).

Pronotum: glistening, polished, rather densely set with small,
shallow impressions; groundcolor yellow-brown, suffuse brown areas
and dots occupying centrum; lateral edges smooth, unserrate, usu-
ally with a very few, moderately long hairs; percent of lateral curva-
ture 13% (av. 75:: 10); posterior lateral angles well rounded, dis-
tinctly posterior to thin, posterior, transverse pronotal line. Dorsal
ratios are:

(1) width between anterior angles to greatest pronotal width 68:: 138 (49%)

(2) median length to greatest width 54:: 138 (39%)

(3) width between anterior angles to distance between anterior angle and
posterior baseline of pronotum 68:: 75 (91%).

Scutellum: deep reddish-brown, more reddish than hemelytra;
surface shiny but not polished, shagreened with dense, shallow and
rough punctation; ratio of three sides 90: : 62: : 62.

Hemelytra: generally dark brown with a reddish tinge, some
lightening along emboliar margin; surface shiny but not polished,
shagreened as is scutellum. Embolium long and narrow for the
genus, length-to-width 82:: 23 (28%), with no marginal pilosity.
Hemelytra rather strongly exposing lateral connexival edges, which
bear some pilosity of yellow hairs more or less concentrated at the

La Rivers: The Ambrysus of Mexico 1305

segmental junctures; hemelytra just, or not quite, attaining ab-
dominal tip.

Venter: Connexival posterolateral spines large and prominent on
all segments except I, which is spineless; connexival edges smooth,
unserrate, weakly curved.

Legs: Prolegs — yellow to amber, often with a greenish tinge;
ratio of femoral length to greatest width of ventral surface 89:: 51
(74%); combined tibiotarsus, when closed, slightly but definitely
exceeding adjacent ( proximal ) end of femur.

Mesolegs — yellow to amber, occasionally with a greenish tinge;
ratio of femoral length to median width of ventral surface 70:. -12
(17%) — length 3.0 mm; tibia distally and ventrally with two promi-
nent, complete, transverse rows of spines set solidly across apex —
ratio of length to median width of ventral surface is 116: :11 (9%)
— length 3.0 mm.

Metalegs — yellow to amber, occasionally greenish; ratio of length
to median width of femur 81:: 14 (17%), comparatively stout for the
genus — length 3.50 mm.; tibial apical transverse spination same as
for mesolegs — ratio of length to median width of ventral surface
114: :8 (7%)— length 5.0 mm.

Recorded distribution: southwestern United States.

Type locality data: "Texas."

Location of types: Royal Stockholm Museum, Sweden.

Specwiens examined: Mexico: — Sonora (Alamos) (Buropaco
District), 23(x)34, H. S. Gentry (RLU); (San Antonio Colonial),
15(vii)27, P. A. Readio (UK). United States: — Arizona — Coco-
nino County (Bill Williams Fork, August, F. H. Snow) (UK);
Maricopa County 7(viii)27, P. A. Readio (UK); Yavapai County
(Camp Verde), 2(ix)38, C. L. Hubbs & Family (UM); (Tonto
Creek), 15(ix)36, Hubbs-Schultz (UM). Texas— Bexar County
( Helotes ) , 1 ( vii ) 17, P. A. Munz ( UK ) .

Labels on specimens which have come to me from other parts of
the United States show that A. puncticollis and A. melanopterus
have been commonly confused in the past, and it is no longer certain
how much of that confusion has shown up in the literature in the
way of erroneous distributional listings.

Ambrysus pulchellus Montandon

Ariibrysus pulchellus Montandon 1897, Verh. zool. bot. Ges. Wien, 47:11, 16;

1909, Bull. Soc. Sci. Buc.-Roum. 17(5-6) :319.
Ambrysus pulchellus, Champion, 1900, Biol. Centr.-Amer. Hemip. 2:354-361.
Ambrysus pulchellus, Van Duzee, 1917, Univ. Calif. Publ. Ent. 2:458.
Ambrijsus pulchellus, Hungerford, 1919, Univ. Kans., Sci. Bull. 11:198, 200.
Ambrysus pulchellus, Usinger, 1946, Univ. Kans., Sci. Bull. 31(1):187.
Ambrysus nitidulus Montandon 1909, Bull. Soc. Sci. Buc.-Roum. 17(5-6) :319,

326.

1306 The University Science "Bulletin

Ambrysus nitidulus, Usinger, 1946, Univ. Kans., Sci. Bull. 31(1):187.
Ambrijsus pulchellus pallidulus Montandon 1910, Bull. Soc. Sci. Buc.-Boum. 18

(5-6): 189.
Ambrysus pulchellus pallidulus, Van Duzee, 1917, Univ. Calif. Publ. Ent. 2:458.
Ambrysus pulchellus pallidulus, Hungerford, 1919, Univ. Kans., Sci. Bull.

11:198, 200.
Ambrysus pallidulus, Usinger, 1946, Univ. Kans., Sci. Bull. 31(1):187.
Ambrysus pulchellus, La Bivers, 1951, Univ. Calif. Publ. Ent. 8(7) : 303-306.

General appearance: rather small for the genus, quite narrow —
size 7.0-9.5 mm. long, 4.5-5.5 mm. wide. Dorsum generally lighter
anteriorly, but the contrast with the posterior is not striking or
sharp; glistening, polished anteriorly, shagreened posteriorly. Venter
generally uncontrastingly colored, but slightly darker anteriorly.

Head: groundcolor extremely variable, from light yellow through
darker browns to a bright green; polished, impunctate; front is
advanced before the eyes, but the outline of head-front and outer
eye margins forms a completely smooth semicircle. Eyes absolutely
flush with head surface from behind, with no perceptible protuber-
ance. Head ratios are:

(1) total length to width 66::10 (66%)

(2) anterior distance between eyes to posterior distance 43:: 55 (78%)

(3) anterior distance between eyes to inner eye margin 43:: 40 (93%)

(4) posterior distance between eyes to greatest length of head posterior
to this line 55:: 20 (36%)

(5) anterior distance between eyes to greatest length of head anterior to
this line 43: :5 (12%).

Pronotum: shiny, polished, impunctate; groundcolor yellow
through brown to green, entire centrum, including anterior border,
mottled with brown spots, only the "V" area immediately behind
region of deepest head penetration free from the mottling. Lateral
edges smooth, unserrate, in unrubbed specimens with some moder-
ately long, sparse, yellow hairs along edge, discernible only with
considerable magnification; percent of lateral pronotal curvature
13% (av. 90:: 12); posterolateral angles weak to lacking as definite
entities, always situated posterior to transverse posterior pronotal
line, not evidently at widest point of pronotum. Dorsal ratios are:

(1) width between anterior angles to width between posterior angles 50:: 84

(60%)
( 2 ) medial length to greatest width 34: :84 ( 40% )
(3) distance between anterior angle and posterior angle on same side to

perpendicular distance between anterior angle and baseline of pronotum

44::49 (89%).

The comparative increase in width of the anterior pronotal end
is significant in this group.

Scatellum: reddish-brown to blackish (even in greenish speci-

La Rivers: The Ambrysus of Mexico 1307

mens), shiny but not polished, shagreened with dense, shallow,
rough pnnctation; ratio of three sides 57::44::43.

Hemelytra: deep reddish-brown to black, laterally with some
greenish cast in greenish specimens (but not in all snch speci-
mens); shagreenation as in scutellum. Embolium very long and
narrow for the genus, length to width 115: :31 (27%), to normal,
95:: 30 (32%); lighter in anterior two thirds; in unrubbed specimens
with a few sparse yellow hairs along outer edge. Hemelytra rather
markedly exposing lateral connexiva, and just attaining abdominal
tip.

Venter: Connexival posterolateral angles all slightly produced
except those of segment I, but this weak prolongation detectable
only with considerable magnification; all lateral edges smooth, non-
serrate.

Legs: Prolegs — whitish to amber; ratio of length to greatest width
of ventral femoral surface 76: :53 (70%); combined tibia-tarsus, when
closed, just slightly, if at all, overlapping adjacent (proximal) end
of femur.

Mesolegs — whitish-yellow; femur long, narrow, ratio of length to
median width of ventral surface 78:: 16 (21%) — length 2.0 mm;
tibia long, narrow, distal end ventrally with one prominent, complete
transverse row of spines set solidly across apex; remnants of the
usual second row proximal to the distal row are present in sufficient
quantity to form a partial second row which extends approximately
halfway across tibia, with other half of tibia bare and lacking even
an isolated spine which could be postulated as a remnant of the
same row (as are present in such as A. hungerfordi) — ratio of length
to median width of ventral surface 68: :7 (10%) — length 1.85mm.

Metalegs — whitish-yellow; femur long, narrow, ratio of length to
median width 96::20 (21%) — length 2.80 mm.; tibia long, narrow,
terminal transverse spination as in mesotibia — ratio of length to
median width 119: :9 (8%)— length 2.90 mm.

Recorded distribution: Texas to Guatemala.

Type locality data: "Guatemala."

Location of types: "collection du Musee de Hambourg." (Ger-
many ) .

Specimens examined: Guatemala: — Los Amatos, Kellerman
(MCZ). Mexico:— Chiapas (La Libertad), l(i)38, O. U. Louis
( 500 m. above sea level ) ( UK ) ; Michoacan ( Uruapan ) ( El Sabino ) ,
28(vii)36, H. D. Thomas (UK); Morelos (Rio Amacuza, 133 km.
S. of Mexico [City]), 14(x)36, H. D. Thomas (UK); Tamaulipas
(Forlon, 3% miles W. of), 8(viii)34, Smith & Dunkle (UK), (Vic-

1308 The University Science Bulletin

toria, 5 miles S. of), (xi)36, H. D. Thomas (UK). United States:
— Arizona — Cochise County, 29(vii)27, R. H. Beamer (UK). Texas
—Kerr County 8(iv)39, D. Millspaugh (RLU); Sutton County
(Roosevelt), 21(iv)24, J. O. Martin (RLU); Uvalde County (Con-
can), 6 (vii) 36, M. B. Jackson (UK); Valverde County (Del Rio),
8(vii)38, R. I. Sailer (UK).

As would be expected of a species with as wide a range as this
one, there is considerable variation in the population, which has
been by no means adequately sampled.

Ambrysus vandnzeei Usinger
Ambrysus vanduzeei Usinger 1946, Univ. Kans., Sci. Bull. 31(1):207.

General appearance: somewhat less than medium size, with a
moderate amount of contrasting mottling; size 8.0-9.5 mm. long and
5.5-6.0 mm. wide. Dorsum moderately mottled with brown and
yellow, lighter anteriorly. Venter light yellow with no conspicuous
darkening.

Head: groundcolor yellow with a variable development of brown-
ish suffusions; shiny, smooth, minutely punctulate; front slightly
protuberant before eyes, and with a suggestion of truncation.
Viewed posteriorly, eyes flush with head surface, no protuberance
evident. Head ratios are:

(1) total length to width 67:: 103 (65%)

(2) anterior distance between eyes to posterior distance 45:: 62 (73%)

( 3 ) width between anterior angles to distance between anterior angle and
posterior baseline of pronotum 52:: 46 (88%).

Scutellum: dark reddish-brown, with or without a median longi-
tudinal stripe; shiny but not polished, shagreened with dense, shallow
punctation; ratio of three sides 64::45::44.

Hcmehjtra: varying from hardly, to moderately contrastingly
mottled with yellow and blackish-brown; surface shagreened as in
scutellum; embolium approximately average in length and width
for the genus (length to width 106:: 36 = 34%), rarely with detect-
able, sparse, marginal pilosity — anterior two thirds light yellowish.
Hemelytra moderately strongly exposing lateral connexival margins,
which have conspicuous marginal pilosity. Posterolateral con-
nexival angles rather weakly angulate-produced, nonspinose; hem-
elytra fully attaining abdominal tip.

Venter: connexival angles can be generally characterized as
weakly angulate-produced, progressing in size and relative develop-
ment anteriorly to posteriorly; all are angulate-produced except
those of segment I, which are weakly right-angulate. Connexival

La Rivers: The Ambrysus of Mexico 1309

margins very strongly serrate on all segments except I, weakening
anteriorly on segment II.

Legs: Prolegs — whitish to yellowish; ratio of length to greatest
width of femoral ventral surface 86:: 54 (63%); combined tibia-
tarsus, when closed, just attaining, or slightly exceeding, adjacent
(proximal) end of femur.

Mesolegs — yellowish-white; ratio of length to median width of
ventral femoral surface 79::15 (19%) — length 2.0 mm.; tibial distal
end ventrally with two prominent transverse rows of spines, the
terminal row set solidly across tibial apex, the secondary or proximal
row incomplete on outer or anterior edge, half, or slightly more than
half the length of terminal row — ratio of length to median width of
ventral surface 69::10 (15%) — length 2.0 mm.

Metalegs — ratio of femoral length to median width of ventral sur-
face 108: :18 (17%) — length 2.90 mm.; tibial apical transverse spina-
tion as in mesotibia — ratio of length to median width of ventral
surface 64:: 6 (9%)— length 3.0 mm.

Recorded distribution: Baja California, Mexico.

Type locality data: "Mulege, Baja California, May 14, 1921, E. P.
Van Duzee" (Usinger 1946).

Location of types: "Holotype, male and allotype, female (Cali-
fornia Academy of Sciences)" (Usinger 1946).

Specimens examined: Mexico: — Baja California (La Purisima
Canon between La Purisima and Comondu), 24(v)47, -LaR (L),
(Mulege), 14(v)21, E. P. Van Duzee (RLU), (San Luis Gonzaga),
22-23 (v)47, -LaR (L). The types have also been examined.

This unique little species is representative of a type seem-
ingly restricted to the long peninsula of Baja California, and has
no counterpart in the rest of the genus except the equally unique
A. ochraceus from South America; both species possess attenuated
and prolonged female subgenital plates, with corresponding valvulae
development. Probably these ovipositors are used to insert eggs
into plant tissues, or hide eggs in crevices or other inaccessible
spots, rather than to merely glue them to underwater surfaces, as
seems to be common with most species of Ambrysus.

Ambrysus buenoi Usinger

Ambrysus buenoi Usinger 1946, Univ. Kans., Sci. Bull. 31(1):199.
Ambrysus buenoi, La Rivers, 1951, Univ. Calif. Publ. Ent, 8(7):310-312.

General appearance: medium-sized species, 8.5-10.0 mm. long
and 5.5-6.0 mm. wide. Dorsum yellowish-brown, varying on the
one hand toward deep reddish, and on the other to greenish, suf-

1310 The University Science Bulletin

fusions; scutellum may stand out somewhat more darkly than re-
mainder, and head and prothorax may be somewhat lighter than
hemelytra in general. Venter dark reddish-brown, darkening an-
teriorly and medially.

Head: amber to yellowish with greenish tints, smooth, shiny,
polished, microscopically very irregularly punctulate. Front of
head very weakly protuberant between eyes. Eyes, viewed pos-
teriorly, weakly but distinctly protuberant above head surface.
Head ratios are:

( 1 ) total length to width 66: : 103 ( 64%)

(2) anterior distance between eyes to posterior distance 45:: 65 (75%)

(3) anterior distance between eyes to inner eye-margin 45:: 41 (89%)

(4) posterior distance between eyes to greatest length of head posterior to
this line 65:: 22 (34%).

Pronotam: glistening but not polished, densely, shallowly punc-
tulate; color amber to light-yellow, in the latter case often with a
tinge of green; lateral edges smooth, unserrate, in unrubbed speci-
mens with a sparse line of yellow hairs present. Percent of curva-
ture 14% (av. 98:: 14), posterolateral angles well rounded. Dorsal
ratios are:

( 1 ) width between anterior angles to greatest pronotal width 54 : : 102 ( 53% )

( 2 ) median length to greatest width 33 : : 102 ( 32% )

( 3 ) width between anterior angles to distance between anterior angle and
posterior baseline of pronotum 54::51 (94%).

Scutellum: deep reddish, either same color as hemelytra or
darker, but not contrastingly so; surface shiny but not polished,
shagreened with dense, shallow, punctation; ratio of three sides
70::50::50.

Hcincli/tra: uniformly reddish-brown, often with a greenish cast;
surface shiny but not polished, shagreened as is scutellum. Em-
bolium somewhat narrow, length to width 61:: 18 (30%), a faint
marginal pilosity usually present. Hemelytra rather markedly ex-
posing lateral connexival edges, which have some marginal pilosity,
and not quite attaining abdominal tip.

Venter: Abdominal venter deep reddish-brown, covered with a
golden pelt of short, dense hydrofuge hairs; meso- and meta-sterna
darker due to lack of the pelt. Connexival posterolateral angles
mediumly spinose except segment I, spines progressively increasing
in size posteriorly; connexival edges smooth, unserrate, moderately
pilose.

Legs: Prolegs — light yellow, amber or greenish; ratio of length
to greatest width of femur 84:: 54 (64%); combined tibia-tarsus,

La Rivers: The Ambrysus of Mexico 1311

when closed, just attaining, or slightly exceeding adjacent (proxi-
mal ) end of femur.

Mesolegs — amber to green; ratio of femoral length to median
width of ventral surface 90:: 18 (20%)— length 2.25 mm.; tibia with
distal end ventrally with one prominent, transverse row of spines
set solidly across tip — remnants of the usual second, proximal row
present as a half row on inner or posterior half of tibia — ratio of
length to median width of ventral surface 72: :8 (11%) — length
1.95 mm.

Metalegs — amber to green; ratio of length to median width of
ventral femoral surface 59::10 (17%) — length 2.75 mm.; distal end
of tibia ventrally with the same type of transverse spination as
mesotibia — ratio of length to median width of ventral surface 130: :9
(7%)— length 3.25 mm.

Recorded distribution: southern Texas.

Type locality data: "Rio Grande, Rrewster Co., Texas, June 13-
17, 1908, Mitchell and Cushman collectors" (Usinger 1946).

Location of types: "Holotype, male (U. S. National Museum).
. . . Allotype, female, and a male and female paratype, same
data as type" (Usinger 1946) in the Usinger collection.

Specimens examined: Mexico: — San Antonio, 15(vii)28, R. H.
Reamer (RLU). United States: — the allotype and the two para-
types ( see above ) .

Ambrysus mormon australis subsp. nov.

General appearance: rather large subspecies, 10.0-12.0 mm. long
and 6.5-7.5 mm. wide. Dorsum nearly always lighter anteriorly,
often with a pronounced mottling of anterior part of hemelytra;
shiny but not polished, variously shagreened. Venter generally
entirely yellowish to deep-amber.

Head: light-yellow through brown, some specimens with a
greenish tint, usually with some longitudinal brownish markings in
center; minutely, densely and shallowly punctulate, shiny; front of
head slightly protuberant before eyes, and with a suggestion of
truncation. Eyes, viewed posteriorly, only slightly protuberant
above general head surface. Ratios are:

(1) total length to width 41:: 62 (66%)

(2) anterior distance between eyes to posterior distance 28:: 38 (74%)

(3) posterior distance between eyes to greatest length of head posterior to
this line 38:: 14 (37%).

Pronotum: shiny but not polished, densely, shallowly and roughly
punctate; groundcolor yellow to brownish, occasionally with green

53_6490

1312 The University Science Bulletin

developing laterally, usually with a variable development of brown
mottling and suffusions on disc; lateral edges smooth, unserrate,
with an occasional sparse yellow pilosity; percent of lateral pronotal
curvature 13% (av. 67: :9); posterolateral angles well rounded.
Dorsal ratios are:

( 1 ) width between anterior angles to greatest pronotal width 62: : 127 (49%)

(2) median length to greatest width 38:: 127 (30%)

(3) width between anterior angles to distance between anterior angle and
posterior baseline of pronotum 62:: 58 (94%).

Scutettum: brown, often with a reddish cast, to black, with a
variable development of yellowing posteriorly; shiny, markedly
rough-shagreened with dense, shallow punctation; ratio of three
sides 84:: 63:: 62.

Hemelytra: deep brownish to jet-black, usually with enough
yellow-spotting to give a mild mottled effect; some greenish lights
evident occasionally; shiny, shagreened as is scutellum. Embolium
comparatively longer and narrower than that of the typical sub-
species, ratio of length to width always less than 35% (av. 72:: 24 =
33%) (in A. m. mormon, length to width varies from 36% [55::20]
to 39% [72::28]). Hemelytra rather strongly exposing lateral con-
nexival edges, which have a variable amount of marginal, rather
short, pilosity; posterolateral connexival angles always well de-
veloped, generally so strongly so as to represent one of the maxima
for the genus in this respect; hemelytra not quite, to just, attaining
abdominal tip.

Venter: connexival posterolateral angles are strongly spinose on
all segments except I, which is entirely lacking in any angular pro-
longation; margins generally quite smooth, occasionally somewhat
rough, but never distinctly dentate or serrate; connexival border
bearing spines increases progressively in width posteriorly in contra-
distinction to another condition so commonly found in the genus,
viz., that in which the border is more or less uniform in width over
most of its length, then bends rather abruptly inward under shadow
of spine of adjacent segment. Female subgenital plate-apex is
distinctively different from that of the typical subspecies, with a
much greater development of the two lateral angles into long, more
or less parallel-sided processes several times longer than wide.
Also, in general, A. m. australis is a slimmer form than A. m. mor-
mon.

Legs: Prolegs — yellow, amber or greenish; ratio of length to
greatest width of ventral femoral surface 54:: 33 (62%); combined
tibia-tarsus, when closed, just attaining, or slightly exceeding, ad-
jacent ( proximal ) end of femur.

La Rivers: The Ambrysus of Mexico 1313

Mesolegs — yellow, amber or greenish; ratio of length to median
width of ventral femoral surface 58:: 10 (17%) — length 2.90 mm.;
tibial distal end ventrally with two prominent, transverse rows of
spines, the terminal row set solidly across apex, the secondary or
proximal row incomplete on outer or anterior edge, about two thirds
length of complete, terminal row — ratio of length to median width
of ventral surface 48: :6 ( 13%)— length 2.30 mm.

Metalegs — yellow, amber or greenish; ratio of length to median
width of ventral femoral surface 74:: 13 (18%) — length 3.45 mm.;
tibial distal transverse spination as in mesotibia — ratio of length to
median width of ventral surface 83: :6 (7%) — length 4.30 mm.

Distribution: see types.

Type locality data: Mexico: — Chihuahua (Rio San Pedro be-
tween Chihuahua and Naica), 22(vi)34, Smith and Dunkle (UK).

Location of types: Holotype male, allotype and two paratypes in
the Snow Museum, University of Kansas, Lawrence; one paratype
each in the collections of Robert L. Usinger, Berkeley, California,
and the writer, Reno, Nevada.

Ambrysus scalenus sp. nov.

General appearance: a strongly contrastingly mottled species, of
rather more than medium size, 10.0-12.5 mm. long, 6.5-8.0 mm. wide.
Dorsum strongly mottled with yellow and brown, shiny, conspicu-
ously lighter anteriorly. Venter uniformly very light, almost whitish-
yellow, with no darkening except vague spots at lateral connexival
abdominal angles.

Head: groundcolor whitish yellow to amber with a variable de-
velopment of brownish or reddish spotting and streaking; shiny,
smooth, minutely punctulate. Head front slightly protuberant be-
fore eyes, latter absolutely flush with head surface, when viewed
posteriorly. Head ratios are:

(1) total length to width 82:: 134 (61%)

(2) anterior distance between eyes to posterior distance 68:: 80 (85%)

(3) posterior distance between eyes to greatest length of head posterior to
this line 80:: 21 (26%).

Pronotum: shiny, smooth, minutely punctate; groundcolor whitish
yellow to yellow, often with a faint roseate tinge; a variable develop-
ment of brown mottling and dotting on disc; lateral edges smooth,
unserrate, nonpilose; percent of lateral curvature 16% (av. 63:: 10),
posterolateral angles well rounded. Dorsal ratios are:

(1) width between anterior angles to width between posterior angles 70::
133 (53%)

(2) median length to greatest width 44:: 133 (33%)

1314 The University Science Bulletin

(3) width between anterior angles to distance between anterior angle and
posterior baseline of pronotum 70 : : 62 ( 89% ) .

Scutellum: dark reddish-brown to blackish, conspicuously marked
with yellow along lateral edges; shiny, shagreened with dense, shal-
low punctation; ratio of three sides 85::62::61.

Hemelytra: strongly and contrastingly mottled with brown and
yellow, shiny but not polished, shagreened as is scutellum. Em-
bolium about normal in width for the genus, length to width 144: :45
(31%), with no noticeable marginal pilosity. Hemelytra very
strongly exposing lateral connexival margins; posterolateral con-
nexival angles rather inconspicuous, but present, angulate-produced.
Hemelytra just attaining abdominal tip.

Venter: Connexival posterolateral angles markedly angulate-
produced, but hardly truly spinose, except angles of segment I;
angles increasing slightly but progressively in size posteriorly; con-
nexival margins distinctly but minutely serrate from segment III
posteriorly, occasionally weakly evident posteriorly on segment II,
lacking on I.

Legs: Prolegs — whitish yellow to amber; ratio of length to great-
est femoral width of ventral surface 99:: 67 (69%); combined tibia-
tarsus, when closed, just attaining adjacent (proximal) end of fe-
mur.

Mesolegs — whitish yellow to amber; ratio of length to greatest
width of ventral femoral surface 105:: 23 (22%)— length 2.50 mm.;
tibia with distal end ventrally with two prominent, transverse rows
of spines, the terminal row set solidly across apex, the secondary
or proximal row incomplete on outer or anterior edge, slightly more
than half the length of terminal row — ratio of length to median
width of ventral surface 90: :11 ( 12%)— length 2.20 mm.

Metalegs — whitish to yellowish; ratio of length to median width
of ventral femoral surface 140: :25 (18%) — length 3.50 mm.; tibia
with distal end ventrally with two prominent, transverse rows of
spines as in mesotibia — ratio of length to median width of ventral
surface 152:: 13 (9%)— length 4.0 mm.

Distribution: see types.

Type locality data: Mexico: — Michoacan (Uruapan, ElSabino),
28(vii)36, H. D. Thomas (UK).

Location of types: Holotype male, allotype and a large series of
paratypes in the Snow Museum, University of Kansas, Lawrence;
paratypes in the collections of Robert L. Usinger, Berkeley, Cali-
fornia and the writer, Reno, Nevada.

La Rivers: The Ambrysus of Mexico 1315

Ambrysus convexus Usinger
Ambrysus convexus Usinger 1946, Univ. Kans., Sci. Bull. 31(1) :196.

General appearance: small, very convex, not conspicuously mot-
tled, compact, species — size 9.0-10.0 mm. long, 6.0 mm. wide. Dor-
sum deep blackish-brown, with vague, uncontrasting mottling; shiny,
lighter anteriorly. Venter deep amber to blackish, darker anteri-
orly, legs of some specimens outstandingly green.

Head: groundcolor light to deep-yellow, with conspicuous deep
brownish areas forming a definite pattern of brownish streaks and
suffusions; shiny, glistening, comparatively strongly punctate; front
slightly protuberant before eyes, and suggestively truncate. Eyes
slightly but distinctly protuberant above head surface when viewed
posteriorly. Head ratios are:

(1) total length to width 66:: 104 (63%)

(2) anterior distance between eyes to posterior distance 46:: 64 (72%)

(3) posterior distance between eyes to greatest length of head posterior
to this line 64:: 19 (30%).

Fronotum: shiny but rough, conspicuously, coarsely, densely and
shallowly punctate; groundcolor yellow-brown, darkening on disc
except centrally, but with no pattern development. Lateral edges
smooth, unserrate, nonpilose; percent of lateral curvature 17% (av.
96:: 16), posterolateral angles well rounded. Dorsal ratios are:

(1) width between anterior angles to width between posterior angles 52::
101 (51%)

(2) median length to greatest width 33::101 (33%)

(3) width between anterior angles to distance between anterior angle and
posterior baseline of pronotum 52:: 50 (96%).

Scutellum: blackish, generally with a very deep reddish suffusion;
shiny, rough-shagreened with dense, coarse, shallow punctation;
ratio of three sides 71: : 51: : 50.

Hemelytra: deep-brown, membrane black, some lightening in
color laterally, but no mottling of any kind; shiny, shagreened as
in scutellum. Embolium normal in proportions, length to width
108:: 35 (32%), no marginal pilosity; hemelytra strongly exposing
lateral connexival margins, which have very short, sparse, marginal
pilosity; posterolateral connexival angles somewhat bluntly pro-
duced, distinctly nonspinose; hemelytra attaining, to not quite
attaining, abdominal tip.

Venter: Connexival posterolateral angles nonspinose, I, II, III
right-angulate, IV a slightly smaller angle ( i. e., in degrees, not
over-all size), V about the same (in degrees only), but larger in
size; all angles increase progressively in size posteriorly, and extend

1316 The University Science Bulletin

laterally a little more prominently with progression caudally; con-
nexival margins weakly microdentate on posterior segments, smooth
on anterior segments.

Legs: Prolegs — yellowish-green to deep amber; ratio of length to
greatest width of ventral femoral surface 80:: 52 (65%); combined
tibia-tarsus, when closed, just attaining adjacent (proximal) end of
femur.

Mesolegs — yellow-green to amber; ratio of length to greatest
width of femoral ventral surface 84::15 (18%) — length 2.10 mm.;
tibia with distal end ventrally with two transverse rows of spines,
the terminal row prominent, set solidly across apex, the secondary
or proximal row very incomplete, only half or less the length of
terminal row (i. e., extending half or less across tibial apex) — ratio
of length to median width of ventral surface 72: :8 (11%) — length
2.00 mm.

Metalegs — yellow-green to amber; ratio of length to greatest
width of femoral ventral surface 109: :17 (16%) — length 2.85 mm.;
distal tibial transverse spination as in mesotibia — ratio of length to
median width of ventral surface 70: :7 ( 10%) — length 3.25 mm.

Recorded distribution: central Mexico.

Type locality data: "Real de Arriba, District of Temascaltepec,
Mexico, May 25, 1933, H. E. Hinton and R. L. Usinger" (Usinger
1946).

Location of types: "Holotype, male and allotype, female (Cali-
fornia Academy of Sciences )" ( Usinger 1946 ) .

Specimens examined: the types and several paratypes, the latter
in the Usinger collection, Berkeley, California.

Ambry sus inflatus sp. nov.

General appearance: a well-mottled, robust, large, species — 10.0-
11.5 mm. long, 7.25-8.25 mm. wide. Dorsum generally well and
contrastingly mottled, lighter anteriorly; shiny but not polished,
variously shagreened. Venter entirely yellow, with no dark mark-
ings.

Head: groundcolor light yellow, variously mottled with brownish
suffusions and dots; glistening, but minutely roughened; front
slightly protuberant before eyes, with a suggestion of truncation.
Eyes, viewed from behind, slightly but distinctly protuberant above
general head surface. Head ratios are:

(1) total length to width 42:: 65 (65%)

(2) anterior distance between eyes to posterior distance 33:: 39 (85%)

(3) posterior distance between eyes to greatest length of head posterior to
this line 39:: 11 (28%).

La Rivers: The Ambrysus of Mexico 1317

Pronotum: shiny but not polished, conspicuously, densely, shal-
lowly and roughly punctate; groundcolor light yellow, no green
tints of any kind evident on the specimens at my disposal, usually
with a variable development of brown mottling on the disc varying
from a brownish suffusion to a definite pattern of brown dots.
Lateral edges smooth, unserrate, with no detectable pilosity; per-
cent of lateral curvature 13% (av. 62: :8), posterolateral angles
rather prominent, usually tending more towards angularity than to
rounding, definitely set at widest point of pronotum and slightly
caudad of transverse pronotal posterior line. Dorsal ratios are:

( 1 ) width between anterior angles to width between posterior angles 65 : :
128 (51%)

(2) median length to greatest width (width between posterior angles in
this case) 42:: 128 (33%)

(3) width between anterior angles to distance between anterior angle and
posterior baseline of pronotum 65:: 56 (86%).

Scutellum: shiny, markedly rough-shagreened with dense, shal-
low, punctation; color reddish-brown, with varying amounts of light
yellow mottling invading disc from the three angles; ratio of three
sides 80:: 58:: 57.

Hemclytra: deep reddish-brown usually with enough yellow-
spotting to give a mild mottled effect; shiny but not polished,
shagreened as in scutellum. Embolium with the most unusual
type of inflation in the entire genus, furnishing the diagnostic
characteristic of this species — embolium exceptionally stout for the
genus (although not quite attaining the maximum in this respect),
length to width 75:: 30 (40%); outer edge, instead of exhibiting the
usual smooth curvature, is almost angulate, an abrupt and pro-
nounced change of direction occurring at about three fifths of the
distance from the cephalad end (see illustration); no significant
pilosity along margin. Hemelytra strongly exposing lateral con-
nexival edges, and attaining abdominal tip.

Venter: Connexival posterolateral angles strongly spinose on all
segments except I; angle of segment I, however, is generally sharply
right-angulate and projects weakly laterad in contrast to most such
angles in other species with this spinal pattern in which segment I
angle is either rounded and/or completely obliterated in the general
contour of the lateral body margin; connexival margins with some
semidentate irregularities visible only under high magnification,
but never actually serrate or dentate.

Legs: Prolegs — light yellow to amber; ratio of length to greatest
width of ventral femoral surface 52:: 34 (65%); combined tibia-

1318 The University Science Bulletin

tarsus, when closed, generally slightly but distinctly exceeding ad-
jacent (proximal) end of femur.

Mesolegs — whitish to yellow; ratio of length to greatest width of
ventral femoral surface 55:: 12 (22%) — length 2.65 mm.; tibial distal
end ventrally with two prominent, transverse rows of spines, the
terminal row set solidly across apex, the secondary or proximal row
incomplete on outer or anterior edge, about two thirds length of
complete, terminal row — ratio of length to median width of ventral
surface 95: :9 (9%)— length 2.35 mm.

Metalegs — whitish to yellow; ratio of length to greatest width of
ventral femoral surface 76:: 13 (17%)— length 3.55 mm.; tibial distal
transverse spination as in mesotibia — ratio of length to greatest
width of ventral surface 82: :7 (9%)— length 4.00 mm.

Distribution: see types.

Type locality data: Mexico:— Jalisco (Cofwnatldn), 9(ix)38,
H.D.Thomas (UK).

Location of types: Holotypic male, allotype and several para-
types in the Snow Museum, University of Kansas, Lawrence; one
paratype each in the collections of Robert L. Usinger, Berkeley,
California, and the writer, Reno, Nevada.

Amhrysus magniceps sp. nov.

General appearance: well-mottled, compact, strongly convex,
rather large species— size 11.0-12.5 mm. long and 7.0-7.5 mm. wide.
Dorsum rather strongly yellow and brown mottled, shiny, polished,
lighter anteriorly. Venter nearly entirely yellow to amber.

Head: groundcolor yellow to yellow-brown with brownish streaks,
suffusions and dotting; surface glistening, microscopically, insignifi-
cantly punctate; front of head slightly protuberant before eyes, with
a suggestion of truncation. Eyes, viewed posteriorly, perfectly
flush with head surface. Head ratios are:

(1) total length to width 48:: 75 (64%)

(2) anterior distance between eyes to posterior distance, 35:: 45 (78%)

( 3 ) posterior distance between eyes to greatest length of head posterior to
this line 45:: 12 (27%).

Pronotum: shiny but not polished, minutely, almost smoothly
microreticulate mediolaterally, micropunctulate medially; ground-
color yellow-brown with a variable development of brown mottling
and dotting on disc. Lateral edges smooth, unserrate, with no
detectable pilosity; percent of lateral curvature 14% (av. 72:: 10),
posterolateral angles well rounded. Dorsal ratios are:

(1) width between anterior angles to width between posterior angles 74::
133 (56%)

La Rivers: The Ambrysus of Mexico 1319

(2) median length to greatest width 50:: 133 (38%)

(3) width between anterior angles to distance between anterior angle and
posterior baseline of pronotum 74:: 68 (92%).

Scutellum: shiny but not polished, shagreened with dense, shal-
low punctation; color brown to deep reddish-brown, with varying
amounts of yellow invading disc from the three angles; ratio of
three sides 87:: 62:: 60.

Hemelytra: black -brown, often with deep red tints, with enough
yellow spotting to give a moderately strong mottled effect; shiny
but not polished, shagreened as in scutellum. Embolium long and
narrow, length to width 80:: 23 (29%), no conspicuous marginal
pilosity. Hemelytra strongly exposing lateral connexival margins,
which are very sparsely pilose; hemelytra not quite, or just, attain-
ing abdominal tip.

Venter: connexival posterolateral angles nonspinose, very slightly
produced laterally from the general body contour as quite sharp,
slightly less than 90° angles, angle of segment I the least definite
and least produced; connexival margins minutely serrate, serration
absent from segment I, weak on II, well developed on remaining
connexiva.

Legs: Prolegs — yellow to amber; ratio of length to greatest width
of femoral ventral surface 57:: 40 (70%); combined tibia-tarsus,
when closed, just about attaining adjacent (proximal) end of femur.

Mesolegs — yellow, greenish or amber; ratio of length to greatest
median width of ventral femoral surface 58:: 13 (22%) — length
2.85 mm.; tibial distal end ventrally with two prominent, transverse
rows of spines, the terminal row set solidly across apex, the sec-
ondary or proximal row incomplete on outer or anterior edge, about
two-thirds length of complete, terminal row — ratio of length to
median width of ventral surface 104:: 11 (11%) — length 2.55 mm.

Metalegs — yellow, greenish to amber; ratio of length to median
width of ventral femoral surface 75:: 14 (20%) — length 3.50 mm.;
distal tibial transverse spination as in mesotibia — ratio of length to
median width of ventral surface 86: : 8 (9%) — length 4.20 mm.

Distribution: see types.

Type locality data: Mexico: — Mexico (Tejupilco), 15-17 (vi) 33,
H. E. Hinton and R. L. Usinger (RLU).

Location of types: Holotype male and allotype in collections of
California Academy of Sciences, San Francisco; paratypes in the
collections of Robert L. Usinger, Berkeley, California, and the
writer, Reno, Nevada.

1320 The University Science Bulletin

Ambnjsus portheo sp. nov.

General appearance: a large, well-mottled species; 12.5-13.5 mm.
long and 8.25-9.25 mm. wide. Dorsum usually contrastingly mottled
with yellows and browns, generally somewhat lighter anteriorly,
shiny. Venter light yellowish without conspicuous dark areas, often
with a greenish tinge anteriorly.

Head: smooth, shiny, minutely punctulate; color yellow to yellow-
brown with a variable development of brown suffusions and dot-
tings; head front slightly protuberant before eyes, and distinctly
truncate. Eyes, viewed posteriorly, absolutely flush with head sur-
face. Head ratios are:

(1) total length to width 51:: 78 (65%)

(2) anterior distance between eyes to posterior distance 41::47 (87%)

(3) posterior distance between eyes to greatest length of head posterior
to this line 47:: 14 (30%).

Pronotum: shiny, smooth, minutely punctulate; groundcolor light
yellow to yellow with a variable development of brown suffusions
and dots. Lateral pronotal edges smooth, unserrate, in unrubbed
specimens with some sparse pilosity; percent of lateral curvature
14% (av. 77::11), posterolateral angles weakly distinct, not quite
lost in the general rounding of pronotum. Dorsal ratios are:

(1) width between anterior angles to width between posterior angles 79::
157 (50%)

(2) median length to greatest width 53:: 157 (34%)

(3) width between anterior angles to distance between anterior angle and
posterior baseline of pronotum 79::71 (90%).

Scntclhim: light reddish-brown, generally lined laterally with
yellow; shiny but not polished, shagreened with dense, shallow
punctation; ratio of three sides 103: :73: :72.

Hemclytra: reddish- to blackish-brown, generally rather well
mottled with yellow; shiny but not polished, shagreened as is
scutellum. Embolium slightly short and stout for the genus, length
to width 85:: 30 (35%), with some sparse, marginal pilosity.
Hemelytra rather strongly exposing lateral connexiva, which have
some marginal pilosity; hemelytra attaining abdominal tip.

Venter: connexival posterolateral angles sharply, distinctly spi-
nose except those of segment I (which are, however, in contra-
distinction to the usual condition for the genus, minutely and
sharply right-angulate, distinctly produced laterad of the general
body outline), all spines slightly but progressively increasing in
size posteriorly; connexival margins distinctly but minutely serrate
on all segments except I, occasionally weakening considerably on
segment II.

La Rivers: The Ambrysus of Mexico 1321

Legs: Prolegs — whitish to amber; ratio of length to greatest width
of ventral femoral surface 64:: 43 (67%); combined tibia-tarsus,
when closed, generally slightly exceeding adjacent (proximal) end
of femur.

Mesolegs — whitish to yellow; ratio of length to median width of
ventral femoral surface 65:: 13 (20%)— length 3.0 mm.; tibial distal
end ventrally with two prominent, transverse rows of spines, the
terminal row set solidly across apex, the secondary or proximal row
incomplete on outer or anterior margin, approximately half or more
the length of complete, terminal row — ratio of length to median
width of ventral surface 60: :7 ( 12%)— length 2.55 mm.

Metalegs — whitish to yellow; ratio of length to median width of
ventral femoral surface 87:: 15 (17%) — length 4.0 mm.; tibial ter-
minal transverse spination as in mesotibia — ratio of length to median
width of ventral surface 99: :7 (7%) — length 5.0 mm.

Distribution: see types.

Ti/pe locality data: Mexico: — Coahuila (Saltillo, H. D. Thomas)
(UK).

The relationship between this species and A. lundbladi (see the
discussion following A. fuscus) cannot be determined with cer-
tainty until the male characteristics of A. lundbladi are known.
With more representative material to base judgment on, it may be
necessary to combine the two species. Also concerned in this im-
mediate picture is the still unrecognized A. signoreti. When the
collecting of Mexican Ambrysus material has been comprehensive
enough to provide an adequate taxonomic sampling of the popula-
tions of these three species, it may be found that the name signoreti
may have to be applied to the entire assemblage; the problem
hinges primarily upon the limits of variability involved, which are
still unknown.

Ambrysus signoreti Stal

Ambrysus signoreti Stal 1862, Stet. Ent. Zeit. 23:450; 1876, Enum. Hemipt.

5:143.
Ambrysus signoreti, Uhler, 1872, Hayden Surv. Terr., Rept. for 1871:423; 1876,

Bull. U.S.G.S. 1:337; 1877, Wheeler Rept. Chief Eng. for 1877:1331;

1884, Stand. Nat. Hist. 2:260; 1894, Proc. Calif. Acad. Sei., ser. 2, 4:291.
Ambrysus signoreti, Gillette and Baker, 1895, Hemipt. Colo., p. 63.
Ambrysus signoreti, Montandon, 1897, Verh. zool-bot. Ges. Wien. 47:13, 23;

1909, Bull. Soc. Sei. Buc.-Roum. 17:323.
Ambrysus signoreti, Champion, 1900, Biol. Centr.-Amer., Hemipt. 2:358.

1322 The University Science Bulletin

Ambrysus signoreti, Snow, 1906, Trans. Kans. Acad. Sci. 20(1):180.
Ambrysus signoreti, Van Duzee, 1917, Univ. Calif. Publ. Ent. 2:459.
Ambrysus signoreti, Hungerford, 1919, Univ. Kans., Sci. Bull. 11:202.
Ambrysus signoreti, Usinger, 1946, Univ. Kans., Sci. Bull. 31(1) :185, 203.
Ambrysus signoreti, La Rivers, 1949, Bull. S. Calif. Acad. Sci. 47(3) :108.

This species has been consistently misrepresented in the United
States literature dealing with Ambrysi, and its status in European
literature dealing with the Mexican fauna is hardly better. In the
large amount of material gathered for this study, I have been un-
able to place it definitely, and the present unavailability of the
original Stal description leaves me no recourse at present but to
reproduce a Montandon description of the species for what it is
worth. Fortunately, Dr. Usinger was able to make camera lucida
drawings of the male and female accessory genitalic structures,
the former from the type in Vienna, the latter from an associated
female in Vienna and another in the British Museum. His notes
on the Vienna and Stockholm specimens may be useful to others
pursuing the same problem:

Stockholm Museum — "Ambrysus signoreti Stal, 1 male, Mexico.
Stal. Type? . . . this should be the type, according to the
Enumeratio. Puncticollis, pudicus and guttatipennis are all repre-
sented by types' with red labels. This differs from the others in
that it bears no label in Stal's handwriting. It is closest to luncl-
blodi . . . but is larger, flatter above, almost perfectly oval,
the pronotum very broad. The connexival angles are only slightly
produced, about as in lundbladi. Of the genital plates figured, it is
most like occidentalis."

Vienna museum — "Ambrysus signoreti Stal. The type is a male
with no genital capsule. It is labelled 'Mexico, coll. Signoret,'
'Pocyi Guer. det. Signoret,' 'Signoreti Stal,' 'Am. signoreti Stal det.
Montd.,' and type' in red. A second specimen which appears to
be conspecific is labelled 'Mexico, 1869,' and 'Amb. signoreti Stal,
det. Montd.' This is a female. These are the same as a female in
the British Museum with the following labels: 'Belim, Mexico,
1871, ' and '63,' 'KK Hof. M. Wien, A. Handlirsch,' 'Ambrysus sig-
noreti Stal 1896,' and 'Montandon coll. 1901—233.' These also ap-
pear to be the same as the 'type' in Stockholm."

Dr. Usinger's camera lucida drawings of the accessory genitalic
apparati make it possible to include the species in the key with
reasonable accuracy, but the best available description of the ani-
mal at the moment is that of Montandon ( 1897) :

"Plus grand et de forme plus elargie que les autres especes; 13.7
mm de longueur sur 8.8 mm de largeur. Sa teinte jaunatre claire

La Rivers: The Ambrysus of Mexico 1323

avec de grandes taches brunes sur les elytres lui donne un aspect
tres reconnaissable a premiere vue. Le clavus jaunatre pale avec
une petite tache brune vers las base et une vers l'extremite. Le
marge elytrale est obtusement sinuee derriere I'embolium, ce dernier
tres elargi, jaunatre sur toute sa largeur sur les trois quarts basi-
laires, brunatre sur le quart posterieur; les segments du connexivum,
brims sur le tiers basilaire, jaunatres ensuite. Le sillon transversal
du pronotum bien visible, mais assez superficiel et un peu inter-
rompu au milieu; la partie posterieure du pronotum derriere le
sillon, tres-pale, lisse a ponctuation tres fine, concolore et peu dense."
There is little doubt that the records of A. signoreti for the United
States have been based on other species (A. occidentalis, A. arizonus
and probably A. mormon), since there is no evidence that it occurs
there. In Mexico proper, from the sparse available evidence,
A. signoreti seems to be closest to A. portheo, and future material
may show the latter to be a variable unit of the A. signoreti popu-
lation per se. The accessory genitalic structures of the two, how-
ever, denote two separable entities with present material.

Ambrysus hydor sp. nov.

General appearance: a strongly contrastingly mottled species;
10.5-12.75 mm. long and 7.0-8.5 mm. wide. Dorsum strongly mottled
with yellow and brown, shiny, lighter anteriorly. Venter more or
less uniformly light yellow to greenish, vaguely darker at lateral
connexival angles of abdomen.

Head: groundcolor whitish-yellow to reddish-amber with a vari-
able development of deep brownish or blackish spotting and streak-
ing; shiny, smooth, minutely punctulate. Front of head slightly
protuberant before eyes, with a suggestion of truncation. Eyes,
viewed posteriorly, essentially flush with head surface. Dorsal head
ratios are:

( 1 ) total length to width 45: :70 ( 64%)

(2) anterior distance between eyes to posterior distance 34:: 42 (81%)

(3) posterior distance between eyes to greatest length of head posterior to
this line 42:: 11 (26%).

Pronotum: shiny, smooth, minutely punctulate; groundcolor
whitish-yellow to amber with a variable development of brownish
mottling and dotting on disc. Lateral edges smooth, unserrate, very
weakly pilose marginally in unrubbed specimens; percent of pro-
notal lateral curvature 14% (av. 66: :9), posterolateral angles well-
rounded. Dorsal ratios are:

( 1 ) width between anterior angles to width between posterior angles 72 : :
138 (52%)

1324 The University Science Bulletin

(2) median length to greatest width 50:: 138 (36%)

(3) width between anterior angles to distance between anterior angle and
posterior baseline of pronotum 72:: 62 (86%).

Scutellum: dark reddish to blackish-brown, with yellow mark-
ings; shiny but not polished, shagreened with dense, shallow punc-
tation; ratio of three sides 94: :69: :67.

Hemelytra: strongly and contrastingly mottled with black and
yellow, the black often showing reddish lights laterally. Embolium
somewhat short and stout for the genus, length to width 151: :54
(36%), rarely with detectable, sparse, marginal pilosity. Hemelytra
very strongly exposing lateral connexival margins, which are con-
spicuously pilose; hemelytra just, or not quite, attaining abdominal
tip.

Venter: Connexival posterolateral angles strongly and grossly
spinose, except segment I, which is, at most, but slightly right-
angulate, generally quite blunt, not at all produced; connexival
margins distinctly, strongly but somewhat minutely dentate or
serrate on all segments except I and occasionally II.

Legs: Prolegs — white, amber or green; ratio of length to greatest
width of ventral femoral surface 105: :73 (70%); combined tibia-
tarsus, when closed, just attaining adjacent (proximal) end of
femur.

Mesolegs— white, amber or green; ratio of length to greatest
width of ventral femoral surface 62:: 12 (19%)— length 2.80 mm.;
tibial distal end ventrally with two prominent, transverse rows of
spines, the terminal row set solidly across apex, the secondary or
proximal row incomplete on outer or anterior edge, half, or slightly
more than half, the length of the terminal, complete, row — ratio of
length to median width of ventral surface 98: :9 (9%)— length
2.50 mm.

Metalegs — white, amber or green; ratio of length to greatest
width of ventral femoral surface 80:: 14 (18%)— length 3.75 mm.;
tibial distal transverse spination as in mesotibia — ratio of length to
median width of ventral surface 86: :7 (8%) — length 4.0 mm.

Distribution: see types.

Type locality data: Holotype male, Mexico:— Morelos (Rio Ama-
cuza, 133 kilom. S. of Mexico [City] ), 14(x)36, H. D. Thomas (UK).
Allotype female, Mexico:— Jalisco {Union de Tula), 16(ix)38,
H.D.Thomas (UK).

Location of types: Holotype, allotype and paratypes from
Mexico:— Guerrero {La Sabana, 226 kilom. S. of Mexico [City]),
20(x)36, H. D. Thomas; (Puente de Ixtla), 12(vii)36, H. D.

La Rivers: The Ambrysus of Mexico 1325

Thomas; Jalisco (Union de Tula), 16(ix)38, H. D. Thomas;
Michoacan (Uruapan) (El Sabino), 28(vii)36, H. D. Thomas in
the Snow Museum, University of Kansas, Lawrence; paratypes from
Mexico: — Mexico (Tejupilco), 19(vi)33, H. E. Hinton and R. L.
Usinger in the collection of Robert L. Usinger, Rerkeley, California;
and Mexico: — Michoacan (Uruapan) (El Sabino), 28(vii)36, H. D.
Thomas in the author's collection, Reno, Nevada.

Ambrysus lunatus lunatus Usinger

Ambrysus lunatus Usinger 1946, Univ. Kans., Sci. Bull. 31(1) :303.
Ambrysus lunatus La Rivers, 1951, Univ. Calif. Publ. Ent, 8(7) :326-328.

General appearance: a widely variable subspecies, both in size
and color; 8.5-11.5 mm. long and 6.0-8.0 mm. wide. Dorsum strongly
and contrastingly mottled with yellow and brown, moderately shiny,
generally not conspicuously lighter anteriorly. Venter light yellow
or greenish to amber, without dark markings, generally lighter pos-
teriorly.

Head: groundcolor whitish yellow to amber, often with a greenish
tinge, with a variable development of deep brown streaking; shiny,
glistening, micropunctate. Front of head slightly protuberant be-
fore eyes, either smoothly rounded, or with a faint suggestion of
truncation. Eyes, viewed posteriorly, practically flush with head
surface. Head ratios are:

( 1 ) total length to width 74: : 117 (63%)

(2) anterior distance between eyes to posterior distance 60:: 71 (85%)

(3) posterior distance between eyes to greatest length of head posterior to
this line 71:: 21 (30%).

Pronotinn: shiny, smooth, conspicuously but minutely punctate;
groundcolor golden yellow to light yellowish-green with a variable
development of brown mottling and dotting on disc. Lateral edges
smooth, unserrate, occasionally with weakly discernible marginal
pilosity; percent of lateral curvature 13% (av. 62: :8), posterolateral
angles well rounded. Dorsal ratios are:

(1) width between anterior angles to width between posterior angles 62::
120 (52%)

(2) median length to greatest width 41:: 120 (34%)

(3) width between anterior angles to distance between anterior angle and
posterior baseline of pronotum 62:: 58 (94%).

Scutellum: conspicuously light and dark banded (neither color
predominating) with light yellow and light to dark brown, often
with a greenish tinge; shiny but not polished, shagreened with
dense, shallow punctation; ratio of three sides 74: :54: :53.

Hemelytra: strongly and contrastingly mottled with brown and

1326 The University Science Bulletin

yellow and green; shiny, but not polished, shagreened as in scu-
tellum. Embolium rather wide and short for the genus, length to
width 132:: 46 (35%), with sparse inconspicuous marginal pilosity.
Hemelytra very strongly exposing lateral connexival margins, and
just, to not quite, attaining abdominal tip.

Venter: Connexival posterolateral angles markedly spinose ex-
cept those of segment I which is, at most, but slightly right-angulate-
produced; connexival margins distinctly but very minutely serrate
on posterior segments, smooth anteriorly.

Legs: Prolegs — yellow, amber or greenish; ratio of length to
greatest width of ventral femoral surface 95:: 61 (64%); combined
tibia-tarsus, when closed, generally slightly, but definitely, exceed-
ing adjacent (proximal) end of femur.

Mesolegs — yellowish, amber or green; ratio of length to greatest
width of ventral femoral surface 96:: 18 (19%) — length 2.45 mm.;
tibial distal end ventrally with two prominent, transverse rows of
spines, the terminal row set solidly across apex, the secondary or
proximal row incomplete on outer or anterior edge, about half the
length of complete, terminal row — ratio of length to median width
of ventral surface 82: :10 ( 12%)— length 2.25 mm.

Metalegs — yellowish, amber or greenish; ratio of length to greatest
width of ventral femoral surface 133: :22 (16%)— length 3.20 mm.;
tibial distal transverse spination as in mesotibia — ratio of length to
greatest width of ventral surface 148:: 13 (9%) — length 3.90 mm.

Recorded distribution: southern Texas.

Type locality data: "Tom Greene County, Texas, July 15, 1928,
R. H. Beamer, collector" (Usinger 1946).

Location of types: "Holotype, male, and allotype, female (Snow
Museum, University of Kansas)" (Usinger 1946).

Specimens examined: United States: — New Mexico — Eddy
County 12(vii)27, R. H. Beamer (UK). Texas— Menard County
19(vii)28, R. H. Beamer (UK); Kerr County 9(iv)39, D. Mills-
paugh (UK); Travis County 10(iv)39, D. Millspaugh (UK); Val-
verde County (Del Rio), 8(vii)38, D. W. Craik, 8(vii)38, R. I. Sailer
(UK).

Future collecting will undoubtedly extend the range of this sub-
species into northern Mexico.

Ambrysus lunatus menoides subsp. nov.

General appearance: one of the most strongly and contrastingly
mottled units of the genus, medium-sized, 8.5-12.0 mm. long and
6.0-7.0 mm. wide.

La Rivers: The Ambrysus of Mexico 1327

Head: essentially as in the typical subspecies. Ratios are:

( 1 ) total length to width 67: : 108 ( 62%)

(2) anterior distance between eyes to posterior distance 56:: 68 (82%)

(3) posterior distance between eyes to greatest length of head posterior
to this line 68::21 (31%).

Pronotum: essentially as in the typical subspecies. Ratios are:

(1) width between anterior angles to width between posterior angles 57::
105 (54%)

(2) median length to greatest width, 35:: 105 (33%)

(3) width between anterior angles to distance between anterior angle and
posterior baseline of pronotum 57::52 (91%).

Hemelytra: essentially as in A. I. lunatus. Embolium rather wide
and short for the genus, length to width 123: :42 (34%).

Venter: connexival posterolateral angles weakly spinose except
those of segment I; angle I right-angulate, slightly produced laterad;
angle II prominent, but hardly truly spinose, being more in the na-
ture of angulate-produced, the angle approximately 45°; angle III
slightly larger than II, approximately transitional between angulate-
produced and true spinosity; IV spinose, larger than III; connexival
margins generally distinctly but very minutely serrate on all margins
except those of segment I, which are smooth.

Legs: essentially as in A. I. lunatus.

Distribution: see types.

Type locality data: Mexico: — Puebla (Tehuacan, 18-25 ( vii ) 37,
H. D. Thomas) (UK).

Location of types: Holotypic male, allotype, a large series of para-
types from the type locality, and paratypes from Mexico: — Puebla
(Puebla, 18 (vii) 37, H. B. Thomas) (UK) in the Snow Museum,
University of Kansas, Lawrence; paratypes from the type locality
in the collections of Robert L. Usinger, Berkeley, California, and
the writer, Reno, Nevada; paratypes from Mexico: — Chiapas (Ix-
tapa, 27(iv)41, I. J. Cantrall) (UM) in the Museum of Zoology,
University of Michigan, Ann Arbor.

The distinctive differences between A. I. lunatus and A. I. menoides
lie in the terminal aspects of the female subgenital plates and in the
male genital processes, as illustrated.

Ambrysus dilatus Montandon

Ambrysus dilatus Montandon 1910, Bull. Soc. Sci. Buc.-Roum. 18(5-6) :190.
Ambrysus dilatus, Usinger, 1946, Univ. Kans., Sci. Bull. 31(1) :198, 199.
Ambrysus hintoni Usinger 1946, Univ. Kans., Sci. Bull. 31(1):206.

General appearance: medium-sized, moderately well-mottled spe-
cies; 9.5-11.0 mm. long and 6.0-6.5 mm. wide. Dorsum usually well
54_6490

1328 The University Science Bulletin

mottled, but often uncontrastingly so, and varying from distinctly
lighter anteriorly than posteriorly to nearly uniformly mottled over-
all, shiny. Venter yellowish to deep reddish with no distinctive
dark areas, somewhat lighter posteriorly.

Head: light-yellowish to deep reddish-brown with a variable
development of brown dotting and suffusions; shiny, minutely but
slightly roughened and punctulate. Front of head slightly pro-
tuberant before eyes, and weakly truncate. Eyes, viewed pos-
teriorly, absolutely flush with head surface. Head ratios are:

(1) total length to width 71:: 117 (61%)

(2) anterior distance between eyes to posterior distance 54:: 67 (81%)

(3) posterior distance between eyes to greatest length of head posterior
to this line 67:: 20 (30%).

Pronotum: shiny, smooth, minutely punctulate; color yellowish
to reddish-brown with a variable development of brown suffusions
and dots; lateral edges smooth, unserrate, in unrubbed specimens
with some sparse marginal pilosity; percent of lateral curvature 14%
(av. 58: :8), posterolateral angles quite evident, but somewhat
rounded, not distinctly angulate. Dorsal ratios are:

( 1 ) width between anterior angles to width between posterior angles 63 : :
117 (54%)

(2) median length to greatest width 39:: 117 (33%)

(3) width between anterior angles to distance between anterior angle and
posterior baseline of pronotum 63:: 55 (87%).

Scutellum: deep reddish-brown, often with a blackish aspect,
nearly always lighter in color at the angles, and occasionally bi-
sected by a longitudinal, median yellow line; shiny but not polished,
shagreened with dense, shallow punctation; ratio of three sides
78::56::55.

Hemelytra: largely blackish-brown to brownish, generally un-
contrastingly mottled; shiny, shagreened as is scutellum. Embolium
about average in proportions for the genus, length to width 68:: 22
(32%), usually with some marginal pilosity. Hemelytra rather
strongly exposing lateral connexival margins, which have some mar-
ginal pilosity; posterolateral connexival angles rather minutely and
distinctly angulate-produced, but not truly spinose; hemelytra fully
attaining abdominal tip.

Venter: connexival posterolateral angles, except those of seg-
ment I, distinctly and sharply, minutely, angulate-produced, but
hardly to be considered truly spinose, slightly but progressively
increasing in size caudally; connexiva generally discernibly serrate
with high magnification, at least on posterior segments.

La Rivers: The Ambrysus of Mexico 1329

Legs: Prolegs — whitish to amber; ratio of length to greatest width
of ventral femoral surface 96: :68 (69%); combined tibia-tarsus, when
closed, either just attaining adjacent (proximal) end of femur, or
distinctly exceeding it.

Mesolegs — whitish to deep amber; ratio of length to greatest
width of ventral femoral surface 95:: 19 (20%) — length 2.50 mm.;
tibial distal end ventrally with two prominent, transverse rows of
spines, the terminal row set solidly across apex, the secondary or
proximal row incomplete on outer or anterior edge, approximately
two thirds the length of the complete row — ratio of length to median
width of ventral surface 86: :11 ( 13%)— length 2.10 mm.

Metalegs — whitish to amber; ratio of length to greatest width of
ventral femoral surface 126: :29 (23%)— length 3.10 mm.; tibial distal
transverse spination as in mesotibia — ratio of length to greatest
width of ventral surface 140: : 12 (9%) — length 3.50 mm.

Recorded distribution: central Mexico, "Mexico." Montandon
gave only the broad designation "Mexico" when describing the
species, but Usinger's type locality data for A. hintoni provides
more specific data: "Tejupilco, District of Temescaltepec, Mexico,
June 30, 1933. . . . H. E. Hinton and R. L. Usinger" (Usinger
1946). The types of A. hintoni are "Holotype, male and allotype,
female (California Academy of Sciences)" (Usinger 1946).

Specirnens examined: Mexico: — Michoacan (near Chinapa),
5(ix)38, H. D. Thomas (UK); Mexico (Tejupilco), 15(vi)33, H. E.
Hinton and R. L. Usinger, paratypes (RLU); (Corriente), 24(viii)
37, H. D. Thomas (UK). The types of A. hintoni have also been
examined. "Mexico:" a "cotype" of A. dilatus in the collection of
the U. S. National Museum, simply labelled "Mexico," established
the synonymy of A. hintoni. The terminal structure of the female
subgenital plate is so distinctive in the species that it was a simple
matter to recognize the synonymy.

Ambrysus cosmius sp. nov.

General appearance: a large, rather well-mottled species; 12.0-14.5
mm. long and 8.0-9.25 mm. wide. Dorsum generally noticeably but
not contrastingly mottled with light and dark browns, usually
lighter anteriorly, and shiny. Venter whitish to amber, with no
distinctive dark areas, slightly lighter posteriorly.

Head: light-yellow to yellowish-brown, often with a greenish
cast and with a variable development of brown dotting and suffu-
sions; shiny, micropunctulate. Head front slightly protuberant be-

1330 The University Science Bulletin

fore eyes, and suggestively truncate. Eyes, viewed from behind,
absolutely flush with head surface. Head ratios are:

(1) total length to width 51:: 80 (64%)

(2) anterior distance between eyes to posterior distance 42:: 52 (81%)

(3) posterior distance between eyes to greatest length of head posterior
to this line 52:: 14 (27%).

Pronotum: shiny, smooth, micropunctulate; groundcolor whitish-
yellow to yellow-brown with a variable development of brown
suffusions and dots. Lateral edges smooth, unserrate, in unrubbed
specimens with some sparse pilosity. Percent of lateral curvature
14% (av. 78::11), posterolateral angles weakly evident, but nearly
lost in the general rounding of pronotum. Dorsal ratios are:

( 1 ) width between anterior angles to width between posterior angles 80 : :
158 (51%)

(2) median length to greatest width 58:: 158 (31%)

(3) width between anterior angles to distance between anterior angle and
posterior baseline of pronotum 80:: 75 (94%).

Scutellum: yellow-brown to deep reddish-brown, mottled with
light yellows; shiny but not polished, shagreened with dense, shallow
punctation; ratio of three sides 105::74::74.

Hemelytra: reddish-brown to blackish, generally uncontrastingly
mottled; shiny, shagreened as in scutellum. Embolium about aver-
age in proportions for the genus, length to width 93:: 29 (32%), in
unrubbed specimens usually with some marginal pilosity. Hemely-
tra rather strongly exposing lateral connexival margins, and not
quite, to fully, attaining abdominal tip.

Venter: Connexival posterolateral angles somewhat variable in
spinosity, but always with strong, well-developed spines; as is usual,
the angles of segment I are entirely nonspinose and nonangulate-
produced; spines slightly but progressively increasing in size cau-
dad; connexival margins distinctly serrate, most strongly so cau-
dally, never so on segment I, and occasionally only weakly so on
segment II.

Legs: Prolegs — white, yellow or greenish; ratio of length to great-
est width of ventral femoral surface 59:: 39 (66%); combined tibia-
tarsus, when closed, generally slightly exceeding adjacent (proxi-
mal) end of femur.

Mesolegs — white, yellow, amber or greenish; ratio of length to
median width of ventral femoral surface 59:: 13 (22%) — length 2.50
mm.; tibial distal end ventrally with two prominent, tranverse rows
of spines, the terminal row set solidly across tibial apex, the second-
ary or proximal row incomplete on outer or anterior edge, approxi-
mately three fourths or more the length of complete row — ratio of

La Rivers: The Ambrysus of Mexico 1331

length to median width of ventral surface 54: :7 (13%) — length 2.50
mm.

Metalegs — whitish to amber, often with a greenish tinge; ratio of
length to median width of ventral femoral surface 93:: 18 (20%) —
length 4.5 mm.; tibial distal transverse spination as in mesotibia —
ratio of length to median width of ventral surface 102: :9 (9%) —
length 5.0 mm.

Distribution: see types.

Type locality data: Mexico: — Mexico (Tejupilco, 19(vi)33, H.
E. Hinton and R. L. Usinger) (RLU).

Location of types: Holotypic male, and allotype in the collections
of the California Academy of Sciences, San Francisco; paratypes
from the type locality in the collection of Robert L. Usinger, Berke-
ley, California; paratypes from Mexico: — Guerrero (Chilapa, 29(x)
30, L. Schultze) (UK), (Tecpan, 2(iv)30, L. Schultze) (UK);
Morelos (Acatlipa, 88 km. on highway from Mexico [City] to Aca-
pulco, 6(v)45, J. G. Shaw) (UK), (Morelos, 14(vii)36, H. D.
Thomas ) ( UK ) ; Oaxaca ( Oaxaca, 25 ( viii ) 37, H. D. Thomas ) ( UK ) ,
in the Snow Museum, University of Kansas, Lawrence.

A. cosmius stood in the Usinger collection for many years as A.
signoreti, and its true status was not determined until Dr. Usinger
was able to examine the European types of A. signoreti.

Ambrysus guttatipennis Stal

Ambrysus guttatipennis Stal 1862, Enum. Hemipt. 5:143.

Ambrysus guttatipennis, Montandon, 1897, Verh. zool.-bot. Ges. Wien. 47:13,

22; 1909, Bull. Soc. Sci. Buc.-Roum. 17:322.
Ambrysus guttatipennis, Champion, 1900, Biol. Centr.-Amer. Hemipt. 2:356.
Ambrysus guttatipennis, Van Duzee, 1917, Univ. Calif. Publ. Ent. 2:459.
Ambrysus guttatipennis, Hungerford, 1919, Univ. Kans., Sci. Bull. 11:203.
Ambrysus guttatipennis, Usinger, 1946, Univ. Kans., Sci. Bull. 31(1) :202.
Ambrysus guttatipennis, La Rivers, 1949, Bull. S. Calif. Acad. Sci. 47(3) :108.

General appearance: a large, well-mottled species; 10.5-13.5 mm.
long and 7.0-8.5 mm. wide. Dorsum rather well and contrastingly
mottled with yellowish and brown or black, usually lighter ante-
riorly. Venter yellow to amber, darker about thoracic centrum,
lighter posteriorly.

Head: yellow-brown with a variable development of brown dot-
ting and suffusion; shiny, micropunctulate. Head front slightly pro-
tuberant before eyes, and quite truncate. Eyes, viewed posteriorly,
absolutely flush with head surface. Head ratios:

(1) total length to width 45:: 73 (62%)

(2) anterior distance between eyes to posterior distance 32:: 43 (74%)

(3) posterior distance between eyes to greatest length of head posterior to
this line 43:: 13 (30%).

1332 The University Science Bulletin

Pronotum: shiny, smooth, micropunctulate; groundcolor yellow-
brown with a variable development of brown suffusions and dots;
lateral edges smooth, unserrate, in unrubbed specimens with some
sparse pilosity; percent of lateral curvature 16% (av. 76:: 12), pos-
terolateral angles well rounded. Dorsal ratios are:

(1) width between anterior angles to width between posterior angles 73::
145 (50%)

(2) median length to greatest width 50:: 145 (34%)

(3) width between anterior angles to distance between anterior angle and
posterior baseline of pronotum 73:: 71 (97%).

Scutellum: deep reddish-brown, often with a blackish tinge; shiny
but not polished, shagreened with dense, shallow punctation; ratio
of three sides 98:: 69:: 68.

Hemelytra: reddish to blackish-brown, generally rather well-
mottled with yellow; shiny, shagreened as is scutellum. Embolium
about average in proportions, length to width 82:: 26 (32%), usually
with some marginal pilosity. Hemelytra moderately exposing lateral
connexival margins, which have prominent pilosity. Hemelytra just
attaining abdominal tip.

Venter: connexival posterolateral angles rather shortly but dis-
tinctly angulate-produced, but not spinose, all angles slightly but
progressively increasing in size caudally; there is even a suggestion
of this angular-production in the angle of segment I, which angle
seldom shows the spinal formula of the remaining angles but is
nearly always more or less contoured in with the general outline of
the body; connexival margins distinctly serrate on all segments ex-
cept I, most strongly so posteriorly.

Legs: Prolegs — amber; ratio of length to greatest width of ventral
femoral surface 64:: 42 (66%); combined tibia-tarsus, when closed,
distinctly exceeding adjacent (proximal) end of femur.

Mesolegs — amber; ratio of length to greatest width of ventral fe-
moral surface 61::11 (18%)— length 2.90 mm.; distal tibial end ven-
trally with two prominent, transverse rows of spines, the terminal
row set solidly across tibial apex, the secondary or proximal row in-
complete on outer or anterior margin, approximately two thirds or
more the length of the complete, terminal row — ratio of length to
median width of ventral surface 51: :8 (16%) — length 2.50 mm.

Metalegs — amber; ratio of length to greatest width of ventral
femoral surface 83::14 (17%)— length 4.0 mm.; tibial distal trans-
verse spination as in mesotibia — ratio of length to greatest width
of ventral surface 94: :7 (8%) — length 4.5 mm.

Recorded distribution: Arizona and Mexico.

Type locality data: "Mexico."

La Rivers: The Ambrysus of Mexico 1333

Location of types: Royal Stockholm Museum, Sweden.

Specimens examined: Mexico: — Mexico (Real de Arriba), 25 (v)
33, H. E. Hinton and R. L. Usinger (RLU).

The United States records for this species (Arizona) are very
probably based on examples of A. arizonus and A. occidentalis, and
possibly A. mormon. There is no evidence at the present time that
A. guttatipennis even approaches the southwestern United States.

Ambrysus fuscus Usinger
Ambrysus fuscus Usinger 1946, Univ. Kans., Sci. Bull. 31(1):198.

General appearance: a very dark, blackish, totally immaculate
species of rather small size, 8.75-9.5 mm. long and 5.5-6.0 mm. wide.
Dorsum rough, blackish-brown, more or less unicolorous, shiny.
Venter blackish-brown, lighter posteriorly.

Head: deep blackish-brown, generally with a readily discernible,
subdermal pair of thin, black, median streaks; shiny but minutely
roughened and punctulate. Front of head slightly protuberant
before eyes, and distinctly truncate. Eyes, viewed posteriorly,
slightly but definitely protuberant above head surface. Ratios are:

( 1 ) total length to width 73: : 107 ( 68%)

(2) anterior distance between eyes to posterior distance 53:: 67 (79%)

(3) posterior distance between eyes to greatest length of head posterior to
this line 67:: 25 (37%).

Pronotum: shiny, but conspicuously roughened and punctate;
dark reddish and blackish brown, occasionally showing a faint,
vague lightening laterally. Lateral edges smooth, unserrate, in
unrubbed specimens with rather conspicuous, but comparatively
sparse marginal pilosity. Percent of lateral curvature 19% (av. 105::
20), posterolateral angles well rounded. Dorsal ratios are:

(1) width between anterior angles to greatest pronotal width 56:: 103 (54%)

(2) median length to greatest width 35:: 103 (34%)

(3) width between anterior angles to distance between anterior angle and
posterior baseline of pronotum 56::51 (91%).

Scutellum: blackish, occasionally reddish around edges and often
becoming light yellow at the angles; shiny but not polished,
shagreened with dense, shallow punctation; ratio of three sides
70::50::50.

Hemelytra: uniformly immaculate blackish-brown, always with
discernible, although sometimes vague, lightening in emboliar
region; shiny, shagreened as in scutellum. Embolium slightly less
than normal in proportions for the genus, length to width 129:: 40
(31%), usually with some marginal pilosity. Hemelytra moderately
exposing lateral connexival margins, which are conspicuously pilose.
Hemelytra fully attaining, or exceeding, abdominal tip.

1334 The University Science Bulletin

Venter: connexival posterolateral angles nonspinose, increas-
ing progressively in size and slight laterad projection from an-
terior to posterior; all angles except those of segment I distinctly
but minutely projecting laterad due to undercutting of adjacent
margin of posterior connexival margin, those of I more or less form-
ing a smooth contour with anterolateral angle of segment II; all
remaining posterolateral connexival angles right-angulate except
those of IV which are acute angles; connexival margins weakly and
irregularly serrate, such serration decreasing anteriorly and totally
undemonstrable on margin of segment I.

Legs: Prolegs — yellow-brown; ratio of length to greatest width
of ventral femoral surface 93:: 62 (67%); combined tibia-tarsus,
when closed, distinctly exceeding adjacent (proximal) end of femur.

Mesolegs — yellow to brown; ratio of length to greatest width of
ventral femoral surface 87:: 17 (20%) — length 2.0 mm.; tibial distal
end ventrally with two prominent, transverse rows of spines, the
terminal row set solidly across apex, the secondary or proximal row
incomplete on outer or anterior edge, half, or slightly more than
half the length of terminal row — ratio of length to greatest width of
ventral surface 78: : 11 ( 14% ) — length 2.0 mm.

Metalegs — yellow to reddish-brown; ratio of length to greatest
width of ventral femoral surface 56:: 10 (18%) — length 2.70 mm.;
tibial distal transverse spination as in mesotibia — ratio of length to
greatest width of ventral surface 68: :6 (9%) — length 3.10 mm.

Recorded distribution: central Mexico.

Type locality data: "Real de Arriba, District of Temescaltepec,
Mexico, July 10, 1933, H. E. Hinton and R. L. Usinger" (Usinger
1946).

Location of types: "Holotype, male, and allotype, female (Cali-
fornia Academy of Sciences )" ( Usinger 1946 ) .

Specimens examined: the types and several paratypes from the
type locality, the latter in the Usinger collection, Berkeley, Cali-
fornia.

It is not practicable, at present, to incorporate the two Usingerian
species, A. sonorensis and A. lundbladi, into the above structure,
since both are known only from a single type specimen, a male in
the case of the former, a female in the latter case, both described
in 1946. A. sonorensis was described from "San Bernardo, Rio Mayo,
State of Sonora, Mexico, March 2, 1935, H. S. Gentry collector"
(Usinger 1946), while the type locality for A. lundbladi is "State of
Morelos, Mexico, May 30, 1897, Koebele collection" (Usinger 1946).
For detailed descriptions of these, the reader is referred to Usinger's
1946 paper.

La Rivers: The Ambrysus of Mexico 1335

REFERENCES
Champion, G. C.

1900. Family Naucoridae, Rhynchota, Hemiptera-Heteroptera. Biologia
Centrali-Americana, Insecta, 2:xvi + 416 (354-361).

HUNGERFORD, H. B.

1919. The biology and ecology of aquatic and semiaquatic Hemiptera.
Univ. of Kan. Sci. Bull. (Entomology Number), 11:1-341 (198-
210).

KlRKALDY, G. W.

1906. List of the genera of the Pagiopodous Hemiptera-Heteroptera, with
their type species, from 1758 to 1904 (and also of the aquatic and
semiaquatic Trochalopoda ) . Trans. Amer. Ent. Soc, 32:117-156.

Kirkaldy, G. W., and J. R. de la Torre-Bueno.

1909. A catalogue of the American aquatic and semiaquatic Hemiptera.
Proc. Ent. Soc. Washington, 10(3-4) : 173-215.
La Rivers, Ira.

1949. A new species of Pelocoris from Nevada, with notes on the genus
in the United States (Hemiptera: Naucoridae). Ann. Ent. Soc.
America, 41(3) :371-376. (1948.)

1949. A new species of Ambrysus from Death Valley, with notes on the
genus in the United States (Hemiptera: Naucoridae). Bull. South.
California Acad, of Sci., 47(3) :103-109. (1948.)

1950. The meeting point of Ambrysus and Pelocoris in Nevada (Hemip-
tera: Naucoridae). Pan-Pacific Ent, 26(1):19-21.

1951. A revision of the genus Ambrysus in the United States (Hemip-
tera: Naucoridae). Univ. of California Publ. in Ent., 8(7):277-
338.

MONTANDON, A. L.

1897a. Hemipteres nouveaux des collections du Museum de Paris. Bull.

Paris Mus. d'Hist. Nat., 3:124-131.
1897b. Hemiptera cryptocerata. Fam. Naucoridae. — Sous-fam. Crypto-

cricinae. Verhandl. d k.-k. Zool.-Bot. Ges. in Wien, 47:6-23.
1898. Hemiptera cryptocerata: notes et descriptions d'especes nouvelles.

Bull Soc. Sci. Bucarest-Roumanie, 7(3-4): 282-290.
1909a. Naucoridae. Descriptions d'especes nouvelles. Ibid., 18(1):43-61.
1909b. Tableau synoptique des Ambrysus et descriptions d'especes nou-
velles. Ibid., 17(5-6): 316-330.
1910a. Trois especes nouvelles de la Famile Naucoridae. Ibid., 19(3):

438-444.
1910b. Hydrocorises de l'Amerique du nord; notes et descriptions d'especes

nouvelles. Ibid., 18(5-6) : 180-191.
Stal, C.

1862. Hemiptera mexicana enumeravit speciesque novas descriptsit. Stet-

tiner ent. Zeit, 23:437:462.
1876. Enumeratio Hemipterorum. 4to. Stockholm, pt. V, in vol. 14,

No. 4.

1336 The University Science Bulletin

USINGER, R. L.

1941. Key to the subfamilies of Naucoridae with a generic synopsis of

the new subfamily Ambrysinae (Hemiptera). Ann. Ent. Soc.

America, 34(1):5-16.
1946. Notes and descriptions of Ambrysus Stal with an account of the

life history of Ambrysus mormon Montd. ( Hemiptera, Naucoridae ) .

Univ. Kansas, Sci. Bull. 31(1):185-210.

Van Duzee, E. P.

1917. Catalogue of the Hemiptera of America north of Mexico excepting
the Aphididae, Coccidae and Aleurodidae. Univ. of California
Publ. Ent., 2:xiv+ 902.

La Rivers: The Ambrysus of Mexico 1337

EXPLANATION OF FIGURE 35, PLATE XCIV

Ventral surface of female abdominal tip region

A — subgenital plate.

B — subgenital plate anterolateral angles.

C — laterocaudal connexival angle (secondary or interior posterolateral

connexival angle ) .
D — posterolateral connexival angles (primary or exterior posterolateral

connexival angles).
E — connexival margins.

Ill, IV, V — connexival areas of abdominal segments of these numbers.
Fig. 33, pi. XCIV is guttatipennis.

1338 The University Science Bulletin

PLATE XCV

(A) — Ambrysus pygmaeus La Rivers, X 6.
(B) — Ambrysus melanopterus Stal, X 6.
(C) — Ambrysus puncticollis Stal, X 6.
(D) — Ambrt/sus parviceps Montandon, X 6.

La Rivers : The Ambrysus of Mexico

1339

PLATE XCV

1340 The University Science Bulletin

PLATE XCVI

(E) — Ambrysus pudicus pudicus Stal, X 6.
(F) — Ambrysus pudicus barbed Usinger, X 6.
(G) — Ambrysus abortus La Rivers, X 6.
(H) — Ambrysus hungerfordi Usinger, X 6.

La Rivers: The Ambrysus of Mexico

1341

PLATE XCVI

1342 The University Science Bulletin

PLATE XCVII

(J) — Amhrysus pulchellus Montandon, X 6.
(K) — Ambrijsus vanduzeei Usinger, X 6.
(L) — Amhrysus buenoi Usinger, X 6.
(M) — Amhrysus mormon australis La Rivers, X 6.

La Rivers: The Ambrysus of Mexico

1343

PLATE XCVII

55—6490

1344 The University Science Bulletin

PLATE XCVIII

(N) — Ambrysus scalenus La Rivers, X 6.
(O) — Ambrysus convexus Usinger, X 6.
(P) — Ambrysus inflatus La Rivers, X 6.
(Q) — Ambrysus magniceps La Rivers, X 6.

La Rivers: The Ambrysus of Mexico

1345

PLATE XCVIII

1346 The University Science Bulletin

PLATE XCIX

(R) — Ambry sus portheo La Rivers, X 6.

( S ) — Ambrysus hydor La Rivers, X 6.

(T) — Ambrysus lunatus lunatus Usinger, X 6.

(U) — Ambrysus lunatus menoides La Rivers, X 6.

La Rivers: The Ambrysus of Mexico

1347

PLATE XCIX

1348 The University Science Bulletin

PLATE C

(V) — Ambnjsus dilatus Montandon, X 6.
(W) — Ambnjsus cosmius La Rivers, X 6.
(X) — Ambnjsus guttatipenni-s Stal, X 6.
(Y) — Ambnjsus fuscus Usinger, X 6.

La Rivers : The Ambrysus of Mexico

1349

PLATE C

■

THE UNIVERSITY OP KANSAS

SCIENCE BULLETIN

Vol. XXXV, Pt. II] September 10, 1953 [No. 11

A Revision of the Genus Buenoa ( Hemiptera
Notonectidae )

Fred S. Truxal
TABLE OF CONTENTS

PAGE

I. Introduction 1353

II. Acknowledgments 1353

III. Biology of the Buenoa 1354

Habitat 1354

Activity 1354

Food 1354

Mating 1355

Oviposition 1355

Immature Stages 1355

IV. Distribution and Phylogeny 1355

V. Geographical Variation 1361

VI. Taxonomy of the Buenoa 1364

Brief History of the Taxonomy of Buenoa 1364

Technique of Identification 1365

Family Characteristics of Notonectidae 1366

Key to the Genera of Notonectidae 1367

Description of the Genus Buenoa 1367

Key to the species of Buenoa 1369

Buenoa antigone ( Kirkaldy ) 1374

Buenoa antigone antigone (Kirkaldy) 1376

Buenoa antigone carinata (Champion) 1379

Buenoa femoralis ( Fieber ) 1382

Buenoa ida Kirkaldy 1383

Buenoa crassipes (Champion) 1385

Buenoa arizonis Bare 1387

Buenoa absidata n. sp. 1391

Buenoa tarsalis n. sp 1392

Buenoa rostra n. sp 1395

Buenoa margaritacea Torre-Buenoa 1397

Buenoa scimitra Bare 1404

(1351)

1352 The University Science Bulletin-

page

Buenoa uhleri n. sp 1409

Buenoa albida ( Champion ) 1412

Buenoa pattens (Champion) 1414

Buenoa pallipes (Fabricius) 1418

Buenoa platycnemis (Fieber) 1421

Buenoa omani n. sp 1426

Buenoa macrotrichia n. sp 1429

Buenoa nitida n. sp 1430

Buenoa mutabilis n. sp 1432

Buenoa arida n. sp 1435

Buenoa speciosa n. sp 1437

Buenoa gracilis n. sp 1439

Buenoa communis n. sp 1442

Buenoa artafrons n. sp 1444

Buenoa macrotibialis Hungerford 1446

Buenoa limnocastoris Hungerford 1450

Buenoa confusa n. sp 1453

Buenoa fuscipennis ( Berg) 1460

Buenoa amnigenus (White) 1462

Buenoa incompta n. sp 1466

Buenoa oculata n. sp 1467

Buenoa salutis Kirkaldy 1469

Buenoa thomasi n. sp 1473

Buenoa alterna n. sp 1474

Buenoa unguis n. sp 1476

Buenoa excavata n. sp 1479

Buenoa macrophthalma (Fieber) 1480

Buenoa hungerfordi n. sp 1483

Buenoa distincta n. sp 1485

Buenoa paranensis Jaczewski 1486

VII. Literature Cited 1488

VIII. Index 1490

IX. Illustrations 1491

Abstract: This paper constitutes a revision of the genus Buenoa (Noto-
nectidae, Hemiptera). Although it is primarily taxonomic in nature, material
on the biology of the genus is included.

Prior to this work, twenty-two species were known. Twenty-two new spe-
cies are described in this paper and all old species are redescribed. Two new
subspecies have been designated. Three species are placed in synonomy.

The new species are: B. absidata (South America), B. tarsalis (South
America), B. rostra (West Indies, South America), B. uhleri (United States,
Mexico), B. omani (United States, Mexico), B. macrotrichia (South America),
B. nitida (South America), B. mutabilis (West Indies, South America),
B. arida (United States), B. speciosa (United States, Mexico), B. gracilis
( Mexico, Central America, West Indies, South America ) , B. communis ( South
America), B. artafrons (United States), B. confusa (Canada, United States,
Mexico, West Indies), B. incompta (South America), B. oculata (South Amer-
ica), B. thomasi (Mexico), B. alterna (Mexico), B. unguis (South America),

A Revision of the Genus Buenoa 1353

B. excavata (South America), B. hungerfordi (United States, Mexico), B.
distincta (Mexico).

New subspecies designated: B. antigone antigone (Kirkaldy), B. antigone
carinata (Champion).

Species placed in synonomy are: B. naias (Kirkaldy), B. dentipes Jaczewski,
B. mallochi Jaczewski.

Up to and including this paper (exclusive of synonyms) there have been
described forty species and two subspecies of Buenoa.

INTRODUCTION

This paper is the result of a taxonomic study of the genus Buenoa.
Its primary purpose is to compile and revise previous knowledge
concerning the genus and to present additional information made
available by the present study. The genus now includes forty
species, twenty-two of which are new.

Approximately 28,000 specimens of Buenoa were examined during
the study. The types of all the species were studied except those
of well-known species and those which have been lost or destroyed.
Completion of this paper was made possible only by the full co-
operation of a number of institutions and entomologists who made
available their type material. I was extremely fortunate in having
at my disposal material compared with types in European museums
by Dr. H. B. Hungerford in 1928, as well as notes and sketches con-
cerning these species.

Large series of many species were of great value in alleviating
confusion concerning species with both brachypterous and macrop-
terous forms and polytypic species where geographic variation
occurs.

ACKNOWLEDGMENTS

I am deeply indebted to Dr. H. B. Hungerford for suggesting
this study and for the encouragement and guidance he so readily
gave. I sincerely appreciate the helpful criticisms and numerous
suggestions so willingly offered by Dr. C. D. Michener, Dr. R. H.
Beamer, Dr. Kathleen Doering, and Dr. R. E. Beer, all of the Uni-
versity of Kansas. I wish to thank my wife, Mrs. Margaret D.
Truxal, for typing much of this paper.

A great majority of the material studied is located in the Francis
Snow Entomological Collections. In addition, I am deeply grate-
ful for material made available by the following museums and col-
lections, without which this study would have been impossible:
United States National Museum; American Museum of Natural
History; British Museum (Natural History); Museum National

1354 The University Science Bulletin

D'Histoire Naturelle, Paris; Zoologisches Museum der Humboldt-
Universitat, Berlin; Das Zoologische Staatsinstitut und Zoologische
Museum, Hamburg; Naturhistorisches Museum Zoologische Abteil-
ung, Vienna; Rijksmuseum van Natuurlijke Historie, Leiden; Museo
Argentino de Ciencias Naturales, Buenos Aires; University of Michi-
gan; Tulane University; California Academy of Sciences; J. C. Lutz
Collection, Philadelphia, Pennsylvania; C. J. Drake Collection, Ames,
Iowa.

BIOLOGY OF THE GENUS BUENOA

The information concerning the biology of Buenoa has been
based upon B. margaritacea studied in the vicinity of Lawrence,
Kansas, unless otherwise indicated.

Habitat
Buenoa is found for the most part in freshwater pools, lakes, and
ponds. Some species are found along the shores of slow and fast
flowing streams. Most members of this genus inhabit the open
water although many are found in water with aquatic vegetation.
Only seven species have been found that have brachypterous forms
which are incapable of flying from temporary pools or ponds.

Activity

Members of this genus in their various stages of development
have been taken in temperate climates during every month of the
year. Upon breaking the ice from ponds during the month of
December, adult Buenoa have been seen swimming sluggishly in
the water below. The Buenoa swim gracefully on their backs in
almost perfect equilibrium with the water. However, immediately
after replenishing their air supply at the surface, the insect must
use vigorous strokes with its hind legs to descend. Soon thereafter,
it is able to again attain its equilibrium with the water.

Food
As to food habits, Buenoa are predaceous, feeding largely upon
Entomostraca and mosquito larvae which they hold in the crib
formed by the bristles arming the fore legs. When these small
animals pass within view, the Buenoa makes a sudden dash to cap-
ture them. If missing on the first attempt, no further effort appears
to be made. Bare (1928) reports that upon dissecting the abdo-
men of an adult female, he found what appeared to be Spirogyra
chloroplasts in the mid-gut. He states that Buenoa, therefore, may
feed upon algae when other food is not available.

A Revision of the Genus Buenoa 1355

Mating
As in the case of Anisops, adult male Buenoa possess various
stridulatory areas which are used in courtship prior to mating.
Hungerford (1924) described the chirping sounds made by
B. limnocastoris in the laboratory at Douglas Lake, Michigan. The
serenade is similar to the sound produced when one draws a nail
quickly across the teeth of a rubber comb. The stridulating male
follows beneath and behind the female for a few moments and
when within a half inch or so of the female, the chirping changes
to a hum and is followed by a sudden dash to embrace her. If she
eludes him, the male begins all over again or transfers his attentions
to another.

Oviposition

Oviposition in temperate areas of the United States appears to
commence as early as the latter part of April and continues into
August. The eggs are placed individually within the stems of
aquatic plants with a portion of the surface exposed. The female
is equipped with a pair of ovipositor valves armed with rows of
sharp teeth for the purpose of excavating holes in plant tissue. It
is not yet known whether the same female lays more than one egg
in a day. The total number of eggs that a female lays has not been
determined but as many as twelve eggs have been dissected from
the abdomen of a single individual. The eggs are elongate and at
least a portion of the surface is covered with fine hexagonal reticula-
tions. The eggs hatch in slightly less than two weeks.

Immature Stages
Buenoa undergoes five nymphal instars as do the majority of
aquatic Hemiptera. Wing pads appear first in the third stage
nymph. Some indication of sex is noted in the form of the seventh
abdominal sternum in the third, fourth, and fifth stage nymphs.
Adulthood is reached in slightly less than two months in the labora-
tory. There is more than one generation a year. The insect over-
winters as an adult.

DISTRIBUTION AND PHYLOGENY

The insects belonging to the genus Buenoa are limited to the
Neotropical region and southern portions of the Nearctic region.
The geographical range extends from Canada, through the United
States, Mexico, Central America, South America, and the West
Indies. Its counterpart in the Eastern Hemisphere, Anisops, ex-
tends through Africa, Madagascar and neighboring islands, the

1356 The University Science Bulletin

Mediterranean area, extending eastward to the islands of the South
Pacific, thence northward into China and Japan. The relationship
between these two genera is very close. Buenoa appears to be the
more primitive. This conclusion is based primarily on the fact that
the males of Buenoa possess an extra segment in the front tarsus.
Anisops is the only genus of the Notonectidae in which the males
have only a single segment in the front tarsus. The antiquity of
the differentiation between Buenoa and Anisops is unknown.

Because of the abundance and diversity of Buenoa in the tropics
and the smaller number of species in America North of Mexico,
one might speculate as to a Neotropical origin with subsequent
invasion of the Nearctic area. There are several possibilities as to
the period and manner of this invasion.

The present distribution pattern of the genus suggests that these
insects invaded North America from the south, possibly late enough
in Tertiary times so that temperate climates no longer reached the
Bering Strait area. If this is the case, the geographical isolation of
Buenoa and Anisops must have resulted from migration over water
and the dating of the differentiation is impossible. It is desirable
to point out that there are species in both genera that must re-
peatedly migrate over water to maintain their specific identity.
Examples of Anisops illustrating this are A. nasuta Fieber, known
from New Guinea, Celebes, Friendly Islands, Australia, and Guam;

A. batillifrons Lundblad, known from Formosa, Iriomote Island,
Hainan Island, Burma, Assam, India, Philippine Islands, and Oki-
nawa Island; A. tahitiensis Lundblad, known from Guadalcanal,
New Guinea, Andaman Islands, New Hebrides, Philippine Islands,
and Okinawa Island. Examples of Buenoa illustrating this are

B. gracilis, known from Mexico, Honduras, Panama, Cuba, Jamaica,
Puerto Rico, St. Croix Island, Grenada Island, and Peru; B. scimitra,
known from the United States, Mexico, Cuba, Jamaica, and Puerto
Rico; B. albida, known from the United States, Mexico, and Puerto
Rico. If repeated transfer over a few hundred miles of water is
possible, then transfer over thousands of miles of water once in
millions of years is likely.

If one argues against the migration over water to explain the
differentiation of Buenoa and Anisops another suggestion is pos-
sible. Buenoa might have occurred in North America early in the
Tertiary (Eocene) when a warm climate extended over most of the
North American continent. At this time, one might assume that
the genus migrated across the Bering Strait region to later give rise

A Revision of the Genus Buenoa 1357

to Anisops. The adaptability to warm climates would have allowed
the Buenoa to occur in the Bering Strait region and to cross to the
Old World at a time in the early Tertiary when this area was warm.
At a later period these forms isolated by continental separation and
cold northern climates diverged to form the Anisops of the Eastern
Hemisphere and the Buenoa of the Western Hemisphere.

It is desirable to point out that although Buenoa is more primitive
than Anisops, there is no certainty that America is where the Buenoa-
Anisops group arose. On the contrary, the possibility exists that
Buenoa or a common ancestor, was once holarctic and was replaced
in the Old World by a more specialized type, Anisops.

The most widely distributed species within the genus is B, mar-
garitacea. Its range extends from Manitoba, Canada, in the north
to Tamaulipas, Mexico, in the south and from New York in the east
to California in the west. The greatest North-South distribution
is shown by two closely related species, B. Hmnocastoris and B. con-
fusa. These species range from Manitoba, Canada, in the north to
Florida and Grand Cayman Island in the south.

It is interesting to note the distribution pattern of the species
within the genus. The range of the genus has been divided into
eight geographical subdivisions as follows: (1) Canada, (2)
Northern half of the United States (further divided into Northeast
and Northwest), (3) Southern half of the United States (further
divided into Southeast and Southwest), (4) Mexico, (5) Central
America, (6) West Indies, (7) Tropical South America, and (8)
Temperate South America. The area boundaries have been arbi-
trarily indicated in some cases. The United States has been divided
into North and South by the parallel of 40° N. latitude. This same
area has been further divided into East and West by the eastern
borders of Montana, Wyoming, Colorado, and New Mexico. South
America has been divided into tropical and temperate areas by
arbitrarily selecting the parallel of 30° S. latitude. The number of
species in each area is indicated as follows: (1) Canada, 4 sp.; (2)
Northern half of the United States, 4 sp. (Northeast, 4 sp.; North-
west, 0 sp.); (3) Southern half of the United States, 11 sp. (South-
east, 7 sp.; Southwest, 6 sp. ); (4) Mexico, 19 sp.; (5) Central
America, 8 sp.; (6) West Indies, 11 sp.; (7) Tropical South America,
23 sp.; (8) Temperate South America, 2 sp. The above informa-
tion indicates that the area of greatest abundance is Tropical South
America. As will be shown below, the area of greatest structural
diversity is also Tropical South America.

1358

The University Science Bulletin

Buenoa is a morphologically monotonous group with few char-
acters. These characters vary within definitely circumscribed
limits. It is the combinations of these few characters which make
recognition of the various species possible. Tables 1, 2, and 3 show
the way in which certain of these characters are correlated. Actu-
ally, the striking fact is the lack of correlation. No two characters
are correlated better than those indicated in the tables. They are
seemingly meaningless recombinations and, therefore, the establish-
ment of a phylogeny is difficult or impossible.

Table 1. Scatter diagram showing the relationship between the number of
sclerotized ridges in the femoral strididatory area and the ratio of the width
of the synthlipsis to the anterior width of the vertex in Buenoa. Figures
within grid indicate numbers of species.

<
S3
O

o"
2

70
65
60
55
50
45
40
35
30
25
20
15
10
5

75
70
65
60
55
50
45
40
35
30
25
20
15
10
5

1 2 3 4 5 6 7 8 9 10 11 12 13 14 15
15 15 15 15 15 15 15 15 15 15 15 15 15 15 15

Width of Synthlipsis: Width of Vertex

A Revision of the Genus Buenoa

1359

Table 2. Scatter diagram showing the relationship between the number of
teeth in the tibial comb and the number of sclerotized ridges in the femoral
stridulatory area in Buenoa. Figures within grid indicate numbers of species.

<
w

K
<

H
CO

J
O

W
U

70
65
60
55
50
45
40
35
30
25
20
15
10
5

10 15 20 25 30 35 40 45 50 55 60 65 70 75

No. Teeth in Tibial Comb

Table 3. Scatter diagram shoiving the relationship between the ratio of the
width of the head to the width of the vertex and the ratio of the width of the
synthlipsis to the width of the vertex in Buenoa. Figures within grid indi-
cate numbers of species.

a
H

X
H

-
<
w

fa
o

X
H

9
8
7
6
5
4
3
2
1

0 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15
15 15 15 15 15 15 15 15 15 15 15 15 15 15 15

Width of Synthlipsis: Width of Vertex

56—6490

1360 The University Science Bulletin

Some features, however, can be established as probably primitive.
Some examples: the moderate width of the synthlipsis, a moder-
ate number of sclerotized teeth in the stridulatory areas, and
possession of a gland-like structure located laterally on the first ab-
dominal segment. On the other hand, other features can almost
certainly be termed specialized. Some examples are: holoptic eyes,
a wide or narrow synthlipsis, intermediate tarsal emargination, the
bizarre shape of the rostral prong, and a large spine on the caudo-
sinistral margin of the seventh abdominal tergum which lies hori-
zontally.

In the absence of fossils and due to the few useful characters
and the baffling way in which they are combined and recombined,
no complete scheme of relationships for the species of Buenoa can
be devised at present. However, certain forms stand out as ob-
viously specialized branches of the tree. These branches are as

follows :

1 2 3 4

hungerfordi amnigenus excavata unguis

distincta salutis

macrophthalma oculata

On the other hand, there are forms which are presumably rela-
tively primitive. These groups are as follows:

12 3

antigone pallens margaritacea

femoralis pallipes uhleri

ida platycnemis scimitra

crassipes omani albida

absidata nitida

arizonis mutabilis

tarsalis macrotrichia

The greatest abundance of the specialized forms is found in
tropical South America. This is shown by the following distribu-
tion of these forms: ( 1 ) Southwest United States, 1 sp.; (2) Mexico,
2 sp.; (3) West Indies, 2 sp.; (4) Tropical South America, 7 sp.;
( 5 ) Temperate South America, 1 sp. The distribution of the species
with the most primitive combination of characters is as follows:
(1) Canada, 1 sp.; (2) Northeast United States, 1 sp.; (3) South-
east United States, 2 sp.; (4) Southwest United States, 1 sp.; (5)
Mexico, 9 sp.; (6) Central America, 6 sp.; (7) West Indies, 7 sp.;
( 8 ) Tropical South America, 10 sp.

The distribution pattern of Buenoa in the United States appears
to substantiate the hypothesis that the group invaded this region
by way of the Mexican Plateau. The abundance of species in New
Mexico and Arizona and the absence of species in the Northwest

A Revision of the Genus Buenoa 1361

compared to the greater numbers in the Southwest is an interesting
problem. This distribution pattern is found in many groups of
plants and animals, a flora and fauna largely peculiar to the old
Mexican Plateau ranging northward into Arizona and New Mexico.
Seemingly the Buenoa adapted to the environmental conditions of
this old plateau region have remained, for no obvious reason, in
this area of the United States and Mexico. These species are un-
doubtedly adapted to an arid environment, whereas those species
found in the southeastern United States are obviously adapted to
humid conditions. The latter forms are also found, for the most
part, in the humid areas of Mexico, Central America, and the West
Indies. Since it is not clear how humidity would limit the dis-
tribution of aquatic insects, other environmental factors are prob-
ably involved. In any case, it is interesting that an aquatic group
shows the same distributional pattern as many terrestrial groups.
It is obvious that Buenoa is influenced in its distribution by cold.
The present northern limit of the genus is approximately 52° N.
latitude. Buenoa was undoubtedly more widespread in the past
when a warm climate extended over most of North America. It is
logical to assume that as the cold climate progressed southward
during the Tertiary, these insects were also pushed southward.
Thus, one might have the explanation of the present northern limit
of this genus in Canada.

GEOGRAPHICAL VARIATION

It is interesting to note that in spite of wide ranges, a striking
feature in Buenoa is the lack of geographical variation in most
species studied. There are only three good examples that are ob-
vious. One case is primarily an example of size variation. In the
two species with the greatest North-South distribution, B. limno-
castoris and B. confusa, one finds a size variation from a large
northern form to a small southern form as indicated in tables 4
and 5. This type of geographical variation is comparable to that
found in warm-blooded vertebrates where the populations that
live in a cooler climate tend to be larger than those that live in a
warm district ( Bergmann's rule ) .

Although too few specimens of B. limnocastoris were available
for this study, it is interesting to note that it parallels B. confusa,
where many specimens were available, in its size variation pattern.
The difference in the mean body lengths between widely separated
localities is statistically significant.

1362

The University Science Bulletin

Table 4. North-South distribution of B. confusa with variations of body
lengths of brachypterous specimens.

Collection localities

Total
specimens

Range and mean body lengths
in mm. (mean in bold-face)

(North-South)

Males

Females

Manitoba & Alberta, Can.
South Dakota

5 . 65—5 . 78—5 . 85
5 . 26—5 . 39—5 . 52
5.07—5.32—5.78
4 . 74—5 . 01—5 . 39
5.20—5.52—5.26

4.81— 5 07— 5.26
4 . 94—5 . 05—5 . 20

4 . 87

4.42—4.51—4.61
4.16—4.48—4.68
4.16—4.58—5.00

6.04—6.51—6.82
5.39—6.07—6.76

Minnesota

Michigan

Connecticut

New York & New Jersey

Kansas

Texas

Mississippi & Alabama. . .

Georgia

Florida

5 31

4.87—5.19—5.65
5 . 20—5 . 46—5 . 72

4.94—5.11—5.39
5 . 13—5 . 28—5 . 52

4.94

4 . 55—4 . 59—4 . 68
4.42—4.60—4.87
4.29—4.61—5.33

Table 5. North-South distribution of B. limnocastoris with variations of
body lengths of brachypterous specimens.

Collection localities

Total
specimens

Range and mean body lengths
in mm. (mean in bold-face)

(North-South)

Males

Females

Quebec, Canada

Minnesota

6.17

5.85—6.01—6.17
5 . 65—5 . 88—5 . 98

5.52

5 59
4.74—4.99—5.07

6 37

5.95—6.05—6.43

Michigan

Maine & New Jersey

Virginia

Georgia & Florida

5.72—6.00—6.30

5 65

5.98

4.81—5.48—5.86

Within these two species, one also finds a variation in the develop-
ment of flight wings with consequent changes in the thorax and
hemelytra. The wings are either fully developed or very short,
there being no intermediate condition. This feature is indicated
in tables 6 and 7 and the differences in percentage from any one
locality to another is statistically highly significant. There is no
correlation between the development of the flight wings and the
sex or season of the year. It is interesting to note that the long-
winged forms have been recorded only in the northern and southern
portions of the geographic ranges of these species. A possible ex-
planation of this phenomenon is not available at the present time.

A Revision of the Genus Buenoa

1363

Table 6. North-South distribution of B. confusa with variation in
development of flight wings.

Collection localities
(North-South)

Manitoba and Alberta, Canada . .
North Dakota and South Dakota

Minnesota

Michigan

Connecticut

New York and New Jersey

Kansas

Texas

Louisiana

Mississippi and Alabama

Georgia

Florida

Grand Cayman Island, W. I

Total no.
specimens

210

239

Percentage of
short-winged forms

32%

25%

89%

100%

40%

72%

88%

71%

Table 7. North-South distribution of B. limnocastoris with variation in

development of flight wings.

Collection localities
(North-South)

Quebec, Canada

Minnesota

Michigan

Maine, New Jersey and Virginia

Georgia

Florida

Total no.

Percentage of

specimens

short-winged forms

100%

72%

148

99%

100%

50%

85%

A third case of geographical variation is an example of a poly-
typic species B. antigone, which has broken up into geographical
races, each differing in minor morphological characteristics. There
is very little difference between the neighboring races, but the
peripheral forms or the two extremes are very distinct. Not only
does one find a size difference but also variations in the divergence
of the lateral margins of the notocephalon anterior to the synthlipsis,
and slight differences in the form of the rostral prong, pronotum,
femoral stridulatory area, and hind femur. Intergrades between
the two extremes are evident. The large form of B. antigone is con-
fined to the southern portions of the United States, Mexico, and
south as far as British Honduras. The small form ranges from
Mexico, the West Indies, and throughout Central and South Amer-
ica south to Argentina. More detailed data concerning the distri-
bution pattern are given under the species description.

1364 The University Science Bulletin

TAXONOMY OF THE GENUS BUENOA

Much confusion has existed in the classification of the Buenoa
as a result of poor descriptions based primarily on color patterns
and inaccurate determinations. Most of the papers dealing with
this genus are isolated descriptions of species. Little work of a
general nature has been published.

Brief History of the Taxonomy of the Buenoa

The genus Buenoa was erected by Kirkaldy in 1904. He included
in this genus all the species formerly included in Anisops that in-
habit the Western Hemisphere and possess a two-segmented front
tarsus in the male. He retained the name Anisops for the Eastern
Hemisphere forms that possess a one-segmented front tarsus in the
male. In this paper Kirkaldy recognized ten former Anisops species
as belonging to the Buenoa, placed two species in synonomy, and
described two species as new. He designated B. antigone (Kir-
kaldy) as the genotype.

The first Buenoa species was described in 1803 by Fabricius in
"Systema Rhyngotorum" as Notonecta pallipes. Almost forty-eight
years later, in 1851, Fieber described the next three species of
Buenoa to be made known as Anisops platycnemis, A. femoralis,
and A. macrophthalmus. In the same publication Fieber described

A. elegans, a name which has for a generation been erroneously
applied to Buenoa confusa described below.

In 1879, twenty-seven years later, Berg and White added two
more species. These were A. fuscipennis Berg and A. amnigenus
White. Some time later, 1899, Kirkaldy described A. antigone
which he later ( 1904 ) designated as the genotype of his new genus
Buenoa, and A. naias which the present writer has relegated to
synonomy under B. fuscipennis (Berg).

In 1901, Champion introduced, with some success, the use of
structural characteristics for species determination in describing
four new species, A. albidus, A. crassipes, A. pollens, and A. cari-
natus. All of these were Buenoa. A. carinatus was later placed in
synonomy with B. antigone by Kirkaldy (1904) but is here con-
sidered a subspecies of that species.

Kirkaldy in 1904 established the genus Buenoa, described two
new species, B. ida and B. salutis, and placed B. carinata and

B. macrophthalma in synonomy with B. antigone. B. macroph-
thalma, however, is easily distinguished from B. antigone and is a
good species, while B. carinata should, as already indicated, be
recognized as a subspecies of B. antigone.

A Revision of the Genus Buenoa 1365

In 1908, J. R. de la Torre-Bueno found that the Buenoa known
to a generation of entomologists as B. platycnemis was actually an
undescribed species which he designated B. margaritacea.

Not until fourteen years later, in 1923, do additional new species
appear in the literature. In this year Hungerford described
B. limnocastoris. One year later, 1924, he described another species,
B. macrotibialis.

Bare in 1925 described a widespread species, B. scimitra, and
three years later, in 1928, Jaczewski described three new species,
B. dentipes, B. mallochi, and B. paranensis, all from the state of
Parana, Brazil. The present writer has found that B. dentipes is in
synonomy with B. fuscipennis, and B. mallochi is in synonomy with
B. sahitis. Finally in 1931, B. arizonis was described by Bare.

The present study is the first generic revision since Kirkaldy's
paper in 1904. Every species except B. paranensis Jaczewski has
been accounted for. The types of this species have been destroyed.

Technique of Identification

To facilitate accurate and rapid interpretation of the species
descriptions, it is desirable to mention certain techniques and
terminology used by this writer. As indicated previously, in most
species only the males possess adequate structural characters for
specific separation. The female, however, can usually be correlated
with the male by general resemblance to the male, and various
structural details.

The characters and terminology used in the key to the species
are, for the most part, self-explanatory or indicated in the illustra-
tions. The reader, however, should be familiar especially with the
method used in making the various measurements.

The measurements of the head and pronotum have been made
from above the dorsal surface of the insect when held in a hori-
zontal position with the transverse and longitudinal axes horizontal.
The length of the rostral prong is determined by placing the insect
in such a position that the rostral prong is horizontal. The measure-
ment is then taken from the base of the rostral prong to the apex
along a median longitudinal line. The length of the third rostral
segment is measured along the frontal surface.

Seldom will it be necessary to make observations of the stridula-
tory area or chaetotaxy of the front leg to make specific determina-
tions. When such a study is desired the front leg must be removed
and cleared. Removal of the leg is best accomplished by placing
the bent point of a fine dissecting needle at the base of the coxa or
trochanter on its inner surface and rotating the needle outward.

1366 The University Science Bulletin

The writer finds that this procedure works better on a dry speci-
men than on one that has been relaxed. It is desirable on occasions
to observe only the femoral stridulatory area. This is easily accom-
plished by relaxing the insect and pulling the leg to a position
perpendicular to the body. Best results in this relaxing procedure
are obtained by touching the specimen with a camel's-hair brush
that has been dipped in five per cent alcohol or a relaxing fluid
consisting of alcohol (95%), 106 cc; distilled water, 98 cc; benzol,
14 cc; ethylacetate, 38 cc.

The illustrations in this paper show only those characters that
the writer feels are of particular taxonomic importance. For ex-
ample, the thick covering of hair-like setae on the inner surface of
the leg is omitted so as not to obscure the larger characteristic setae.

Family Characteristics of the Notonectidae

The family Notonectidae is composed of aquatic forms differing
from all other such insects, except Pleidae and Helotrephidae, in
the habit of swimming on their backs. They are deep-bodied, flat
ventrally, and convex dorsally. The eyes are large, reniform, and
twice sinuate. Ocelli are absent in this family. The antennae,
which are usually hidden from above, consist of three or four seg-
ments, and arise on the latero-ventral surface of the head. The
beak is four-segmented. The hind pair of legs is more or less flat-
tened and heavily fringed for swimming, while the front and middle
pairs are adapted for grasping. The tarsi are two or three seg-
mented except for the one-segmented fore tarsi of the male Anisops.
The tarsi possess two apical claws which are prominent except on
the hind legs where they are greatly reduced and inconspicuous.
The pronotum is transverse, convex, and usually narrower an-
teriorly. The abdomen possesses a prominent longitudinal mid-
ventral keel, having hairs at least along its lateral margins. To-
gether with the hairs along the sides of the venter, they cover the
two longitudinal troughs forming air chambers.

Key to the Genera, of Notonectidae

(Modified from Hungerford, 1933)

The family Notonectidae consists of eight genera which may be classified by
the following key: *

A. Hemelytral commissure without definite hair-lined pit at anterior

end (Subfamily Notonectinae )

B. Intermediate femur with anteapical pointed protuberance

and antennae 4-segmented ( Tribe Notonectini )

C. Anterolateral margins of prothorax not foveate.

Notonecta

A Revision of the Genus Buenoa 1367

CC. Anterolateral margins of prothorax foveate .... Enithares
BB. Intermediate femur without anteapical pointed protuber-
ance and antennae 3- or 4-segmented ( Tribe Nychini )

C. Sides of prothorax not foveate, the lateral ledge
straight. Infracoxal plates bare but margined with
hair. Intermediate tarsus with two well-defined
segments and a very small basal one in both sexes.

Neonychia

CC. Sides of prothorax foveate, the lateral ledge curving

downward to embrace the fovea. Infracoxal plates

covered with hair. Intermediate tarsus with one

well-defined segment, except in males of Nychia.

D. Antennae 3-segmented Nychia

DD. Antennae 4-segmented Martarega

AA. Hemelytral commissure with definite hair-fined pit at anterior

end ( Subfamily Anisopinae )

B. Ventral abdominal keel not extending onto last abdominal
segment. Male genital capsule cleft behind. Males with-
out stridular protuberance on front tibia. Females with

short gonapophyses Paranisops

BB. Ventral abdominal keel extending onto last abdominal seg-
ment. Male genital capsule closed behind. Males with
stridular protuberance on front tibia. Females with long
subspatulate gonapophyses.

C. Male with anterior tarsus 2-segmented Buenoa

CC. Male with anterior tarsus 1-segmented Anisops

Description of the Genus Buenoa

(Genotype Buenoa antigone Kirkaldy, 1904)

1901. Anisops, Champion, G. C. Biologia Centrali Americana, Heteroptera,

vol. II, p. 371.

1904. Buenoa Kirkaldy, G. W. Wiener Ent. Zeitung, vol. XXIII, pp. 120-135.

1909. Buenoa, Torre-Bueno, J. R. de la. Jr. New York Ent. Soc, vol. XVII,

pp. 74-75.

1917. Buenoa, Hungerford, H. B. Ent. News, vol. XXVIII, pp. 175-176.

1919. Buenoa, Hungerford, H. B. Univ. Kansas Sci. Bull., vol. XI, pp. 41-42,

173 and 177.

1923. Buenoa, Hale, H. M. Records South Australian Mus., vol. II, p. 399.

1924. Buenoa, Hungerford, H. B. Ann. Ent. Soc. Am., vol. XVII, p. 225.
1926. Buenoa, Blatchley, W. S. Heteroptera Eastern N. Am., p. 1056.
1928. Buenoa, Bare, C. O. Univ. Kansas Sci. Bull., vol. XVIII, pp. 265-349.
1928. Buenoa, Jaczewski, T. Ann. Mus. Zool. Polonici, vol. VII, p. 123.

This genus is very similar to its counterpart in the Old World,
the genus Anisops. It is readily distinguishable from all other
genera of the Notonectidae by the combination of characters noted
in the key. In addition it differs from its close relative, Anisops,
by possessing in all males (except B. fuscipennis, B. hungerfordi,
B. distincta, and B. macrophthalma) a small gland-like struc-

8 See 1949. Truxal, F. S. Jr. Kansas Ent. Soc, vol. XXII, p. 2. Key under A-BB-C,
should read "sides of prothorax not foveate."

1368 The University Science Bulletin

ture located laterally on the first abdominal segment. Male geni-
talic characters as indicated by Truxal (1952) and the femoral
stridulatory area on the front femur of most males also aid in dis-
tinguishing Buenoa from Anisops.

The members of this genus are long, slender insects, and more or
less conical in cross-section. The widest portion of the body is
usually at a point midway of the longitudinal axis. The head is
somewhat rounded anteriorly and closely connected to the thorax.
The posterior margins of the eyes invade the pronotum. The eyes
are large and not holoptic [except B. amniginus (White)]. As seen
from above, the notocephalon is usually sulcate medianly. The
labrum is small, triangular, and covered with many fine hair-like
setae. The rostrum is four-segmented, the third segment of the
male prolonged laterally into prongs which aid in stridulation. The
pronotum is trapezoidal in shape and wider than long. The meta-
thorax is the largest segment of the entire body. The legs each have
the same number of segments. All have a large coxa, small tro-
chanter, femur, tibia, two distinct tarsal segments, and two tarsal
claws at the end of the distal tarsal segment. The hemelytra are
generally hyaline and distinctly divided into clavus, corium, and
membrane, except in brachypterous forms.

The male genital capsule, as in other members of the family, is
composed of the ninth abdominal segment. Its tergum is narrow
and bridgelike while the sternum is strongly developed, boat-like
in shape, and fused posteriorly. The genital claspers are asym-
metrical, the left one being larger and hooked at the apex. The
genitalic characters are not important in species determination.

An interesting phenomenon of the internal anatomy of this genus
is the presence of haemoglobin cells in the abdomen as found by
Hungerford (1922). They are grouped about the tracheal trunks
which have their connections with the spiracles of the third to the
seventh abdominal segments, inclusive. Similar cells were found
by Poisson ( 1925 ) in the genus Anisops.

To the worker particularly interested in the morphology of the
Buenoa, reference should be made to Bare (1928). This study
applies to B. margaritacea Torre-Bueno in particular. Bare's text
figures, illustrating both internal and external structures, are well
done.

Key to the Males of Buenoa

1. Synthlipsis wide, one half or more the anterior width of

vertex 2

Synthlipsis narrow, less than one half the anterior width of
vertex 24

A Revision of the Genus Buenoa 1369

2. ( 1 ) Rostral prong longer than third rostral segment 3

Rostral prong equal to or shorter than third rostral segment . . 20
3.(2) Rostral prong with base originating laterally at or near distal

end of third rostral segment. ( Pi. CVI, fig. 47b ) 4

Rostral prong with base not originating laterally at or near
distal end of third rostral segment. (Pi. CXIII, fig. 66b) . . 12

4.(3) Over 7 mm. in length 5

Less than 7 mm. in length 7

5.(4) Tylus strongly inflated with shallow median depression form-
ing two lateral protuberances (Pi. CVI, fig. 47b); pro-

notum strongly convex B. arizonis Bare (p. 1387)

Tylus slightly inflated and smoothly rounded; pronotum not

strongly convex 6

6.(5) Rostral prong with posterior margin distinctly sinuate (Pi.
XII, fig. 63c); interocular cephalic space narrow. (Pi.

CII, fig. 16) B. speciosa n. sp. (p. 1437)

Rostral prong with posterior margin almost straight; inter-
ocular cephalic space relatively wide. (Pi. CII, fig. 15),

B. crassipes (Champion) (p. 1385)
7. ( 4 ) Head approximately half to more than half the length of pro-
notum along median longitudinal axis 8

Head approximately one third to less than one third the

length of pronotum along median longitudinal axis 10

8.(7) Tylus slightly inflated, with wide median depression; lateral
margins of frons distinctly divergent anteriorly and pos-
teriorly B. albida (Champion) (p. 1412)

Tylus inflated, smoothly rounded; lateral margins of frons

almost parallel 9

9.(8) Pronotum distinctly tricarinate; head equal to or wider than
humeral width of pronotum,

B. macrotibialis Hungerford (p. 1446)
Pronotum unimpressed, not tricarinate; head distinctly nar-
rower than humeral width of pronotum,

B. macrotrichia n. sp. (p. 1429)
10.(7) Rostrum robust with posterior margin of rostral prong dis-
tinctly sinuate (PI. CVII, fig. 50b); tylus only slightly

inflated B. rostra n. sp. (p. 1395)

Rostrum not robust and posterior margin of rostral prong

almost straight; tylus distinctly inflated 11

11.(10) Greatest width of head approximately five times the anterior
width of vertex; as viewed from above, vertex protuberant,

B. absidata n. sp. (p. 1391)
Greatest width of head approximately six times the anterior
width of vertex; as viewed from above, vertex slightly

indented B. pallipes (Fabricius) (p. 1418)

12. ( 3 ) Posterior margin of hind femur with more than forty setae in

ventral row 13

Posterior margin of hind femur with forty or less setae in
ventral row 14

1370 The University Science Bulletin

13.(12) Greatest width of head more than seven times the anterior
width of vertex; interocular cephalic space narrow,

B. artafrons n. sp. (p. 1444)
Greatest width of head less than seven times the anterior
width of vertex; interocular cephalic space relatively wide,

B. pattern (Champion) (p. 1414)

14. ( 12) Over 7.75 mm. in length; posterior margin of hind femur with

less than thirty, large setae in ventral row 15

Less than 7.75 mm. in length; posterior margin of hind femur
with thirty or more, small setae in ventral row 17

15. ( 14 ) Greatest width of head approximately seven times the anterior

width of vertex; head at least half the length of pronotum
along median longitudinal axis,

B. femoralis (Fieber) (p. 1382)
Greatest width of head six times or less the anterior width of
vertex; head one third the length of pronotum along

median longitudinal axis 16

16.(15) Pronotum almost unimpressed, not tricarinate; length of fore
femur two times the width at apex; approximately twenty-
five teeth in tibial comb B. ida Kirkaldy (p. 1383)

Pronotum tricarinate; length of fore femur three times or
more the width at apex; forty to fifty teeth in tibial comb,

B. antigone (Kirkaldy) (p. 1367)
17.(14) Pronotum distinctly tricarinate; fore femur narrowed at apex.

(PL CVII, fig. 49a) 18

Pronotum almost unimpressed, not distinctly tricarinate; fore

femur widened at apex. (PL CXI, fig. 60a) 19

18.(17) Intermediate leg with first tarsal segment deeply emarginate
on inner margin. (PL CVII, fig. 49b),

B. tarsalis n. sp. (p. 1392)
Intermediate leg with margins of first tarsal segment straight,

B. uhleri n. sp. (p. 1409)
19.(17) Rostrum with frontal surface protruding at base of rostral
prong (PL CXI, fig. 60b); interocular cephalic space rela-
tively wide; approximately thirty teeth in tibial comb and
seven sclerotized ridges in femoral stridulatory area,

B. nitida n. sp. (p. 1430)
Rostrum with frontal surface flat, not protruding; interocular
cephalic space narrow; approximately sixteen teeth in
tibial comb and four sclerotized ridges in femoral stridula-
tory area B. amani n. sp. (p. 1426)

20. ( 2 ) Stridulatory area absent on inner surface of fore femur 21

Stridulatory area present on inner surface of fore femur 22

21.(20) Over 8 mm. in length; greatest width of head less than five
times the anterior width of vertex; spine from caudo-
sinistral margin of seventh abdominal tergite large and
sickle-shaped, horizontal. (PL CII, fig. 14),

B. hungerfordi n. sp. (p. 1483)
Less than 8 mm. in length; greatest width of head six or more
times the anterior width of vertex; spine from caudo-

A Revision of the Genus Buenoa 1371

sinistral margin of seventh abdominal tergite small and
straight, vertical. (Pi. CII, fig. 11),

B. alterna n. sp. (p. 1474)
22.(20) Synthlipsis more than half the anterior width of vertex;
greatest width of head more than seven and one half times

the anterior width of vertex B. arida n. sp. ( p. 1435 )

Synthlipsis approximately half to less than half the anterior
width of vertex; greatest width of head less than seven and

one half times the anterior width of vertex 23

23. ( 22 ) Over 8)2 mm. in length; posterior margin of hind femur with
less than thirty, large setae in ventral row,

B. ida Kirkaldy (p. 1383)
Less than 8/2 mm. in length; posterior margin of hind femur
with more than thirty, small setae in ventral row,

B. margaritacea Torre-Bueno (p. 1397)
24.(1) Stridulatory area present on inner surface of fore femur. ... 25

Stridulatory area absent on inner surface of fore femur 42

25.(24) Fore femur narrowed at apex, with length more than three

times the width at apex. (Pi. CXII, fig. 64a) 26

Fore femur widened at apex, with length three times or less

the width at apex. (Pi. CXI, fig. 60a) 34

26. ( 25 ) Rostral prong equal to or shorter than third rostral segment . . 27

Rostral prong longer than third rostral segment 29

27.(26) Synthlipsis less than one third the anterior width of vertex;
fore tibia with four short, peg-like setae on inner surface
at apex (Pi. CXII, fig. 64d); femoral stridulatory area
with six to nine sclerotized ridges . B. gracilis n. sp. ( p. 1439 )
Synthlipsis one third or more the anterior width of vertex;
fore tibia without short, peg-like setae on inner surface at
apex; femoral stridulatory area with fifteen or more sclero-
tized ridges 28

28.(27) Fore femur with long, conspicuous, sword-shaped stridula-
tory area consisting of more than forty sclerotized ridges;
fore femur not accuminate at apex. ( PI. CVIII, fig. 52a ) ,

B. scimitra Bare (p. 1404)
Fore femur with small, subtriangular to oval stridulatory area
consisting of less than twenty-five sclerotized ridges; fore
femur accuminate at apex. (Pi. CVIII, fig. 51a),

B. margaritacea Torre-Bueno (p. 1397)

29. ( 26 ) Pronotum distinctly tricarinate 30

Pronotum almost unimpressed, not distinctly tricarinate 33

30.(29) Synthlipsis less than one third the anterior width of vertex;
femoral stridulatory area with more than twenty-five

sclerotized ridges B. communis n. sp. (p. 1442)

Synthlipsis more than one third the anterior width of vertex;
femoral stridulatory area with less than twenty-five sclero-
tized ridges 31

31.(30) Tylus slightly inflated with wide median depression,

B. alhida (Champion) (p. 1412)
Tylus slightly inflated, smoothly rounded 32

1372 The University Science Bulletin

32.(31) Over 8 mm. in length; greatest width of head more than six
and one half times the anterior width of vertex,

B. femoralis (Fieber) (p. 1382)
Less than 8 mm. in length; greatest width of head less than
six and one half times the anterior width of vertex,

B. uhleri n. sp. (p. 1409)
33.(29) Posterior margin of hind femur with less than forty setae in
ventral row; greatest width of head six and one half times

the anterior width of vertex B. mutabilis n. sp. (p. 1432)

Posterior margin of hind femur with more than forty setae
in ventral row; greatest width of head six times the
anterior width of vertex . B. pattens (Champion) (p. 1414)
34.(25) Head equal to or slightly wider than humeral width of pro-

notum 35

Head narrower than humeral width of pronotum 39

35.(34) Pronotum strongly inflated, with median length equal to
humeral width; greatest width of head less than six times
the anterior width of vertex,

B. limnocastaris Hungerford (p. 1450)
Pronotum not strongly inflated, with median length three
fourths to less than three fourths the humeral width; great-
est width of head six or more times the anterior width of

vertex 36

36.(35) Pronotum almost unimpressed, not tricarinate; rostral prong
with base originating laterally midway to near proximal

end of third rostral segment 37

Pronotum tricarinate, median carina distinct; rostral prong
with base originating laterally at distal end of third rostral

segment 38

37.(36) Interocular cephalic space very narrow (Pi. CII, fig. 16);
femoral stridulatory area with approximately sixty, fine,

sclerotized ridges B. arida n. sp. (p. 1435)

Interocular cephalic space relatively wide (Pi. CII, fig. 15);
femoral stridulatory area with six to nine, thick, sclerotized

ridges B. nitida n. sp. (p. 1430)

38.(36) Greatest width of head seven or more times the anterior

width of vertex B. confusa n. sp. (p. 1453)

Greatest width of head six and one half times the anterior

width of vertex B. macrotibialis Hungerford (p. 1446)

39.(34) Rostrum robust, with posterior margin of rostral prong dis-
tinctly sinuate. (Pi. CVII, fig. 50b) . B. rostra n. sp. (p. 1395)
Rostrum not robust and posterior margin of rostral prong

almost straight 40

40.(39) Pronotum distinctly tricarinate,

B. platycnemis (Fieber) (p. 1421)
Pronotum almost unimpressed, not distinctly tricarinate. ... 41
41.(40) Less than 6 mm. in length; greatest width of head more than
six times the anterior width of vertex; fore femur with
large, spatulate setae on inner posterior margin. ( PL CX,
fig. 59a ) B. macrotrichia n. sp. ( p. 1429 )

A Revision of the Genus Buenoa 1373

Over 6 mm. in length; greatest width of head six times or
less the anterior width of vertex; fore femur with small
setae on inner posterior margin. (Pi. CXI, fig. 60a),

B. nitida n. sp. (p. 1430)
42.(24) Greatest width of head five times or less the anterior width

of vertex 43

Greatest width of head more than five times the anterior

width of vertex 49

43. ( 42 ) Rostral prong equal to or longer than third rostral segment ... 44

Rostral prong shorter than third rostral segment 47

44.(43) Over 7 mm. in length; tylus strongly inflated with median
depression forming two lateral protuberances. (Pi. CXVI,

fig. 79b) B. macrophthalma (Fieber) (p. 1480)

Less than 7 mm. in length; tylus flat or slightly inflated and

smoothly rounded 45

45.(44) Eyes holoptic or synthlipsis extremely narrow, less than one

tenth the anterior width of vertex 46

Eyes not holoptic, synthlipsis wide, one third to more than
one third the anterior width of vertex,

B. thomasi n. sp. (p. 1473)
46.(45) Eyes holoptic; greatest width of head less than three times
the anterior width of vertex; over 4 mm. in length,

B. amnigenus (White) (p. 1462)
Eyes not holoptic, synthlipsis extremely narrow; greatest
width of head more than three times the anterior width of
vertex; less than 4 mm. in length,

B. salutis Kirkaldy (p. 1469)
47.(43) Front leg with inner tarsal claw shield-like at base (PL XV,
fig. 78a); synthlipsis narrow, approximately one fifth the

anterior width of vertex B. unguis n. sp. (p. 1476)

Front leg with inner tarsal claw gradually tapering from base
to apex; synthlipsis wide, approximately half the anterior

width of vertex 48

48. ( 47 ) Less than 8 mm. in length; trochanter of front leg with prom-
inent tubercle on posterior margin (PL CXIV, fig. 71a);
spine from caudo-sinistral margin of seventh abdominal
tergite small and straight, vertical. (PL CII, fig. 7),

B. fuscipennis (Berg) (p. 1460)
Over 8 mm. in length; trochanter of front leg with posterior
margin smoothly rounded; spine from caudo-sinistral mar-
gin of seventh abdominal tergite large and sword-shaped,

horizontal. (Pi. CII, fig. 13) B. distincta n. sp. (p. 1485)

49.(42) Synthlipsis one fourth to more than one fourth the anterior

Synthlipsis less than one fourth the anterior width of vertex . . 53

width of vertex 50

50.(49) Over 8 mm. in length; tylus strongly inflated with median
depression forming two lateral protuberances (PL CXVI,
fig. 79b); rostral prong extremely long. (PL CXVI, fig.
79b) B. macrophtlialma (Fieber) (p. 1480)

1374 The University Science Bulletin

Less than 8 mm. in length; tylus not inflated or inflated and
smoothly rounded; rostral prong relatively short. (Pi.

CXVI, fig. 71b) 51

51.(50) Trochanter of front leg with prominent tubercle on posterior
margin (PL XIV, fig. 71a); fore tibia with base and apex
approximately same width. . B. fuscipennis (Berg) (p. 1460)

Trochanter of front leg with posterior margin smoothly
rounded; fore tibia with base distinctly wider than apex. . 52
52.(51) Greatest width of head less than six times the anterior width
of vertex; fore tibia with approximately seventeen stout,
club-shaped setae on inner surface at apex. (PL CXV,
fig. 76d) B. thomasi n. sp. (p. 1473)

Greatest width of head six times or more the anterior width of
vertex; fore tibia without area of club-shaped setae on

inner surface at apex B. alterna n. sp. ( p. 1474 )

53.(49) Tylus strongly excavate with short anteromedial ridge. (PL

CXV, fig. 75b) B. excavata n. sp. (p. 1479)

Tylus not excavate, smoothly rounded 54

54.(53) Synthlipsis one fifteenth the anterior width of vertex; pro-
notum not tricarinate; intermediate leg with first tarsal
segment emarginate on inner surface. (PL CXIV, fig.
72d ) B. oculata n. sp. ( p. 1467 )

Synthlipsis one fifth the anterior width of vertex; pronotum
tricarinate; intermediate leg with margins of first tarsal
segment straight B. incompta n. sp. (p. 1466)

Buenoa antigone (Kirkaldy)
(PL CIII, fig. 30; PL CIV, figs. 42, 43)

This variable species is divisible into two subspecies as indicated
in the following pages.

Size: Male, length 7.80 mm. to 8.97 mm., greatest body width
2.46 mm. to 2.79 mm.; female, length 8.25 mm. to 9.75 mm., greatest
body width 2.60 mm. to 3.12 mm.

Color: General facies testaceous to dark brown. Head, pro-
notum, thoracic venter, and limbs sordid white to testaceous. Scu-
tellum rufo-testaceous to testaceous with base, brown to black.
Abdomen usually black except ventral keel and portions of con-
nexivum and dorsum, testaceous to light brown. This species
somewhat variable in color with some specimens entirely sordid
white to testaceous.

Male Structural Characteristics: As viewed from above, outline
of head laterally rounded, anteriorly truncate with vertex indented;
greatest width of head five and one-half to slightly more than six
times the anterior width of vertex and less than humeral width of

A Revision of the Genus Buenoa 1375

pronotum; synthlipsis one-half to two-thirds the anterior width of
vertex; along median longitudinal axis, head is approximately one-
third the length of pronotum; notocephalon slightly sulcate, with
lateral margins variable in amount of divergence anterior to synth-
lipsis; tylus slightly inflated; labrum with basal width not quite
twice its median length and apex bluntly rounded; rostral prong
(pi. CIV, figs. 42b, 43b) distinctly longer than third rostral seg-
ment, with base originating laterally midway to near proximal end
of third rostral segment, and with apex bluntly rounded. Pronotum
with its median length approximately three-fifths its humeral width;
disk with two elongate depressions toward the middle and a large
subtriangular depression on each side, thus appearing tricarinate;
lateral margins divergent; posterior margin convex, medianly con-
cave. Scutellum large, with median length greater than that of
pronotum. Fore femur (pi. CIV, figs. 42a, 43a) neither wide nor
greatly thickened at apex; triangular stridulatory area somewhat
variable, consisting of seventeen to thirty-two sclerotized ridges,
located medianly. Fore tibia (pi. CIV, figs. 42a, 43a) with stridu-
latory comb (pi. CIV, figs. 42c, 43c) consisting of forty-one to fifty-
one teeth; apical teeth thicker and slightly narrower than basal.
Chaetotaxy of male front leg as shown on Plate CIV. Male genital
claspers normal. Spine from caudo-sinistral margin of seventh
abdominal tergite with apical half very narrow and apex strongly
acuminate.

Female Structural Characteristics: As viewed from above, out-
line of head laterally rounded, anteriorly truncate with vertex
indented at least at lateral margins; greatest width of head six to
seven times the anterior width of vertex and distinctly less than
humeral width of pronotum; synthlipsis one-half to two-thirds the
anterior width of vertex; along median longitudinal axis, head is
one-fourth to one-third the length of pronotum; notocephalon
slightly sulcate, with lateral margins variable in amount of diver-
gence anterior to synthlipsis; tylus slightly inflated. Pronotum with
its median length approximately half its humeral width; disk usu-
ally with two shallow, elongate depressions toward the middle
forming a faint median carina, not tricarinate; lateral margins
divergent; posterior margin convex, medianly concave. Scutellum
large, with median length distinctly greater than that of pronotum.
Female ovipositor of normal shape (pi. CIII, fig. 30) with teeth
arranged in two longitudinal rows, one inner row of few, large

57—6490

1376 The University Science Bulletin

teeth and one long outer row of smaller teeth; approximately four
to nine small, lateral, toothlike setae near apex.

N omenclatorial Notes: On examination of large series of speci-
mens of the species known as B, antigone (Kirkaldy) and B. carinata
(Champion), one finds many intergrades between the two. This
fact was primary in assisting the author to conclude that we are
dealing with one variable species divisible into two subspecies.

Comparative Notes: Superficially this species somewhat resem-
bles B. tarsalis n. sp. Examination of the male, however, will show
distinct differences. This species differs from B. tarsalis in having
synthlipsis wider, first tarsal segment of intermediate leg not
emarginate, hind femur more robust, and differences in the rostral
prong, femoral stridulatory area, and tibial comb.

Buenoa antigone antigone (Kirkaldy)
(PI. CIV, fig. 42)

1899. Anisops antigone Kirkaldy, G. W. The Entomologist, vol. XXXII, p. 30.
1904. Buenoa antigone, Kirkaldy, G. W. Wiener Ent. Zeit, vol. XXIII,

pp. 120, 122, and 134 (description establishing B. antigone as genotype

and A. carinatns Champion as nov. syn. ).
1906. Buenoa antigone, Kirkaldy, G. W. Trans. Am. Ent. Soc., vol. XXXII,

p. 153 (listed).
1909. Buenoa antigone, Kirkaldy, G. W. and Torre-Bueno, J. R. de la. Proc.

Ent. Soc. Wash., vol. X, p. 200 (catalogue).

1916. Buenoa antigone, Van Duzee, E. P. New York Ent. Soc, p. 51 (check
list).

1917. Buenoa antigone, Van Duzee, E. P. Cat. Hemiptera Am. North of
Mexico, p. 454 (catalogue).

1919. Buenoa antigone, Hungerford, H. B. Univ. Kansas Sci. Bull., vol. XI,

p. 174 (brief note on synonomy).
1923. Buenoa antigone, Torre-Bueno, J. R. de la. Univ. of Iowa Studies in

Nat. Hist., 10:3, p. 35.
1940. Buenoa antigone, Hungerford, H. B. Ent. Month. Mag., vol. LXXVI,

p. 256 (ecological note).

Size: Male, length 7.80 mm. to 7.99 mm., greatest body width
2.46 mm. to 2.60 mm.; female, length 8.25 mm. to 8.51 mm., greatest
body width 2.60 mm. to 2.79 mm.

Color: General facies testaceous. Head, pronotum, thoracic
venter, and limbs testaceous. Scutellum usually ochraceous, brown
at base. Abdominal dorsum testaceous to brown; venter brown to
black with connexivum usually testaceous.

Male Structural Characteristics: As viewed from above, greatest
width of head more than six times the anterior width of vertex;
synthlipsis one half to two thirds the anterior width of vertex;
notocephalon with margins almost parallel to moderately divergent
anterior to synthlipsis; rostral prong (pi. CIV, fig. 42b) with base
originating laterally at proximal end of third rostral segment. Pro-

A Revision of the Genus Buenoa 1377

notum with its median length approximately three fifths its humeral
width; disk with two elongate, shallow depressions toward the mid-
dle and a large subtriangular, shallow depression on each side, thus
appearing faintly tricarinate. Fore femur (pi. CIV, fig. 42a) with
triangular stridulatory area consisting of approximately fifteen to
twenty-three sclerotized ridges. Fore tibia (pi. CIV, fig. 42a) with
stridulatory comb (pi. CIV, fig. 42c) consisting of approximately
forty-one teeth.

Female Structural Characteristics: As viewed from above, great-
est width of head approximately six times the anterior width of
vertex; synthlipsis slightly more than half the anterior width of
vertex; along median longitudinal axis, head is approximately one
third the length of pronotum; notocephalon with margins almost
parallel to moderately divergent anterior to synthlipsis. Female
ovipositor with approximately four or five very smalj, lateral tooth-
like setae near apex.

Nomenclatorial Notes: Mr. C. O. Bare labeled a long series of
B. antigonc as types and paratypes, using a manuscript name based
on the country of origin, Paraguay. As such paratypes may have
been widely distributed, it seems desirable to point out that the
name has not been, and should not be, validated by publication.

Comparative Notes: This subspecies differs from B. antigone cari-
nata ( Champion ) in being less robust, margins of notocephalon less
divergent anterior to synthlipsis, and in the shape and number of
sclerotized ridges in femoral stridulatory area.

Location of Types: The type female and two other female speci-
mens labeled "St. Andrew, Jamaica, C. B. Taylor, 20/IX, 98" (type
locality and determined by Kirkaldy) from the G. W. Kirkaldy Col-
lection, are now in the Francis Huntington Snow Entomological
Collections, University of Kansas.

Data on Distribution: Recorded from Mexico, Guatemala, West
Indies (Cuba, Cayman Islands, Haiti, Jamaica, Puerto Rico, St.
Croix Island), Brazil, Ecuador, Peru, Bolivia, Paraguay, and Argen-
tina. Specimens from the following localities have been examined:

MEXICO: Tamaulipas: 5 m. S. Ciudad Victoria, Nov. 5, 1936,
H. D. Thomas, 35 males, 49 females.

Veracruz: Minatitlan, Sept. 22, 1936, H. D. Thomas, 20 males, 18
females.

Chiapas: Hda. La Libertad, Sept. 1, 1937, H. D. Thomas, 2
males, 3 females; La Libertad, Jan. 1, 1938, Octavio Utrilla Louis,
2 males, 4 females; Comitan, Jan. 18, 1938, Octavio Utrilla L.,
25 males, 30 females.

1378 The University Science Bulletin

Campeche: Hda. Encarnation, 12 m. S. Pital, Oct. 15, 1936, H. M.
Smith, 11 males, 13 females.

GUATEMALA: El Salto, Escuintla, 1934, F. X. Williams, 1 male.

WEST INDIES: Cuba: Camaguey, July 2, 1923, J. Acuda, 1
male, 2 females; Soledad, Feb. 14, 1925, J. G. Myers, 1 male; Havana,
Bot. Garden, Jan. 1925, P. J. Bermudez, 10 males; Havana Prov.,
Catalina, Nov. 27, 1933, P. J. Bermudez, 23 males, 16 females.

Cayman Islands: Cayman Brae, Earthquake hole, May 22, 1938,
Lewis and Thompson (Oxford Univ. Bio. Exp.), 6 males, 6 females;
Grand Cayman, Booby Creek, June 23, 1938, Lewis and Thompson
(Oxford U. Bio. Exp.), 1 male; Grand Cayman, Cow-well, near
Pedro Castle, Aug. 4, 1938, Lewis and Thompson (Oxford U. Bio.
Exp.), 1 male, 1 female.

Haiti: Port au Prince, March, 1927, G. N. Wolcott, 1 male, 4 fe-
males.

Jamaica: St. Andrew, Sept. 20, 1898, C. B. Taylor, 3 females;
Claremont, Feb., 1928, Lilly G. Perkins, 41 males, 49 females; Baron
Hill Trelawny, Feb., 1928, L. G. Perkins, 130 males, 196 females;
Claremont, Baron Hill Trelawny, March and April, 1928, L. G.
Perkins, 139 males, 248 females; Baron Hill Trelawny, Nov. and
Dec, 1928, L. G. Perkins, 1 male, 1 female; Lumsden Tydenbam,
St. Ann, Feb., 1928, L. G. Perkins, 4 males, 2 females.

Puerto Rico: Camuy-Isabela, May 12, 1935, Julio Garcia Diaz,
2 males, 4 females; Cabo Bojo, June 9, 1937, J. A. Bamos, 1 male,
2 females.

St. Croix: Christiansted, 1941, H. A. Beatty, 8 males, 5 females.

BRAZIL: Rio Grande do Norte: Ouro Branco, No. 258, Stillman
Wright, 1 female; Caico, No. 327, Stillman Wright, 1 female; Caico,
No. 12637, Stillman Wright, 7 females.

Sao Paulo: Rio S. Paulo, Dec. 26, 1944, Wygd., 1 male, 11 fe-
males; Oct., 1947, Fritz Plaumann, 1 male, 2 females.

Santa Catarina: Nova Teutonia, Dec, 1946, Fritz Plaumann, 1
male, 2 females; Nova Teutonia, May, 1948, Fritz Plaumann, 93
males, 113 females.

PEBU: Dept. Junin, Prov. Tarma, Palmapaca Jungle, Oct. 19 to
26, 1940, F. Woytkowski, 17 males, 18 females; Dept. San Martin,
Feb. 16, 1947, Felix Woytkowski, 101 males, 62 females, 3 nymphs.

PARAGUAY: Villarrica, May 16, 1923, Fran. Schade, 3 females;
Sept. to Dec, 1923, Fran. Schade, 57 males, 108 females; Jan. to
March, 1924, Fran. Schade, 40 males, 43 females; April 16, 1924,
Fran. Schade, 10 females; July 8, 1924, Fran. Schade, 13 males, 6
females; Sept. 21, 1924, Fran. Schade, 23 males, 19 females; Jan.,

A Revision of the Genus Buenoa 1379

1926, Fran. Schade, 1 male, 1 female; Oct. 29, 1929, Fran. Schade,
1 male, 1 female; Caraveni, June 15, 1924, F. Schade, 4 males,
6 females; Feb. 15, 1925, F. Schade, 3 males; Oct. 30, 1924, F.
Schade, 4 males, 6 females; Feb. 16, 1925, F. Schade, 1 male, 12
females; Estero Grande, Nov. 1, 1924, F. Schade, 6 males; Meli-
nesque, June 20 to 28, 1935, F. Schade, 2 males, 9 females.

ARGENTINA: Prov. de Salta, Dept. Metau, 13 males, 9 fe-
males; Salta, 1 male, 1 female.

All specimens listed above are in the Francis Huntington Snow
Entomological Collections, University of Kansas, Lawrence, Kansas.

The specimens collected in the States of Tamaulipas, Veracruz,
and Chiapas, Mexico, overlap the area of B. antigone carinata.
However, this region along the Gulf of Mexico south through the
State of Chiapas, shows a varied ecology and these records are not
surprising.

Buenoa antigone carinata (Champion)

(PI. CIV, fig. 43)

1901. Anisops carinatus Champion, G. C. Biologia Centrali Americana, Heter-

optera, vol. II, p. 372, pi. 22, fig. 12.
1904. Anisops carinatus, Uhler, P. R. Proc. U. S. Nat. Mus., vol. XXVII,

p. 364 (listed).
1904. Buenoa carinata, Kirkaldy, G. W. Wiener Ent. Zeit., vol. XXIII, pp. 122

and 134 (listed as synonym of B. antigone) .
1909. Buenoa carinata, Kirkaldy, G. W. and Torre-Bueno, J. R. de la. Proc.

Ent. Soc. Wash., vol. X, p. 200 (catalogue).
1909. Buenoa carinata, Torre-Bueno, J. R. de la. Jr. New York Ent. Soc,

vol. XVII, p. 75 (listed).
1914. Buenoa carinata, Barber, H. G. Bull. Am. Mus. Nat. Hist., vol. XXXIII,

p. 499 (listed).

1916. Buenoa carinata, Van Duzee, E. P. New York Ent. Soc, p. 51 (check
list).

1917. Buenoa carinata, Van Duzee, E. P. Cat. Hemiptera Am. North of
Mexico, p. 454 (catalogue).

1919. Buenoa carinata, Hungerford, H. B. Univ. Kansas Sci. Bull., vol. XI,
pp. 174-175 (key and description).

1925. Buenoa carinata, Bare, C. O. Ent. News, vol. XXXVI, p. 228 (key).

1926. Buenoa carinata, Blatchley, W. S. Heteroptera or True Bugs of Eastern
North America, pp. 1057-1058 (key and description).

1928. Buenoa carinata, Bare, C. O. Univ. Kansas Sci. Bull., vol. XVIII, No. 3,

p. 268 (key).
1948. Buenoa carinata, Hvnes, H. B. N. Trans. Roy. Ent. Soc. London, vol.

XCIX, p. 354 ( distribution note ) .

Size: Male, length 7.94 mm. to 8.97 mm., greatest body width
2.60 mm. to 2.79 mm.; female, length 8.77 mm. to 9.75 mm., greatest
body width 2.92 mm. to 3.12 mm.

Color: General facies testaceous to dark brown. Head, pronotum,
thoracic venter, and limbs sordid white to testaceous. Scutellum
rufo-testaceous to testaceous with base brown to black; metathoracic
dorsum brown to black. Abdomen usually black except ventral

1380 The University Science Bulletin

keel, margins of connexivum, and last two or three segments, testa-
ceous. This subspecies somewhat variable in color with some speci-
mens entirely sordid white to testaceous.

Male Structural Characteristics: As viewed from above, greatest
width of head approximately five and one half times the anterior
width of vertex; synthlipsis approximately two thirds the anterior
width of vertex; notocephalon with margins distinctly divergent
anterior to synthlipsis; rostral prong (pi. CIV, fig 43b) with base
originating laterally midway of third rostral segment. Pronotum
with its median length approximately five-ninths its humeral width;
disk with two elongate depressions toward the middle and a large
subtriangular depression on each side, thus appearing distinctly
tricarinate. Fore femur (pi. CIV, fig. 43a) with subtriangular to
oblong stridulatory area consisting of approximately twenty-nine to
thirty-three sclerotized ridges. Fore tibia (pi. CIV, fig. 43a) with
stridulatory comb (pi. CIV, fig. 43c) consisting of approximately
fifty to fifty-two teeth.

Female Structural Characteristics: As viewed from above, great-
est width of head approximately seven times the anterior width of
vertex; synthlipsis slightly less than two thirds the anterior width
of vertex; along median longitudinal axis, head is approximately
one fourth the length of pronotum; notocephalon with margins dis-
tinctly divergent anterior to synthlipsis. Female ovipositor with
approximately eight or nine small, lateral, toothlike setae near apex.

Comparative Notes: This subspecies differs from B. antigone anti-
gone (Kirkaldy) in being more robust, margins of notocephalon
more divergent anterior to synthlipsis, and in the shape and number
of scletorized ridges in femoral stridulatory area.

Location of Types: Type series is located at the British Museum,
London. Homotype male, compared with type by H. B. Hunger-
ford, labeled "Colima, Vulcano, Mex., L. Conrad", now in the
Francis Huntington Snow Entomological Collections, University of
Kansas.

Data on Distribution: Recorded from United States (fide Uhler
and Blatchley and doubtful), Mexico, Guatemala, and British Hon-
duras (fide Champion). Specimens from the following localities
have been examined:

MEXICO: Sonora: 2 mi. E. of Guirecoba, Apr. 30, 1939, C. Sib-
ley, 1 male, 3 females.

Sinaloa: Badiraguato, Mar. 30, 1937, A. Dampf, 1 male, 1 fe-
male.

A Revision of the Genus Buenoa 1381

Durango: San Antonio near El Salto, June 10, 1937, Meldon Em-
bury, 2 males.

Tamaulipas: 5 mi. S. Ciudad Victoria, Nov. 5, 1936, H. D.
Thomas, 1 female.

Jalisco: 20 mi. S. Guadalajara, Sept. 10, 1938, H. D. Thomas, 2
males, 2 females; Chapala, Sept. 11, 1938, H. D. Thomas, 1 male,
1 female; Guadalajara-Tequila Rd., 28 mi. N. Jalisco, Sept. 13, 1938,
H. D. Thomas, 1 female; 15 mi. S. W. Lake Chapala, Sept. 14, 1938,
H. D. Thomas, 2 females; 15 mi. down Autlan Rd., Sept. 14, 1938,
H. D. Thomas, 1 female; Tecolotlan, Sept. 15, 1938, H. D. Thomas,
5 males, 8 females; Union de Tula, Sept. 16, 1938, H. D. Thomas,
1 female; Jet. Guadalajara Hwy. and road to Autlan, Sept. 17, 1938,
H. D. Thomas, 11 males, 11 females.

Guanajuato: 10 mi. N. E. Leon, Aug. 17, 1932, Hobart Smith,
3 females.

Veracruz: Jalapa, May 18, 1930, Creaser-Gordon, 1 male.

Michoacdn: 10 mi. down Chinapa Rd., Sept. 5, 1938, H. D.
Thomas, 6 males, 4 females; Zamora, Sept. 8, 1938, H. D. Thomas,
19 males, 20 females.

Colima: Vulcano, L. Conrad, 1 male.

Guerrero: Rio Balsas, Jet. Acapulco Mex. Hwy., June 24, 1932,
Hobart Smith, 2 females; Puente de Ixtla, July 12, 1937, H. D.
Thomas, 4 females; Iguala, Oct. 7, 1936, H. D. Thomas, 9 males,
15 females; Palo Blanco, kil. 338 S. Mex. City, Oct. 10, 1936, H. D.
Thomas, 1 female; Petaquillas, kil. 320 S. Mex. City, Oct. 21, 1936,
H. D. Thomas, 1 male; Salto de Valadez, kil. 325 S. Mex. City,
Oct. 30, 1936, H. D. Thomas, 12 males, 13 females; Tierra Colo.,
kil. 377 S. Mex. City, Oct. 31, 1936, H. D. Thomas, 5 males, 6 fe-
males; Rio Balsas, kil. 259 S. Mex. City, Oct. 31, 1936, H. D. Thomas,
23 males, 16 females; Acapulco, kil. 442 S. Mex. City„ Nov. 1, 1936,
H. D. Thomas, 50 males, 63 females.

Morelos: Rio Amacuzac, kil. 133 S. Mex. City, Oct. 14, 1936,
H. D. Thomas, 6 males, 13 females; Cuernavaca, Oct. 1-17, 1936,
H. D. Thomas, 1 male, 3 females; Acatlipa, kil. 84 S. Mex. City,
Oct. 17, 1936, H. D. Thomas, 2 females.

Chiapas: Hda. La Libertad, Sept. 1, 1937, H. D. Thomas, 49
males, 39 females; La Libertad, Jan. 1, 1938, Octavio Utrilla Louis,
83 males, 96 females; San Vicente, Jan. 4, 1938, Octavio Utrilla
Louis, 4 females.

All specimens listed above are in the Francis Huntington Snow
Entomological Collections, University of Kansas, Lawrence, Kansas.

1382 The University Science Bulletin

Buenoa femoralis (Fieber)
(PI. CII, fig. 4; pi. CV, fig. 44)

1851. Anisops femoralis Fieber, F. X. Abhandlungen Kongl. Bohmischen

Gesellschaft Wissenschaften, vol. VII, p. 483.
1904. Buenoa femoralis, Kirkaldy, G. W. Wiener Ent. Zeit., vol. XXIII,

pp. 120 and 134 (listed).
1909. Buenoa femoralis, Kirkaldy, G. W. and Torre-Bueno, J. R. de la. Proc.

Ent. Soc. Washington, vol. X, p. 200 (listed).
1928. Buenoa femoralis, Jaezewski T. Ann. Mus. Zool. Polonici, vol. VII,

pp. 123-125 ( description and illustrations ) .
1939. Buenoa femoralis, Barber, H. G. New York Acad. Sci., vol. XIV, p. 420.

Size: Male, length 8.32 mm. to 8.64 mm., greatest body width
2.66 mm. to 2.73 mm.; female, length 8.72 mm. to 8.90 mm., greatest
body width 2.60 mm. to 2.72 mm.

Color: General facies pale testaceous to nigro-violaceous. Head
portions of pronotum, thoracic venter, and limbs sordid white to
testaceous. Scutellum, and portions of metathoracic dorsum and
abdomen, black. Ventral abdominal keel, most of connexivum, and
lateral portions of last two segments, testaceous. Posterior half of
hemelytron, except for membrane, black. This species varies in
color from specimens that are entirely testaceous to specimens that
are almost entirely nigro-violaceous.

Male Structural Characteristics: As viewed from above, outline
of head laterally rounded, anteriorly truncate with vertex indented;
greatest width of head approximately seven times the anterior width
of vertex and less than humeral width of pronotum; synthlipsis ap-
proximately half the anterior width of vertex; along median longi-
tudinal axis, head is slightly less than half the length of pronotum;
tylus inflated; labrum with basal width almost twice its median
length and apex bluntly rounded; rostral prong (pi. CV, fig. 44b)
much longer than third rostral segment, with base originating later-
ally midway of third rostral segment, and with apex moderately
rounded. Pronotum with its median length less than three fifths its
humeral width; disk with two elongate depressions toward the
middle and a large subtriangular depression on each side, thus ap-
pearing tricarinate; lateral margins divergent; posterior margin con-
vex, medianly concave. Scutellum large with median length dis-
tinctly greater than that of pronotum. Fore femur (pi. CV, fig. 44a)
slightly widened at apex; triangular stridulatory area consisting of
approximately seventeen sclerotized ridges. Fore tibia (pi. CV,
fig. 44a) with stridulatory comb (pi. CV, fig. 44c) consisting of
approximately thirty-four or thirty-five teeth which are of same
size and thickness. Chaetotaxy of male front leg as shown on
Plate CV. Male genital claspers normal. Spine from caudo-sinis-

A Revision of the Genus Buenoa 1383

tral margin of seventh abdominal tergite (pi. CII, fig. 4) with apical
half narrow and apex acuminate.

Female Structural Characteristics: As viewed from above, outline
of head laterally rounded, anteriorly truncate with vertex indented;
greatest width of head approximately seven times the anterior width
of vertex and less than humeral width of pronotum; synthlipsis ap-
proximately half the anterior width of vertex; along median longi-
tudinal axis, head is less than half the length of pronotum; tylus
inflated. Pronotum with its median length slightly less than half
its humeral width; disk only slightly impressed and occasionally not
at all; lateral margins divergent; posterior margin convex, medianly
concave. Scutellum large with median length much greater than
that of pronotum. Female ovipositor of normal shape with teeth
arranged in two longitudinal rows; one inner row of few, large teeth
and one long outer row of smaller teeth; approximately seven or
eight small, lateral, toothlike setae near apex.

Comparative Notes: Superficially this species closely resembles
B. crassipes (Champion). Examination of the male, however, will
show distinct differences. This species differs from B. crassipes
in the shape and armature of the fore and hind femora, the shape
of the tibial comb, and the lateral origin of the base of the rostral
prong. Buenoa femoralis is a little larger than B. crassipes.

Location of Types: The type specimen, a somewhat damaged
male, labeled "Porto Rico, Moritz", is located at the Berlin Museum.

Data on Distribution: Recorded from Puerto Rico, Brazil, and
Peru. Specimens from the following localities have been examined:

PERU: Vicinity Sani Beni, muddy pool in shady jungle, Oct. 16,
1935, F. Woytkowski, 3 males, 4 females; vicinity Sani Beni, muddy
pool in shady jungle, Oct. 17, 1935, F. Woytkowski, 38 males, 56
females; Oct. 24, 1935, F. Woytkowski, 111 males, 128 females; Dept.
Ayacucho, Prov. La Mar., Sivia jungle, stagnant boggy pool, June
20-23, 1941, F. Woytkowski, 2 males, 1 female.

All specimens listed above are in the Francis Huntington Snow
Entomological Collections, University of Kansas, Lawrence, Kansas.

Buenoa ida Kirkaldy
(PI. CV, fig. 45)

1904. Buenoa ida Kirkaldy, G. W. Wiener Ent. Zeit., vol. XXIII, pp. 121, 122

and 134.
1909. Buenoa ida, Kirkaldy, G. W. and Torre-Bueno, J. R. de la. Proc. Ent.

Soc. Washington, vol. X, p. 200 (catalogue).

Size: Male, length 9.16 mm. to 9.62 mm., greatest body width
2.73 mm. to 2.82 mm.; female, length 9.29 mm. to 9.94 mm., greatest
body width 3.12 mm. to 3.25 mm.

1384 The University Science Bulletin

Color: General facies fuscous. Head, pronotum, thoracic venter,
and limbs testaceous. Scutellum brown to black with apex testa-
ceous; metathoracic dorsum testaceous to brown. Abdomen black
except ventral keel, margins of connexivum, and last one or two
segments, testaceous. Posterior half of hemelytra, except mem-
brane, usually black.

Male Structural Characteristics: As viewed from above, outline
of head laterally rounded, anteriorly truncate with vertex usually
slightly indented; greatest width of head slightly more than five
times the anterior width of vertex and less than humeral width of
pronotum; synthlipsis approximately half the anterior width of ver-
tex; along median longitudinal axis, head is approximately one
third the length of pronotum; notocephalon slightly sulcate; tylus
not inflated; labrum with basal width one third greater than its
median length and apex bluntly rounded; rostral prong (pi. CV,
fig. 45b) longer than third rostral segment, with base originating
laterally near proximal end of third rostral segment, and with apex
bluntly rounded. Pronotum with its median length approximately
one half its humeral width; disk with two elongate, shallow de-
pressions toward the middle and a large, subtriangular, shallow
depression on each side, thus appearing faintly tricarinate; lateral
margins divergent; posterior margin convex, medianly concave.
Scutellum large, with median length distinctly greater than that of
pronotum. Fore femur (pi. CV, fig. 45a) wide and somewhat
thickened at apex; long, curved, triangular stridulatory area con-
sisting of approximately thirty-four or thirty-five sclerotized ridges.
Fore tibia (pi. CV, fig. 45a) with stridulatory comb (pi. CV, fig.
45c) consisting of approximately twenty-two to twenty-four thick
teeth; apical teeth slightly thicker than basal. Chaetotaxy of male
front leg as shown on Plate CV. Male genital claspers normal.
Spine from caudo-sinistral margin of seventh abdominal tergite
with apical half narrow and apex strongly acuminate.

Female Structural Characteristics: As viewed from above, head
is laterally rounded, anteriorly truncate with vertex indented only
at lateral margins; greatest width of head approximately five and
one half times the anterior width of vertex and less than humeral
width of pronotum; synthlipsis slightly more than half the anterior
width of vertex; along median longitudinal axis, head is approxi-
mately one fourth the length of pronotum; notocephalon sulcate;
tylus not inflated. Pronotum with its median length slightly more
than half its humeral width; disk usually unimpressed, occasionally
with a feeble median carina; lateral margins divergent; posterior

A Revision of the Genus Buenoa 1385

margin convex, medianly concave. Scutellum large, with median
length distinctly greater than that of pronotum. Female ovipositor
of normal shape with teeth arranged in two longitudinal rows; one
inner row of few, large teeth and one long outer row of smaller
teeth; approximately eight or nine small, lateral, toothlike setae
near apex.

Comparative Notes: Superficially this species resembles B. anti-
gone carinata (Champion). Examination of the male, however,
will show distinct differences. This species differs from B. antigone
carinata in the shape of the fore femur, the femoral stridulatory
area, and the shape and number of teeth of the tibial comb. The
pronotum of B. antigone carinata is more distinctly tricarinate than
that of B. ida, and the size and coloration are different in the two
species.

Location of Types: Dr. G. W. Kirkaldy states that the type is in
his collection. The Kirkaldy Collection is now located in the
Francis Huntington Snow Entomological Collections, University of
Kansas, but the type has been lost or destroyed. A series of three
specimens, 2 males and 1 female, were located in the remnants of
the Kirkaldy Collection. Two of these are labeled "Uruguay," the
type locality, and one is labeled "Guatemala." I have selected a
male from Uruguay as a neoholotype.

Data on Distribution: Recorded from Mexico, Guatemala, and
Uruguay. Specimens from the following localities have been ex-
amined :

MEXICO: Chiapas: L. Tepancuapan, Aug. 28, 1937, H. D.
Thomas, 3 males, 7 females; Comitan, Jan. 18-20, 1938, Octavio
Utrilla L., 12 males, 7 females.

GUATEMALA: No locality, Breddin (Kirkaldy Coll.), 1 female.

URUGUAY: No locality, E. Autran (Kirkaldy Coll.), 1 male, 1
female.

All specimens listed above are in the Francis Huntington Snow
Entomological Collections, University of Kansas, Lawrence, Kansas.

Buenoa crassipes (Champion)

(PI. CII, fig. 9; pi. CVI, fig. 46)

1901. Anisops crassipes Champion, G. C. Biologia Centrali Americana, Heter-

optera, vol. II, p. 374.
1904. Buenoa crassipes, Kirkaldy, G. W. Wiener Ent. Zeit., vol. XXIII,

pp. 121 and 134 (listed).
1909. Buenoa crassipes, Kirkaldy, G. W. and Torre-Bueno, J. R. de la. Proc.

Ent. Soc. Washington, vol. X, p. 200 (catalogue).
1928. Buenoa crassipes, Jaczewski, T. Annales Musei Zoologici Polonici, vol.

VII, pp. 125-126 (description).

1386 The University Science Bulletin

Size: Male, length 7.15 mm. to 7.67 mm., greatest body width 2.27
mm. to 2.40 mm.; female, length 7.28 mm. to 8.12 mm., greatest
body width 2.27 mm. to 2.47 mm.

Color: General facies testaceous to a shining nigro-violaceous.
Head, portions of pronotum, thoracic venter, and limbs sordid
white to testaceous. Scutellum, metathoracic dorsum, and abdomen
black, except for ventral keel, portions of connexivum and lateral
portions of last two segments, testaceous. Basal portion of hemely-
tral membrane usually black. This species variable in color.

Male Structural Characteristics: As viewed from above, outline of
head laterally rounded, anteriorly truncate with vertex indented;
greatest width of head approximately six and one half times the an-
terior width of vertex and slightly less than humeral width of pro-
notum; synthlipsis approximately half the anterior width of vertex;
along median longitudinal axis, head is approximately two fifths the
length of pronotum; notocephalon medianly sulcate; tylus slightly
inflated; labrum with basal width slightly greater than its median
length, and apex bluntly rounded; rostral prong ( pi. CVI, fig. 46b )
much longer than third rostral segment, with base originating later-
ally near distal end of third rostral segment, and with apex moder-
ately rounded. Pronotum with its median length approximately
three fifths its humeral width; disk almost unimpressed, not tricari-
nate; lateral margins slightly divergent; posterior margin convex,
medianly concave. Scutellum large, with median length greater
than that of pronotum. Fore femur (pi. CVI, fig. 46a) wide and
somewhat thickened at apex; triangular stridulatory area variable
consisting of approximately ten wide or sixteen narrow sclerotized
ridges. Fore tibia (pi. CVI, fig. 46a) with wide stridulatory comb
(pi. CVI, figs. 46c, 46d) consisting of approximately thirty to thirty-
four thick teeth; apical teeth much wider than basal, with width of
comb somewhat variable. Chaetotaxy of male front leg as shown
on Plate CVI. Male genital claspers normal. Spine from caudo-
sinistral margin of seventh abdominal tergite (pi. CII, fig. 9) with
apical two thirds very narrow and apex extremely acuminate.

Female Structural Characteristics: As viewed from above, outline
of head laterally rounded, anteriorly truncate with vertex indented;
greatest width of head approximately five and one half times the
anterior width of vertex and less than humeral width of pronotum;
synthlipsis approximately half the anterior width of vertex; along
median longitudinal axis, head is slightly more than one third the
length of pronotum; notocephalon medianly sulcate; tylus slightly
inflated. Pronotum with its median length approximately half its

A Revision of the Genus Buenoa 1387

humeral width; disk unimpressed; lateral margins divergent; poste-
rior margin convex, medianly concave. Scutellum large with median
length distinctly greater than that of pronotum. Female ovipositor
of normal shape with teeth arranged in two longitudinal rows; one
inner row of large teeth, one outer row of small teeth, and a few
small teeth intermingled with the two rows medianly; approximately
seven or eight small, lateral, toothlike setae near apex.

Comparative Notes: Superficially this species closely resembles
B. femoralis ( Fieber). Examination of the male, however, will show
distinct differences. This species differs from B. femoralis in the
shape and armature of the fore and hind femora, the shape of the
tibial comb, and the lateral origin of the base of the rostral prong.
Buenoa crassipes is a little smaller than B. femoralis.

Location of Types: The original type series from Guatemala, is
located at the British Museum, London.

Data on Distribution: Recorded from Guatemala, Costa Rica,
Ecuador, and Brazil. Specimens from the following localities have
been examined:

COSTA RICA: San Jose, Purchased June 1931, Heinrich Schmidt,
6 males, 20 females; San Jose, June & July 1931, Heinrich Schmidt,
10 males, 11 females; San Jose, Purchased 1932, Heinrich Schmidt,
58 males, 82 females; Rio Virilla, Dec. 26, 1931, Heinrich Schmidt,
21 males, 33 females; Rio Torres, Feb. 10, 1932, Heinrich Schmidt,
1 male, 1 female, 1 nymph; Rio Sarapiqui, Heinrich Schmidt, 8
males, 8 females.

ECUADOR: Tungurahua Vale Banos, Jan. 1, 1923, F. X. Wil-
liams, 4 females, 22 nymphs; Tena, Feb. 28, 1923, F. X. Williams,
3 nymphs.

BRAZIL: Santa Catarina: Nova Teutonia, Dec. 1946, Fritz Plau-
mann, 34 males, 20 females; Nova Teutonia, May 1948, Fritz Plau-
mann, 23 males, 42 females.

All specimens listed above are in the Francis Huntington Snow
Entomological Collections, University of Kansas, Lawrence, Kansas.

Buenoa arizonis Bare
(PI. CII, fig. 10; PI. CIII, figs. 29, 32; PI. CVI, fig. 47)

1928. Buenoa arizonis Bare, C. O. Univ. Kansas Sci. Bull., vol. XVIII, p. 342,

pi. 54 (figured only).
1931. Buenoa arizonis, Bare, C. O. Pan-Pacific Ent, vol. VII, No. 3, pp. 115-

118 (description).
1942. Buenoa arizonis, Hutchinson, G. E. American Jr. Sci., vol. CCXL,

p. 336 (morphology note).

Size: Male, length 8.45 mm. to 8.80 mm., greatest body width

1388 The University Science Bulletin

2.56 mm. to 2.73 mm.; female, length 8.45 mm. to 8.83 mm., greatest
body width 2.68 mm. to 2.81 mm.

Color: General facies testaceous to dark brown. Head, pro-
notum, thoracic venter, and limbs testaceous. Scutellum black with
lateral and hind margins yellowish; metathoracic dorsum brown to
black. Abdomen usually entirely black except ventral keel, mar-
gins of connexivum, and last two to three segments, testaceous.
Some specimens entirely sordid white to testaceous.

Male Structural Characteristics: As viewed from above, outline
of head laterally rounded, anteriorly truncate with vertex indented;
greatest width of head approximately six times the anterior width
of vertex and less than humeral width of pronotum; synthlipsis
approximately as wide as anterior width of vertex; along median
longitudinal axis, head is approximately one fifth the length of
pronotum; notocephalon usually slightly sulcate; tylus strongly in-
flated with shallow median depression forming two lateral protuber-
ances; labrum short, basal width almost twice its median length
with apex bluntly rounded; rostral prong (pi. CVI, fig. 47b) dis-
tinctly longer than third rostral segment, with base originating
laterally near distal end of third rostral segment, and with apex
bluntly rounded. Pronotum strongly convex with its median length
approximately one fifth greater than humeral width; disk with two
elongate depressions toward the middle and a subtriangular de-
pression on each side, thus appearing tricarinate; lateral margins
divergent; posterior margin convex, very slightly concave medianly.
Scutellum with median length approximately one fifth less than
that of pronotum. Fore femur (pi. VI, fig. 47a) wide and some-
what thickened at apex; oblong stridulatory area consisting of
approximately thirteen to fourteen sclerotized ridges. Fore tibia
(pi. VI, fig. 47a) with stridulatory comb (pi. VI, fig. 47c) con-
sisting of approximately forty-two to fifty teeth; apical teeth thicker
and slightly narrower than basal. Chaetotaxy of male front leg as
shown on Plate VI. Male genital claspers normal (pi. Ill, fig. 29).
Spine from caudo-sinistral margin of seventh abdominal tergite
(pi. II, fig. 10) with apical half very narrow and apex strongly
acuminate.

A Revision of the Genus Buenoa 1389

median longitudinal axis, head is slightly more than one fifth the
length of pronotum; notocephalon usually slightly sulcate; tylus
not strongly inflated and without median depression. Pronotum
strongly convex with its median length approximately two thirds
its humeral width; disk only slightly impressed and occasionally
not at all; lateral margins divergent; posterior margin convex, very
slightly concave medianly. Scutellum with median length slightly
less than that of pronotum. Female ovipositor (pi. Ill, fig. 32) of
normal shape with teeth arranged in two longitudinal rows; one
inner row of few, large teeth and one long outer row of smaller
teeth; approximately seven or eight small, lateral, tooth-like setae
near apex.

Comparative Notes: This species is nearest in general appear-
ance to B. antigone (Kirkaldy). Buenoa arizonis Bare differs from
this species in the number and size of setae on hind femur, the
form of the male pronotum, and in having the tylus of the male
strongly inflated. Differences also occur in the fore leg of the male.

Location of Types: Holotype male (on slides) and allotype fe-
male, Superstition Mts., Arizona, Nov. 7, 1922, P. A. Glick, in the
Francis Huntington Snow Entomological Collections. Paratypes
distributed as follows: (1) 1 male, Superstition Mts., Arizona,
Nov. 7, 1922, P. A. Glick; 3 males, 5 females, Baboquivari Mts.,
Arizona, F. H. Snow; 17 males, 49 females, Santa Cruz Co., Arizona,
Aug. 4, 1927, P. A. Readio, R. H. Beamer, & L. D. Anderson; 11
males, 20 females, Gila Co., Arizona, July 6, 1927, P. A. Readio,
R. H. Beamer, & L. D. Anderson; 1 female, Cochise Co., Arizona,
July 29, 1927, R. H. Beamer; 1 male, Santa Rita Mts., Arizona, July
25, 1927, R. H. Beamer, in Francis Huntington Snow Entomological
Collections, University of Kansas; (2) 19 specimens in the C. O.
Bare Collection; (3)3 specimens in California Academy of Sciences
Collection, and (4)5 specimens in the U. S. National Museum.

Data on Distribution: Recorded from the United States and
Mexico. In addition to type series, specimens from the following
localities have been examined:

U. S. A.: Arizona: Tucson, July 20, 1932, R. H. Beamer, 3 males,
1 female; Baboquivari Mts., Oct., 1934, Frank Blanchord, 1 male;
Baboquivari Mts., July 18, 1932, R. H. Beamer & R. H. Beamer, Jr.,
4 males, 11 females; Baboquivari Mts., July 24, 1941, B. Hodgden,
27 males, 39 females; Arabaca, July 6, 1941, B. Hodgden, 1 female;
Miami, July 6, 1941, E. L. Todd, 1 female; Sunnyside Canyon,
Huachuca Mts., July 9, 1940, L. C. Kuitert, 2 males; Ruby, July 13,

1390 The University Science Bulletin

1940, L. C. Kuitert, 18 males, 35 females; Ruby, July 27-28, 1941,
B. Hodgden, 6 males, 28 females.

MEXICO: Sonora: San Bernardo, Rio Mayo, Oct. 14, 1934, H. S.
Gentry, 15 males, 5 females; Buropaco Dist, Alamos, Oct. 23, 1934,
H. S. Gentry, 4 males; Conejos Dist., Alamos, Oct. 26, 1934, H. S.
Gentry, 32 males, 28 females; Arroyo de los Mescales, Rio Mayo,
Feb. 16, 1935, H. S. Gentry, 44 males, 85 females; Salitial, Rio Mayo,
Feb. 23, 1935, H. S. Gentry, 26 males, 56 females; Tepoca S. Charibo,
Mar. 9, 1935, H. S. Gentry, 34 females; 2 mi. E. of Guerocoba,
Apr. 30, 1939, C. Sibley, 5 females.

Chihuahua: Carimechi, Rio Mayo, Dec. 12, 1934, H. S. Gentry,
63 males, 126 females; San Luis Babarocos, Dec. 30, 1934, H. S.
Gentry, 35 males, 26 females.

Jalisco: Guadalajara, Tequila Rd., 28 mi. N. Jalisco, Sept. 13,
1938, H. D. Thomas, 28 females; 15 mi. S. W. of Lake Chapala,
Sept. 14, 1938, H. D. Thomas, 1 male; Tecolotlan, Sept. 15, 1938,
H. D. Thomas, 4 males, 41 females; 20 mi. S. Tecolotlan, Sept. 16,
1938, L. J. Lipovsky, 2 males, 5 females; Sept. 17, 1938, H. D.
Thomas, 1 female.

Michoacdn: 10 mi. down Chinapa Rd., Sept. 5, 1938, H. D.
Thomas, 2 females; S. side L. Cuitzeo, July 7, 1947, T. H. Hubbell,
1 male, 1 female.

Puebla: Near Zapotitlan, July 26, 1932, Hobart Smith, 1 male,
1 female.

Morelos: Cuernavaca, Oct. 1-17, 1936, H. D. Thomas, 92 males,
185 females; Acatlipa, kil. 84 S. of Mex. City, Oct. 17, 1936, H. D.
Thomas, 23 males, 34 females.

Guerrero: Rio Balsas, Jet. Acapulco Mex. Hwy., June 24, 1932,
Hobart Smith, 2 males, 11 females; Iguala, Oct. 7, 1936, H. D.
Thomas, 42 males, 121 females; Palo Blanco, kil. 338 S. Mex. City,
Oct. 10, 1936, H. D. Thomas, 1 female; Taxco, kil. 175 S. Mex. City,
Oct. 10, 1936, H. D. Thomas, 26 males, 27 females; Salto de Valadez,
kil. 325 S. Mex. City, Oct. 30, 1936, H. D. Thomas, 13 males, 20
females; Rio Balsas, 259 kil. S. Mex. City, Oct. 31, 1936, H. D.
Thomas, 161 males, 212 females; Tierra Colo., kil. 377 S. Mex. City,
Oct. 31, 1936, H. D. Thomas, 13 males, 19 females; Acapulco, kil.
442 S. of Mex. City, Nov. 1, 1936, H. D. Thomas, 20 males, 15
females; Sacacorjuca, kil. 216 S. Mex. City, H. D. Thomas, 15 males,
33 females.

All specimens listed above are in the Francis Huntington Snow
Entomological Collections, University of Kansas, Lawrence, Kansas.

A Revision of the Genus Buenoa 1391

Buenoa absidata n. sp.
(PI. CVII, fig. 48)

Size: Male, length 5.85 mm. to 6.56 mm., greatest body width
1.56 mm to 1.75 mm.; female, length 5.78 mm. to 6.95 mm., greatest
body width 1.56 mm. to 2.01 mm.

Color: General facies sordid white to gray. Head and pronotum
sordid white to yellowish white. Thoracic venter and limbs pale
testaceous to light brown; scutellum varies from entirely sordid
white, to black with apex and lateral margins yellowish white; meta-
thoracic dorsum yellowish white to mostly black. Abdomen black
except ventral keel and portions of connexivum and dorsum, sordid
white to testaceous. Some specimens entirely sordid white except
abdomen, mostly black. This species variable in color.

Male Structural Characteristics: As viewed from above, outline
of head rounded with vertex slightly protuberant; greatest width
of head approximately five times the anterior width of vertex and
slightly less than humeral width of pronotum; synthlipsis half the
anterior width of vertex; along median longitudinal axis, head is
approximately one third the length of pronotum; notocephalon dis-
tinctly sulcate; tylus inflated; labrum with basal width slightly more
than twice its median length and apex bluntly rounded; rostral
prong ( pi. CVII, fig. 48b ) long, distinctly longer than third rostral
segment, with base originating laterally at distal end of third rostral
segment, and with apex moderately rounded. Pronotum long, with
its median length two thirds to three fourths its humeral width; disk
with two elongate depressions toward the middle and a large sub-
triangular depression on each side, thus appearing distinctly tri-
carinate; lateral margins slightly divergent; posterior margin convex,
medianly concave. Scutellum with median length distinctly less
than that of pronotum. Fore femur (pi. CVII, fig. 48a) wide and
somewhat thickened at apex; long, triangular stridulatory area con-
sisting of approximately seventeen to twenty-two sclerotized ridges.
Fore tibia (pi. CVII, fig. 48a) with stridulatory comb (pi. CVII, fig.
48c) consisting of approximately thirty-nine to forty-two teeth;
apical teeth thicker and slightly narrower than basal. Chaetotaxy
of male front leg as shown on Plate VII. Male genital claspers
normal. Spine from caudo-sinistral margin of seventh abdominal
tergite normal, tapering gradually from base to strongly acuminate
apex.

Female Structural Characteristics: As viewed from above, outline
of head rounded with vertex slightly protuberant; greatest width of

58—6490

1392 The University Science Bulletin

head four and one half to five times the anterior width of vertex
and less than humeral width of pronotum; synthlipsis approximately
half the anterior width of vertex; along median longitudinal axis,
head is one third to two fifths the length of pronotum; notocephalon
distinctly sulcate; tylus inflated. Pronotum with its median length
slightly more than half its humeral width; disk usually with two
shallow, elongate depressions toward the middle forming a faint
median carina, not tricarinate; lateral margins divergent; posterior
margin convex, medianly concave. Scutellum large, with median
length equal to or greater than that of pronotum. Female ovipositor
of normal shape with teeth arranged in two irregular, longitudinal
rows which merge in proximal third of ovipositor valve; one inner
row of large teeth and one outer row of smaller teeth; approximately
six or seven small, lateral, toothlike setae near apex.

Comparative Notes: Superficially this species somewhat resem-
bles B. arizonis Bare. Examination of the male, however, will show
distinct differences. This species differs from B. arizonis in its dis-
tinctly smaller size, in having head smaller in proportion to pro-
notum, tylus less inflated and not medianly depressed, and distinct
differences in rostral prong and femoral stridulatory area.

Location of Types: Holotype male, allotype female, 26 male and
26 female paratypes, Dept. Lima, Lagunas Villa, Peru, June 8 to
July 1, 1934, F. Woytkowski. The type series is in the Francis
Huntington Snow Entomological Collections, University of Kansas.

Data on Distribution: Known only from Peru. In addition to
type series, specimens from the following localities have been ex-
amined:

PERU: Dept. Lima: Lagunas Villa, June 8 to July 15, 1934, F.
Woytkowski, 37 males, 43 females; Lurin, Nov. 3-5, 1934, F. Woyt-
kowski, 128 males, 146 females; Vicinity Pacasmayo, May 19-20,
1936, F. Woytkowski, 26 males, 20 females; Chilca, Jan. 31, 1937,
F. Woytkowski, 6 males, 42 females.

All specimens listed above are in the Francis Huntington Snow
Entomological Collections, University of Kansas, Lawrence, Kansas.

Buenoa tarsalis n. sp.
(PI. CVII, fig. 49)

Size: This species varies considerably in size within the same
population. Male, length 6.24 mm. to 7.15 mm., greatest body width
1.62 mm. to 1.95 mm.; female, length 6.30 mm. to 8.06 mm., greatest
body width 1.95 mm. to 2.53 mm.

Color: General facies sordid white to testaceous. Head, pro-

A Revision of the Genus Buenoa 1393

notum, and most of thoracic venter and limbs, sordid white, por-
tions of thoracic venter and limbs sometimes light brown to black.
Scutellum sordid white to pale testaceous, occasionally with two
anterolateral brown to black areas; metathoracic dorsum sordid
white to light brown. Abdomen black except ventral keel and por-
tions of connexivum and dorsum, sordid white.

Male Structural Clwracteristics: As viewed from above, outline
of head rounded with anterior margin of vertex slightly indented
to continuous with that of eyes; greatest width of head five and
one half to six times the anterior width of vertex and slightly less
than humeral width of pronotum; synthlipsis approximately half
the anterior width of vertex; along median longitudinal axis, head
is two fifths to one half the length of pronotum; notocephalon
slightly sulcate; tylus inflated; labrum with its basal width more
than twice its median length and apex bluntly rounded; rostral
prong ( pi. CVII, fig. 49c ) slightly longer than third rostral segment,
with base originating laterally near proximal end of third rostral
segment, and with apex moderately to bluntly rounded. Pronotum
with its median length approximately three fifths its humeral width;
disk with two elongate depressions toward the middle and a large
subtriangular depression on each side, thus appearing tricarinate;
lateral margins divergent; posterior margin convex, medianly con-
cave. Scutellum large, with median length slightly greater than
that of pronotum. Fore femur (pi. CVII, fig. 49a) relatively narrow
at apex, not greatly thickened; oblong stridulatory area consisting
of approximately seventeen to twenty-three sclerotized ridges. Fore
tibia (pi. CVII, fig. 49a) with stridulatory comb (pi. CVII, fig. 49d)
consisting of approximately thirty-two to forty teeth; apical teeth
thicker and slightly narrower than basal. Intermediate leg with
first tarsal segment ( pi. CVII, fig. 49b ) deeply emarginate on inner
margin. Chaetotaxy of male front leg as shown on Plate CVII.
Male genital claspers normal. Spine from caudo-sinistral margin
of seventh abdominal tergite with apical half very narrow and apex
strongly acuminate.

Female Structural Characteristics: As viewed from above, outline
of head rounded with anterior margin of vertex usually continuous
with that of eyes; greatest width of head approximately five times
the anterior width of vertex and less than humeral width of prono-
tum; synthlipsis slightly less than half the anterior width of vertex;
along median longitudinal axis, head is approximately two fifths
the length of pronotum; notocephalon sulcate; tylus slightly inflated.
Pronotum with its median length slightly less than half its humeral

1394 The University Science Bulletin

width; disk with two elongate depressions toward the middle
forming a median carina, occasionally with a shallow, subtriangular
depression on each side, thus appearing faintly tricarinate; lateral
margins divergent; posterior margin convex, medianly concave.
Scutellum large, with median length distinctly greater than that
of pronotum. Female ovipositor of normal shape with teeth ar-
ranged in two longitudinal rows which merge in proximal third
of ovipositor valve; one inner row of large teeth and one outer
row of smaller teeth; approximately seven to nine small, lateral,
toothlike setae located in a row extending from apex to midway of
ovipositor valve.

Comparative Notes: Superficially this species resembles B. anti-
gone antigone (Kirkaldy). Examination of the male, however, will
show distinct differences. This species differs from B. antigone
antigone in having synthlipsis narrower, first tarsal segment of
intermediate leg strongly emarginate, hind femur less robust, and
differences in the rostral prong, femoral stridulatory area, and tibial
comb.

Location of Types: Holotype male, allotype female, 18 male and
20 female paratypes, Rio Sao Paulo Road, State of Rio de Janeiro,
Brazil, June 19, 1945, Wygodzinsky. The type series is in the
Francis Huntington Snow Entomological Collections, University of
Kansas.

Data on Distribution: Known only from Brazil. In addition to
type series, specimens from the following localities have been ex-
amined :

BRAZIL: Para: Marco Belem, Jan. 1, 1947, L. & M. Deane, 16
males, 21 females, 5 nymphs.

Ceard: Agua Verde, July 6, 1937, S. Wright, 17 males, 38 females;
Fortaleza, Aug. 3, 1937, S. Wright, 1 female; Lavras, Artificial lake,
Aug. 23, 1937, S. Wright, 1 male; Lagoa Frexeiras, Nr. Maranguape,
Sept., 1937, S. Wright, 11 males, 5 females; Primavera, Oct. 28, 1937,
S. Wright, 4 males, 8 females; Choro, Oct. 29, 1937, S. Wright, 2
males, 4 females; Maranguape, Nov. 3-4, 1937, S. Wright, 4 males,
8 females.

Rio Grande do Norte: Ouro Branco, No. 258, S. Wright, 3 females.

Parahiba: Campina Grande, Nos. 43 & 212, S. Wright, 3 males,
12 females; Pocinhos, Nos. 197 & 5582, S. Wright, 1 male, 6 females;
Santa Luzia, No. 260, S. Wright, 1 male, 5 females; Souza, No. 5530,
S. Wright, 14 males, 3 females.

Pernambuco: Caruaru, No. 403, S. Wright, 8 males, 14 females;
Pesqueira, No. 434, S. Wright, 3 females; Rio Branco, No. 445, S.

A Revision of the Genus Buenoa 1395

Wright, 4 females; Belem, No. 643, S. Wright, 5 males, 10 females;
Itaparica, No. 818, S. Wright, 4 males, 3 females.

Rio de Janeiro: Rio Sao Paulo Road, June 19, 1945, Wygodzinsky,
19 females; Rio Sao Paulo Road, Feb., 1945, P. Wygodzinsky, 10
males, 26 females.

All specimens listed above are in the Francis Huntington Snow
Entomological Collections, University of Kansas, Lawrence, Kansas.

Buenoa rostra n. sp.
(PI. CVII, fig. 50)

Size: Male, length 5.78 mm. to 6.50 mm., greatest body width
1.62 mm. to 1.85 mm.; female, length 5.80 mm. to 6.21 mm., greatest
body width 1.62 mm. to 1.88 mm.

Color: General fades sordid white to testaceous. Head, pro-
notum, thoracic venter, and limbs sordid white to pale testaceous.
Scutellum sordid white to yellowish-white; metathoracic dorsum
varies from entirely yellowish white to mostly dark brown. Abdo-
men black except ventral keel and portions of connexivum and
dorsum, yellowish white. This species variable in color.

Male Structural Characteristics: As viewed from above, outline
of head laterally rounded, anteriorly truncate with vertex indented;
greatest width of head five and one half to six and one half times
the anterior width of vertex and slightly less than humeral width
of pronotum; synthlipsis one third to one half the anterior width
of vertex; along median longitudinal axis, head is one fourth to
one third the length of pronotum; notocephalon sulcate; tylus
slightly inflated; labium with basal width not quite twice its median
length and apex bluntly rounded; rostrum very robust; rostral prong
(pi. VII, fig. 50b) distinctly longer than third rostral segment, with
base originating laterally at distal end of third rostral segment, and
with apex bluntly rounded. Pronotum large, with its median length
approximately two thirds its humeral width; disk usually with two
elongate depressions posteriorly and toward the middle and a sub-
triangular depression on each side, thus appearing tricarinate pos-
teriorly; lateral margins almost parallel; posterior margin convex,
medianly concave. Scutellum large but with median length less
than that of pronotum. Fore femur (pi. CVII, fig. 50a) wide and
somewhat thickened at apex; subtriangular stridulatory area con-
sisting of approximately fourteen to seventeen sclerotized ridges.
Fore tibia (pi. CVII, fig. 50a) with stridulatory comb (pi. CVII,
fig. 50c) consisting of approximately twenty-seven to twenty-eight
teeth; apical teeth thicker and slightly wider and taller than basal.

1396 The University Science Bulletin

Chaetotaxy of male front leg as shown on Plate CVII. Male genital
claspers normal. Spine from caudo-sinistral margin of seventh
abdominal tergite with apical third very narrow and apex strongly
acuminate.

Female Structural Characteristics: As viewed from above, out-
line of head laterally rounded, anteriorly truncate with vertex
slightly indented; greatest width of head five and one half to six
and one half times the anterior width of vertex and less than hu-
meral width of pronotum; synthlipsis one third to one half the
anterior width of vertex; along median longitudinal axis, head is
approximately two fifths the length of pronotum; notocephalon
sulcate; tylus slightly inflated. Pronotum with its median length
approximately half its humeral width; disk unimpressed; lateral
margins divergent; posterior margin convex, medianly concave.
Scutellum large, with median length distinctly greater than that of
pronotum. Female ovipositor of normal shape with teeth arranged
in two longitudinal rows which merge proximally to form a single
irregular row; one inner row of large teeth and one outer row of
smaller teeth; approximately seven or eight small, lateral, tooth-
like setae near apex.

Variation Within Species: This species varies considerably in the
proportional size of the head, anterior width of vertex, and synth-
lipsis. The greatest contrast is shown between a series from Vene-
zuela and one from Trinidad. The former is larger and usually
darker than the latter.

Comparative Notes: Superficially this species somewhat resem-
bles the pale form of B. crassipes (Champion). Examination of
the male, however, will show distinct differences. This species
differs from B. crassipes in its smaller size, in having the pronotum
longer and more convex, and in the distinctly different rostral prong
and tibial comb ( pi. C, figs. 50b, 50c ) .

Location of Types: Holotype male, allotype female, 3 male and
6 female paratypes, Trinidad, British West Indies, Sept. 27, 1931,
W. E. Broadway. The type series is in the Francis Huntington
Snow Entomological Collections, University of Kansas.

Data on Distribution: Known only from the West Indies (Trini-
dad) and Venezuela. In addition to type series, specimens from
the following localities have been examined:

VENEZUELA: No locality, July 3, 1897, A. Speyer, 1 male, 1 fe-
male (F. H. Snow Coll.), 4 males, 8 females (Hamburg Mus.);
San Esteban, Nov. 22, 1939, Pablo J. Anduze, 1 male, 3 females
(F. H. Snow Coll.).

A Revision of the Genus Buenoa 1397

Buenoa margaritacea Torre-Bueno
(PL CHI, fig. 33; pi. CVIII, fig. 51)

1882. Anisops platycnemis, Uhler, P. R. Standard Nat. Hist., vol. II, p. 253.
1891. Anisops platycnemis, Summers, H. E. Bull. Agr. Exp. Sta. Tennessee,

vol. IV, p. 82.
1902. Anisops platycnemis, Torre-Bueno, J. R. de la. Jr. New York Ent. Soc,

vol. X, p. 236 (description).
1904. Anisops platycnemis, Kirkaldy, G. W. Wiener Ent. Zeit., vol. XXIII,

p. 123 (description).

1908. Buenoa margaritacea, Torre-Bueno, J. R. de la. Jr. New York Ent. Soc,
vol. XVI, p. 238 ( B. margaritacea nom. nov. = platycnemis, Uhler, Bueno,
et auct., nee. Fieber).

1909. Buenoa margaritacea, Torre-Bueno, J. R. de la. Jr. New York Ent. Soc,
vol. XVII, pp. 75-77 (key, notes, and synonomy).

1910. Buenoa margaritacea, Smith, J. B. Cat. Insects New Jersey, edn. 3,
p. 170 (catalogue).

1910. Buenoa margaritacea, Torre-Bueno, J. R. de la. Jr. New York Ent. Soc,

vol. XVIII, p. 33 (listed and ecological note).
1914. Buenoa margaritacea, Barber, H. G. Bull. American Mus. Nat. Hist.,

vol. XXXIII, p. 499 (fisted).

1916. Buenoa margaritacea, Van Duzee, E. P. New York Ent. Soc, p. 51
(check list).

1917. Buenoa margaritacea, Van Duzee, E. P. Cat. Hemiptera Am. North of
Mexico, p. 454 (catalogue).

1917. Buenoa margaritacea, Hungerford, H. B. Ent. News, vol. XXVIII,

pp. 174-183 (biological notes).
1917. Buenoa margaritacea, Hungerford, H. B. Ent. News, vol. XXVIII,

p. 271 (biological note).
1917. Buenoa margaritacea, Parshley, H. M. Occasional Papers of Boston Soc

Nat. Hist., vol. VII, p. 113 (listed).
1919. Buenoa margaritacea, Hungerford, H. B. Univ. Kansas Sci. Bull., vol.

XI, pp. 1-328 (biology, taxonomy, and key).
1919. Buenoa margaritacea, Hungerford, H. B. Univ. Kansas Sci. Bull., vol.

XI, pp. 329-333 (morphological note).
1922. Buenoa margaritacea, Parshley, H. M. South Dakota State College

Tech. Bull., No. 2, p. 22.

1922. Buenoa margaritacea, Hungerford, H. B. Canadian Ent., vol. LXIV,
pp. 262-263 (morphological note).

1923. Buenoa margaritacea, Torre-Bueno, J. R. de la. Guide to Insects of
Connecticut, part 4, p. 407.

1923. Buenoa margaritacea, Hungerford, H. B. Ent. News, vol. XXXIV,

pp. 150-151 (biological note).
1923. Buenoa margaritacea, Hale, H. M. Records South Australian Mus., vol.

II, p. 399 (notes).

1923. Buenoa margaritacea, Torre-Bueno, J. R. de la. Connecticut State Geol.
and Nat. Hist. Survey Bull., No. 34, p. 407 (key and notes).

1924. Buenoa margaritacea, Hungerford, H. B. Ann. Ent. Soc. Am., vol.
XVII, pp. 223 and 325 (biology and taxonomic notes).

1925. Buenoa margaritacea, Bare, C. O. Ent. News, vol. XXXVI, pp. 225-228
( key and taxonomic notes ) .

1925. Buenoa margaritacea, Hungerford, H. B. and Beamer, R. H. Ent. News,
vol. XXXVI, pp. 264 and 297 (fisted and notes).

1926. Buenoa margaritacea, Bare, C. O. Ann. Ent. Soc. Am., vol. XIX, pp. 93-
101 (biological notes).

1926. Buenoa margaritacea, Blatchley, W. S. Heteroptera or True Bugs of
Eastern North America, pp. 1057-1058 (description and key).

1926. Buenoa margaritacea, Clark, L. B. Canadian Ent., vol. LVIII, pp. 203-
204 (listed and distributional note).

1928. Buenoa margaritacea, Bare, C. O. Univ. Kansas Sci. Bull., vol. XVIII,
pp. 265-349 (taxonomy, biology, and morphology).

1398 The University Science Bulletin

1928. Bucnoa margaritacea, Torre-Bueno, J. R. de la. Cornell Univ. Agr. Exp.

Station, Memoir 101, p. 139 (listed).
1939. Buenoa margaritacea, Millspaugh, D. D. Field and Laboratory, vol. VII,

p. 78.
1942. Buenoa margaritacea, Rice, L. A. Tennessee Acad. Sci., vol. XVII,

pp. 55, 62, and 63 (listed and biology).
1942. Buenoa margaritacea, Hutchinson, G. E. American Jr. Sci., vol. CCXL,

p. 336 (morphological note).

Size: This species varies considerably in size within the same
population. Male, length 6.01 mm. to 7.36 mm., greatest body
width 1.50 mm. to 2.14 mm.; female, length 6.63 mm. to 8.25 mm.,
greatest body width 1.95 mm. to 2.40 mm.

Color: General facies sordid white to dark brown. Head, pro-
notum, most of thoracic venter and usually metathoracic dorsum,
and limbs, sordid white to testaceous; portions of thoracic venter
and metathoracic dorsum often light brown to black; scutellum
yellowish to testaceous, occasionally with two anterolateral black
spots. Abdominal dorsum varying from testaceous anteriorly and
black posteriorly to almost entirely black; abdominal venter black
except keel, portions of connexivum, and sometimes last one or two
segments, testaceous. This species varies considerably in color.

Male Structural Characteristics: As viewed from above, outline
of head rounded with vertex occasionally indented at lateral mar-
gins; greatest width of head five and one half to six and one half
times the anterior width of vertex and distinctly less than humeral
width of pronotum; synthlipsis approximately half the anterior
width of vertex; along median longitudinal axis, head is two fifths
to three fifths the length of pronotum; notocephalon usually slightly
sulcate dorsally; tylus not inflated; labrum with basal width ap-
proximately twice its median length and apex bluntly rounded;
rostral prong (pi. CVIII, fig. 51c) short, shorter than third rostral
segment, with base originating laterally at proximal end of third
rostral segment, and with apex bluntly rounded. Pronotum with
its median length approximately three fifths its humeral width; disk
often only laterally impressed, usually with two very shallow, elon-
gate depressions toward the middle and a large subtriangular
depression on each side, thus appearing faintly tricarinate; lateral
margins divergent; posterior margin convex, medianly concave.
Scutellum large, with median length equal to or slightly greater
than that of pronotum. Fore femur (pi. CVIII, fig. 51a) narrow,
not thickened at apex; subtriangular to oval stridulatory area con-
sisting of approximately fifteen to eighteen sclerotized ridges. Fore
tibia (pi. CVIII, fig. 51a) with stridulatory comb (pi. CVIII, fig.
51b) consisting of approximately twenty-three to twenty-six teeth

A Revision of the Genus Buenoa 1399

which increase slightly in thickness from base to apex. Chaetotaxy
of male front leg as shown on Plate CVIII. Male genital claspers
normal. Spine from caudo-sinistral margin of seventh abdominal
tergite with apical half very narrow and apex strongly acuminate.

Female Structural Characteristics: As viewed from above, out-
line of head rounded with vertex occasionally indented at lateral
margins; greatest width of head five to six times the anterior width
of vertex and less than humeral width of pronotum; synthlipsis
approximately half the anterior width of vertex; along median
longitudinal axis, head is approximately two fifths the length of
pronotum; notocephalon not sulcate; tylus not inflated. Pronotum
with its median length approximately three fifths its humeral width;
disk usually unimpressed, occasionally with a shallow, subtriangular
depression on each side, not tricarinate; lateral margins divergent;
posterior margin convex, medianly concave. Scutellum large, with
median length distinctly greater than that of pronotum. Female
ovipositor (pi. CIII, fig. 33) of normal shape with teeth arranged
in two longitudinal rows; one inner row of large teeth which merges
proximally with long outer row of small teeth; approximately four
small, lateral, toothlike setae near apex.

Comparative Notes: Superficially this species closely resembles
B. scimitra Bare. Examination of the male, however, will show
distinct differences. This species differs from B. scimitra in the
shape of the fore femur, the femoral stridulatory area, and the
rostral prong. In general, B. margaritacea is slightly larger in size
than the above mentioned species.

Nomenclatorial Notes: Mr. J. R. de la Torre-Bueno (1908) in
designating B. margaritacea as a new name for "platycnemis, Uhler,
Bueno, et auct,, nee Fieber" states that, "The Buenoa known to a
generation of American entomologists as Anisops platycnemis was
an undescribed species." He does not proceed to describe B. mar-
garitacea but states that the descriptions by Uhler (1882), Bueno
( 1902 ) and Kirkaldy ( 1904 ) "will enable anyone to identify this
species with certainty." The main distinguishing factor in the
above mentioned descriptions is the size. Buenoa platycnemis
Fieber is a small species whereas B. margaritacea is a relatively
large species.

1400 The University Science Bulletin

Location of Types: To the knowledge of this author, no types
were designated by J. R. de la Torre-Bueno for B. margaritacea.
If types were established, they have subsequently been lost or
destroyed. A series of three specimens, 2 males, 1 female, were
located in the remnants of the Bueno collection now in the Francis
Huntington Snow Entomological Collections of the University of
Kansas. These specimens are from Maryland, U. S. A., in the gen-
eral vicinity of Bueno's collecting, and where Uhler states, "I have
found it late in October." I have selected one of these males as a
neoholotype and the female as a neoallotype.

Data on Distribution: Recorded from Canada, United States,
and Mexico. Specimens from the following localities have been
examined:

U.S.A.: Minnesota: St. Paul, Sept. 23, 1918, R. F. Hussey, 1
female (U. of Mich.); St. Paul, June 19, 1921, H. B. Hungerford,
3 females; St. Paul, July 28-31, 1921, H. B. Hungerford, 4 males, 3
females.

Michigan: Berrien Co., Sept. 2, 1919, R. F. Hussey, 5 males, 8
females ( U. of Mich. ) ; Livingston Co., Aug. 9, 1938, I. J. Cantrall,
1 female ( U. of Mich. ) .

South Dakota: Blunt, July 19, 1937, C. L. Johnston, 1 female;
Brookings Co., Aug. 11, 1939, H. C. Severin and W. Eakins, 2
females.

New York: West Point, Apr. 19, 1909, W. Robinson, 2 females
(U.S.N.M.).

Pennsylvania: Philadelphia, Oct. 18, 1908, G. M. Greene, 1 fe-
male (U.S.N.M.).

Illinois: Lake Forest, S. G. Needham, 2 females (U.S.N.M.); P.
R. Uhler Collection, 1 male, 1 female (U.S.N.M.).

Maryland: Patuxent, Aug. 2, 1879, P. R. Uhler, 1 male, 1 female
(Bueno Coll.); Baltimore, Sept. 4, 1897, P. R. Uhler, 1 male (Bueno
Coll.).

Colorado: Lamar, July 22, 1927, P. A. Readio, 2 males, 8 females;
Lamar, Aug. 20, 1927, P. A. Readio, 9 males; Hadley, Sept. 22, 1927,
P. A. Readio, 1 male; Denver, P. R. Uhler collection, 1 female
(U.S.N.M.).

California: Santa Ana, July 30, 1932, J. D. Beamer, 1 female, 1
nymph; Laguna Beach, July 25, 1933, R. H. Beamer, 3 males, 4 fe-
males; Idyllwild, Aug. 3, 1935, Jack Beamer, 1 female; Campo, June
18, 1940, R. H. Beamer, 1 male; Stanford University, March 16, 1915,
1 female.

A Revision of the Genus Buenoa 1401

Kansas: Doniphan Co., Aug. 23, 1921, W. J. Brown, 5 males, 1 fe-
male; Doniphan Co., Aug. 24-25, 1921, Robert Guntert, 2 males, 1 fe-
male; July 23, 1924, E. P. Breakey, 1 female; Atchison Co., July 15-
17, 1924, E. P. Breakey, 8 males, 16 females; Atchison Co., July 15,

1924, Beamer, 4 males, 7 females; Pottawatomie Co., Crevecoeur, 1
female (U.S.N.M.); Leavenworth Co., June 1923, E. P. Breakey, 3
females; Shawnee Co., May 26, 1923, H. B. Hungerford, 3 females;
Shawnee Co., June 27, 1923, H. B. Hungerford, 3 males; Douglas Co.,
April 7, 1909, 1 male; Douglas Co., Aug. 2, 1909, 3 males, 5 females;
Douglas Co., 1916, H. B. Hungerford, 1 male, 1 female; Douglas Co.,
May 17, 1920, H. B. Hungerford, 5 males, 2 females; Douglas Co.,
Feb. 23, 1921, H. B. Hungerford, 1 female; Douglas Co., Apr. 8, 1921,
H. B. Hungerford, 1 male, 4 females; Douglas Co., June 3, 1921, W.
J. Brown, 1 male, 1 female; Douglas Co., Nov. 3, 1922, H. B. Hunger-
ford, 1 female; Douglas Co., May 20, 1923, C. O. Bare, 15 males, 5 fe-
males; Douglas Co., Sept. 28, 1924, C. O. Bare, 59 males, 55 females;
Douglas Co., Oct. 25, 1924, C. O. Bare, 11 males, 17 females; Douglas
Co., Nov. 19-20, 1924, C. O. Bare, 85 males, 78 females; Douglas Co.,
Apr. 5. 1925, C. O. Bare, 4 males, 48 females; Douglas Co., May 6,

1925, C. O. Bare, 8 males, 2 females; Douglas Co., Beamer, 2 males,
48 females; Osage Co., June 1923, Beamer, 2 males, 5 females; Saline
Co., July 14, 1923, L. C. Woodruff, 4 females; Saline Co., July 15-18,
1923, R. H. Beamer, 2 males, 8 females; Franklin Co., March 20,

1926, Wesley Clanton, 1 female (U. of Mich.); Lyon Co., June 14,
1923, C. O. Bare, 1 female; Lyon Co., June 15, 1923, W. J. Brown, 1
female; Lyon Co., June 18, 1923, H. Darby, 1 female; Linn Co., Apr.
14, 1923, H. B. Hungerford, 1 female; Bourbon Co., 1915, R. H.
Beamer, 1 male; Reno Co., Aug. 27, 1925, W. J. Brown, 2 males, 4
females; Reno Co., July 2, 1927, P. A. Readio, 1 male, 2 females;
Reno Co., July 3, 1927, L. D. Anderson, 1 male; Gray Co., July 9-15,
1917, 1 female; Cherokee Co., Aug. 15, 1920, H. B. Hungerford, 1
female; Cherokee Co., Aug. 18, 1920, R. H. Beamer, 3 females;
Montgomery Co., 1916, R. H. Beamer, 3 females; Comanche Co.,
June 19, 1927, H. B. Hungerford, 3 males, 2 females; Morton Co.,
Aug. 3, 1924, C. O. Bare, 1 male, 15 females.

Missouri: St. Louis, July, 1910, J. F. Abbott, 1 male; St. Louis,
Oct., 1911, F. J. Abbott, 1 female.

Virginia: Great Falls, Jan. 9, 1906, D. H. Clemons, 1 female
(U.S.N.M.); Vienna, Sept. 19, 1931, P. W. Oman, 58 males, 61 fe-
males; Vienna, Sept. 19, 1931, P. W. Oman, 1 male, 3 females (U.S.
N.M.).

1402 The University Science Bulletin

Tennessee: Murfreesboro, Aug. 29, 1929, Greaser Becker, 1 fe-
male; Knoxville, 1 female (U. S. N. M.).

Oklahoma: Cimarron Co., July 5, 1926, T. H. Hubbell, 2 males,
24 females (U. of Mich.); Ardmore, Apr. 14, 1923, H. B. Hunger-
ford, 1 male, 17 females; Osage Co., June 23, 1936, W. F. Blair,
1 female ( U. of Mich. ) .

New Mexico: Torrance Co., June, 1925, C. H. Martin, 9 males,
31 females; Torrance Co., July 19, 1925, C. H. Martin, 4 females;
Torrance Co., Sept., 1925, C. H. Martin, 18 males, 16 females;
Estancia, Aug. 25 to Sept. 6, 1925, C. H. Martin, 21 males, 25 fe-
males; Chaves Co., July 8, 1927, R. H. Beamer, 1 male, 2 females;
Santa Cruz, Aug. 4, 1927, R. H. Beamer, 1 male, 3 females; Santa
Cruz, Aug. 4, 1927, P. A. Readio, 6 males, 7 females; Santa Cruz,
Aug. 20, 1927, L. D. Anderson, 11 males, 4 females; Socorro Co.,
Aug. 8, 1927, P. A. Readio, 1 male, 11 females; Otero Co., June 28,
1931, L. K. Gloyd, 2 males, 5 females (U. of Mich.); Wagon Mound,
July 18, 1936, M. B. Jackson, 2 females; Santa Fe, July 20, 1936,
J. D. Beamer, 1 female.

Arizona: Cochise Co., July 29, 1927, R. H. Beamer, 7 males, 30
females; Gila Co., Aug. 5, 1927, L. D. Anderson, 2 males, 4 females;
Gila Co., Aug. 5, 1927, R. H. Beamer, 5 males, 19 females; Gila Co.,
Aug. 6, 1927, P. A. Readio, 6 males, 3 females; Navajo Co., Aug. 15,
1927, Anderson and Readio, 6 males, 10 females; Navajo Co., Aug.
15, 1927, R. H. Beamer, 2 males, 1 female; Apache Co., Aug. 16,
1927, R. H. Beamer, 9 males, 11 females; Apache Co., Aug. 16, 1927,
P. A. Readio, 10 males, 4 females; Apache Co., Aug. 16, 1927, L. D.
Anderson, 2 females; Coconino Co., July 1, 1929, L. D. Anderson,
1 female; Baboquivari Mts., July 16, 1932; R. H. Beamer, Jr., 5 males,
18 females; Baboquivari Mts., July 24, 1941, B. Hodgden, 12 males,
57 females; Tuscon, July 20, 1932, R. H. Beamer, 2 males, 12 fe-
males; Douglas, Apr. 21, 1933, W. W. Jones, 1 female (U. S. N. M.);
Douglas, Aug., F. H. Snow, 1 male; Yavapai Co., June 9, 1937,
L. K. Gloyd, 1 female (U. of Mich.); Ruby, July 13, 1940, D. E.
Hardy, 1 male, 1 female; Ruby, July 27, 1941, B. Hodgden, 2 fe-
males; Miami, Aug. 6, 1941, E. L. Todd, 45 males, 20 females;
Miami, Aug. 6, 1941, B. Hodgden, 20 males, 9 females; Fort Grant,
H. G. Hubbard, 1 female (U. S. N. M.).

Arkansas: Scott Co., Aug. 23, 1928, R. H. Beamer, 3 females;
Fayetteville, Apr. 23-28, 1930, 4 females; Saline Co., July 7, 1950,
R. H. Beamer, 1 male.

South Carolina: Organsburg, Sept. 2, 1914, W. J. Brown, 1 fe-
male.

A Revision of the Genus Buenoa 1403

Georgia: Atlanta, March 14, 1933, P. W. Fattig, 1 female (U. S.

N.M.).

Mississippi: Agricultural College, Oct., 1895, W. E. Weed, 1 fe-
male (P. R. Uhler Coll.); Agricultural College, June 20, 1919, J. B.
Ray, 1 female; Fulton, July 14, 1930, R. H. Beamer, 1 female.

Louisiana: Baton Rouge, March 9, 1929, R. M. DeCoursey, 1
female.

Texas: Brownsville, June 5, 1904, H. S. Barber, 2 males, 1 female
(U. S. N. M.); Brownsville, Aug. 1919, 2 females (U. S. N. M.);
Kerrville, Apr. 12, 1907, F. C. Pratt, 1 female (U. S. N. M.); Vic-
toria, Dec. 27, 1910, J. D. Mitchell, 1 female (U.S.N.M.); East-
land Co., May 12-26, 1921, Grace Wiley, 26 males, 29 females;
Colorado Co., March 30, 1922, 4 females; Colorado Co., Apr. 3-24,
1922, Mrs. Grace Wiley, 3 males, 39 females; Colorado Co., May
5-19, 1922, Grace Wiley, 1 male, 7 females; Tulsa Co., March 16-21,
1922, Grace Wiley, 1 male, 4 females; Randall Co., July 7, 1927,
R. H. Beamer, 4 males, 23 females; Valentine, July 13, 1927, R. H.
Beamer, 13 males, 15 females; Valentine, July 12, 1938, D. W. Craik,
1 male, 1 female; Presidio Co., July 16, 1927, R. H. Beamer, 6 males,
17 females; El Paso Co., July 17, 1927, L. D. Anderson, 1 female;
Sulton Co., July 20, 1928, J. G. Shaw, 3 males; Sulton Co., Aug. 20,
1928, A. M. James, 3 males, 5 females; Kendall Co., July 22, 1928,
R. H. Beamer and J. G. Shaw, 1 male, 5 females; Jim Wells Co.,
July 24, 1928, R. H. Beamer, 6 males, 6 females; Alfred, July 24,
1928, R. H. Beamer, 1 male; Bee Co., July 25, 1928; R. H. Beamer,

1 male; Hidalgo Co., July 28, 1928, J. G. Shaw, 3 males; Hidalgo
Co., July 30, 1928, R. H. Beamer, 1 male, 4 females; Hidalgo Co.,
Aug. 3, 1928, J. G. Shaw, 14 males, 9 females; Starr Co., July 30,
1928, J. G. Shaw, 33 males, 32 females, 3 nymphs; Cameron Co.,
Aug. 13, 1928, A. M. James, 1 female; Bowie Co., Aug. 16, 1928,
R. H. Beamer, 1 male; Brewster Co., Apr. 15, 1930, O. C. Poling,

2 males (U. of Mich.); Fulfurrias, Nov. 2, 1932, L. D. Tuthill, 2
males; Sinton, Nov. 8, 1932, L. D. Tuthill, 1 male, 1 female; Mc-
Allen, Nov. 20, 1932, L. D. Tuthill, 1 male; McAllen, Dec. 30, 1945,
R. H. Beamer, 2 females; Marathon, July 9, 1938, R. I. Sailer, 4
males, 6 females; Davis Mts., July 12, 1938, D. W. Craik, 2 males,
1 female; Cypress Mills, Chittenden, 1 female (U. S. N. M.); Texas,
P. R. Uhler Collection, 2 females ( U. S. N. M. ).

MEXICO: Chihuahua: Juarez, June 18, 1931, Smith and Dunkie,
1 male.

Coahuila: Satillo, Nov. 21, 1932, L. D. Tuthill, 7 males, 14 fe-
males.

1404 The University Science Bulletin

Tamaulipas: San Jose, Apr., 1910, Bueno Collection, 1 male, 1
female.

All specimens listed above are in the Francis Huntington Snow
Entomological Collections, University of Kansas, Lawrence Kansas,
unless otherwise indicated.

Buenoa scimitra Bare
(PL CII, fig. 6; pi. CVIII, fig. 52)

1925. Buenoa scimitra Bare, C. O. Ent. News, vol. XXXVI, pp. 226-228.

1926. Buenoa scimitra, Bare, C. O. Ann. Ent. Soc. America, vol. XIX, p. 93
(biological note).

1926. Buenoa scimitra, Blatchley, W. S. Heteroptera or True Bugs of Eastern

North America, pp. 1057-1059 (key and description).
1928. Buenoa scimitra, Bare, C. O. Univ. Kansas Sci. Bull., vol. XVIII, p. 268

(key).
1948. Buenoa scimitra, Hynes, H. B. N. Trans. Roy. Ent. Soc. London, vol.

XCIX, p. 354 (distributional note).

Size: This species varies considerably in size within the same
population. Male, length 5.46 mm. to 6.50 mm., greatest body
width 1.56 mm. to 1.82 mm.; female, length 5.85 mm. to 7.50 mm.,
greatest body width 1.69 mm. to 2.15 mm.

Color: General facies sordid white to fuscous. Head, anterior
portion of pronotum, most of thoracic venter, and limbs sordid white
to testaceous. Posterior portion of pronotum white to hyaline; scu-
tellum orange to reddish yellow, occasionally with anterolateral
portions black; metathoracic dorsum with lateral portions sordid
white to testaceous, remaining area light brown to black. Abdomen
black except ventral keel and portions of connexivum and dorsum,
testaceous. Some specimens entirely sordid white to testaceous
except most of abdomen, black. This species variable in color.

Male Structural Characteristics: As viewed from above, outline
of head rounded with vertex indented, often at lateral margins only;
greatest width of head six to six and one half times the anterior
width of vertex and less than humeral width of pronotum; synth-
lipsis slightly more than one third the anterior width of vertex;
along median longitudinal axis, head is two fifths to one-half the
length of pronotum; notocephalon sulcate dorsally; tylus inflated;
labrum with basal width approximately twice its median length and
apex bluntly rounded; rostral prong (pi. CVIII, figs. 52b, 52c)
shorter than third rostral segment, with base originating laterally
at a point midway to near proximal end of third rostral segment, and
with apex bluntly rounded. Pronotum with its median length ap-
proximately three fifths its humeral width; disk with two elongate
depressions toward the middle and a large, subtriangular depres-

A Revision of the Genus Buenoa 1405

sion on each side, thus appearing tricarinate; lateral margins di-
vergent; posterior margin convex, medianly concave. Scutellum
large, with median length equal to or greater than that of pronotum.
Fore femur (pi. CVIII, fig. 52a) wide and somewhat thickened at
apex; large, sword-shaped stridulatory area consisting of approxi-
mately sixty fine, sclerotized ridges. Fore tibia (pi. CVIII, fig. 52a)
with stridulatory comb (pi. CVIII, fig. 52d) consisting of approxi-
mately nineteen to twenty-two teeth; all teeth approximately same
thickness. Chaetotaxy of male front leg as shown on Plate CVIII.
Male genital claspers normal. Spine from caudo-sinistral margin of
seventh abdominal tergite (pi. CII, fig. 6) with apical half narrow
and apex strongly acuminate.

Female Structural Characteristics: As viewed from above, outline
of head rounded with vertex indented at lateral margins; greatest
width of head five to five and one half times the anterior width
of vertex and less than humeral width of pronotum; synthlipsis
approximately two fifths the anterior width of vertex; along median
longitudinal axis, head is two fifths to one half the length of pro-
notum; notocephalon sulcate dorsally; tylus slightly inflated. Pro-
notum with its median length approximately three fifths its humeral
width; disk with two shallow, elongate depressions toward the
middle and a shallow, subtriangular depression on each side, thus
appearing faintly tricarinate; lateral margins divergent; posterior
margin convex, medianly concave. Scutellum large, with median
length greater than that of pronotum. Female ovipositor of normal
shape with teeth arranged in two longitudinal rows; one inner row
of few, large teeth and one long outer row of smaller teeth; ap-
proximately three or four small, lateral, toothlike setae near apex.

Variation Within Species: As is indicated under measurements
of length, this species varies a great deal in size; there is also con-
siderable variation in the proportional size of the pronotum. This
variation is not due to distributional factors since it occurs within
the same population. Due to the fact that all characters used in
species determination are identical in these variable forms, no
specific separation appears justified.

Comparative Notes: Superficially this species closely resembles
B. margaritacea Torre-Bueno. Examination of the male, however,
will show distinct differences. This species differs from B. margari-
tacea in having the fore femur distinctly wider at apex, the femoral
stridulatory area long and sword-shaped, and the rostral prong
longer.

1406 The University Science Bulletin

Location of Types: Holotype male, allotype female, 20 male para-
types, Douglas Co., Kansas, May 20, 1923, Oct. 25, 1924, Nov. 19,
1924, C. O. Bare; other paratypes: 4 males, Colorado Co., Texas,
May 19, 1922, Grace Wiley. The type series is in the Francis
Huntington Snow Entomological Collections, University of Kansas.

Data on Distribution: Recorded from the United States, Mexico,
and the West Indies ( Cuba, Jamaica, Puerto Rico ) . In addition to
type series, specimens from the following localities have been ex-
amined:

U. S. A.: Kansas: Doniphan Co., Aug. 14, 1921, R. Guntert, 1 fe-
male; Doniphan Co., Aug. 23, 1921, W. J. Brown, 1 female; Atchi-
son Co., July 15-17, 1924, E. P. Breakey, 2 males, 1 female; Shawnee
Co., May 6, 1923, H. B. Hungerford, 2 females; Douglas Co., May
17, 1920, 1 male; Douglas Co., Nov. 3-7, 1922, H. B. Hungerford,
14 males, 44 females; Douglas Co., Nov. 3, 1922, R. Guntert, 5 males,
35 females; Douglas Co., May 20, 1923, C. O. Bare, 3 females;
Douglas Co., Sept. 28, 1924, C. O. Bare, 115 males, 40 females;
Douglas Co., Oct. 25, 1924, C. O. Bare, 60 males, 87 females; Doug-
las Co., Nov. 19-20, 1924, C. O. Bare, 38 males, 36 females; Douglas
Co., Oct. 10, 1925, C. O. Bare, 1 male, 1 female; Saline Co., July 18,
1923, R. H. Beamer, 1 female; Cherokee Co., Aug., 1920, H. B.
Hungerford and R. H. Beamer, 4 females; Cherokee Co., Dec. 1922,
R. H. Beamer, 1 male, 1 female; Montgomery Co., 1916, R. H.
Beamer, 1 male; Comanche Co., June 19, 1927, H. B. Hungerford,

2 males; Morton Co., July 20, 1924, C. O. Bare, 3 males, 3 females.
Virginia: New Church, July 15, 1934, L. D. Anderson, 1 male,

1 female.

California: Palo Alto, March 17, 1892, G. W. Kirkaldy Coll., 4
females; Lagoon Lake, Reade, Dec. 24, 1922, J. G. Needham, 6
females; Lagoon Lake, Aug. 24, 1925, J. G. Needham, 1 male;
Lagoon Lake, Dec. 24, 1925, J. G. Needham, 10 males, 3 females;
Calipatria, Apr. 4, 1924, Warewick Benedict, 4 females; Tehama
Co., Apr. 14, 1928, Jean Linsdale, 1 male, 8 females; Holtville,
July 2, 1929, Beamer and Anderson, 1 male, 3 females; San Diego
Co., July 7, 1929, L. D. Anderson, 1 male; San Diego Co., Apr. 19,
1930, C. & D. Martin, 1 female; Marin Co., Aug. 3, 1929, L. D.
Anderson, 12 males, 4 females; Campo, Aug. 25, 1932, H. W. Capps,

3 males, 3 females; Berkeley, Apr. 26, 1933, Jean Linsdale, 5 males,
6 females; Laguna Beach, July 25, 1933, R. H. Beamer, 58 males,
85 females; Red Bluff, June 27, 1935, Jack Beamer, 13 males, 8 fe-
males; Red Bluff, June 27, 1935, Jean Russell, 2 males, 4 females;
Anza, Aug. 6, 1935, R. H. Beamer, 1 female; El Centro, July 24,

A Revision of the Genus Buenoa 1407

1938, R. H. Beamer and D. W. Craik, 1 male, 6 females; Arroyo
Seco, Aug. 8, 1938, R. I. Sailer, 2 males, 5 females; Jamesburg,
Aug. 11, 1938, D. W. Craik, 2 males, 1 female.

Oklahoma: Tuko Co., Mar. 21, 1922, Grace Wiley, 2 males, 2
females.

Tennessee: Fentress Co., Aug. 17, 1922, T. H. Hubbell, 5 males,
4 females ( U. of Mich. ) .

Arizona: Pima Co., July 27, 1927, R. H. Beamer, 1 male; Cochese
Co., July 29, 1927, R. H. Beamer, 35 males, 41 females; Santa Cruz
Co., Aug. 4, 1927, R. H. Beamer and L. D. Anderson, 5 males, 4
females; Santa Cruz Co., Aug. 4, 1927, P. A. Readio, 12 males, 5
females; Gila Co., Aug. 6, 1927, P. A. Readio, 1 male, 1 female:
Yavapai Co., June 3, 1937, L. K. Gloyd, 2 females (U. of Mich.);
Ruby, July 27, 1941, B. Hodgden, 2 males, 1 female; Arivaca, July
10, 1947, L. D. Beamer, 2 males, 1 female.

Neiv Mexico: Socorro Co., Aug. 8, 1927, L. D. Anderson, 14 males,
7 females; Socorro Co., Aug. 18, 1927, P. A. Readio, 9 males, 9 fe-
males; Belen, July 20, 1936, W. D. Field, 3 males, 3 females.

Arkansas: Scott Co., Aug. 23, 1928, R. H. Beamer, 1 male, 21 fe-
males; Fayetteville, Apr. 24, 1930, 9 females; Arkansas Co., Sept. 4,
1930, D. Isely, 3 males, 7 females.

South Carolina: Orangeburg, Sept. 2, 1914, 1 male, 1 female.

Texas: Brownsville, 1875, 1 male, 3 females (Berlin Mus.);
Brownsville, Feb. 27, 1895, C. H. T. Townsend, 1 male (U.S.N.M.);
Brownsville, Dec. 29, 1945, R. H. Beamer, 1 male, 1 female; Victoria,
July 7, 1915, J. D. Mitchell, 3 males, 4 females (U.S.N.M.); East-
land Co., May 23-24, 1921, Grace Wiley, 5 males, 9 females; East-
land Co., June 16, 1921, Grace Wiley, 1 male; Eastland Co., May
14-25, 1927, Grace Wiley, 4 males, 2 females; Colorado Co., Apr.
3-24, 1922, Grace Wiley, 38 males, 30 females; Colorado Co., May
16-19, 1922, Grace Wiley, 11 females; Presidio Co., July 16, 1927,
P. A. Readio, 2 males, 3 females; Valentine, July 13, 1927, R. H.
Beamer, 3 males, 2 females; Palo Pinto Co., July 14, 1928, R. H.
Beamer, 1 female; Sutton Co., July 20, 1928, J. G. Shaw, 4 males,
1 female; Sutton Co., Aug. 20, 1928, A. M. James, 3 males, 11 fe-
males; Sutton Co., Aug. 20, 1928, J. G. Shaw, 1 female; Kendall
Co., July 22, 1928, R. H. Beamer, 1 female; Kendall Co., July 22,
1928, J. G. Shaw, 6 males, 2 females; Jim Wells Co., July 24, 1928,
R. H. Beamer, 19 males, 20 females; Alfred, July 24, 1928, R. H.
Beamer, 5 males, 7 females; Brooks Co., July 25, 1928, R. H. Beamer,
1 female; Bee Co., July 25, 1928, R. H. Beamer, 3 females; Hidalgo

59—6490

1408 The University Science Bulletin

Co., July 28, 1928, J. G. Shaw, 3 females; Hidalgo Co., Aug. 3, 1928,
J. G. Shaw, 1 male; Hidalgo Co., July 30, 1929, R. H. Beamer, 1 male,
6 females; Hidalgo Co., Nov. 22, 1932, L. D. Tuthill, 1 male; Starr
Co., July 30, 1928, J. G. Shaw, 8 males, 15 females; Starr Co., July 5,
1938, R. I. Sailer, 13 males, 10 females; Cameron Co., Aug. 3, 1928,
J. G. Shaw, 2 females; Cameron Co., Aug. 13, 1928, A. M. James, 3
males, 3 females; Brazoria Co., Aug. 12, 1928, L. D. Beamer, 25
males, 16 females; Brown Co., Aug. 16, 1928, L. D. Beamer, 5 males,
9 females; Leon Co., July 12, 1931, Delevan, 1 male (U. of Mich.);
Falfurrias, Nov. 2, 1932, L. D. Tuthill, 34 males, 17 females; Fal-
furrias, Jan. 1, 1946, L. D. Beamer, 11 males, 10 females; Beasley,
Nov. 7, 1932, L. D. Tuthill, 1 male, 4 females; Sinton, Nov. 8,
1932, L. D. Tuthill, 5 males, 5 females; McAllen, Nov. 20, 1932,
L. D. Tuthill, 6 males, 7 females; Del Rio, 1937, H. D. Thomas,
2 males, 10 females; Peeler, June 22, 1938, D. W. Craik, 1 female;
Progress, July 1, 1938, R. I. Sailer, 1 male; Marathon, July 9, 1938,
R. I. Sailer, 1 male, 1 female.

Louisiana: Winn Co., July 14, 1918, G. R. Pilate, 8 males, 6
females, 6 nymphs (U. of Mich.); St. Tammany Co., Feb. 25, 1923,
T. H. Hubbell, 1 female (U. of Mich.); Creole, June 18, 1948, E. L.
Todd, 1 male, 2 females.

Mississippi: Vicksburg, July 19, 1921, C. J. Drake, 1 male, 1 fe-
male; Woodville, July 26, 1921, C. J. Drake, 1 male, 1 female.

Alabama: Crawford, July 24, 1930, Paul W. Oman, 2 males, 3
females.

Georgia: Baker Co., Feb. 12, 1928, C. H. Martin, 8 males, 8 fe-
males; Baker Co., Dec. 23, 1946, L. W. Morgan, 41 males, 39 fe-
males; Okefenokee Swp., July 30, 1934, P. M. McKinstry, 1 male,
18 females; Okefenokee Swp., Aug. 3, 1934, M. E. Griffith, 49 males,
52 females; Okefenokee Swp., Aug. 3, 1934, R. H. Beamer, Jr., 3
males, 10 females; Okefenokee Swp., Aug. 3, 1934, R. H. Beamer,
22 males, 20 females; Okefenokee Swp., July 25-27, 1939, R. H. and
J. D. Beamer, 6 males, 20 females; Newton, Mar. 19, 1947, R. H.
Beamer, 5 males, 12 females.

Florida: Cocoanut Grove, Aug. 9, 1930, R. H. Beamer and P. W.
Oman, 35 males, 30 females; Ft. Meade, Aug. 13, 1930, R. H. Beamer,
8 males, 14 females; Wakulla Sprs. July 14, 1934, R. H. Beamer, 3
males, 7 females; Hilliard, July 28, 1934, R. H. Beamer, 1 male;
Sanford, Aug. 8, 1939, J. D. Beamer, 2 males, 3 females; L. Mate-
cumba Key, Mar. 14, 1947, R. H. Beamer, 8 males, 6 females.

MEXICO: Tamaulipas: San Jose, Apr. 1910, 6 males, 2 females;
Victoria, Nov. 5, 1936, H. D. Thomas, 2 males, 15 females.

A Revision of the Genus Buenoa 1409

Mexico: Mexico, July 11, 1938, L. J. Lipovsky, 2 males, 1 female.

WEST INDIES: Cuba: Havana, Jan. 25, 1932, P. J. Bermudez,
2 females.

Jamaica: Baron Hill Trelawny, Feb,, 1928, L. G. Perkins, 5 males,
20 females; Baron Hill Trelawny, Dec. 24, 1928, L. G. Perkins, 1
female; Claremont, Baron Hill Trelawny, Mar. 4, 1928, L. G. Perkins,
1 female.

Puerto Rico: Cabo Rojo, June 9, 1937, J. A. Ramos, 4 males, 2
females.

All specimens listed above are in the Francis Huntington Snow
Entomological Collections, University of Kansas, Lawrence, Kansas,
unless otherwise indicated.

Buenoa uhleri n. sp.
(PI. CIII, fig. 31; pi. CVIII, fig. 53)

Size: This species varies considerably in size within the same
population. Male, length 6.50 mm. to 7.67 mm., greatest body
width 1.82 mm. to 2.27 mm.; female, length 6.95 mm. to 8.19 mm.,
greatest body width 1.88 mm. to 2.47 mm.

Color: General facies sordid white to gray. Head, pronotum,
most of thoracic venter, and limbs sordid white to pale testaceous.
Scutellum usually orange with an irregular area of black at base;
metathoracic dorsum black with lateral areas yellowish white. Ab-
dominal venter light brown to black except keel and portions of
connexivum, yellowish white; abdominal dorsum varies from black
to mostly yellowish white with small light brown to black areas.
Some specimens entirely sordid white to pale testaceous except
most of abdominal venter and portions of abdominal dorsum, light
brown to black. This species variable in color.

1410 The University Science Bulletin

width; disk with two elongate depressions toward the middle and
a large subtriangular depression on each side, thus appearing dis-
tinctly tricarinate; lateral margins slightly divergent; posterior
margin convex, medianly concave. Scutellum with median length
distinctly less than that of pronotum. Fore femur (pi. CVIII, fig.
53a) neither wide nor greatly thickened at apex; oblong stridula-
tory area consisting of approximately nineteen to twenty-four
sclerotized ridges. Fore tibia (pi. CVIII, fig. 53a) with stridula-
tory comb (pi. CVIII, fig. 53c) consisting of approximately thirty-
five to thirty-eight teeth; apical teeth narrower and slightly taller
than basal. Chaetotaxy of male front leg as shown on Plate CVIII.
Male genital claspers normal. Spine from caudo-sinistral margin
of seventh abdominal tergite normal, tapering gradually from broad
base to acuminate apex.

Female Structural Characteristics: As viewed from above, out-
line of head rounded with anterior margin of vertex continuous
with that of eyes, occasionally vertex slightly protuberant; greatest
width of head approximately five times the anterior width of vertex
and distinctly less than humeral width of pronotum; synthlipsis
slightly less than half the anterior width of vertex; along median
longitudinal axis, head is approximately one third the length of
pronotum; notocephalon sulcate; tylus slightly inflated. Pronotum
with its median length approximately three fifths its humeral width;
disk usually with two shallow, elongate depressions toward the
middle and a shallow, subtriangular depression on each side, thus
appearing faintly tricarinate, occasionally with median carina only;
lateral margins divergent; posterior margin convex, medianly trun-
cate to slightly concave. Scutellum with median length less than
that of pronotum. Female ovipositor (pi. CIII, fig. 31) of normal
shape with teeth arranged in two longitudinal rows which merge
proximally; one inner row of few, large teeth and one long, outer
row of smaller teeth; approximately two small, lateral, toothlike
setae near apex.

Comparative Notes: Superficially this species somewhat resem-
bles B. margaritacea Torre-Bueno. Examination of the male, how-
ever, will show distinct differences. This species differs from
B. margaritacea in having the fore femur wider and more robust,
and in the form of the pronotum, rostral prong, femoral stridulatory
area, and tibial comb. Btienoa uhleri is usually larger and more
robust than B. margaritacea.

Nomenclatorial Notes: This species was first recognized as new
by Mr. C. O. Bare who labeled a series as types and paratypes

A Revision of the Genus Buenoa 1411

using the manuscript name B. uhleri. As such paratypes may have
been widely distributed, it seems desirable to point out that the
name was not validated by publication. However, to avoid con-
fusion, the name suggested by C. O. Bare has been retained by
this author.

Location of Types: Holotype male, allotype female, 27 male and
33 female paratypes, Michoacan, Mexico, Sept. 1-8, 1938, H. D.
Thomas and L. J. Lipovsky, in the Francis Huntington Snow Ento-
mological Collections, University of Kansas; other paratypes: 1
male and 1 female, Penon, Mexico, D. F., Oct. 27, 1898, P. R. Uhler
Collection, in the U. S. National Museum.

Data on Distribution: Known only from the United States and
Mexico. In addition to type series, specimens from the following
localities have been examined:

U.S.A.: California: Lagoon, Lake Reade, Aug. 17, 1925, J. G.
Needham, 1 male, 1 female.

Texas: Sanderson, Sept., 1937, H. D. Thomas, 18 males, 18 fe-
males.

MEXICO: Durango: Durango City, May 30, 1937, Meldon
Embury, 1 female.

Tamaulipas: Ciudad Victoria, Nov. 5, 1936, H. D. Thomas, 1 fe-
male.

Zacatecas: Los Potosi, Aug. 8, 1944, Henry Thomas, 9 males, 53
females.

San Luis Potosi: Cerritos, June 2, 1930, Creaser-Gordon, 1 male,
3 females (U. of Mich.); San Luis Potosi, Aug. 7-8, 1944, H. D.
Thomas, 7 males, 6 females.

Aguascalientes: 5 mi. S. Aguascalientes, July 16, 1934, Smith and
Dunkle, 2 males, 4 females; Aguascalientes, Aug. 9, 1944, Henry
Thomas, 2 males, 30 females.

Jalisco: Jalisco, Sept. 14, 1938, H. D. Thomas, 11 males, 11 fe-
males, Tecolotan, Sept. 5-17, 1938, H. D. Thomas, 2 males, 6 females.

Guanajuato: 10 mi. N. E. Leon, Aug. 17, 1932, Hobart Smith,
1 male, 32 females.

Veracruz: Bilimek, 1883, 13 females ( Berlin Mus. ) .

Hidalgo: Agua Fria, Aug. 27, 1944, Henry Thomas, 14 males, 16
females; Real del Monte, Sept. 23, 1938, H. D. Thomas, 2 males.

Michoacan: Patzcuaro, Aug. 31, 1938, H. D. Thomas, 3 males, 4
females; Zacapu, Sept. 1, 1938, H. D. Thomas and L. J. Lipovsky, 1
male, 45 females; Morelia, Sept. 3-4, 1938, H. D. Thomas, 27 males,
57 females; 10 mi. down Chinapa road, Sept. 5, 1938, H. D. Thomas,

1412 The University Science Bulletin

1 male, 6 females; Carapan, Sept. 2-8, 1938, H. D. Thomas, 4 males,
80 females; L. Cuitzeo, July 7, 1947, T. H. Hubbell, 8 males, 9 fe-
males.

Federal District: Mexico, Apr. 22-25, 1910, Bueno Collection,
3 males; Lago de Chapultepec, 1933, L. Ancona H., 1 male, 2 fe-
males; Xochimilco, June 21, 1934, H. Hinton, 5 females; Lake Tex-
coco, July 26, 1937, 5 females; Mexico, Sept. 17, 1938, 1 male, 1 fe-
male; Mexico, A. Dampf, 1 male, 1 female.

Puebla: Rio Frio, July 26, 1932, Hobart Smith, 1 female; Tehua-
can, July 18-25, 1937, H. D. Thomas, 2 males, 4 females; Tehuacan,
Aug. 5-15, 1937, H. D. Thomas, 1 male, 2 females; Cacaloapan,
July 22, 1937, H. D. Thomas, 14 males, 40 females; Puebla, July 25,
1937, H. D. Thomas, 3 males, 2 females; Puebla, Aug. 16, 1937, H. D.
Thomas, 2 males, 7 females.

Guerrero: Petaquillas, Oct. 21, 1936, H. D. Thomas, 1 male, 3
females; Salto de Valadez, Oct. 30, 1936, H. D. Thomas, 10 males,
10 females.

Oaxaca: Posita, Aug. 24, 1937, H. D. Thomas, 4 females.

Chiapas: Hda. La Libertad, Sept. 1, 1937, H. D. Thomas, 1 male,
3 females; Tuxtla Gutierrez, Aug. 27, 1939, H. D. Thomas, 1 male, 1
female; San Vicente, Jan. 4, 1938, Octavio Utrilla L., 1 male, 3 fe-
males.

All specimens listed above are in the Francis Huntington Snow
Entomological Collections, University of Kansas, Lawrence, Kansas,
unless otherwise indicated.

Buenoa albida (Champion)
(PI. CIX, fig. 54)

1901 Anisops olbidus Champion, G. C. Biologia Centrali Americana, Heter-

optera, vol. II, pp. 371 and 373, pi. 22, fig. 14.
1904. Buenoa albida, Kirkaldy, G. W. Wiener Ent. Zeit., vol. XXIII, pp. 121

and 134 (listed and states "wahrseheinlich mit A. platycnetms identiseh )
1909 Buenoa albida, Kirkaldy, G. W. and Torre-Bueno, J. R. de la. Proc.

Ent. Soc. Washington, vol. X, p. 200 (catalogue).
1909. Buenoa albida, Torre-Bueno, J. R. de la. Jr. New York Ent. Soc, vol.

1916. Buenoa albida, Van Duzee, E. P. New York Ent. Soc, p. 51 (check

1917. Buenoa albida, Van Duzee, E. P. Cat. Hemiptera America North of
Mexico, p. 454 (catalogue).

1919. Buenoa albida, Hungerford, H. B. Univ. Kansas Sci. Bull., vol. XI,

pp. 174-175 (key and description). #

1923. Buenoa albida, Torre-Bueno, J. R. de la. Univ. of Iowa Studies in Nat.

19351S Buenoa albida, Bare, C. O. Ent. News, vol. XXXVI, No. 8, p. 228 (key).
1939 Buenoa albida, Millspaugh, D. D. Field and Laboratory, vol. VII, No. 2,

p. 78.

Size: Male, length 5.85 mm. to 6.04 mm., greatest body width

A Revision of the Genus Buenoa 1413

1.69 mm. to 1.75 mm.; female, length 5.85 mm. to 6.50 mm., greatest
body width 1.62 mm. to 1.82 mm.

Color: General facies sordid white. Head, thoracic venter, and
limbs pale testaceous. Scutellum usually rufo-testaceous. Ab-
dominal venter black with keel and connexivum pale testaceous;
abdominal dorsum testaceous with transverse black bands or com-
pletely black with last two segments pale testaceous. Some speci-
mens entirely sordid white.

Male Structural Characteristics: As viewed from above, outline
of head rounded with vertex slightly indented; greatest width of
head more than six times the anterior width of vertex and less than
humeral width of pronotum; synthlipsis approximately half the
anterior width of vertex; along median longitudinal axis, head is
approximately half the length of pronotum; tylus slightly inflated,
with wide median depression forming two short, lateral carinae;
labrum short, basal width twice its median length with apex bluntly
rounded; rostral prong (pi. CIX, fig. 54b) longer than third rostral
segment, with base originating laterally at distal end of third rostral
segment, and with apex sharply rounded. Pronotum with its
median length slightly less than two thirds its humeral width; disk
with two elongate depressions toward the middle and a large sub-
triangular depression on each side, thus appearing tricarinate;
lateral margins divergent; posterior margin convex, medianly con-
cave. Scutellum large, with median length slightly greater than
that of pronotum. Fore femur (pi. CIX, fig. 54a) neither wide nor
greatly thickened at apex; oblong stridulatory area consisting of
approximately thirty-five to forty sclerotized ridges. Fore tibia
(pi. CIX, fig. 54a) with stridulatory comb (pi. CIX, fig. 54c) con-
sisting of approximately thirty-four to thirty-six teeth which increase
slightly in height and thickness from base to apex. Chaetotaxy of
male front leg as shown on Plate CIX. Male genital claspers nor-
mal. Spine from caudo-sinistral margin of seventh abdominal
tergite normal, tapering gradually from broad base to acuminate
apex.

Female Structural Characteristics: As viewed from above, out-
line of head rounded; greatest width of head approximately five
times the anterior width of vertex and less than humeral width of
pronotum; synthlipsis less than half the anterior width of vertex;
along median longitudinal axis, head is less than half the length of
pronotum; tylus very slightly inflated and without median longi-
tudinal depression. Pronotum with its median length less than two
thirds its humeral width; disk only slightly impressed and occa-

1414 The University Science Bulletin

sionally not at all; lateral margins divergent; posterior margin con-
vex, medianly concave. Scutellum large with median length greater
than that of pronotum. Female ovipositor of normal shape with
teeth arranged in two longitudinal rows; one inner row of larger
teeth and one long outer row of smaller teeth; approximately four
very small, lateral, toothlike setae near apex.

Comparative Notes: Superficially this species closely resembles
B. scimitra Bare. Examination of the male, however, will show
distinct differences. This species differs from B. scimitra in the
shape of the rostral prong, in having the tylus medianly depressed,
and in the shape of the stridulatory area on the fore femur.

Location of Types: The original type series is located at the
British Museum, London. Holotype male now on slides. Homo-
type male, compared with type by Dr. W. E. China of the British
Museum, labeled "Mata Capestra, Ver., Mex., 10/1926, M. F. 1077,
received from Dr. Dampf 1932," now in the Francis Huntington
Snow Entomological Collections, University of Kansas.

Data on Distribution: Recorded from the United States, Mexico,
and Puerto Rico. Specimens from the following localities have
been examined:

MEXICO: Sonora: Salitial Rio Mayo, Feb. 23, 1935, H. S. Gen-
try, 1 male.

Sinaloa: Mazatlan, May, 1934, H. Hinton, 5 males, 8 females,
2 nymphs.

Veracruz: Mata Capestra, Oct., 1926, 1 male, 1 female.

Hidalgo: Agua Fria, Aug. 27, 1944, H. D. Thomas, 1 male.

Morelos: Cuernavaca, Oct. 5, 1936, H. D. Thomas, 1 male.

Guerrero: Salto de Valadez, Oct. 30, 1936, H. D. Thomas, 1 male,

1 female.

PUERTO RICO: Cabo Rojo, June 9, 1937, J. A. Ramos, 3 males,

2 females; Isabela, May 12, 1935, Julio Garcia Diaz, 17 males, 6
females; Rio Piedras exp. sta., May 23, 1935, Julio Garcia Diaz, 5
nymphs; Cartagena Lagoon, Aug. 10, 1935, Julio Garcia Diaz, 1
male, 6 females.

All specimens listed above are in the Francis Huntington Snow
Entomological Collections, University of Kansas, Lawrence, Kansas.

Buenoa pallens (Champion)
(PI. CII, figs. 5, 11; pi. CIX, fig. 55)

1901. Anisops pallens Champion, G. C. Biologia Centrali Americana, Heter-

optera, vol. II, p. 374.
1904. Buenoa %mllens, Kirkaldy, G. W. Wiener Ent. Zeit, vol. XXIII, pp. 121

and 134 (listed).

A Revision of the Genus Buenoa 1415

1909. Buenoa pallens, Kirkaldy, G. W. and Torre-Bueno, J. R. de la. Proc.

Ent. Soc. Washington, vol. X, p. 201 (catalogue).
1940. Buenoa pallens, Hungerl'ord, H. B. Ent. Monthly Mag., vol. LXXVI,

p. 256 ( listed and ecological note ) .

Size: This species varies considerably in size. Male, length 5.52
mm. to 6.89 mm., greatest body width 1.49 mm. to 1.56 mm.; female,
length 5.98 mm. to 7.15 mm., greatest body width 1.75 mm. to
2.01 mm.

Color: General facies pale testaceous to nigro-violaceous. In
pale specimens, head, thorax, and limbs sordid white to testaceous
with abdomen black except ventral keel, portions of connexivum,
and last one or two segments, testaceous. In dark specimens, head,
anterior portion of pronotum, most of thoracic venter, and limbs
sordid white to testaceous. Posterior portion of pronotum black;
scutellum usually entirely black, occasionally with apex testaceous;
metathoracic dorsum black. Abdomen black except ventral keel,
portions of connexivum and occasionally last one or two segments,
testaceous. This species variable in color.

Male Structural Characteristics: As viewed from above, outline
of head laterally rounded, anteriorly truncate with vertex indented;
greatest width of head approximately six times the anterior width
of vertex and less than humeral width of pronotum; synthlipsis two
fifths to three fifths the anterior width of vertex; along median
longitudinal axis, head is two fifths to three fifths the length of
pronotum; notocephalon sulcate dorsally at least; tylus slightly in-
flated; labrum with basal width not quite twice its median length
and apex bluntly rounded; rostral prong (pi. CIX, figs. 55b, 55c)
slightly variable, longer than third rostral segment, with base origi-
nating laterally midway of third rostral segment, and with apex
moderately rounded. Pronotum with its median length approxi-
mately half its humeral width; disk usually unimpressed, occasion-
ally with a shallow, subtriangular depression on each side, very
seldom appearing tricarinate; lateral margins divergent; posterior
margin convex, medianly concave. Scutellum large, with median
length distinctly greater than that of pronotum. Fore femur (pi.
CIX, fig. 55a ) neither wide nor thickened at apex; triangular to sub-
triangular stridulatory area consisting of approximately sixteen to
twenty-two sclerotized ridges. Fore tibia (pi. CIX, fig. 55a) with
stridulatory comb (pi. CIX, figs. 55e, 55f) consisting of approxi-
mately twenty-four to thirty-eight teeth; apical teeth thicker than
basal. Chaetotaxy of male front leg as shown on Plate CIX. Male
genital claspers normal. Spine from caudio-sinistral margin of

1416 The University Science Bulletin

seventh abdominal tergite (pi. CII, figs. 5, 11) small, tapering
gradually from base to strongly acuminate apex.

Female Structural Characteristics: As viewed from above, out-
line of head laterally rounded, anteriorly truncate with vertex in-
dented; greatest width of head approximately five and one half
times the anterior width of vertex and less than humeral width of
pronotum; synthlipsis approximately half the anterior width of
vertex; along median longitudinal axis, head is one third to one
half the length of pronotum; notocephalon sulcate dorsally; tylus
not inflated. Pronotum with its median length approximately half
its humeral width; disk unimpressed or occasionally with feeble
median carina; lateral margins divergent; posterior margin convex,
slightly concave medianly. Scutellum large, with median length
distinctly greater than that of pronotum. Female ovipositor of
normal shape with teeth arranged in two longitudinal rows; one
inner row of few, large teeth and one outer row of smaller teeth;
approximately three or four small and obscure, lateral, toothlike
setae near apex.

Variation Within Species: As is indicated under measurements of
length, this species varies a great deal in size, there is also some
variation in the proportional size of the pronotum and head, and in
the rostral prong and femoral stridulatory area. The greatest con-
trast is shown between a series from Costa Rica and one from
Ecuador. The former is a small form, the males seldom more than
5.70 mm. in length; the males of the latter form are approximately
6.80 mm. in length. It is the opinion of this author that these forms
all belong to one variable species.

Comparative Notes: Superficially the dark form of this species
closely resembles B. pallipes (Fabricius) and B. mutabilis n. sp.
Examination of the male, however, will show distinct differences.
This species differs from B. pallipes in having the fore femur narrow
and not thickened at apex, rostral prong shorter and originating
laterally midway of the third rostral segment, and pronotum shorter
and not distinctly tricarinate. Buenoa pollens differs from B. muta-
bilis in having the notocephalon distinctly wider, and fore femur
narrower and less thickened at apex. B. mutabilis is a smaller
species.

Location of Types: The original type series from Guatemala, is
located in the British Museum, London.

Data on Distribution: Recorded from Mexico, Guatemala, Costa
Rica, West Indies (Islands of St. Thomas, St. Croix, Guadeloupe,
Dominica, Grenada, and Trinidad), Colombia, Ecuador, Brazil,

A Revision of the Genus Buenoa 1417

Peru, and Chile. Specimens from the following localities have been
examined:

MEXICO: Colima: Colima, 1 female.

Morelos: Cuernavaca, Oct. 5-17, 1936, H. D. Thomas, 40 males,
126 females.

Oaxaca: Posita, Aug. 24, 1937, H. D. Thomas, 5 males, 6 females;
Oaxaca, Aug. 25, 1937, H. D. Thomas, 11 males, 30 females.

Chiapas: Mt. Obando, Apr. 15, 1940, H. M. Smith, 2 males, 3 fe-
males.

Yucatan: Motul, July 26, 1932, E. R. Creaser, 1 female, (U. of
Mich.); Yunca, July 29, 1932, E. P. Creaser, 1 male, 1 female, (U.
of Mich.).

GUATEMALA: Peten, San Andres Lake, Dec. 10, 1925, Dampf, 1
male; El Salto Escuintla, 1934, F. X. Williams, 3 males, 3 females.

COSTA RICA: San Jose, Purchased June, 1931, Heinrich
Schmidt, 4 males, 20 females; San Jose, June and July, 1931, Hein-
rich Schmidt, 6 males, 12 females; San Jose, Purchased 1932, Hein-
rich Schmidt, 6 males, 4 females; Rio Virilla, Dec. 26, 1931, Hein-
rich Schmidt, 25 males, 14 females.

PANAMA: Tabernilla, June 20, 1907, Aug. Busck (U.S.N.M.),
1 female; Panama, Jan. 31, 1911, 1 male (U.S.N.M.); La Chorrera,
May 15, 1912, Aug. Busck, 1 female (U.S.N.M.); Sona, May, 1914,
J. Zetek, 1 female.

WEST INDIES: St. Thomas: March 11, 1925, F5029, 2 females
(A.M.N.H.).

St. Croix: Christiansted, June, 1941, H. A. Beatty, 3 males, 12
females; 1941, H. A. Beatty, 4 males, 3 females (U.S.N.M.).

Guadeloupe: A Kirkaldy remnant, 1 female.

Dominica: Laudet, June 13, 1911, 7 males, 2 females (A.M.N.H.).

Grenada: Mount Gay Est., H. H. Smith, 3 males ( U.S.N.M. ) .

Trinidad: January, Aug. Busck, 1 male (U.S.N.M.).

COLOMBIA: Cali, 3 males, 2 females (U.S.N.M. ).

ECUADOR: Bafios, March, 1936, Clarke Mclntyre, 1 male, 3
females; Barios, Runtun Lake, June, 1936, Clarke Mclntyre, 3 males,
12 females.

BRAZIL: Amazonas: Ireng R. to Roraima, Aug. 13, 1911, 1 male
(A.M.N.H.).

Minas Geraes: 1897, Fruhstorfer, 1 male (Berlin Mus.).

PERU: Dept. Cajamarca, May 26 to June 17, 1936, F. Woyt-
kowski, 117 males, 127 females; Dept. Amazones, San Ildefonso,
July 29, 1936, F. Woytkowski, 12 males, 11 females; Dept. Amazonas,

1418 The University Science Bulletin

Vic. Chachapoyas, Aug. 4-10, 1936, F. Woytkowski, 44 males, 46
females.

CHILE: Magdalena I., May 19, 1925, H. H. Keifer, 3 males, 1 fe-
male (Cal. Acad. Sci.).

All specimens listed above are in the Francis Huntington Snow
Entomological Collections, University of Kansas, Lawrence, Kansas,
unless otherwise indicated.

Buenoa pallipes (Fabricius)

(PL CIX, fig. 56)

1803. Notonecta pallipes Fabricius, J. C. Systema Rhyngotorum, p. 103.
1868. Anisops pallipes, Stal, C. Kongliga Svenska Vetenskaps-Akademien

Handlingar, vol. VII, p. 137 ( description ).
1901. Anisops pallipes, Champion, G. C. Biologia Centrali Americana, Heter-

optera, vol. II, pp. 371-372 (description and gives B. platycnemis as synonym

of A. pallipes).
1904. Buenoa pallipes, Kirkaldy, G. W. Wiener Ent. Zeit., vol. XXIII, pp. 123

and 134 (listed).
1909. Buenoa pallipes, Kirkaldy, G. W. and Torre-Bueno, J. R. de la. Proc.

Ent. Soc. Washington, vol. X, p. 201 (catalogue).
1909. Buenoa pallipes, Torre-Bueno, J. R. de la. Jr. New York Ent. Soc, vol.

XVII, p. 75 (listed).
1913. Buenoa pallipes, Perkins, R. C. L. Fauna Hawaiiensis, vol. I, p. cciii

( ecology and distributional note ) .
1913. Buenoa pallipes, Kirkaldy, G. W. Fauna Hawaiiensis, vol. II, p. 555

( listed ) .
1939. Buenoa pallipes, Barber, H. G. New York Acad. Sci., vol. XIV, p. 421.
1939. Buenoa pallipes, Hungerford, H. B. Ann. Ent. Soc. America, vol.

XXXII, p. 588 (recorded from Costa Rica).
1944. Buenoa pallipes, Williams, F. X. Proc. Hawaiien Ent. Soc, vol. XII,

pp. 193-194 ( biology note ) .
1948. Buenoa pallipes, Zimmerman, E. C. Insects of Hawaii, vol. Ill, pp. 232-

233 (biology note).

Size: Male, length 5.52 mm. to 6.22 mm., greatest body width
1.56 mm. to 1.95 mm.; female, length 5.62 mm. to 6.50 mm., greatest
body width 1.69 mm. to 1.95 mm.

Color: General facies sordid white to black. Head, pronotum,
thoracic venter, and limbs sordid white to testaceous; pronotum
occasionally black with anterior portion testaceous and carinae
rufescent. Scutellum usually black or fuscous, with apex more
or less testaceous; metathoracic dorsum testaceous to black. Ab-
domen usually black except ventral keel, portions of connexivum,
and terminal segment, testaceous. Pale specimens entirely sordid
white to testaceous except abdominal venter, black. This species
variable in color.

Male Structural Characteristics: As viewed from above, outline
of head laterally rounded, anteriorly truncate with vertex slightly
indented, strongly indented just above tylus; greatest width of head
six to six and one half times the anterior width of vertex and less

A Revision of the Genus Buenoa 1419

than humeral width of pronotum; synthlipsis quite narrow but
approximately half the anterior width of vertex; along median
longitudinal axis, head is slightly less than one third the length of
pronotum; notocephalon sulcate dorsally; tylus inflated; labrum
with basal width distinctly greater than its median length and apex
bluntly rounded; rostral prong (pi. CIX, fig. 56b) distinctly longer
than third rostral segment, with base originating laterally near distal
end of third rostral segment, and with apex moderately rounded.
Pronotum with its median length approximately two-thirds its
humeral width; disk with two elongate depressions toward the
middle and a large, subtriangular depression on each side, thus
appearing distinctly tricarinate; lateral margins slightly divergent;
posterior margin convex, medianly concave. Scutellum with median
length distinctly less than that of pronotum. Fore femur (pi. CIX,
fig. 56a) wide and somewhat thickened at apex; oblong to sub-
triangular stridulatory area consisting of approximately seventeen
sclerotized ridges. Fore tibia (pi. CIX, fig. 56a) with narrow
stridulatory comb (pi. CIX, fig. 56c) consisting of approximately
thirty-four teeth, apical teeth thicker than basal. Chaetotaxy of
male front leg as shown on Plate CIX. Male genital claspers nor-
mal. Spine from caudo-sinistral margin of seventh abdominal
tergite tapering gradually from base to strongly acuminate apex.

Female Structural Characterisitics: As viewed from above, out-
line of head laterally rounded, anteriorly truncate with vertex
slightly indented, strongly indented just above tylus; greatest width
of head approximately six times the anterior width of vertex and
less than humeral width of pronotum; synthlipsis approximately
half the anterior width of vertex; along median longitudinal axis,
head is one fourth to one third the length of pronotum; noto-
cephalon slighty sulcate; tylus usually not inflated. Pronotum with
its median length approximately half its humeral width; disk usu-
ally unimpressed, occasionally with a faint median carina; lateral
margins divergent; posterior margin convex, medianly concave.
Scutellum large, with median length distinctly greater than that
of pronotum. Female ovipositor of normal shape with teeth ar-
ranged in two irregular, longitudinal rows intermingling medianly;
one inner row of large teeth and one outer row of smaller teeth;
approximately six or seven small, lateral, toothlike setae near apex.

Variation Within Species: As is indicated under measurements
of length, this species varies somewhat in size; there is also some
variation in the proportional size of the pronotum.

Comparative Notes: Superficially this species closely resembles

1420 The University Science Bulletin

B. platycnemis (Fieber) and B. pollens (Champion). Examina-
tion of the male, however, will show distinct differences. This
species differs from B. pollens in having the fore femur wide and
thickened at apex, and the pronotum distinctly longer and tri-
carinate. Buenoa pallipes differs from B. platycnemis in having
the pronotum longer with posterior margin more concave medianly,
the frons narrower, and slight differences in the fore femur and
rostral prong. Buenoa platycnemis is less robust.

Location of Types: The type is located at the Museum of Lund,
Sweden.

Data on Distribution: Recorded from Hawaii (fide Zimmerman
and Perkins), Mexico, Honduras, Costa Rica, Panama, West Indies
(Jamaica, Puerto Rico, St. Thomas, Guadeloupe, St. Vincent),
Colombia, Peru, and Paraguay. Specimens from the following
localities have been examined:

MEXICO: Oaxaca: Papaloapan, Mar. 4, 1939, M. & E. Gordon,

1 male, 2 females ( U. of Mich. ) .

Chiapas: San Vicente, Jan. 4, 1938, Octavio Utrilla L., 1 male,

2 females.

HONDURAS: Tela, Apr. 1, 1923, T. H. Hubbell, 4 males, 1 fe-
male (U. of Mich.); Tela, March 1-15, 1936, John Deal, 76 males,
93 females.

COSTA RICA: San Isidro del Gen., Feb., 1939, Dean L. Rounds,
1 male, 1 female.

WEST INDIES: Jamaica: Montego Bay, Mar. 11, 1911, 1 male
(A.M.N. H.); Lumsden Tydenham, St. Ann, Feb., 1928, L. G.
Perldns, 3 males, 7 females; Baron Hill Trelawny, Feb., 1928, L. G.
Perkins, 2 males, 2 females; Claremont, Feb., 1928, L. G. Perkins,
1 male, 1 female; Bath St. Thomas, Mar. 29, 1937, Chester Roys,
8 males, 19 females; St. Andrew, Dec. 3, 1946, G. B. Thompson,

3 males, 2 females; St. Andrew, Apr. 15-16, 1947, G. B. Thompson,
1 male, 1 female, 1 nymph.

Puerto Bico: Coamo Springs, July 17-19, 1914, 3 males (A.M.
N.H.).

St. Thomas: Sulphur River, Apr. 3, 1937, Chester Roys, 15 males,
40 females.

Guadeloupe: St. Anne, Louis Mesmin, 1 male, 1 female.

COLOMBIA: Cali, 1 male, 1 female (U. S. N. M.).

PERU: Dept. Amazonas, Vic. Guayabamba, Aug. 14-19, 1936,
F. Woytkowski, 67 males, 70 females.

PARAGUAY: Villarrica, Dec. 6, 1923, Fran. Schade, 4 males;
Villarrica, Oct. 9, 1924, Fran. Schade, 1 male; Villarrica, Dec. 16,

A Revision of the Genus Buenoa 1421

1924, Fran. Schade, 19 males; Villarrica, Nov. 20, 1929, Fran. Schade,
1 male, 1 female; Caraveni, June 15, 1924, Fran. Schade, 2 females;
Estero Grande, Nov. 1, 1924, Fran. Schade, 3 males; Melinesque,
Dept. Caruga, Dec, 1925, Fran. Schade, 3 males; Melinesque, June
28, 1935, Fran. Schade, 3 males, 2 females.

All specimens listed above are in the Francis Huntington Snow
Entomological Collections, University of Kansas, Lawrence, Kansas,
unless otherwise indicated.

Buenoa platycnemis (Fieber)
(PI. X, fig. 57)

1851. Anisops platycnemis Fieber, F. X. Abhandlungen Kongl. Bohmischen

Gesellschaft Wissenschaften, vol. VII, Series 5, p. 485.
1899. Anisops platycnemis, Kirkaldy, G. W. The Entomologist, vol. XXXII,

p. 30.
1901. Anisops platycnemis, Champion, G. C. Biologia Centrali Americana,

Heteroptera, vol. II, pp. 371-372 (gives A. platycnemis as synonym of B.

pallipes ) .

1904. Buenoa platycnemis, Kirkaldy, G. W. Wiener Ent. Zeit., vol. XXIII,
p. 134 (listed and description of B. margaritacea under name B. platy-
cnemis).

1905. ? Buenoa platycnemis, Snow, F. H. Trans. Kansas Acad. Sci., vol. XX,
p. 153 (recorded from Texas).

1908. Buenoa platycnemis, Torre-Bueno, J. B. de la. Jr. New York Ent. Soc,
vol. XVI, p. 238 (listed).

1909. Buenoa platycnemis, Torre-Bueno, J. R. de la. Jr. New York Ent. Soc,
vol. XVII, pp. 75-77 (key, notes, and synonomy).

1914. Buenoa platycnemis, Van Duzee, E. P. Trans. San Diego Soc. Nat.
Hist, vol. II, p. 33 (listed).

1916. Buenoa platycnemis, Van Duzee, E. P. New York Ent. Soc, p. 51
(check list).

1917. Buenoa platycnemis, Van Duzee, E. P. Cat. Hemiptera America North
of Mexico, p. 455 (catalogue).

1917. Buenoa platycnemis, Hungerford, H. B. Ent. News, vol. XXVIII, p. 176

(key).
1919. Buenoa platycnemis, Hungerford, H. B. Univ. Kansas Sci. Bull., vol.

XI, pp. 174 and 176 (description, key, and notes).
1923. Buenoa platycnemis, Torre-Bueno, J. R. de la. Connecticut State Geol.

and Nat. Hist. Survey Bull., No. 34, p. 407 (key and notes).

1923. Buenoa platycnemis, Hungerford, H. B. Ent. News, vol. XXXIV, p. 151
( note ) .

1924. Buenoa platycnemis, Hungerford, H. B. Ann. Ent. Soc. America, vol.
XVII, p. 225 (note on mistaken identification).

1925. Buenoa platycnemis, Bare, C. O. Ent. News, vol. XXXVI, p. 228 (dis-
tributional note).

1925. Buenoa platycnemis, Hungerford, H. B. and Beamer, R. H. Ent. News,

vol. XXXVI, p. 297 (note).
1928. Buenoa platycnemis, Bare, C. O. Univ. Kansas Sci. Bull., vol. XVIII,

p. 268 (note).
1928. ? Buenoa platycnemis, Torre-Bueno, J. R. de la. Cornell Univ. Agr.

Experiment Station, Memoir 101, p. 139 (listed).
1939. ? Buenoa platycnemis, Millspaugh, D. D. Field and Laboratory, vol.

VII, p. 78.

Size: This species varies considerably in size. Male, length
4.55 mm. to 5.35 mm., greatest body width 1.36 mm. to 1.62 mm.;

1422 The University Science Bulletin

female, length 5.00 mm. to 5.43 mm., greatest body width 1.49 mm.
to 1.75 mm.

Color: General facies sordid white to black. Head, pronotum,
thoracic venter, and limbs sordid white to testaceous. Scutellum
sordid white to testaceous with base brown to black; metanotum
usually brown to black with portions testaceous. Abdominal dor-
sum usually brown to black with portions testaceous; abdominal
venter black except keel, portions of connexivum, and last one or
two segments, testaceous. Pale specimens entirely sordid white
except most of abdomen, black. This species variable in color.

Male Structural Characteristics: As viewed from above, outline
of head laterally rounded, anteriorly truncate with vertex usually
slightly indented; greatest width of head five and one half to six
times the anterior width of vertex and less than humeral width of
pronotum; synthlipsis slightly less than half the anterior width of
vertex; along median longitudinal axis, head is approximately two
fifths the length of pronotum; notocephalon slightly sulcate; tylus
slightly inflated; labrum with basal width not quite twice its median
length and apex bluntly rounded; rostral prong (pi. CX, fig. 57b)
long, much longer than third rostral segment, with base originating
laterally and protruding anteriorly at distal end of third rostral
segment, and with apex moderately rounded. Pronotum with its
median length approximately two thirds its humeral width; disk
with two elongate depressions toward the middle and a large sub-
triangular depression on each side, thus appearing tricarinate; lat-
eral margins divergent; posterior margin convex, medianly truncate.
Scutellum with median length less than that of pronotum. Fore
femur (pi. CX, fig. 57a) wide and somewhat thickened at apex;
oblong to subtriangular stridulatory area consisting of approxi-
mately eleven to fourteen sclerotized ridges. Fore tibia (pi. CX,
fig. 57a) wide with stridulatory comb (pi. CX, fig. 57c) consisting
of approximately thirty to thirty-five teeth; apical teeth thicker and
narrower than basal. Chaetotaxy of male front leg as shown on
Plate CX. Male genital claspers normal. Spine from caudo-sinis-
tral margin of seventh abdominal tergite with apical one third very
narrow and apex strongly acuminate.

Female Structural Characteristics: As viewed from above, out-
line of head laterally rounded, anteriorly truncate with vertex often
indented at lateral margins; greatest width of head five to five
and one half times the anterior width of vertex and less than
humeral width of pronotum; synthlipsis slightly less than half the

A Revision of the Genus Buenoa 1423

anterior width of vertex; along median longitudinal axis, head is
one third to one half the length of pronotum; notocephalon sulcate;
tylus slightly inflated. Pronotum with its median length approxi-
mately half its humeral width; disk unimpressed, occasionally with
faint median carina; lateral margins divergent; posterior margin
convex, usually medianly truncate. Scutellum large, with median
length greater than that of pronotum. Female ovipositor of normal
shape with teeth arranged in two longitudinal rows; one inner row
of large teeth and one outer row of small teeth; approximately
three small, lateral, toothlike setae near apex.

Variation Within Species: As is indicated under measurements of
length, this species varies considerably in size; there is also some
variation in the proportional size of head and pronotum.

Comparative Notes: Superficially this species closely resembles
B. paJlipes (Fabricius) and B. 7iitida n. sp. Examination of the
male, however, will show distinct differences. This species differs
from B. pallipcs in having the pronotum shorter with posterior
margin more truncate medianly, the frons wider, and slight differ-
ences in the fore femur and rostral prong. Buenoa platycnemis
differs from B. nitida in having a tricarinate pronotum, more sclero-
tized ridges in the femoral stridulatory area, and in the form of the
rostral prong, fore femur, and tibial comb. Buenoa platycnemis
is smaller and less robust than the above mentioned species.

Nomenclatorial Notes: One finds in the literature and collections,
many species masquerading under the name Buenoa platycnemis.
This situation is primarily the result of misidentifications by several
distinguished hemipterists. The confusion surrounding this species
serves to illustrate the necessity for the worker to have access to
type material.

Location of Types: The type, a male, labeled "Portorico St.
Thomas Moritz", is located at the Berlin Museum.

Data on Distribution: Many United States records for this species
have been erroneously cited in the literature. Buenoa platycnemis
is, for the most part, a Neotropical species. From the study of a
vast amount of material, this author finds that for the United States,
only Texas and Florida are represented in its distribution. Recorded
also from Mexico, Panama, Canal Zone, Costa Rica, West Indies
(Cuba, Grand Cayman, Haiti, Jamaica, Mona, Puerto Rico, St.
Thomas, St. Croix, and Martinique), Pearl Island, Darien, Colom-
bia, Venezuela, Brazil, and Peru. Specimens from the following
localities have been examined:

60—6490

1424 The University Science Bulletin

U. S. A.: Texas: McAllen, Nov. 20, 1932, L. D. Tuthill, 6 males,
4 females; Brownsville, June 29, 1938, R. I. Sailer, 1 male; Progress,
July 1, 1938, R. I. Sailer, 7 males, 5 females; Star Co., July 5, 1938,
R. I. Sailer, 3 males, 4 females; Falfurrias, Jan. 1, 1946, L. D. Beamer,

2 males, 2 females.

Florida: L. Matecumba Key, Mar. 14, 1947, R. H. Beamer and
L. D. Beamer, 5 males, 6 females.

MEXICO: Sonora: Rio Mayo, Arroyo de los Mescales, Feb. 16,
1935, H. S. Gentry, 8 males, 19 females.

Jalisco: Guadalajara, Sept. 13, 1938, H. D. Thomas, 1 female;
15 mi. down Autlan Rd., Sept. 14, 1938, H. D. Thomas, 2 males,
1 female; 15 mi. S. W. Lake Chapala, Sept. 14, 1938, H. D. Thomas,

1 male.

Veracruz: Carrizal, Aug. 6, 1932, A. Dampf, 1 female; Minatitlan,
Sept. 22, 1936, H. D. Thomas, 7 females.

Michoacdn: El Sabino Uruapan, July 30, 1936, H. D. Thomas,
19 males, 19 females; Zamora, Sept. 8, 1938, H. D. Thomas, 1 male;
L. Cuitzeo, July 7, 1947, T. H. Hubbell, 2 males, 3 females.

Federal District: Xochimilco, June 21, 1934, H. Hinton, 1 male,

3 females; Mexico City, July 7, 1937, H. D. Thomas, 6 males, 16
females; Mexico City, 1937, H. D. Thomas, 6 males, 2 females.

Morelos: Cuernavaca, Oct. 5, 1936, H. D. Thomas, 4 males, 3
females.

Guerrero: Iguala, Oct. 7, 1936, H. D. Thomas, 8 males, 14 fe-
males; Palo Blanco, Oct. 10, 1936, H. D. Thomas, 1 male, 2 females;
Tierra Colo., Oct. 31, 1936, H. D. Thomas, 4 males, 3 females; Rio
Agua, Oct. 31, 1936, H. D. Thomas, 1 male; Acapulco, Nov. 1, 1936,
H. D. Thomas, 6 males, 19 females.

Chiapas: Huixtla, Nov. 9, 1932, A. Dampf, 1 female; Suchiate,
Nov. 16-17, 1932, A. Dampf, 1 male, 3 females.

Campeche: Ciudad del Carmen, Sept. 18, 1936, H. D. Thomas,
6 males, 13 females; Hda. Encarnation, Oct. 15, 1936, H. D. Thomas,
6 males, 5 females.

Yucatan: Chichen-Itza, June 6-27, 1932, E. P. Creaser, 1 male,

2 females, 2 nymphs (U. of Mich.); Chichen-Itza, Aug. 29, 1936,
H. D. Thomas, 2 males; Piste, June 22, 1932, E. P. Creaser, 2 fe-
males (U. of Mich.); Merida, Jalal Aguada, July 22, 1932, E. P.
Creaser, 3 males, 5 females, 1 nymph (U. of Mich.); Merida, July
28, 1932, E. P. Creaser, 10 males, 5 females, 2 nymphs (U. of Mich.).

PANAMA: Old Panama, Jan. 31, 1911, Aug. Busck, 3 females
(U. S. N. M.); Sona, May, 1914, J. Zetek, 3 males, 7 females (U. S.
N. M.); Cano Saddle, Gatun L., Aug. 6, 1923, R. C. Shannon, 1 male

A Revision of the Genus Buenoa 1425

(U. S. N. M.); Las Palmas, Dec. 21, 1944, A. W. Lindquist, 1 male,
2 females (U. S.N. M.); San Miguel, 1 male, 2 females (U.S.
N.M.).

CANAL ZONE: Ft. Clayton, 1933, R. F. Edwards, 5 males, 2 fe-
males.

COSTA RICA: San Isidro del Gen., Feb., 1939, Dean L. Rounds,
4 males, 1 female.

WEST INDIES: Cuba: Soledad, Feb. 14, 1925, J. G. Myers, 2
males, 5 females; Habana Bot. Garden, Jan. 25, 1932, P. J. Bermudez,
6 males, 8 females; Habana, Casa Blanca, Dec. 20, 1933, P. J. Ber-
mudez, 7 males, 11 females; Habana, 1933, P. J. Bermudez, 31 males,
14 females; Havana Prov., Catalina, Nov. 27, 1933, P. J. Bermudez,
1 female; Matanzas, Yumuri Valley, Dec. 9, 1933, P. J. Bermudez,

1 male; P. R. Uhler Collection, 1 male (U. S. N. M.).

Grand Cayman: Cow well near Pedro Castle, Oxford U. Bio.
Exp., Aug. 4, 1938, Lewis and Thompson, 11 males, 11 females.

Haiti: Attelye, Oct. 22, 1925, W. A. Hoffman, 1 male, 1 female
(U. S. N. M.); Jacmel, W. A. Hoffman, 2 females (U. S. N. M.).

Jamaica: Palm Beach, Montego Bay, March 11, 1911, 6 males,

2 females (A. M. N. H.); Montego Bay, Mar. 15, 1911, 2 males
(A. M.N.H.).

Mona: Feb. 21-26, 1914, 1 male (A. M. N. H.).

Puerto Rico: Culebra, Feb., 1899, Aug. Busck, 2 males ( U. S.
N. M.); Coamo Springs, July 17-19, 1914, 2 males, 2 females
(A. M. N. H.); Mayaguez, Oct. 16, 1930, J. Landron, 1 male (U. S.
N. M.); Luquillo, July 7-8, 1932, J. Blanch, 2 males, 1 female;
Luquillo, July 9, 1932, J. Blanch, 1 male, 1 female (U.S.N.M.);
Iuebrada, Feb. 21, 1935, J. G. Diaz, 1 male, 2 females; Florida Road,
Feb. 28, 1935, J. G. Diaz, 5 males, 10 females; Almirante Rd.,
March 9, 1935, J. G. Diaz, 5 males, 8 females; Tortuguera L., Mar.
20, 1935, J. G. Diaz, 3 males, 8 females; Near Isabela, May 12, 1935,
J. G. Diaz, 7 males, 2 females; Exp. Sta. Rio Piedras, May 23, 1935,
J. G. Diaz, 4 females; Cartagena Lagoon, Aug. 10, 1935, J. G. Diaz,
4 females; Luquillo Mts., Nov. 18, 1935, J. G. Diaz, 4 males, 8 fe-
males.

St. Thomas: Charlotte Amalie, June 2, 1917, H. Morrison, 2
males, 3 females (U. S. N. M.); St. Thomas, Mar. 11, 1925, 1 male
(A.M.N.H.); St. Thomas, May 15, 1937, Chester Roys, 3 males,

3 females; St. Thomas, Feb., Aug. Busck, 1 female (U. S. N. M.);
St. Thomas, Klug, 2 males ( Leiden Mus. ) .

St. Croix: St. Croix, Apr. 4, 1925, F 5145C and F 5022, 1 male,
2 females (A.M.N.H.); St. Croix, 1941, H. A. Beatty, 2 females

1426 The University Science Bulletin

(U. S. N. M.); St. Croix, No. 744, H. A. Beatty, 1 male, 2 females
(U.S.N.M.).

Martinique: Fort de France, June 27, 1911, 1 female (A.M.
N.H.).

COLOMBIA: Villavieja, 1944, R. A. Stirton, 22 males, 26 fe-
males; Darien, Laguna de Pita, D. Festa, 1 male (U. S. N. M.).

VENEZUELA: San Esteban, Nov. 22, 1939, Pablo J. Anduze,
4 males, 5 females.

BRAZIL: Maranhdo: Chapada, No. 2966, 2 males, 2 females.

PERU: Vic. Guayabamba, Aug. 18, 1936, F. Woytkowski, 28
males, 23 females; Vic. Rioja, Dept. San Martin, Sept. 9 to Oct. 3,
1936, F. Woytkowski, 11 males, 11 females; Satipo, Nov., 1942,
Pedro Paprzyki, 6 females.

All specimens listed above are in the Francis Huntington Snow
Entomological Collections, University of Kansas, Lawrence, Kansas,
unless otherwise indicated.

Buenoa omani n. sp.

(PL CIII, fig. 37; pi. CX, fig. 58)

Size: Male, length 5.72 mm. to 6.56 mm., greatest body width
1.62 mm. to 1.88 mm.; female, length 6.17 mm. to 6.69 mm., greatest
body width 1.82 mm. to 2.08 mm.

Color: General facies sordid white to nigro-violaceous. Head,
pronotum, most of thoracic venter, and limbs sordid white to pale
testaceous. Scutellum black with apex and lateral margins yellow
to orange; metathoracic dorsum, black. Abdomen black except
ventral keel, and portions of connexivum and dorsum, sordid white
to pale testaceous. Some specimens entirely sordid white except
most of abdomen, black. This species variable in color.

Male Structural Characteristics: As viewed from above, outline
of head laterally rounded, somewhat truncate anteriorly, with ver-
tex slightly indented; greatest width of head seven and one half
to eight times the anterior width of vertex and equal to or slightly
less than humeral width of pronotum; synthlipsis approximately
half the anterior width of vertex; along median longitudinal axis,
head is one half to three fifths the length of pronotum; notocepha-
lon narrow, sulcate dorsally; frons just above tylus, very narrow;
tylus inflated; labium with basal width approximately twice its
median length and apex bluntly rounded; rostral prong (pi. CX,
fig. 58b) slightly longer than third rostral segment, with base
originating laterally at proximal end of third rostral segment, and
with apex bluntly rounded. Pronotum with its median length one

A Revision of the Genus Buenoa 1427

half to four sevenths its humeral width; disk unimpressed, occa-
sionally with a faint median carina; lateral margins divergent; pos-
terior margin convex, slightly concave medianly. Scutellum large,
with median length greater than that of pronotum. Fore femur (pi.
CX, fig. 58a) wide and thickened at apex; subtriangular stridulatory
area consisting of four wide, sclerotized ridges. Fore tibia (pi. CX,
fig. 58a ) with stridulatory comb ( pi. CX, fig. 58c ) consisting of ap-
proximately sixteen teeth; apical teeth slightly thicker and usually
taller than basal. Chaetotaxy of male front leg as shown on Plate
CX. Male genital claspers normal. Spine from caudo-sinistral
margin of seventh abdominal tergite with apical half very narrow
and apex strongly acuminate.

Female Structural Characteristics: As viewed from above, outline
of head rounded with vertex indented at lateral margins; greatest
width of head approximately six and one half times the anterior
width of vertex and less than humeral width of pronotum; synthlip-
sis half the anterior width of vertex; along median longitudinal
axis, head is two fifths to three fifths the length of pronotum;
notocephalon narrow, sulcate dorsally; frons just above tylus, nar-
row; tylus slightly inflated. Pronotum with its median length
approximately half its humeral width; disk unimpressed; lateral
margins divergent; posterior margin convex, medianly truncate to
slightly concave. Scutellum large, with median length distinctly
greater than that of pronotum. Female ovipositor (pi. CIII, fig. 37)
of normal shape with teeth arranged in two irregular, longitudinal
rows which merge in proximal third of ovipositor valve; one inner
row of large teeth and one outer row of smaller teeth; approximately
seven or eight small, lateral, toothlike setae near apex.

Comparative Notes: Superficially this species resembles B. muta-
bilis n. sp. and B. alterna n. sp. Examination of the male, however,
will show distinct differences. This species differs from B. mutabilis
in having a narrower frons, wider synthlipsis, and distinctly dif-
ferent femoral stridulatory area and tibial comb. Buenoa omani
differs from B. alterna in having fore femur wider at apex, femoral
stridulatory area present, and different rostral prong and tibial comb.

N omenclatorial Notes: This species was first recognized as new
by Mr. C. O. Bare who labeled a series as types and paratypes using
the manuscript name B. omani. As such paratypes may have been
widely distributed, it seems desirable to point out that the name
was not validated by publication. However, to avoid confusion, the
name suggested by Bare has been retained by this author.

1428 The University Science Bulletin

Location of Types: Holotype male, allotype female, 2 male and

4 female paratypes, San Diego Co., California, July 4, 1929, L. D.
Anderson; other paratypes: 1 male and 1 female, Santa Ana Co.,
California, July 30, 1932, J. D. Beamer; 1 male and 1 female, Alpine,
California, July 9, 1929, Paul W. Oman; 1 male, Laguna Beach,
California, July 25, 1933, R. H. Beamer; 1 male and 1 female,
Durango, Mexico, May 30, 1937, Meldon Embury. The type series
is in the Francis Huntington Snow Entomological Collections, Uni-
versity of Kansas.

Data on Distribution: Known only from United States and Mex-
ico. In addition to type series, specimens from the following locali-
ties have been examined:

U. S. A.: California: Miramar, July 28, 1938, R. I. Sailer, 6 males,

5 females; Claremont, P. R. Uhler Collection, 3 males, 1 female
(U.S.N.M.).

MEXICO: Sonora: Pocito, Ciudad Las Casas, Sept. 4, 1937, H. D.
Thomas, 32 males, 25 females.

Tamaulipas: 5 m. N. of Ciudad Victoria, Nov. 5, 1936, H. D.
Thomas, 2 males.

Jalisco: Guadalajara, 20 mi. on Tequilla Rd., Sept. 13, 1938, H. D.
Thomas, 1 female.

Hidalgo: Real del Monte, Sept. 23, 1938, H. D. Thomas, 2 males;
Aguas Fria, Aug. 27, 1944, H. D. Thomas, 1 male.

Michoacdn: Patzcuaro, Aug. 31, 1938, H. D. Thomas, 2 males,

6 females; Patzcuaro, Sept. 2, 1938, H. D. Thomas, 2 males, 2 fe-
males; Carapa, Sept. 2, 1938, H. D. Thomas, 2 males, 1 female; L.
Cuitzeo, July 7, 1947, T H. Hubbell, 2 males.

Federal District: Mexico, A. Dampf, 1 male, 1 female.

Morelos: Cuernavaca, May 21, 1898, P. R. Uhler Collection, 1
female (U.S.N.M.).

Puebla: Tehuacan, July 18-25, 1937, H. D. Thomas, 2 males, 1
female; Puebla, July 25, 1937, H. D. Thomas, 3 males, 11 females.

Oaxaca: Posita, Aug. 24, 1937, H. D. Thomas, 6 males, 8 females;
Oaxaca, Aug. 25, 1937, H. D. Thomas, 1 male, 1 female.

Chiapas: San Cristobal, Aug. 30, 1937, H. D. Thomas, 74 males,
43 females; San Cristobal, Sept. 2, 1937, H. D. Thomas, 22 males,
13 females; San Vicente, Jan. 4, 1938, Octavio Utrilla L., 1 male.

All specimens listed above are in the Francis Huntington Snow
Entomological Collections, University of Kansas, Lawrence, Kansas,
unless otherwise indicated.

A Revision of the Genus Buenoa 1429

Buenoa macrotrichia n. sp.
(PI. CX, fig. 59)

Size: Male, length 5.52 mm. to 5.85 mm., greatest body width
1.43 mm. to 1.62 mm.; female, length 5.59 mm. to 6.37 mm., greatest
body width 1.49 mm. to 1.75 mm.

Color: General facies black. Head, anterolateral portions of
pronotum, thoracic venter, and limbs sordid white to testaceous.
Posterior portion of pronotum, scutellum, metathoracic dorsum, and
abdomen black except ventral keel, portions of connexivum, and
usually terminal segment, testaceous.

Male Structural Characteristics: As viewed from above, outline
of head laterally rounded, anteriorly truncate with vertex indented;
greatest width of head six and one half to seven times the anterior
width of vertex and less than humeral width of pronotum; synthlip-
sis narrow, slightly less than half the anterior width of vertex; along
median longitudinal axis, head is slightly less than half the length
of pronotum; notocephalon narrow, sulcate dorsally; tylus slightly
inflated; labrum with basal width not quite twice its median length
and apex bluntly rounded; rostral prong (pi. CX, fig. 59b) long,
distinctly longer than third rostral segment, with base originating
laterally and protruding anteriorly at point midway to near distal
end of third rostral segment, and with apex moderately rounded.
Pronotum with its median length approximately four sevenths its
humeral width; disk occasionally with two shallow, elongate de-
pressions toward the middle forming a faint median carina, not
tricarinate; lateral margins divergent; posterior margin convex,
medianly concave. Scutellum large, with median length greater
than that of pronotum. Fore femur (pi. CX, fig. 59a) wide and
thickened at apex, with several large setae on inner posterior margin;
triangular stridulatory area consisting of approximately seven to
nine sclerotized ridges. Fore tibia (pi. CX, fig. 59a) with stridula-
tory comb (pi. CX, fig. 59c) consisting of approximately thirty-one
to thirty-four teeth; apical teeth thicker than basal. Chaetotaxy of
male front leg as shown on Plate CX. Male genital claspers normal.
Spine from caudo-sinistral margin of seventh abdominal tergite with
apical half very narrow and apex strongly acuminate.

Female Structural Characteristics: As viewed from above, out-
line of head rounded with vertex usually indented at lateral margins
only; greatest width of head approximately five and one half times
the anterior width of vertex and distinctly less than humeral width
of pronotum; synthlipsis narrow, approximately two fifths the an-

1430 The University Science Bulletin

terior width of vertex; along median longitudinal axis, head is
slightly less than two fifths the length of pronotum; notocephalon
narrow, slightly sulcate; tylus slightly inflated. Pronotum with its
median length two fifths to one half its humeral width; disk unim-
pressed, occasionally with faint median carina; lateral margins di-
vergent; posterior margin convex, medianly concave. Scutellum
large, with median length distinctly greater than that of pronotum.
Female ovipositor of normal shape with teeth arranged in two longi-
tudinal rows; inner row of large teeth merges proximally with outer
row of smaller teeth; approximately six or seven small, lateral, tooth-
like setae near apex.

Comparative Notes: Superficially this species closely resembles
B. mutabilis n. sp. and B. nitida n. sp. Examination of the male,
however, will show distinct differences. This species differs from
B. mutabilis in having the fore femur wider at apex, possessing sev-
eral large setae on inner, posterior margin of fore femur, and in
having fewer sclerotized ridges in femoral stridulatory area. Buenoa
macrotrichia differs from B. nitida in having a narrower synthlipsis
and notocephalon, a less robust pronotum, and in possessing the
large setae on fore femur.

Location of Types: Holotype male, allotype female, 50 male and
50 female paratypes, Vicinity of San Pedro, Peru, May 15-29, 1935,
F. Woytkowski. The type series is in the Francis Huntington Snow
Entomological Collections, University of Kansas.

Data on Distribution: Known only from Peru. In addition to
type series, specimens from the following localities have been ex-
amined.

PERU: Vic. Pampa Hermosa, May 1-5, 1935, F. Woytkowski, 21
males, 14 females; Vic. San Pedro, May 15-29, 1935, F. Woytkowski,
166 males, 215 females; Vic. Sani Beni, Oct. 24, 1935, F. Woyt-
kowski, 1 female; Dept. Ayacucho, Prov. La Mar. Sivia, June
15-28, 1941, 33 males, 34 females.

All specimens listed above are in the Francis Huntington Snow
Entomological Collections, University of Kansas, Lawrence, Kansas.

Buenoa nitida n. sp.
(PI. CXI, fig. 60)

Size: Male, length 6.17 mm. to 6.82 mm., greatest body width
1.75 mm. to 1.82 mm.; female, length 6.50 mm. to 7.02 mm., greatest
body width 1.95 mm. to 2.08 mm.

Color: General facies sordid white to black. Dark form with
head, anterolateral portions of pronotum, most of thoracic venter,

A Revision of the Genus Buenoa 1431

and limbs sordid white to testaceous. Posterior portion of pro-
notum, scutellum, metathoracic dorsum, and abdomen black, ex-
cept ventral keel, portions of connexivum, and terminal segment,
testaceous. Hemelytron colorless with a black band at base of
membrane. Pale form usually entirely pale testaceous except most
of abdomen, black. This species variable in color.

Male Structural Characteristics: As viewed from above, outline
of head laterally rounded, anteriorly truncate with vertex indented;
greatest width of head approximately six times the anterior width
of vertex and slightly less than humeral width of pronotum; synth-
lipsis slightly less than half the anterior width of vertex; along
median longitudinal axis, head is approximately three fifths the
length of pronotum; notocephalon slightly sulcate; tylus usually
not inflated; labrum with basal width not quite twice its median
length and apex bluntly rounded; rostral prong (pi. CXI, fig. 60b)
long, almost twice as long as third rostral segment, with base origi-
nating laterally and protruding anteriorly at point midway of third
rostral segment, and with apex bluntly rounded. Pronotum wide
and somewhat inflated, with its median length approximately three
fifths its humeral width; disk with two shallow, elongate depressions
toward the middle forming a faint median carina, not tricarinate;
lateral margins in pale form only slightly divergent, more so in
dark form; posterior margin convex, medianly truncate to slightly
concave. Scutellum with median length less than that of pronotum
in pale form, greater than that of pronotum in dark form. Fore
femur (pi. CXI, fig. 60a) wide and thickened at apex; triangular
stridulatory area consisting of six to nine wide, sclerotized ridges.
Fore tibia (pi. CXI, fig. 60a) with stridulatory comb (pi. CXI, fig.
60c) consisting of approximately thirty-one to thirty-three teeth;
apical teeth thicker than basal. Chaetotaxy of male front leg as
shown on Plate CXI. Male genital claspers normal. Spine from
caudo-sinistral margin of seventh abdominal tergite tapering grad-
ually from base to strongly acuminate apex.

Female Structural Characteristics: As viewed from above, out-
line of head laterally rounded, anteriorly truncate with vertex in-
dented at least at lateral margins; greatest width of head approxi-
mately six times the anterior width of vertex and less than humeral
width of pronotum; synthlipsis half the anterior width of vertex;
along median longitudinal axis, head is approximately half the
length of pronotum; notocephalon slightly sulcate; tylus not inflated.
Pronotum with its median length two fifths to one half its humeral
width; disk unimpressed, occasionally with a faint median carina;

1432 The University Science Bulletin

lateral margins divergent; posterior margin convex, medianly trun-
cate to slightly concave. Scutellum large, with median length
distinctly greater than that of pronotum. Female ovipositor of
normal shape with teeth arranged in two longitudinal rows; inner
row of large teeth merges proximally with outer row of smaller
teeth; approximately two or three small, lateral, toothlike setae
near apex.

Variation Within Species: A slight amount of variation exists
between the light and dark forms of this species. They differ pri-
marily in the form of the pronotum and scutellum. The light form
differs from the dark in having a more robust pronotum with lateral
margins less divergent and a smaller scutellum. These two forms
have been taken within the same population.

Comparative Notes: Superficially this species resembles B. macro-
trichia n. sp. and B. platycnemis (Fieber). Examination of the
male, however, will show distinct differences. This species differs
from B. macrotrichia in having a wider synthlipsis and notocephalon,
a more robust pronotum, and in lacking the large setae on the
inner posterior margin of the fore femur. Buenoa nitida differs
from B. platycnemis in having fewer sclerotized ridges in the
femoral stridulatory area, pronotum more robust and not tricarinate,
and in the form of the rostral prong, fore femur, and tibial comb.

Location of Types: Holotype male, allotype female, 13 male and
19 female paratypes, Dept. Amazonas, Peru, Aug. 14-19, 1936,
F. Woytkowski; other paratypes: 5 males and 6 females, Valao de
Sao Pedro Tereza Santa Estado de Esperito Santo, Feb. 2, 1948,
Antenor Leitao de Carvalho. The type series is in the Francis
Huntington Snow Entomological Collections, University of Kansas.

Data on Distribution: Known only from Brazil and Peru. Speci-
mens from the following localities have been examined:

BRAZIL: Valao de Sao Pedro Tereza Santa Estado de Esperito
Santo, Feb. 2, 1948, Antenor Leitao de Carvalho, 5 males, 6 females.

PERU: Dept. Amazonas, Vic. Guayabamba, Aug. 14-19, 1936,
F. Woytkowski, 15 males, 39 females; Dept. San Martin, Vic. Rioja,
Sept. 9-Oct. 3, 1936, F. Woytkowski, 15 males, 25 females.

All specimens listed above are in the Francis Huntington Snow
Entomological Collections, University of Kansas, Lawrence, Kansas.

Buenoa mutabilis n. sp.

(PI. CXI, fig. 61)

Size: Male, length 5.20 mm. to 5.98 mm., greatest body width
1.43 mm. to 1.83 mm.; female, length 5.65 mm. to 6.30 mm., greatest
body width 1.69 mm. to 1.88 mm.

A Revision of the Genus Buenoa 1433

Color: General facies pale testaceous to black. Head, anterior
portion of pronotum, thoracic venter, and limbs sordid white to
testaceous. Posterior portion of pronotum usually black; scutellum
entitrely black, occasionally with apex sordid white to testaceous;
metathoracic dorsum light brown to black. Abdomen black except
ventral keel, portions of connexivum, and last one or two segments,
testaceous. Pale specimens entirely sordid white to testaceous ex-
cept most of abdomen, black. This species variable in color.

Male Structural Characteristics: As viewed from above, outline
of head laterally rounded, anteriorly truncate with vertex indented;
greatest width of head approximately six and one half times the
anterior width of vertex and less than humeral width of pronotum;
synthlipsis narrow, approximately one third the anterior width of
vertex; along median longitudinal axis, head is one half to three
fifths the length of pronotum; notocephalon slightly sulcate; tylus
slightly inflated; labrum with basal width not quite twice its median
length and apex bluntly rounded; rostral prong (pi. CXI, figs. 61b,
61c) variable, distinctly longer than third rostral segment, with
base originating laterally and protruding anteriorly at point mid-
way to near distal end of third rostral segment, and with apex
moderately to sharply rounded. Pronotum with its median length
one half to four sevenths its humeral width; disk usually with two
shallow, elongate depressions toward the middle forming a faint
median carina, not tricarinate; lateral margins divergent; posterior
margin convex, medianly concave. Scutellum large, with median
length greater than that of pronotum. Fore femur (pi. CXI, fig
61a) not greatly thickened at apex; subtriangular stridulatory area
consisting of approximately ten to eighteen sclerotized ridges.
Fore tibia (pi. CXI, fig. 61a) with stridulatory comb (pi. CXI, fig.
61d) consisting of approximately thirty-three to thirty-eight teeth;
apical teeth thicker than basal. Chaetotaxy of male front leg as
shown on Plate CXI. Male genital claspers normal. Spine from
caudo-sinistral margin of seventh abdominal tergite with apical
half very narrow and apex strongly acuminate.

Female Structural Characteristics: As viewed from above, head
laterally rounded, anteriorly truncate with vertex indented; greatest
width of head approximately six times the anterior width of vertex
and distinctly less than humeral width of pronotum; synthlipsis
narrow, slightly less than half the anterior width of vertex; along
median longitudinal axis, head is slightly less than half the length
of pronotum; notocephalon slightly sulcate; tylus not inflated. Pro-
notum with its median length approximately half its humeral width;

1434 The University Science Bulletin

disk only faintly impressed and sometimes not at all, not tricarinate;
lateral margins divergent; posterior margin convex, medianly con-
cave. Scutellum large, with median length distinctly greater than
that of pronotnm. Female ovipositor of normal shape with teeth
arranged in two longitudinal rows; one long inner row of large
teeth and one long outer row of smaller teeth; approximately seven
or eight small, lateral, toothlike setae near apex.

Variation Within Species: This species varies considerably in
the form of the rostral prong; there is also some variation in the
proportional size of the head and pronotum and in the form of the
fore femur. The greatest contrast is shown between a series from
Peru and one from Paraguay. The former is a small, pale form, the
male with a relatively short, straight rostral prong; the latter is a
larger, dark form, the male with a long, curved rostral prong.

Comparative Notes: Superficially this species closely resembles
B. macrotrichia n. sp. and B. alterna n. sp., and somewhat resembles
the dark form of B. pollens (Champion). Examination of the male,
however, will show distinct differences. This species differs from
B. macrotrichia in having the fore femur narrower at apex, lacking
the large setae on inner, posterior margin of fore femur, and having
more sclerotized ridges in femoral stridulatory area. It differs from
B. alterna in having a femoral stridulatory area and a distinctly
different rostral prong. Buenoa mutabilis differs from B. pallens in
having the fore femur wider at apex, the notocephalon distinctly
narrower, and the rostral prong protruding anteriorly on third
rostral segment. Other minor differences also occur.

Location of Types: Holotype male, allotype female, 23 male and
19 female paratypes, Vicinity of Rioja, Dept. San Martin, Peru,
Sept. 9-Oct. 22, 1936, F. Woytkowski. The type series is in the
Francis Huntington Snow Entomological Collections, University
of Kansas.

Data on Distribution: Known from West Indies (Haiti), Vene-
zuela, British Guiana, Peru, and Paraguay. In addition to type
series, specimens from the following localities have been examined:

WEST INDIES: Haiti: Port au Prince, June, 1925, W. H. Hoff-
man, 3 females.

VENEZUELA: Surukum, June 1941, P. J. Anduze, 14 males, 7
females, 5 nymphs.

BRITISH GUIANA: Cartabo, Mrs. Brindly, 5 females.

PERU: Vic. Pampa Hermosa, May 1-5, 1935, F. Woytkowski, 5
males; Vic. San Pedro, May 15-29, 1935, F. Woytkowski, 18 males,

A Revision of the Genus Buenoa 1435

17 females; Vic. San Pedro, May 15-19, 1935, F. Woytkowski, 10
males, 6 females; Vic. Sani Beni, Aug. 5, 1935, F. Woytkowski, 5
males, 2 females; Vic. Sani Beni, Sept.-Oct., 1935, F. Woytkowski, 14
males, 21 females; Rio Negro, Oct. 30, 1935, F. Woytkowski, 3
males, 2 females; Dept. Huanuco, Vic. Tingo Maria, May 10, 1937,
F. Woytkowski, 10 males, 9 females; Dept. Huanuco, Vic. of Afila-
dor, June 3-9, 1937, F. Woytkowski, 29 males, 6 females; Dept.
Huanuco, Vic. Leonpampa, Dec. 11-30, 1937, F. Woytkowski, 37
males, 26 females; Dept. Huanuco, Loc. Shapajilla, June 1-10, 1938,
F. Woytkowski, 25 males, 11 females; Peru, Dec. 12-14, 1937, F.
Woytkowski, 41 males, 30 females; Peru, Oct. 8, 1940, F. Woyt-
kowski, 40 males, 43 females; Dept. Ayacucho, Prov. La Mar. Sivia,
June 15-28, 1941, F. Woytkowski, 16 males, 7 females; Satipo, Oct.,
1942, Pedro Paprzyki, 25 males, 26 females.

PARAGUAY: Villarrica, Jan. 7, 1923, F. Schade, 1 male, 1 female;
Villarrica, June 7-15, 1923, F. Schade, 2 females; Villarrica, July 6,

1923, F. Schade, 3 females; Villarrica, Nov. 10, 1923, F. Schade, 1
female; Villarrica, Dec. 5-6, 1923, F. Schade, 3 males, 4 females;
Villarrica, Jan. 12, 1924, F. Schade, 1 female; Villarrica, Mar.-Apr.,

1924, F. Schade, 1 male, 6 females; Villarrica, July 8, 1924, F.
Schade, 1 male, 4 females; Villarrica, Sept. 9-Nov. 16, 1924, F.
Schade, 9 females; Villarrica, Dec. 16, 1924, F. Schade, 20 males, 50
females; Villarrica, Jan., 1926, F. Schade, 23 males, 31 females;
Villarrica, Mar., 1926, F. Schade, 3 males, 2 females; Estero Grande,
Nov. 1, 1924, F. Schade, 1 male, 2 females; Melinesque, Dec. 1925,
F. Schade, 3 females; Melinesque, June 20-28, 1935, F. Schade, 4
females.

All specimens listed above are in the Francis Huntington Snow
Entomological Collections, University of Kansas, Lawrence, Kansas.

Buenoa arida n. sp.
(PI. CXI, fig. 62)

Size: Male, length 5.33 mm. to 5.91 mm., greatest body width
1.69 mm. to 1.95 mm.; female, length 5.95 mm. to 7.41 mm., greatest
body width 2.01 mm. to 2.21 mm.

Color: General facies sordid white to slate. Head, pronotum,
and limbs sordid white to pale testaceous. Thoracic venter tes-
taceous to black; scutellum black with posterolateral margins and
apex sordid white to pale testaceous; metathoracic dorsum pale
testaceous to black, usually appearing gray through hyalin hemely-
tra. Abdomen black except ventral keel and portions of connexivum

1436 The University Science Bulletin

and dorsum, yellowish white to pale testaceous. Hemelytra hyalin
with posterior third black. Some specimens entirely sordid white
except most of abdomen, black. This species variable in color.

Male Structural Characteristics: As viewed from above, outline
of head rounded, vertex slightly indented; greatest width of head
seven and one half to eight and one half times the anterior width
of vertex and equal to or slightly less than humeral width of pro-
notum; synthlipsis approximately one half the anterior width of
vertex; along median longitudinal axis, head is slightly more than
half the length of pronotum; notocephalon sulcate dorsally; frons
immediately above tylus, very narrow; tylus slightly inflated; labrum
with basal width not quite twice its median length and apex bluntly
rounded; rostral prong (pi. XI, fig. 62b) with length equal to that
of third rostral segment, with base originating laterally near proxi-
mal end of third rostral segment, and with apex bluntly rounded.
Pronotum with its median length slightly more than half its humeral
width; disk with two elongate depressions toward the middle form-
ing, a median carina; lateral margins slightly divergent; posterior
margin convex, slightly concave medianly. Scutellum large, with
median length greater than that of pronotum. Fore femur (pi. CXI,
fig. 62a) wide but not greatly thickened at apex; large, subtriangular
stridulatory area consisting of approximately sixty to sixty-five fine,
sclerotized ridges. Fore tibia (pi. CXI, fig. 62a) with stridulatory
comb (pi. CXI, fig. 62c) consisting of approximately nineteen to
twenty-two teeth; apical teeth thicker and slightly taller than basal.
Chaetotaxy of male front leg as shown on Plate CXI. Male genital
claspers normal. Spine from caudo-sinistral margin of seventh ab-
dominal tergite normal, tapering gradually from base to strongly
acuminate apex.

Female Structural Characteristics: As viewed from above, out-
line of head rounded with vertex usually indented; greatest width
of head six to six and one half times the anterior width of vertex
and slightly less than humeral width of pronotum; synthlipsis one
half to two thirds the anterior width of vertex; along median longi-
tudinal axis, head is approximately three fifths the length of pro-
notum; notocephalon sulcate dorsally; tylus slightly inflated. Pro-
notum with its median length approximately half its humeral width;
disk with two elongate depressions toward the middle forming a
median carina; lateral margins divergent; posterior margin convex,
medianly truncate to slightly concave. Scutellum large, with
median length distinctly greater than that of pronotum. Female
ovipositor of normal shape with teeth arranged in two longitudinal

A Revision of the Genus Buenoa 1437

rows which merge proximally; one inner row of large teeth and one
outer row of smaller teeth; approximately three or four small,
lateral, toothlike setae near apex.

Comparative Notes: Superficially this species somewhat resem-
bles B. nitida n. sp. Examination of the male, however, will show
distinct differences. This species differs from B. nitida in having
head wider in relation to pronotum, frons just above tylus distinctly
narrower, fore femur less robust, and differences in the rostral
prong, femoral stridulatory area, and tibial comb.

Location of Types: Holotype male, Santa Rita Mts., Arizona,
July 10, 1950, W. J. Arnold; allotype female, Santa Rita Mts., Ari-
zona, July 9, 1947, L. D. Beamer; paratypes as follows: 3 males
and 2 females, Oro Blanco Mts., Arizona, Apr. 3, 1937, Owen
Bryant; 1 male, Gila Co., Arizona, Aug. 5, 1927, R. H. Beamer;
1 female, Santa Catalina Mts., Arizona, July 14, 1950, W. J. Arnold.
The type series is in the Francis Huntington Snow Entomological
Collections, University of Kansas.

Data on Distribution: Known only from the United States. In
addition to the type series, specimens from the following localities
have been examined:

U.S.A.: Arizona: Santa Cruz Co., Aug. 4, 1927, P. A. Readio,
1 male; Huachuca Mts., July 8, 1932, R. H. Beamer, 1 female; Oro
Blanco Mts., Apr. 3, 1937, Owen Bryant, 2 males, 1 female; Cata-
lina Mts., Oct. 27, 1941, Victor Potter, 1 male (U. of Mich.); Cata-
lina Mts., Sabino Canyon, Hubbard Coll., 1 male, 1 female (U.S.
N. M.); Arizona, P. R. Uhler Collection, 2 males, 1 female (U.S.
N.M.).

All specimens listed above are in the Francis Huntington Snow
Entomological Collections, University of Kansas, Lawrence, Kansas,
unless otherwise indicated.

Buenoa speciosa n. sp.
(PI. CXII, fig. 63)

Size: Male, length 8.38 mm. to 8.84 mm., greatest body width
2.40 mm. to 2.60 mm.; female, length 8.45 mm. to 9.10 mm., greatest
body width 2.53 mm. to 2.66 mm.

Color: General facies pale testaceous to blackish gray. Head,
pronotum, and limbs sordid white to pale testaceous. Thoracic
venter testaceous to black; scutellum black with posterolateral mar-
gins and apex sordid white to testaceous; metathoracic dorsum tes-
taceous to black, usually appearing gray through hyalin hemelytra.
Abdomen black except ventral keel and portions of connexivum and

1438 The University Science Bulletin

dorsum, yellowish white to testaceous. Hemelytra hyalin with
posterior third brown to black. Some specimens entirely sordid
white to pale testaceous except most of abdomen, black. This spe-
cies variable in color.

Male Structural Characteristics: As viewed from above, outline
of head laterally rounded, anteriorly truncate with vertex indented;
greatest width of head eight to ten times the anterior width of ver-
tex and slightly less than humeral width of pronotum; synthlipsis
three fifths to four fifths the anterior width of vertex; along median
longitudinal axis, head is approximately three fifths the length
of pronotum; notocephalon narrow, sulcate dorsally; tylus inflated;
labrum with basal width not quite twice its median length and
apex bluntly rounded; rostral prong (pi. CXII, fig. 63c) unusual in
shape, distinctly longer than third rostral segment, with base origi-
nating laterally near distal end of third rostral segment, and
with apex moderately rounded. Pronotum with its median length
slightly more than half its humeral width; disk with two shallow,
elongate depressions toward the middle forming a faint median ca-
rina; usually a subtriangular depression on each side but not ap-
pearing tricarinate; lateral margins slightly divergent; posterior
margin convex, medianly truncate to slightly concave. Scutellum
large, with median length usually greater than that of pronotum.
Fore femur (pi. CXII, fig. 63a) wide and thickened at apex; small,
triangular stridulatory area consisting of approximately eleven
sclerotized ridges. Fore tibia (pi. CXII, fig. 63a) with stridulatory
comb (pi. CXII, fig. 63b) consisting of approximately thirty-nine
to forty-two teeth; apical teeth thicker, narrower, and taller than
basal. Chaetotaxy of male front leg as shown on Plate CXII.
Male genital claspers normal. Spine from caudo-sinistral margin
of seventh abdominal tergite normal, tapering gradually from base
to strongly acuminate apex.

Female Structural Characteristics: As viewed from above out-
line of head rounded with vertex indented only at lateral margins;
greatest width of head seven to seven and one half times the an-
terior width of vertex and less than humeral width of pronotum;
synthlipsis three fifths to four fifths the anterior width of vertex;
along median longitudinal axis, head is slightly more than half
the length of pronotum; notocephalon sulcate dorsally; tylus slightly
inflated. Pronotum with its median length three sevenths to
four sevenths its humeral width; disk usually unimpressed, occa-
sionally with a faint median carina; lateral margins divergent;
posterior margin convex, medianly truncate to slightly concave.

A Revision of the Genus Buenoa 1439

Scutellum large, with median length distinctly greater than that
of pronotum. Female ovipositor of normal shape with teeth ar-
ranged in two longitudinal rows which merge proximally; one inner
row of large teeth and one outer row of smaller teeth; approximately
seven or eight small, lateral, toothlike setae near apex.

Comparative Notes: Superficially this species resembles B. cras-
sipes (Champion). Examination of the male, however, will show
distinct differences. This species differs from B. crassipes in having
the head with eyes distinctly larger, frons just above tylus narrower,
and distinct differences in the rostral prong, femoral stridulatory
area and tibial comb.

Location of Types: Holotype male, allotype female, 3 female
paratypes, 20 miles W. of San Luis Potosi, Mexico, Aug. 8, 1944,
H. D. Thomas; other paratypes: 2 males and 1 female, San Luis
Potosi, Mexico, Aug. 5, 1944, H. D. Thomas; 3 males and 1 female,
Real del Monte, Hidalgo, Mexico, Sept. 23, 1938, H. D. Thomas.
The type series is in the Francis Huntington Snow Entomological
Collections, University of Kansas.

Data on Distribution: Known only from United States and Mex-
ico. In addition to type series, specimens from the following locali-
ties have been examined:

U. S. A.: Texas: Ft. Davis, 1914, C. Thompson, 1 female (U. of
Mich.); Alpine, June 5, 1927, 1 male (U.S.N.M.); Davis Mts., July
12, 1938, R. I. Sailer and D. W. Craik, 1 male, 2 females.

All specimens listed above are in the Francis Huntington Snow
Entomological Collections, University of Kansas, Lawrence, Kansas,
unless otherwise indicated.

Buenoa gracilis n. sp.
(PI. CXII, fig. 64)

Size: Male, length 5.39 mm. to 6.04 mm., greatest body width
1.30 mm. to 1.49 mm.; female, length 5.52 mm. to 6.11 mm., greatest
body width 1.36 mm. to 1.69 mm.

Color: General facies sordid white. Head, thoracic dorsum, most
of thoracic venter, and limbs sordid white. Abdominal venter
black except keel and portions of connexivum, sordid white to pale
testaceous; abdominal dorsum yellowish white with irregular black
area posteriorly.

1440 The University Science Bulletin

the anterior width of vertex and slightly less than humeral width of
pronotum; synthlipsis very narrow, approximately one fourth the
anterior width of vertex, often carinate; along median longitudinal
axis, head is approximately half the length of pronotum; noto-
cephalon sulcate dorsally; tylus inflated; labrum with basal width
approximately twice its median length and apex bluntly rounded to
almost truncate; rostral prong (pi. CXII, fig. 64b) equal to or slightly
shorter than third rostral segment, with base originating laterally
near proximal end of third rostral segment, and with apex bluntly
rounded. Pronotum long, with its median length two thirds to
three fourths its humeral width; disk with two shallow, elongate
depressions toward the middle and a deep, subtriangular depres-
sion on each side, thus appearing tricarinate posteriorly; lateral
margins slightly divergent, occasionally parallel; posterior margin
convex, distinctly concave medianly. Scutellum with median length
distinctly less than that of pronotum. Fore femur (pi. CXII, fig.
64a) relatively wide, not greatly thickened at apex; subtriangular
stridulatory area consisting of approximately six to nine wide,
sclerotized ridges. Fore tibia (pi. CXII, fig. 64a) with stridulatory
comb (pi. CXII, fig. 64c) consisting of approximately twenty-five
to twenty-eight teeth; apical teeth slightly thicker and narrower
than basal; approximately four short, peg-like setae (pi. CXII, fig.
64d) on inner surface of tibia at apex. Chaetotaxy of male front
leg as shown on plate CXII. Male genital claspers normal. Spine
from caudo-sinistral margin of seventh abdominal tergite normal,
tapering gradually from base to strongly acuminate apex.

Female Structural Characteristics: As viewed from above, out-
line of head rounded with anterior margin of vertex continuous with
that of eyes, occasionally slightly protuberant; greatest width of
head approximately five times the anterior width of vertex and
slightly less than humeral width of pronotum; synthlipsis narrow,
approximately one fourth the anterior width of vertex; along median
longitudinal axis, head is one half to two thirds the length of pro-
notum; notocephalon sulcate dorsally; tylus slightly inflated. Pro-
notum with its median length slightly less than three fifths its
humeral width; disk with two shallow, elongate depressions toward
the middle and occasionally a large subtriangular depression on
each side, thus sometimes appearing faintly tricarinate; lateral mar-
gins divergent; posterior margin convex, distinctly concave me-
dianly. Scutellum with median length usually greater than that of
pronotum. Female ovipositor of normal shape with teeth arranged
in two longitudinal rows; one inner row of large teeth and one

A Revision of the Genus Buenoa 1441

irregular, outer row of smaller teeth; approximately six to ten small,
lateral, toothlike setae near apex.

Variation Within Species: Occasionally specimens are found with
flight wings not fully developed. These specimens have lateral mar-
gins of pronotum parallel, scutellum smaller, and hemelytra with
claval sutures less evident and membranes smaller than the form
with fully developed flight wings.

Comparative Notes: Superficially this species somewhat resem-
bles B. oculata n. sp. Examination of the male, however, will show
distinct differences. This species is larger than B. oculata, has
synthlipsis wider, femoral stridulatory area present, and first tarsal
segment of intermediate leg not emarginate.

Location of Types: Holotype male, allotype female, 25 male and
25 female paratypes, Region of Tarapoda, Dept. San Martin, Peru,
Feb. 16, 1947, Felix Woytkowski. The type series is in the Francis
Huntington Snow Entomological Collections, University of Kansas.

Data on Distribution: Known from Mexico, Honduras, Panama,
West Indies (Cuba, Jamaica, Puerto Rico, St. Croix, Grenada), and
Peru. In addition to type series, specimens from the following lo-
calities have been examined:

MEXICO: Veracruz: Minatitlan, Sept. 22, 1936, H. D. Thomas,
15 males, 16 females.

Guerrero: Rio Agua, Oct. 31, 1936, H. D. Thomas, 1 male; Aca-
pulco, July 12, 1937, H. D. Thomas, 1 male, 10 females.

Oaxaca: Papaloapan, Feb. 20, 1939, Gordon and Atz, 3 males;
Papaloapan, Feb. 22, 1939, Gordon and Atz, 2 females, 2 nymphs
(U. of Mich.); Arroyo Zacatispan, Mar. 4, 1943, M. & E. Gordon,
1 female ( U. of Mich. ) .

Chiapas: La Libertad, Jan., 1938, Octavio Utrilla L., 1 male;
San Vicente, Jan. 4, 1948, Octavio Utrilla L., 2 females.

Campeche: Ciudad del Carmen, Sept. 1-18, 1936, H. D. Thomas,
7 males, 14 females.

HONDURAS: Tela, La Fragua farm, March and April, 1923,
T. H. Hubbell, 6 males, 3 females ( U. of Mich. ).

PANAMA: Sona, May, 1914, J. Zetek, 1 male.

WEST INDIES: Cuba: Santiago, Sept. 20, 1912, J. M. Espin,
4 males, 3 females, 1 nymph (U. S. N. M.); San Carlos Guantaname,
Oct. 4-8, 1913, 2 males, 1 female, 4 nymphs (U. S. N. M.); Havana,
Bot. Garden, Jan. 25, 1932, P. J. Bermudez, 17 males, 12 females.

Jamaica: Palm Beach, Montego Bay, Mar. 11, 1911, 11 males,
3 females, 1 nymph ( A. M. N. H. ) .

Puerto Rico: Guayabal Reservoir, Feb. 20, 1934, S. Hildebrand,

1442 The University Science Bulletin

6 males, 4 females (U. S. N. M); Guinea Reservoir, Feb. 20, 1934,
S. Hildebrand, 2 females (U. S. N. M.).

St. Croix: Slob Pond, 1937, H. A. Beatty, 1 male, 1 female (U. S.
N.M.); St. Croix, 1941, H. A. Beatty, 2 males, 1 female (U.S.

N.M.).

Grenada: Grenada, H. H. Smith, P. R. Uhler Collection, 1 male,

4 females (U.S.N.M.).

PERU: Vic. Sani Beni, Oct. 9-16, 1935, F. Woytkowski, 15 males,
11 females; Dept. San Martin, Region Tarapoto, Feb. 8, 1947, Felix
Woytkowski, 24 females.

All specimens listed above are in the Francis Huntington Snow
Entomological Collections, University of Kansas, unless otherwise
indicated.

Buenoa communis n. sp.
(PL CXII, fig. 65)

Size: Male, length 5.39 mm. to 6.04 mm., greatest body width
1.49 mm. to 1.75 mm.; female, length 5.91 mm. to 6.50 mm., greatest
body width 1.62 mm. to 1.95 mm.

Color: General facies testaceous to rufescent. Head, pronotum,
thoracic venter, and limbs sordid white to testaceous; pronotum
occasionally with median and basal portions orange. Scutellum
usually orange, often with two anterolateral brown to black areas;
metathoracic dorsum usually light brown to black. Abdomen light
brown to black except ventral keel, portions of connexivum, and
last two segments testaceous. Hemelytron with rufescent area at
humeral angle. This species somewhat variable in color.

Male Structural Characteristics: As viewed from above, outline
of head rounded with anterior margin of vertex continuous with
that of eyes; greatest width of head six to six and one half times
the anterior width of vertex and less than humeral width of pro-
notum; synthlipsis very narrow, one fifth to one fourth the anterior
width of vertex, often carinate; along median longitudinal axis,
head is two fifths to one half the length of pronotum; notocephalon
narrow, slightly sulcate; tylus not inflated; labrum with basal width
more than twice its median length and apex bluntly rounded; rostral
prong (pi. CXII, fig. 65b) slightly longer than third rostral seg-
ment, with base originating laterally near proximal end of third
rostral segment, and with apex bluntly rounded. Pronotum with
its median length approximately three fifths its humeral width;
disk with two elongate depressions toward the middle and a large
subtriangular depression on each side, thus appearing tricarinate;

A Revision of the Genus Buenoa 1443

lateral margins divergent; posterior margin convex, medianly con-
cave. Scutellum with median length equal to or slightly less than
that of pronotum. Fore femur (pi. CXII, fig. 65a) neither wide
nor strongly thickened at apex; oblong to subtriangular stridulatory
area consisting of approximately twenty-six to thirty-one sclerotized
ridges. Fore tibia (pi. CXII, fig. 65a) with stridulatory comb
(pi. CXII, fig. 65c) consisting of approximately thirty-one to thirty-
three teeth; apical teeth thicker and slightly narrower than basal.
Chaetotaxy of male front leg as shown on Plate CXII. Male genital
claspers normal. Spine from caudo-sinistral margin of seventh ab-
dominal tergite with apical one third very narrow and apex strongly
acuminate.

Female Structural Characteristics: As viewed from above, out-
line of head rounded with anterior margin of vertex usually con-
tinuous with that of eyes, occasionally indented at lateral margins;
greatest width of head six to six and one half times the anterior
width of vertex and less than humeral width of pronotum; synth-
lipsis very narrow, approximately one fourth the anterior width of
vertex, often carinate; along median longitudinal axis, head is one
third to one half the length of pronotum; notocephalon narrow,
slightly sulcate; tylus not inflated. Pronotum with its median
length slightly more than half its humeral width; disk with two
shallow, elongate depressions toward the middle and a shallow,
subtriangular depression on each side, thus appearing faintly tri-
carinate, occasionally with median carina only; lateral margins
divergent; posterior margin convex, slightly concave medianly.
Scutellum large, with median length greater than that of pronotum.
Female ovipositor of normal shape with teeth arranged in two
longitudinal rows; one inner row of few, large teeth that merges
proximally with long outer row of smaller teeth; approximately
four to seven small, lateral, toothlike setae near apex.

Comparative Notes: Superficially this species somewhat resem-
bles B. platycnemis (Fieber) and B. mutabilis n. sp. Examination
of the male, however, will show distinct differences. This species
differs from B. platycnemis in having the fore femur narrower at
apex, synthlipsis and notocephalon distinctly narrower, rostral prong
shorter, and minor differences in the femoral stridulatory area and
tibial comb. Buenoa communis differs from B. mutabilis in having
the synthlipsis narrower, pronotum distinctly tricarinate, more
sclerotized ridges in the femoral stridulatory area, and minor differ-
ences in the rostral prong and tibial comb.

1444 The University Science Bulletin

Location of Types: Holotype male, allotype female, 50 male and
50 female paratypes, vicinity of Joao Pessoa, River Jurua, Brazil,
July 10 to Sept. 20, 1936, A. M. Olalla. The type series is in the
Francis Huntington Snow Entomological Collections, University of
Kansas.

Data on Distribution: Known only from Brazil and Bolivia. Spec-
imens from the following localities have been examined:

BRAZIL: Para: Lago Grande, Feb., 1939, A. M. Olalla, 2 males,
2 females.

Parahiba: Vic. Joao Pessoa, River Jurua, July 10 to Sept. 20, 1936,
A. M. Olalla, 117 males, 136 females.

BOLIVIA: Junction of Madre de Dios and Beni Rivers, Victoria,
Oct., 1937, A. M. Olalla, 124 males, 142 females; R. Beni, Puerto
Salinas, Nov., 1937, A. M. Olalla, 4 males, 7 females.

All specimens listed above are in the Francis Huntington Snow
Entomological Collections, University of Kansas, Lawrence, Kansas.

Buenoa artafrons n. sp.
(PI. CXIII, fig. 66)

Size: Male, length 5.20 mm. to 5.88 mm., greatest body width
1.49 mm. to 1.69 mm.; female, length 5.29 mm. to 5.98 mm., greatest
body width 1.56 mm. to 1.75 mm.

Color: General facies sordid white to pale testaceous. Head,
thorax, and limbs usually sordid white to testaceous. Scutellum
occasionally with apex orange and two anterolateral black areas.
Abdomen light brown to black except ventral keel and portions of
connexivum and dorsum, pale testaceous. Thoracic and abdominal
venter and limbs often entirely light brown to black. This species
variable in color.

Male Structural Characteristics: As viewed from above, outline
of head rounded with vertex indented; greatest width of head ap-
proximately nine times the anterior width of vertex and usually
slightly greater than humeral width of pronotum; synthlipsis narrow,
approximately half the anterior width of vertex; along median
longitudinal axis, head is more than two thirds the length of pro-
notum; notocephalon narrow, sulcate dorsally; frons immediately
above tylus, very narrow; tylus slightly inflated; labrum with basal
width twice its median length and apex bluntly rounded; rostral
prong (pi. CXIII, fig. 66b) slightly longer than third rostral segment,
with base originating laterally at a point midway to near proximal
end of third rostral segment, and with apex moderately to bluntly
rounded. Pronotum with its median length approximately three fifths

A Revision of the Genus Buenoa 1445

its humeral width; disk with two elongate depressions toward the
middle and a large subtriangular depression on each side, thus ap-
pearing distinctly tricarinate; lateral margins slightly divergent;
posterior margin convex, medianly concave. Scutellum with median
length equal to that of pronotum. Fore femur (pi. CXIII, fig. 66a)
neither wide nor greatly thickened at apex; subtriangular stridula-
tory area consisting of approximately seven wide, sclerotized ridges.
Fore tibia (pi. CXIII, fig. 66a) with stridulatory comb (pi. CXIII,
fig. 66c) consisting of approximately twenty-five to twenty-seven
teeth; all teeth approximately same size and thickness. Chaetotaxy
of male front leg as shown on Plate XIII. Male genital claspers
normal. Spine from caudo-sinistral margin of seventh abdominal
tergite normal, tapering gradually from base to strongly acuminate
apex.

Occasionally specimens are found with flight wings more fully
developed. These specimens have head narrower than humeral
width of pronotum, lateral margins of pronotum more divergent,
and scutellum larger. Both forms have claval sutures present in
hemelytra but in the form with more fully developed flight wings,
membranes are larger.

Female Structural Characteristics: As viewed from above, outline
of head rounded with anterior margin of vertex continuous with
that of eyes, occasionally indented at lateral margins; greatest
width of head approximately seven times the anterior width of ver-
tex and slightly less than humeral width of pronotum; synthlipsis
narrow, approximately half the anterior width of xertex; along me-
dian longitudinal axis, head is slightly more than three fifths the
length of pronotum; notocephalon narrow, sulcate dorsally; tylus
slightly inflated. Pronotum with its median length approximately
four sevenths its humeral width; disk usually with two shallow,
elongate depressions toward the middle and a shallow, subtriangular
depression on each side, thus appearing faintly tricarinate, occa-
sionally with median carina only; lateral margins divergent; pos-
terior margin convex, slightly concave medianly. Scutellum large,
with median length greater than that of pronotum. Female oviposi-
tor of normal shape with teeth arranged in two longitudinal rows
which merge midway of ovipositor valve; one inner row of large
teeth and one outer row of smaller teeth; approximately four or five
small, lateral, toothlike setae near apex.

Occasionally specimens are found with flight wings more fully de-
veloped. These specimens have head distinctly narrower than
humeral width of pronotum, lateral margins of pronotum more di-

1446 The University Science Bulletin

vergent, and scutellum larger. Both forms have claval sutures
present in hemelytra but in the form with more fully developed
flight wings, membranes are larger.

Comparative Notes: Superficially this species resembles B. albida
(Champion). Examination of the male, however, will show dis-
tinct differences. This species differs from B. albida in having
frons just above tylus narrower, tylus without a median depression,
rostral prong distinctly shorter, and differences in the femoral
stridulatory area and tibial comb.

Location of Types: Holotype male, allotype female, 8 male and
14 female paratypes, Cocoanut Grove, Florida, Aug. 9, 1930, R. H.
Beamer and Paul W. Oman; other paratypes: 11 males and 6 females,
Wildwood, Florida, Aug. 2-9, 1930, R. H. Beamer and Paul W.
Oman. The type series is in the Francis Huntington Snow Entomo-
logical Collections, University of Kansas.

Data on Distribution: Known only from the United States. In
addition to type series, specimens from the following localities have
been examined:

U. S. A.: Georgia: Okefenokee Swp., July 30, 1934, R. H. Beamer,
P. McKinstry, and M. E. Griffith, 15 males, 7 females.

Florida: Sanford, Aug. 4, 1930, Paul W. Oman, 2 males, 1 female;
Cocoanut Grove, Aug. 9, 1930, R. H. Beamer and Paul W. Oman,
1 male, 19 females.

All specimens listed above are in the Francis Huntington Snow
Entomological Collections, University of Kansas, Lawrence, Kansas.

Buenoa macrotibialis Hungerford
(PL CIII, 6g. 38; pi. CXIII, fig. 67)

1924. Buenoa macrotibialis Hungerford, H. B. Ann. Ent. Soc. America, vol.
XVII, No. 2, p. 225.

1925. Buenoa macrotibialis, Bare, C. O. Ent. News, vol. XXXVI, No. 8, p. 228

(key).

1926. Buenoa macrotibialis, Blatchley, W. S. Heteroptera or True Bugs ol
Eastern North America, pp. 1057 and 1060 (key and description).

1928. Buenoa macrotibialis, Bare, C. O. Univ. Kansas Sci. Bui., vol. XVIII,

No. 3, p. 268 (key).
1942. Buenoa macrotibialis, Hutchinson, G. E. American Jr. Sci., vol. CCXL,

p. 336 (morphological note).

Size: This species varies considerably in size within the same
population. Male, length 5.85 mm. to 6.50 mm., greatest body
width 1.49 mm. to 1.69 mm.; female, length 5.91 mm. to 7.00 mm.,
greatest body width 1.75 mm. to 2.08 mm.

Color: General facies sordid white to testaceous. Head and limbs
testaceous to black. Prothorax sordid white with a smoky to black
area on each side above the margin; scutellum testaceous to color-

A Revision of the Genus Buenoa 1447

less; metathoracic dorsum and thoracic venter testaceous with por-
tions light brown to black. Abdominal dorsum testaceous with a
light brown to black area usually located posteriorly; abdominal
venter black with keel, portions of connexivum, and occasionally last
segment, testaceous. Hemelytron sordid white with black band
covering humeral angle and extending along anterior margin of wing
for approximately one fourth its length and occasionally another
very small smoky to black area at tip of corium limited usually to
edge of wing. This species somewhat variable in color.

Male Structural Characteristics: As viewed from above, outline
of head rounded with anterior margin of vertex continuous with
that of eyes; greatest width of head approximately six and one half
times the anterior width of vertex and usually equal to humeral
width of pronotum; synthlipsis half, or slightly less than half, the
anterior width of vertex; along median longitudinal axis, head is
approximately three fifths the length of pronotum; notocephalon
slightly sulcate dorsally; tylus inflated; labrum small, with basal
width more than twice its median length and apex bluntly rounded;
rostral prong (pi. CXIII, fig. 67b) long, much longer than third
rostral segment, with base originating laterally at distal end of third
rostral segment causing this segment to protrude anteriorly over
terminal segment, and with apex bluntly rounded. Pronotum with
its median length approximately three fourths its humeral width;
disk with two elongate depressions toward the middle and a large
subtriangular depression on each side, thus appearing tricarinate;
lateral margins slightly divergent; posterior margin convex, me-
dianly truncate to slightly concave. Scutellum relatively small,
elevated, but depressed near anterior margin by a transverse groove;
median length distinctly less than that of pronotum. Fore femur
(pi. CXIII, fig. 67a) wide and somewhat thickened at apex; obscure,
triangular stridulatory area consisting of approximately seven to
nine irregular, sclerotized ridges. Fore tibia (pi. CXIII, fig. 67a)
with stridulatory comb (pi. CXIII, fig. 67c) consisting of approxi-
mately fifty-four to fifty-six teeth; apical teeth thicker and narrower
than basal. Chaetotaxy of male front leg as shown on Plate CXIII.
Male genital claspers normal. Spine from caudo-sinistral margin of
seventh abdominal tergite normal, tapering gradually from broad
base to acuminate apex.

A single macropterous form has been seen by this author. This
specimen has head distinctly narrower than humeral width of pro-
notum; pronotum with lateral margins more divergent; scutellum

1448 The University Science Bulletin

larger; hemelytra with claval sutures present and large membranes;
fully developed flight wings.

Female Structural Characteristics: As viewed from above, out-
line of head rounded with anterior margin of vertex continuous with
that of eyes or slightly indented at lateral margins; greatest width
of head approximately six times the anterior width of vertex and
usually equal to humeral width of pronotum; synthlipsis slightly
less than half the anterior width of vertex; along median longi-
tudinal axis, head is approximately two thirds the length of pro-
notum; notocephalon slightly sulcate dorsally; tylus inflated. Pro-
notum with its median length approximately three fifths its humeral
width; disk with two elongate depressions toward the middle and a
shallow, subtriangular depression on each side, thus appearing
faintly tricarinate with median carina distinct; lateral margins di-
vergent; posterior margin convex, medianly truncate. Scutellum
relatively small, elevated, but depressed near anterior margin by a
shallow transverse groove; median length less than that of pronotum.
Female ovipositor (pi. CIII, fig. 38) of normal shape with teeth
arranged in two longitudinal rows; one short inner row of few, large
teeth and one long outer row of small teeth with a few teeth lo-
cated medianly and intermingled with the two rows; approximately
six or seven small, lateral, toothlike setae near apex.

No macropterous female forms have been seen by this author.
This form undoubtedly exists, however, and one would assume that
it differs from the brachypterous form in the same characteristics
as mentioned for the male.

Comparative Notes: Superficially this species resembles B. lim-
nocastoris Hungerford and B. confusa n. sp. Examination of the
male, however, will show distinct differences. This species differs
from B. Jimnocastoris and B. confusa in the shape and chaetotaxy of
the fore femur and tibia, the femoral stridulatory area and the shape
and length of rostral prong. It also differs from B. limnocastoris
in having the pronotum not inflated, tylus not as prominent, and
scutellum not as reduced. It differs also, from B. confusa, in that
the eyes are not as close together.

Location of Types: Holotype male, allotype female, Bryant's Bog,
Douglas Lake, Michigan, Aug. 1-3, 1923, H. B. Hungerford; para-
types as follows: 1 male and 6 females, Pelican Rapids, Minne-
sota, Aug. 22, 1922, H. B. Hungerford; 2 males and 6 females,
Bryant's Bog, Douglas Lake, Michigan, Aug. 3-17, 1923, H. B.
Hungerford; 5 males and 6 females, Bryant's Bog, Douglas Lake,

A Revision of the Genus Buenoa 1449

Michigan, July 29, 1923, H. B. Hungerford. The above series is in
the Francis Huntington Snow Entomological Collections, Univer-
sity of Kansas. Other paratypes collected by H. B. Hungerford are
located at University of Michigan, University of Minnesota, U. S.
National Museum, and in the following private collections: J. R.
de la Torre-Bueno (now at the University of Kansas), C. J. Drake,
W. E. Hoffmann, H. B. Hungerford (now in the Francis Huntington
Snow Entomological Collections), R. F. Hussey, and H. M. Parshley.
Data on Distribution: Recorded from Canada and United States.
In addition to the type series, specimens from the following localities
have been examined:

CANADA: Manitoba: Cowan, Aug. 7, 1937, H. T. Peters, 1 male
brachypterous.

Quebec: Aqueduc Lake, July 23, 1949, Howard Loeb, 1 male and
1 female brachypterous.

Nova Scotia: July 28, 1947, D. Livingston, 2 male brachypterous.
U.S.A.: Minnesota: Minnehaha Creek, Hennepin Co., July 9,
1921, H. B. Hungerford, 1 female brachypterous; Pelican Rapids,
Aug. 22, 1922, H. B. Hungerford, 1 female brachypterous; Benson,
Aug. 23, 1922, H. B. Hungerford, 5 males and 2 females brachyp-
terous.

Michigan: Cheboygan Co., July 27, 1918, R. F. Hussey, 3 females
brachypterous (U. of Mich.); Cheboygan Co., Aug. 14, 1918, R. F.
Hussey, 2 females brachypterous (U. of Mich.); Huron Co., Sand
Point, June 22, 1922, R. F. Hussey, 6 males and 4 females brachyp-
terous, 1 nymph (U. of Mich.); Douglas Lake, Bryant's Bog, July
17, 1923, H. B. Hungerford, 1 female brachypterous; Douglas Lake,
Bryant's Bog, Aug. 1, 1923, H. B. Hungerford, 2 females brachyp-
terous; Douglas Lake, Bryant's Bog, Aug. 3, 1923, H. B. Hunger-
ford, 11 females brachypterous; Douglas Lake, Bryant's Bog, July
30, 1924, H. B. Hungerford, 1 male brachypterous; Douglas Lake,
Aug. 6, 1924, H. B. Hungerford, 5 males and 43 females brachyp-
terous; Douglas Lake, Bryant's Bog, July 24, 1925, H. B. Hunger-
ford, 1 male and 9 females brachypterous; Douglas Lake, Bryant's
Bog, Aug. 8, 1925, H. B. Hungerford, 1 male and 9 females brachyp-
terous.

South Dakota: Weta, July 18, 1937, R. H. Beamer and C. L.
Johnston, 1 male macropterous, 1 male brachypterous.

All specimens listed above are in the Francis Huntington Snow
Entomological Collections, University of Kansas, Lawrence, Kansas,
unless otherwise indicated.

1450 The University Science Bulletin

Buenoa limnocastoris Hungerford
(PI. CII, fig. 8; pi. CIII, fig. 40; pi. CXIII, fig. 68)

1923. Buenoa limnocastoris Hungerford, H. B. Ent. News, vol. XXXIV, No. 5,
pp. 150-152.

1924. Buenoa limnocastoris, Hungerford, H. B. Ann. Ent. Soe. America, vol.
XVII, No. 2, pp. 223-227 (discussion of biology and taxonomy).

1925. Buenoa limnocastoris, Bare, C. O. Ent. News, vol. XXXVI, No. 8, p. 228
(key).

1926. Buenoa limnocastoris, Blatchley, W. S. Heteroptera or True Bugs of
Eastern North America, pp. 1057, 1059 and 1060 (key and description).

1928. Buenoa limnocastoris, Bare, C. O. Univ. Kansas Sci. Bull., vol. XVIII,
No. 3, p. 268 (key).

Size: This species varies considerably in size from its most
northern range, Quebec, Canada, to its most southern range, Florida.
Male, length 4.77 mm. to 6.17 mm., greatest body width 1.30 mm. to
1.62 mm.; female, length 5.00 mm. to 6.37 mm., greatest body width
1.50 mm. to 1.95 mm.

Color: General facies sordid white to testaceous. Head and
limbs testaceous to black. Prothorax sordid white with a smoky
to black area on each side above the margin; scutellum sordid
white to colorless; metathoracic dorsum and thoracic venter tes-
taceous with portions light brown to black. Abdominal dorsum
testaceous with a light brown to black area located posteriorly;
abdominal venter testaceous, with portions of connexivum black
or black with keel and portions of connexivum testaceous. Hemely-
tron sordid white with black band covering humeral angle and
extending along anterior margin of wing for approximately one
third its length and another large subtriangular black area at tip
of corium, extending across wing. This species somewhat variable
in color.

Male Structural Characteristics: As viewed from above, outline
of head rounded with lateral margins of vertex slightly indented;
greatest width of head approximately five and one half times the
anterior width of vertex and greater than humeral width of pro-
notum; synthlipsis approximately one third the anterior width of
vertex; along median longitudinal axis, head is slightly less than
half the length of pronotum; notocephalon slightly sulcate dorsally;
tylus inflated; labium very short, basal width approximately three
times its median length with apex bluntly rounded; rostral prong
(pi. CXIII, fig. 6Sb) distinctly longer than third rostral segment,
with base originating laterally at distal end of third rostral seg-
ment, and with apex bluntly rounded. Pronotum with its median
length equal to its humeral width; in lateral view, strongly arched
and inflated with a deep depression laterally near posterior margin;

A Revision of the Genus Buenoa 1451

disk with two shallow, elongate depressions toward the middle
and a shallow, subtriangular depression on each side, thus appear-
ing faintly tricarinate; lateral margins divergent; posterior margin
convex, medianly truncate. Scutellum small, elevated, but de-
pressed near anterior margin by a transverse groove; median length
much less than that of pronotum. Fore femur (pi. CXIII, fig. 68a)
widened but with anterior margin sharply curved at apex; obscure,
subtriangular stridulatory area consisting of approximately six or
seven sclerotized ridges. Fore tibia (pi. CXIII, fig. 68a) with
stridulatory comb (pi. CXIII, fig. 68c) consisting of approximately
forty to forty-five teeth; apical teeth thicker and narrower than
basal. Chaetotaxy of male front leg as shown on Plate CXIII.
Male genital claspers normal. Spine from caudo-sinistral margin
of seventh abdominal tergite normal (pi. CII, fig. 8), tapering
gradually from broad base to acuminate apex; length variable.

Macropterous forms are occasionally found. These specimens
have head narrower than humeral width of pronotum; pronotum
not as strongly inflated; scutellum much larger and without trans-
verse depression near anterior margin; hemelytra with claval su-
tures present and large membranes; fully developed flight wings.

Female Structural Characteristics: As viewed from above, outline
of head rounded with lateral margins of vertex slightly indented;
greatest width of head five to six times the anterior width of vertex
and usually equal to or greater than humeral width of pronotum;
synthlipsis approximately one third the anterior width of vertex;
along median longitudinal axis, head is approximately one half the
length of pronotum; notocephalon slightly sulcate dorsally; tylus
inflated. Pronotum with its median length approximately two thirds
its humeral width; not strongly arched or inflated as in male and
without lateral depression near posterior margin; disk usually faintly
tricarinate, occasionally with only median carina; lateral margins
divergent; posterior margin convex, medianly truncate. Scutellum
small, elevated, but depressed near anterior margin by a shallow
transverse groove; median length distinctly less than that of pro-
notum. Female ovipositor (pi. CIII, fig. 40) of normal shape with
teeth arranged in two longitudinal rows; one very short inner row
of few, large teeth and one long, irregular outer row of small teeth
with a few teeth located medianly and intermingled with the two
rows; one or two small, lateral, toothlike setae near apex.

Macropterous forms are occasionally found. These specimens
have head distinctly narrower than humeral width of pronotum;
pronotum wider and stronger; scutellum much larger with its me-

1452 The University Science Bulletin

dian length greater than that of pronotum and without transverse
depression near anterior margin; hemelytra with claval sutures
present and large membranes; fully developed flight wings.

Variation Within Species: As is indicated under measurements of
length, this species varies a great deal in size; there is also some
variation in the proportional size of the pronotum. The greatest
contrast is shown between several series from Canada, Minnesota,
and Michigan, and a series from Florida. The former is a large form,
the males seldom less than 6 mm. in length; the males of the latter
form are approximately 5 mm. in length.

Comparative Notes: Superficially this species somewhat re-
sembles B. macrotihialis Hungerford. Examination of the male,
however, will show distinct differences. This species differs from
B. macrotihialis in the shape and chaetotaxy of the fore femur and
tibia, the femoral stridulatory area, the shape and length of rostral
prong, and in having the pronotum strongly inflated. The latter
character, especially, will also separate this species from B. confusa
n. sp., which it also somewhat resembles.

Location of Types: Described from a series taken near Maple
Hill, Cook County, Minnesota, Aug. 12, 1922 by H. B. Hungerford.
Holotype male in University of Minnesota Collection, allotype fe-
male in the Francis Huntington Snow Entomological Collections,
University of Kansas, and paratypes at University of Minnesota,
University of Kansas, U. S. National Museum, and the following
private collections: J. R. de la Torre-Bueno (now at University of
Kansas), C. J. Drake, W. E. Hoffmann, H. B. Hungerford (now in
Francis Huntington Snow Entomological Collections ) , R. F. Hussey,
and H. M. Parshley.

Data on Distribution: Recorded from Canada and the United
States. In addition to type series, specimens from the following
localities have been examined:

CANADA: Quebec: Beattie Lake, Aug. 11, 1949, H. Loeb, 1
male and 1 female brachypterous.

U. S. A.: Minnesota: Cook County, Beaver Dam, Aug. 12, 1922,
H. B. Hungerford, 1 male and 1 female macropterous, 5 females
brachypterous; Cook Co., Beaver Dam, Aug. 12, 1922, W. E. Hoff-
mann, 3 male brachypterous; Pelican Rapids, Aug. 22, 1922, H. B.
Hungerford, 1 female macropterous.

Michigan: Cheboygan Co., July 29, 1918, 1 male brachypterous
(U. of Mich.); Cheboygan Co., Aug. 19, 1918, R. F. Hussey, 1 male
and 6 females brachypterous (U. of Mich.); Huron Co., Sand Point,
June 22, 1922, R. F. Hussey, 2 males and 1 female brachypterous, 1

A Revision of the Genus Buenoa 1453

nymph (U. of Mich.); Douglas Lake, June 29, 1923, H. B. Hunger-
ford, 1 male brachypterous; Douglas Lake, Sedge Point Pool, July
3-24, 1923, H. B. Hungerford, 20 males and 38 females brachyp-
terous; Douglas Lake, Bessey Creek, July 18, 1923, H. B. Hunger-
ford, 2 males brachypterous; Douglas Lake, Mud Lake, July 31,
1923, H. B. Hungerford, 1 female brachypterous; Douglas Lake,
Sedge Point Pool, Aug. 15, 1923, H. B. Hungerford, 1 male and 4
females brachypterous, 1 nymph; Douglas Lake, Bryant's Bog, Aug.
17, 1923, H. B. Hungerford, 1 female brachypterous; Douglas Lake,
Nickols pond, July 22, 1924, H. B. Hungerford, 2 females brachyp-
terous; Douglas Lake, July 27 to Aug. 8, 1924, H. B. Hungerford,
19 males and 45 females brachypterous; Douglas Lake, Bryant's Bog,
July 30, 1925, H. B. Hungerford, 1 female brachypterous; Douglas
Lake, July 29, 1926, H. B. Hungerford, 1 male macropterous, 1 male
and 2 females brachypterous.

Maine: Bridgton, Aug. 20, 1934, R. H. Beamer, 1 male brachyp-
terous.

New Jersey: P. R. Uhler Collection, 1 female brachypterous
(U.S.N.M.).

Virginia: Virginia Beach, Aug. 11, 1934, M. E. Griffith, 1 male
and 1 female brachypterous.

Georgia: Okefenokee Swamp, Aug. 3, 1934, M. E. Griffith, 1 male
macropterous; Okefenokee Swamp, July 27, 1931, J. D. Beamer, 1
male brachypterous.

Florida: Ponce de Leon, July 13, 1934, R. H. Beamer, 1 male and
1 female macropterous, 5 males and 12 females brachypterous;
Wakulla Springs, July 14, 1934, P. A. McKinstry and R. H. Beamer,
1 male and 1 female macropterous.

All specimens listed above are in the Francis Huntington Snow
Entomological Collections, University of Kansas, Lawrence, Kansas,
unless otherwise indicated.

Buenoa confusa n. sp.
(PI. CII, fig. 20; pi. CHI, fig. 34; pi. CXIV, fig. 69)

1894. Anisops elegans, Uhler, P. R. Proc. California Acad. Sci., vol. IV, p. 293

( distributional note ) .
1894. Anisops elegans, Uhler, P. R. Proc. Zoological Soc. London, p. 223

(descriptive and distributional note).

1908. Buenoa elegans, Torre-Bueno, J. R. de la. Jr. New York Ent. Soc, vol.

XVI, p. 238 (listed).

1909. Buenoa elegans, Kirkaldy, G. W. and Torre-Bueno, J. R. de la. Proc.
Ent. Soc. Washington, vol. X, p. 200 (catalogue).

1909. Buenoa elegans, Torre-Bueno, J. R. de la. Jr. New York Ent. Soc, vol.

XVII, pp. 75-77 (key and notes).

1910. Buenoa elegans, Smith, J. B. Catalogue Insects New Jersey, edn. 3,
p. 170.

1454 The University Science Bulletin

1912. Buenoa elegans, Torre-Bueno, J. R. de la. Canadian Ent, vol. XLIV,

p. 213 (listed).
1914. Buenoa elegans, Parshley, H. M. Psyche, vol. XXI, p. 140 (listed).

1916. Buenoa elegans, Van Duzee, E. P. New York Ent. Soc., p. 51 (check
list).

1917. Buenoa elegans, Hungerford, H. B. Ent. News, vol. XXVIII, p. 176
(key).

1917. Buenoa elegans, Parshley, H. M. Occasional Papers Boston Soc. Nat.

Hist., vol. VII, p. 113 (listed).
1917. Buenoa elegans, Van Duzee, E. P. Catalogue Hemiptera America North

of Mexico, p. 455.
1919. Buenoa elegans, Hungerford, H. B. Univ. Kansas Sci. Bull., vol. XI,

pp. 173-176 (description and key).
1919. Buenoa elegans, Hungerford, H. B. Univ. Kansas Sci. Bull, vol. XI,

p. 332 (figure).
1921. Buenoa elegans, Parshley, H. M. Proc. British Columbia Ent. Soc,

No. 18, p. 24.
1923. Buenoa elegans, Torre-Bueno, J. R. de la. Guide to Insects of Con-
necticut, part 4, p. 407.
1923. Buenoa elegans, Hungerford, H. B. Ent. News, vol. XXXIV, pp. 150-

152 (notes).

1923. Buenoa elegans, Torre-Bueno, J. R. de la. Connecticut State Geol. and
Nat. Hist. Survey Bull., No. 34, p. 407 (key and notes).

1924. Buenoa elegans, Hungerford, H. B. Ann. Ent. Soc. America, vol. XVII,
pp. 225-226 (notes).

1925. Buenoa elegans, Hungerford, H. B. and Beamer, R. H. Ent. News, vol.
XXXVI, pp. 272 and 297 (notes).

1925. Buenoa elegans, Bare, C. O. Ent. News, vol. XXXVI, pp. 225 and 228
(key and note).

1926. Buenoa elegans, Bare, C. O. Ann. Ent. Soc. America, vol. XIX, pp. 96-
97 ( biological note ) .

1926. Buenoa elegans, Blatchley, W. S. Heteroptera or True Bugs of Eastern

North America, pp. 1057 and 1059 (key and description).
1928. Buenoa elegans, Torre-Bueno, J. R. de la. Cornell Univ. Agricultural

Exp. Station, Memoir 101, p. 139 (listed).
1928. Buenoa elegans, Bare, C. O. Univ. Kansas Sci. Bull., vol. XVIII, pp. 268

and 241-243 (key and figures).
1940. Buenoa elegans, Hungerford, H. B. Ent. Monthly Magazine, vol.

LXXVI, p. 256 (note).
1942. Buenoa elegans, Rice, L. A. Tennessee Acad. Sci., vol. XVII, pp. 55

and 63 (notes).
1942. Buenoa elegans, Hutchinson, G. E. American Jr. Sci., vol. CCXL,

pp. 335-338 (notes and figures).
1948. Buenoa elegans, Hynes, H. B. N. Trans. Royal Ent. Soc. London, vol.

XCIX, p. 354 (distributional note).

Size: This species varies considerably in size from its most
northern range, Manitoba, Canada, to its most southern range,
Florida and the Cayman Islands. Male, length 4.16 mm. to 5.78
mm., greatest body width 1.10 mm. to 1.49 mm.; female, length
4.29 mm. to 7.00 mm., greatest body width 1.30 mm. to 1.62 mm.

Color: General facies sordid white to pale testaceous. Head,
pronotum, scutellum, most of thoracic venter, and limbs sordid
white to testaceous. Metathoracic dorsum usually sordid white
with a light brown to black, longitudinal stripe on each side. Ab-
dominal venter light brown to black except keel and portions of
connexivum, pale testaceous; abdominal dorsum usually sordid

A Revision of the Genus Buenoa 1455

white to testaceous with a lateral, light brown to black area near
posterior end. Hemelytron hyalin, usually with a black band
covering humeral angle and extending along anterior margin of
wing for approximately one fourth its length, and a light brown to
black area at tip of corium. This species somewhat variable in
color.

Male Structural Characteristics: As viewed from above, outline
of head rounded with anterior margin of vertex continuous with
that of eyes; head large with eyes prominent; greatest width of
head seven to seven and one-half times the anterior width of vertex
and greater than humeral width of pronotum; synthlipsis narrow,
approximately one third the anterior width of vertex; along median
longitudinal axis, head is five sevenths to six sevenths the length
of pronotum; notocephalon narrow, slightly sulcate dorsally; tylus
inflated; labrum with basal width approximately twice its median
length and apex bluntly rounded; rostral prong ( pi. CXIV, fig. 69b )
distinctly longer than third rostral segment, with base originating
laterally at distal end of third rostral segment, and with apex
bluntly rounded. Pronotum with its median length approximately
two thirds its humeral width; disk with two elongate depressions
toward the middle and a large subtriangular depression on each
side, thus appearing tricarinate; lateral margins divergent; pos-
terior margin convex, medianly truncate. Scutellum elevated but
depressed near anterior margin by a transverse groove; median
length less than that of pronotum. Fore femur (pi. CXIV, fig. 69a)
wide, somewhat thickened at apex; obscure, subtriangular stridula-
tory area consisting of approximately five or six sclerotized ridges.
Fore tibia (pi. CXIV, fig. 69a) with stridulatory comb (pi. CXIV,
fig. 69c) consisting of forty to fifty teeth; apical teeth thicker and
narrower than basal. Chaetotaxy of male front leg as shown on
Plate CXIV. Male genital claspers (pi. CII, fig. 20) normal. Spine
from caudo-sinistral margin of seventh abdominal tergite normal,
tapering gradually from broad base to acuminate apex; length
variable.

Macropterous forms are occasionally found. These specimens
have head distinctly narrower than humeral width of pronotum;
pronotum with lateral margins more divergent; scutellum larger;
hemelytra with claval sutures present and large membranes; fully
developed flight wings.

Female Structural Characteristics: As viewed from above, out-
line of head rounded with anterior margin of vertex continuous
with that of eyes; eyes prominent; greatest width of head six to

62—6490

1456 The University Science Bulletin

six and one half times the anterior width of vertex and slightly
greater than humeral width of pronotum; synthlipsis narrow, ap-
proximately one third the anterior width of vertex; along median
longitudinal axis, head is approximately five sixths the length of
pronotum; notocephalon narrow, slightly sulcate dorsally; tylus
slightly inflated. Pronotum with its median length approximately
three fifths its humeral width; disk usually with two shallow, elon-
gate depressions toward the middle and a shallow, subtriangular
depression on each side, thus appearing faintly tricarinate; lateral
margins divergent; posterior margin convex, medianly truncate.
Scutellum slightly elevated but depressed near anterior margin by
a shallow, transverse groove; median length equal to or slightly
greater than that of pronotum. Female ovipositor ( pi. CIII, fig. 34 )
of normal shape with teeth arranged in three longitudinal rows;
one very short inner row of few, large teeth, one median row of
normal teeth, and one long outer row of smaller teeth which merges
proximally with median row; approximately four or five small,
lateral, toothlike setae near apex.

Macropterous forms are occasionally found. These specimens
have head distinctly narrower than humeral width of pronotum;
pronotum wider and stronger with lateral margins more divergent;
scutellum much larger with its median length distinctly greater
than that of pronotum and without transverse depression near
anterior margin; hemelytra with claval sutures present and large
membranes; fully developed flight wings.

Variation Within Species: As is indicated under measurements
of length, this species varies a great deal in size; there is also some
variation in the proportional size of the pronotum and head. The
greatest contrast is shown between two series from Manitoba,
Canada, and Storrs, Connecticut, and several series from Florida and
the Cayman Islands. The former is a large form, the male seldom
less than 5.75 mm. in length; the males of the latter form are ap-
proximately 4.50 mm. in length. Due to the fact that all characters
used in species determination are identical in these two forms, no
specific separation appears justified. It is the opinion of this author
that the immediate environment, including such factors as food
source, temperature of water, salinity of water, etc., are responsible
for these variations.

Comparative Notes: Superficially this species resembles B. mac-
rotibialis Hungerford. Examination of the male, however, will
show distinct differences. This species differs from B. rnacrotibialis
in having a narrower synthlipsis, distinctly shorter pronotum, a

A Revision of the Genus Buenoa 1457

much less robust front leg, and differences in the femoral stridula-
tory area and rostral prong.

Nomenclatorial Notes: One finds in the literature and collections
that this species has been masquerading under the name Buenoa
elegans (Fieber). On examining notes and sketches accumulated
on Fieber's type (Berlin Museum) by Dr. H. B. Hungerford in
1928, the validity of B. elegans appeared to be questionable.
Through the kindness of Dr. S. v. Keler of the Berlin Museum, the
type specimen, a male, was made available to me for study. Anisops
elegans Fieber is a good Anisops species. The Buenoa known to a
generation of entomologists as Buenoa elegans ( Fieber ) was, there-
fore, an undescribed species. The confusion surrounding this spe-
cies serves to illustrate the necessity for the worker to have access
to type material. The A. elegans type was sent to Dr. George T.
Brooks who determined the specimen as A. apicalis Stal, an African
species. In Fieber's original description, he gives the locality as
"Aus Amerika." The type is labeled "Brasil Coll. Germ." Due to
the fact that Anisops is an Old World genus and the type has been
determined A. apicalis, an African species, it would seem that the
label on Fieber's type specimen is in error.

Location of Types: Holotype male, allotype female, 23 male and
24 female paratypes, Douglas Co., Kansas, Nov. 3-7, 1922, H. B.
Hungerford and Robert Guntert. The type series is in the Francis
Huntington Snow Entomological Collections, University of Kansas.

Data on Distribution: Recorded from Canada, United States,
Mexico (fide Uhler), and West Indies (Grand Cayman, Grenada).
In addition to type series, specimens from the following localities
have been examined:

CANADA: Alberta: Red Deer R., Aug. 3, 1937, H. T. Peters, 4
males and 5 females brachypterous.

Manitoba: Cowan, Aug. 7, 1937, R. H. Beamer, H. T. Peters, and
C. L. Johnston, 12 males and 7 females macropterous.

U.S.A.: North Dakota: Fargo, July 26, 1937, C. L. Johnston, 1
male macropterous.

South Dakota: Weta, July 18, 1937, R. H. Beamer, H. T. Peters,
and C. L. Johnston, 1 male and 1 female brachypterous, 7 males and
4 females macropterous; Waubay, Blue Dog Lake, Sept. 14, 1939,
H. C. Severin, 1 male and 1 female brachypterous.

Minnesota: Cook Co., Aug. 12, 1922, H. B. Hungerford, 2 males
brachypterous, 1 female macropterous; Benson, Aug. 23, 1922, H. B.
Hungerford, 4 males and 2 females brachypterous.

Michigan: Cheboygan Co., July 27 to Aug. 14, 1918, R. F. Hus-

1458 The University Science Bulletin

sey, 2 males and 3 females brachypterous ( U. of Mich. ) ; Cheboygan
Co., July 27, 1936, H. B. Hungerford, 1 male and 1 female brachyp-
terous; Berrien Co., June 30, 1919, R. F. Hussey, 2 males and 1
female brachypterous (U. of Mich.); Berrien Co., Sept. 2, 1919,
R. F. Hussey, 5 males and 5 females brachypterous ( U. of Mich. ) ;
Ann Arbor, July 30, 1921, R. F. Hussey, 1 male and 1 female
brachypterous (U. of Mich.); Douglas L., Bryant's Bog, July 3-31,
1923, H. B. Hungerford, 28 males and 18 females brachypterous,
2 nymphs; Douglas L., Bryant's Bog, Aug. 1-17, 1923, H. B. Hunger-
ford, 36 males and 57 females brachypterous, 2 nymphs; Douglas L.,
Bryant's Bog, July 30, 1924, H. B. Hungerford, 1 male brachypterous;
Douglas L., Bryant's Bog, H. B. Hungerford, July 24, 1925, 9 males
and 15 females brachypterous; Douglas L., Bryant's Bog, Aug. 12,
1925, H. B. Hungerford, 7 males and 6 females brachypterous;
Douglas L., Sedge Point Pool, Aug. 15, 1923, 1 male brachypterous.

New York: Huntington, May, 1909, Bueno Collection, 1 male
brachypterous; White Plains, Sept. 28, 1924, J. R. de la Torre-Bueno,
1 male and 1 female brachypterous.

Connecticut: Storrs, Aug. 13, 1946, R. H. Beamer, 6 males and
7 females brachypterous.

New Jersey: Ft. Lee Dist, Sept. 18, 1904, 1 male and 2 females
brachypterous; Rancoca, Aug. 29, 1927, E. M. Benton, 2 females
brachypterous, 1 nymph.

District of Columbia: Washington, Aug. 20, 1893, P. R. Uhler, 1
female brachypterous (U.S.N.M.).

Kansas: Douglas Co., Nov. 3-7, 1922, H. B. Hungerford and
Robert Guntert, 1 male and 5 females brachypterous, 2 nymphs;
Douglas Co., Nov. 3, 1922, H. B. Hungerford, 1 male brachypterous;
Douglas Co., Sept. 28, 1924, C. O. Bare, 1 male brachypterous;
Cherokee Co., Larson's Creek, Aug. 18, 1923, R. H. Beamer, 8
males and 10 females brachypterous; Atchison Co., July 15, 1924,
E. P. Breaky, 1 female brachypterous; Doniphan Co., July 20, 1924,
R. H. Beamer, 2 males brachypterous.

Virginia: Great Falls, Jan. 9, 1906, D. H. Clemons, 1 female
macropterous (U. S. N. M.)

Texas: Cedar Lane, Aug. 9, 1928, R. H. Beamer, 1 male and 1
female macropterous; Orange Co., Aug. 14, 1928, R. H. Beamer,
1 female macropterous; Rockport, Jan. 1, 1946, L. D. Beamer, 2
males brachypterous.

Louisiana: Baton Rouge, June, 1905, A. W. Morril, 1 female
macropterous (U.S.N.M.); Creole, June 18, 1948, E. L. Todd,
7 males and 26 females macropterous.

A Revision of the Genus Buenoa 1459

Mississippi: Shuqualak, July 16, 1930, Paul W. Oman, 1 male
brachypterous; Bay St. Louis, July 9, 1934, R. H. Beamer, 3 females
macropterous.

Alabama: Grand Bay, July 11, 1934, R. H. Beamer and P. A. Mc-
Kinstry, 3 males and 4 females brachypterous.

Georgia: Okefenokee Swp., July 30, 1934, J. D. Beamer, M. E.
Griffith, and P. McKinstry, 4 males and 15 females brachypterous,
1 male and 2 females macropterous; Okefenokee Swp., Aug. 3,
1934, R. H. Beamer, 26 males and 15 females brachypterous, 1 male
macropterous, 3 nymphs; Okefenokee Swp., July 25-27, 1939, R. H.
and J. D. Beamer and E. G. Wegenek, 5 males and 9 females
brachypterous, 4 males and 2 females macropterous, 1 nymph;
Okefenokee Swp., Aug. 1, 1939, J. D. Beamer, 1 female brachyp-
terous.

Florida: Archer, July 31, 1930, R. H. Beamer, Paul W. Oman,
and J. Nottingham, 6 males and 12 females brachypterous, 7 males
and 11 females macropterous; Yankeetown, July 31, 1930, R. H.
Beamer, J. Nottingham, and L. D. Tuthill, 3 males and 3 females
brachypterous, 3 males and 3 females macropterous; Inverness,
Aug. 1, 1930, Paul W. Oman and J. Nottingham, 3 males and 4 fe-
males brachypterous, 1 male and 3 females macropterous; Wild-
wood, Aug. 2, 1930, R. H. Beamer, Paul W. Oman, and L. D. Tut-
hill, 34 males and 29 females brachypterous, 3 males and 2 females
macropterous; Sanford, Aug. 4, 1930, R. H. Beamer and Paul W.
Oman, 26 males and 17 females brachypterous, 1 male and 2 fe-
males macropterous, 3 nymphs; Cocoanut Grove, Aug. 9, 1930,
R. H. Beamer and Paul W. Oman, 14 males and 10 females brachyp-
terous, 2 males and 5 females macropterous, 8 nymphs; Ft. Mead,
Aug. 13, 1930, Paul W. Oman, 4 males and 3 females brachypterous,
3 nymphs; Plant City, Aug. 15, 1930, Paul W. Oman, 1 female
brachypterous; Plant City, July 14, 1939, P. B. Lawson and R. H.
Beamer, 6 males and 4 females macropterous; Hilliard, July 28,
1934, R. H. Beamer, 2 males and 3 females macropterous; Old
Town, July 11, 1939, P. B. Lawson, 1 female macropterous; La Belle,
July 16, 1939, P. B. Lawson and R. H. Beamer, 6 males and 3 fe-
males macropterous; Lacoochee, Aug. 9, 1939, J. D. Beamer, 1 fe-
male brachypterous, 1 female macropterous.

WEST INDIES: Grand Cayman: Georgetown, Apr. 17, 1926,
C. B. Lewis and G. H. Thompson, 1 male and 2 females macrop-
terous.

All specimens listed above are in the Francis Huntington Snow
Entomological Collections, University of Kansas, Lawrence, Kansas,
unless otherwise indicated.

1460 The University Science Bulletin

Buenoa fuscipennis (Berg)
(PI. CII, fig. 7; pi. CIH, fig. 28; pi. CXIV, fig. 71)

1879. Anisops fuscipennis Berg, C. Hemiptera Argentina, pp. 198-199 (de-
scribed from Argentina).

1899. Anisops naias Kirkaldy, G. W. Entomologist, vol. XXXII, p. 194 (de-
scribed from Chile). New Synonomy.

1904. Buenoa naias, Kirkaldy, G. W. Wiener Ent. Zeit., vol. XXIII, pp. 120
and 134 (listed). New Synonomy.

1904. Buenoa fuscipennis, Kirkaldy, G. W. Wiener Ent. Zeit., vol. XXIII,
pp. 120 and 134 (listed).

1909. Buenoa naias, Kirkaldy, G. W. and Torre-Bueno, J. B. de la. Proc. Ent.
Soc. Washington, vol. X, p. 200 (catalogue).

1909. Buenoa fuscipennis, Kirkaldy, G. W. and Torre-Bueno, J. B. de la. Proc.
Ent. Soc. Washington, vol. X, p. 200 (catalogue).

1928. Buenoa dentipes Jaczewski., T. Ann. Musei Zoologici Polonici, vol. VII,
pp. 127-129 (described from Brazil). New Synonomy.

1935. Buenoa naias, De Carlo, J. A. Bevista Chilena de Historia Natural,
p. 110 (catalogue).

Size: Male, length 6.82 mm. to 7.21 mm., greatest body width
1.82 mm. to 2.01 mm.; female, length 7.02 mm. to 7.28 mm., greatest
body width 1.95 mm. to 2.08 mm.

Color: General facies testaceous to fuscous. Head, thoracic dor-
sum, portions of thoracic venter, abdominal dorsum, and limbs,
testaceous. Portions of thoracic venter and abdominal dorsum
light brown to black. Abdominal venter black except ventral keel,
posterior margins of segments, and portions of connexivum, tes-
taceous.

Male Structural Characteristics: As viewed from above, outline
of head rounded with lateral margins of vertex indented; greatest
width of head approximately five times the anterior width of vertex
and distinctly less than humeral width of pronotum; synthlipsis
slightly less than half the anterior width of vertex; along median
longitudinal axis, head is approximately half the length of pro-
notum; tylus inflated; labrum short, basal width approximately a
third greater than its median length with apex moderately rounded;
rostral prong (pi. CXIV. fig. 71b) shorter than third rostral seg-
ment, with base originating laterally at proximal end of third rostral
segment, and with apex bluntly rounded. Pronotum with its me-
dian length slightly more than half its humeral width; disk unim-
pressed; lateral margins divergent; posterior margin convex, me-
dianly concave. Scutellum large, with median length distinctly
greater than that of pronotum. Trochanter of fore leg with promi-
nent tubercle on posterior margin. Fore femur (pi. CXIV, fig. 71a)
neither wide nor thickened at apex; without stridulatory area but
with row of short, thick setae on posterior margin of inner surface.
Fore tibia (pi. CXIV, fig. 71a) with stridulatory comb (pi. CXIV,

A Revision of the Genus Buenoa 1461

fig. 71c) consisting of approximately eleven thick teeth of uniform
size. Chaetotaxy of male front leg as shown on Plate CXIV. Inter-
mediate tibia with row of short, thick setae located parallel with
inner, anterior margin in apical half. Male genital claspers (pi.
CIII, fig. 28) abnormal in shape. Spine from caudo-sinistral mar-
gin of seventh abdominal tergite (pi. II, fig. 7) short and thick,
tapering gradually from broad base to acuminate apex.

Female Structural Characteristics: As viewed from above, out-
line of head rounded with vertex slightly protuberant; greatest
width of head approximately four and one half times the anterior
width of vertex and less than humeral width of pronotum; synthlip-
sis approximately half the anterior width of vertex; along median
longitudinal axis, head is more than half the length of pronotum;
tylus slightly inflated. Pronotum with its median length approxi-
mately half its humeral width; disk unimpressed; lateral margins
divergent; posterior margin convex, medianly concave. Scutellum
large with median length distinctly greater than that of pronotum.
Female ovipositor of normal shape with teeth arranged in three
longitudinal rows; one short inner row of larger teeth, one outer
row of smaller teeth, and one median row of relatively large teeth
which originates in apical fourth of valve and intermingles with
outer row proximally; approximately eight or nine small, lateral,
toothlike setae near apex in an irregular row.

Comparative Notes: Superficially this species somewhat resem-
bles B. margaritacea Torre-Bueno and B. scimitra Bare. Examina-
tion of the male, however, will show distinct differences. This spe-
cies differs from B. margaritacea and B. scimitra in having a tubercle
on the trochanter of the front leg, in having a row of short, thick
setae on the intermediate tibia, and in lacking a stridulatory area
on the femur of the front leg.

Nomenclatorial Notes: A small series of specimens from Argen-
tina, compared with the types of B. fuscipennis by Dr. J. A. De
Carlo, assisted the author to conclude that B. naias Kirkaldy and
B. dentipes Jaczewski are identical with this species. Buenoa naias
and B. dentipes are hereby placed in synonomy with B. fuscipennis
(Berg) 1879.

Location of Types: The original type series from Argentina, is
located at the Museo de La Plata, Argentina. Homotype male,
"Prov. Buenos Aires, Oct. 27, 1898. S. Venturi," compared with the
type by Dr. J. A. De Carlo of the Museo Argentino de C. Naturales,
is now in the Francis Huntington Snow Entomological Collections,
University of Kansas.

1462 The University Science Bulletin

Data on Distribution: Recorded from Venezuela (fide Kirkaldy
& Bueno ) , Brazil, Bolivia, Chile, Paraguay, Uruguay, and Argentina.
Specimens from the following localities have been examined:

BRAZIL: Nova Teutonia, May 1948, Fritz Plaumann, 1 male.

BOLIVIA: Villa Montes, Nov. 1917, 1 female.

CHILE: Icario-Rio Salado, Santiago del Estero, 1865, 1 male, 1
female; Vina del Mar, Nov. 15, 1920, 3 females; Vina, 1921, Alfredo
Faz, 1 female; Santiago, Quinto Normal, Aug. 30, 1922, Alfredo
Faz, 20 males, 28 females; Quinta Normal, Oct. 26, 1922, Alfredo
Faz, 1 male, 3 females; Santiago, Laguna de la Quinta Normal,
Dec. 20, 1922, Alfredo Faz, 28 males, 24 females; Santiago, Sept. 2,

1923, Alfredo Faz, 16 males, 10 females; Santiago, Alfredo Faz, 6
males, 4 females; Termas Cauquenes, Dec. 15, 1922, Alfredo Faz,
217 males, 186 females.

PARAGUAY: Villarrica, Jan. 7, 1923, Fran. Schade, 11 females;
Villarrica, June 6-15, 1923, Fran. Schade, 8 males, 10 females; Vil-
larica, Dec. 15, 1923, Fran. Schade, 1 female; Villarrica, Sept. 20,

1924, Fran. Schade, 2 males, 4 females; Villarrica, July 8, 1924,
Fran. Schade, 1 male; Villarrica, Sept. 26, 1931, Fran. Schade, 2
males, 1 female; Estero Grande, Nov. 1, 1924, Fran. Schade, 1
female.

URUGUAY: Maldonado, 1885, C. Darwin, 2 females (British
Mus.); Paso de arriera, Rivera, Jan. 13, 1933, C. S. Carbonell, 1 male
(U.S.N.M.).

ARGENTINA: Argentina, Apr. 13, 1897, G. Wiengreen, 1 female
(Hamburg Mus.); Prov. Buenos Aires, Oct. 27, 1898, S. Venturi,
1 male, 1 female; Buenos Aires, Oct. 15, 1920, P. Frank, 2 males,
1 female, 1 nymph ( Hamburg Mus. ) ; Cordoba, June 5, 1907, 2 males;
Dept. de Lujan San Luis, Sept. 1934, 2 males; Carcarana, 1 male;
Esperanza, Bred., 1 female (Kirkaldy Coll.); Salta, 1 male, 1 female.

All specimens listed above are in the Francis Huntington Snow
Entomological Collections, University of Kansas, Lawrence, Kansas,
unless otherwise indicated.

Buenoa amnigenus (White)
(PI. CIII, fig. 36; pi. CXIV, fig. 70)

1879. Anisops amnigenus White, F. B. Trans. Ent. Soc. London, Pt. IV,

p. 271.
1904. Buenoa amnigenus, Kirkaldy, G. W. Wiener Ent. Zeit., voh XXIII,

pp. 120 and 134 (listed and states "die Type ist verloren gegangen").
1909. Buenoa amnigenus, Kirkaldy, G. W. and Torre-Bueno, J. R. de la. Proc.

Ent. Soc. Washington, vol. X, No. 3-4, p. 200 (listed).

Size: Male, length 4.77 mm. to 5.39 mm., greatest body width

A Revision of the Genus Buenoa 1463

1.26 mm. to 1.40 mm.; female, length 4.81 mm. to 5.85 mm., greatest
body width 1.37 mm. to 1.54 mm.

Color: General facies sordid white. Head, thorax, and abdomi-
nal dorsum testaceous. Limbs testaceous to brown. Abdominal
venter black with connexivum light brown to black.

Male Structural Characteristics: As viewed from above, outline
of head rounded with vertex protuberant; greatest width of head
approximately five times the anterior width of vertex and less
than humeral width of pronotum; inner margins of eyes contiguous
posteriorly; along median longitudinal axis, head is approximately
two thirds the length of pronotum; frons wide; tylus not inflated;
labrum very short, basal width more than twice its median length
with apex bluntly rounded; rostral prong (pi. CXIV, fig. 70c)
slightly longer than third rostral segment, with base originating
laterally at proximal end of third rostral segment, and with apex
sharply rounded. Pronotum with its median length slightly more
than half its humeral width; disk unimpressed; lateral margins di-
vergent; posterior margin convex, medianly concave. Scutellum
large, with median length greater than that of pronotum. Fore
femur (pi. CXIV, fig. 70a) neither wide nor greatly thickened at
apex; stridulatory area absent. Fore tibia (pi. CXIV, fig. 70a) with
stridulatory comb (pi. CXIV, fig. 70b) consisting of approximately
twenty-eight to thirty teeth which increase in height from base to
apex. Chaetotaxy of male front leg as shown on Plate CXIV. Male
genital claspers normal. Spine from caudo-sinistral margin of
seventh abdominal tergite tapering, with apical half very narrow
and apex acuminate.

Brachypterous forms are occasionally found. These specimens
have pronotum narrower, scutellum smaller, and hemelytra that
lack claval sutures.

Female Structural Characteristics: As viewed from above, outline
of head rounded with vertex slightly protuberant; greatest width of
head approximately four times the anterior width of vertex and less
than humeral width of pronotum; synthlipsis very narrow, approxi-
mately one twentieth the anterior width of vertex; along median
longitudinal axis, head is more than half the length of pronotum;
frons wide; tylus not inflated. Pronotum with its median length
approximately half its humeral width; disk unimpressed; lateral
margins divergent; posterior margin convex, medianly concave.
Scutellum large with median length distinctly greater than that
of pronotum. Female ovipositor (pi. CIII, fig. 36) of normal shape

1464 The University Science Bulletin

with teeth arranged in three longitudinal rows; one inner row of
large teeth, one short outer row of smaller teeth, and one median
row of relatively large teeth which originates in apical fourth of
valve rather than at apex; approximately seven small, lateral, tooth-
like setae near apex.

Brachypterous forms are occasionally found. These specimens
have pronotum narrower, scutellum smaller, and hemelytra that lack
claval sutures.

Comparative Notes: This species is nearest in general appearance
to B. gracilis n. sp. Examination of the male, however, will show
distinct differences. This species differs from B. gracilis in having
the eyes contiguous posteriorly in the male, in lacking the stridula-
tory area on the fore femur, and in having the frons immediately
above the tylus wide.

Location of Types: Dr. G. W. Kirkaldy (1904) states that "die
Type ist verloren gegangen", however, Dr. H. B. Hungerford in
1928, examined the holotype male at the British Museum, London.
Dr. Hungerford states that there are two specimens, male and fe-
male, which belong to the Perthshire Museum, Scotland. The holo-
type male bears the label "Manaos, on board at light, Aug. 1875".
Homotype male, compared with type by Dr. H. B. Hungerford of
the University of Kansas, labeled "Paraguy S. A., Villarrica, Loma,
7/6/23, Fran. Schade", now in the Francis Huntington Snow
Entomological Collections, University of Kansas.

Data on Distribution: Recorded from British Guiana, Brazil,
Peru, Bolivia, and Paraguay. Specimens from the following locali-
ties have been examined:

BRITISH GUIANA: Georgetown Demerara, Botanic Gardens,
July 26, 1932, S. Harris, 1 female brachypterous.

BRAZIL: Amazonas: Manaos, Manacapuru, Mar. 1928, S. M.
Klages, 14 males and 26 females macropterous, 10 males and 28
females brachypterous; Manaos region, Rio Negro, Oct. 1935, A. M.
Olalla, 60 males and 36 females macropterous, 18 males and 11
females brachypterous, 2 nymphs; Manaos, July 29, 1924, Flores
(Bueno Coll.), 3 males and 1 female macropterous; Reg. de Itacoa-
tiara, Jan. 3, 1936, A. M. Olalla, 35 males and 19 females macrop-
terous; Reg. de Itacoatiara (Nrte.), Jan., 1936 to Apr., 1936, A. M.
Olalla, 8 females macropterous; Vic. Joao Pessoa (Sao Phelipe),
River Jurua, July 10, 1936 to Sept. 20, 1936, A. M. Olalla, 28 males
and 44 females macropterous, 14 males and 13 females brachyp-
terous; Castanha region, Rio Purus, Sept., 1935, A. M. Olalla, 18

A Revision of the Genus Buenoa 1465

males and 16 females macropterous, 8 males and 30 females brachyp-
terous; Vic. Santo Antonio, River Eiru, Sept. 25, 1936 to Oct. 7,

1936, A. M. Olalla, 27 females macropterous, 8 males and 12 fe-
males brachypterous.

Ceard: Bom Acud Successo nr. Maranguape, Nov. 4, 1937, Still-
man Wright, 3 females brachypterous; Artificial Lake nr. Russas,
July 22, 1937, Stillman Wright, 6 females macropterous, 1 male and
3 females brachypterous; Roadside pool nr. Primavera, Oct. 28,

1937, Stillman Wright, 1 female brachypterous; Roadside pool nr.
Sobral, Oct. 26, 1937, Stillman Wright, 1 female macropterous;
Pacatuba, Acude Piripau, Aug. 1937, Stillman Wright, 1 male and
1 female brachypterous.

Rio Grande do Norte: Caico, No. 327, Stillman Wright, 2 males
and 1 female macropterous, 3 males and 11 females brachypterous.

Parahiba: Souza, Stillman Wright, 2 females macropterous, 1
female brachypterous; Alagoa do Monteiro, Stillman Wright, 2 fe-
males macropterous, 18 females brachypterous; Areia, Stillman
Wright, 10 females brachypterous; Santa Luzia, Stillman Wright, 1
female macropterous, 7 females brachypterous.

Pernambuco: Vila Bela, No. 5467, Stillman Wright, 3 males
brachypterous.

Motto Grosso: Corumba, March, Lowland, 1 male macropterous.

Parana: Lago Grande, Feb. 1939, A. M. Olalla, 4 females macrop-
terous, 24 males and 23 females brachypterous.

PERU: Dept. Huanuco, Loc. Shapajilla, July, 1938, F. Woytkow-
ski, 5 males brachypterous.

BOLIVIA: Victoria, R. Beni, Junction of Madre de Dios and
Beni rivers, Oct., 1937, A. M. Olalla, 1 male and 2 females macrop-
terous; El Consuelo, R. Beni, Jan., 1938, A. M. Olalla, 4 males and
10 females macropterous, 12 males and 16 females brachypterous;
Las Pampas, R. Beni, Mojos, April, 1938, A. M. Olalla, 1 male and
1 female macropterous, 7 males and 3 females brachypterous; Las
Pampas, R. Beni, Mojos, May, 1938, A. M. Olalla, 1 male and 1 fe-
male macropterous, 8 males and 3 females brachypterous; Santa
Ana del Yacuma, Feb., 1938, A. M. Olalla, 1 male macropterous, 2
males and 2 females brachypterous; Prov. del Sara, Nov. 30, 1912,
Steinbach, 1 male macropterous.

PARAGUAY: Villarrica, Loma, July 6, 1923, Fran. Schade, 1 male
macropterous.

All specimens listed above are in the Francis Huntington Snow
Entomological Collections, University of Kansas, Lawrence, Kansas.

1466 The University Science Bulletin

Buenoa incompta n. sp.
(PL CXIV, fig. 73)

Size: Male, length 5.00 mm. to 5.20 mm., greatest body width
1.30 mm. to 1.36 mm.; female, length 5.39 mm. to 5.78 mm., greatest
body width 1.36 mm. to 1.49 mm.

Color: General facies sordid white to pale testaceous. Head and
pronotum sordid white. Thoracic venter and limbs sordid white
to brown. Scutellum sordid white, often with two anterolateral
brown areas; metathoracic dorsum pale testaceous to light brown.
Abdominal venter black except keel and portions of connexivum
sordid white; abdominal dorsum yellowish white with irregular
areas of black.

Male Structural Characteristics: As viewed from above, outline
of head rounded with anterior margin of vertex continuous with
that of eyes; greatest width of head approximately five and one
half times the anterior width of vertex and less than humeral width
of pronotum; synthlipsis very narrow, approximately one fifth the
anterior width of vertex, often carinate; along median longitudinal
axis, head is one half to three fifths the length of pronotum; noto-
cephalon slightly sulcate dorsally; tylus slightly inflated; labrum
with basal width not quite twice its median length and apex moder-
ately rounded; rostral prong (pi. CXIV, fig. 73b) equal to or slightly
longer than third rostral segment, with base originating laterally
near proximal end of third rostral segment, and with apex bluntly
rounded. Pronotum with its median length approximately three
fifths its humeral width; disk with two elongate depressions toward
the middle and a large subtriangular depression on each side, thus
appearing tricarinate; lateral margins divergent; posterior margin
convex, medianly concave. Scutellum large but with median length
less than that of pronotum. Fore femur ( pi. CXIV, fig. 73a ) neither
wide nor greatly thickened at apex; without stridulatory area. Fore
tibia (pi. CXIV, fig. 73a) with stridulatory comb (pi. CXIV, fig.
73c) consisting of approximately twenty-seven to thirty teeth;
apical teeth thick and both apical and basal teeth wider and taller
than those in center. Fore tarsus with claws more slender and
less dissimilar than usual. Chaetotaxy of male front leg as shown
on Plate CXIV. Male genital claspers normal. Spine from caudo-
sinistral margin of seventh abdominal tergite with apical half very
narrow and apex strongly acuminate.

Female Structural Characteristics: As viewed from above, out-
line of head rounded with anterior margin of vertex continuous

A Revision of the Genus Buenoa 1467

with that of eyes; greatest width of head five and one half to six
times the anterior width of vertex and less than humeral width of
pronotum; synthlipsis very narrow, one sixth to one fifth the an-
terior width of vertex, often carinate; along median longitudinal
axis, head is approximately half the length of pronotum; noto-
cephalon slightly sulcate dorsally; tylus not inflated. Pronotum
with its median length slightly more than half its humeral width;
disk usually with two shallow, elongate depressions toward the
middle forming a faint median carina, not tricarinate; lateral mar-
gins divergent; posterior margin convex, medianly concave. Scu-
tellum large, with median length slightly greater than that of pro-
notum. Female ovipositor of normal shape with teeth arranged in
two longitudinal rows which merge proximally; one inner row of
few, large teeth and one long, outer row of smaller teeth; approxi-
mately three or four small, lateral, toothlike setae near apex.

Comparative Notes: Superficially this species resembles B. com-
munis n. sp. Examination of the male, however, will show dis-
tinct differences. This species differs from B. communis in its
smaller size, in having synthlipsis narrower, and in lacking femoral
stridulatory area.

Location of Types: Holotype male, allotype female, 26 male and
26 female paratypes, vicinity of Joao Pessoa, River Jurua, Brazil,
July 10 to Sept. 20, 1936, A. M. Olalla. The type series is in the
Francis Huntington Snow Entomological Collections, University of
Kansas.

Data on Distribution: Known only from Brazil and Bolivia. In
addition to type series, specimens from the following localities
have been examined:

BRAZIL: Amazonas: Vic. Santo Antonio, Sept. 25 to Oct. 17,
1936, A. M. Olalla, 8 males, 9 females.

Parahiba: Vic. Joao Pessoa, River Jurua, July 10 to Sept. 20,
1936, A. M. Olalla, 45 males, 67 females.

BOLIVIA: Victoria, Rio Beni, October, 1937, A. M. Olalla, 40
males, 37 females; Las Pampas, Rio Beni, April, 1938, A. M. Olalla,
1 male, 1 female.

All specimens listed above are in the Francis Huntington Snow
Entomological Collections, University of Kansas, Lawrence, Kansas.

Buenoa oculata n. sp.
(PI. CHI, fig. 26; pi. CXIV, fig. 72)
Size: Male, length 4.74 mm. to 5.13 mm., greatest body width
1.17 mm. to 1.30 mm.; female, length 4.81 mm. to 5.33 mm., greatest
body width 1.23 mm. to 1.36 mm.

1468 The University Science Bulletin

Color: General facies sordid white. Head, most of pronotum,
thoracic venter, and limbs sordid white. Pronotum often with
median and posterior portions orange; scutellum varies from sordid
white to orange; metathoracic dorsum sordid white to pale tes-
taceous. Abdominal venter black except keel, portions of con-
nexivum, and last one or two segments, sordid white; abdominal
dorsum yellowish white with a median, irregular black area.

Male Structural Characteristics: As viewed from above, outline
of head rounded with anterior margin of vertex continuous with
that of eyes; greatest width of head approximately five and one
half times the anterior width of vertex and equal to or slightly
greater than humeral width of pronotum; synthlipsis extremely
narrow, approximately one fifteenth the anterior width of vertex;
along median longitudinal axis, head is approximately three
fourths the length of pronotum; notocephalon slightly sulcate
dorsally; tylus slightly inflated; labrum with basal width not quite
twice its median length and apex bluntly rounded; rostral prong
( pi. CXI V, fig. 72b ) slightly longer than third rostral segment, with
base originating laterally at a point midway to near proximal end
of third rostral segment, and with apex moderately to bluntly
rounded. Pronotum with its median length approximately three
fifths its humeral width; disk with two shallow, elongate depres-
sions toward the middle forming a faint median carina; lateral
margins divergent; posterior margin convex, medianly concave.
Scutellum large, with median length greater than that of pronotum.
Fore femur ( pi. CXIV, fig. 72a ) neither wide nor greatly thickened
at apex; without stridulatory area. Fore tibia (pi. CXIV, fig. 72a)
with stridulatory comb (pi. CXIV, fig. 72c) consisting of approxi-
mately twenty-four to twenty-seven teeth; apical teeth slightly
wider, taller, and thicker than basal. Intermediate leg with first
tarsal segment (pi. CXIV, fig. 72d) slightly emarginate on inner
margin. Chaetotaxy of male front leg as shown on Plate CXIV.
Male genital claspers (pi. CIII, fig. 26) normal. Spine from caudo-
sinistral margin of seventh abdominal tergite normal, tapering
gradually from base to strongly acuminate apex.

Female Structural Characteristics: As viewed from above, out-
line of head rounded with anterior margin of vertex continuous
with that of eyes; greatest width of head five and one half to six
times the anterior width of vertex and less than humeral width of
pronotum; synthlipsis extremely narrow, approximately one fifteenth
the anterior width of vertex; along median longitudinal axis, head
is three fourths to four fifths the length of pronotum; notocephalon

A Revision of the Genus Buenoa 1469

slightly sulcate dorsally; tylus not inflated. Pronotum with its me-
dian length approximately half its humeral width; disk usually un-
impressed, occasionally with a faint median carina; lateral margins
divergent; posterior margin convex, medianly truncate to slightly
concave. Scutellum large, with median length distinctly greater
than that of pronotum. Female ovipositor of normal shape with
teeth arranged in two longitudinal rows; one inner row of few,
large teeth and one long outer row of smaller teeth; approximately
two or three small, lateral, toothlike setae near apex.

Comparative Notes: Superficially this species closely resembles
B. amnigenus (White). Examination of the male, however, will
show distinct differences. This species differs from B. amnigenus
in having the eyes not quite holoptic, frons just above tylus nar-
rower, tylus slightly inflated, and intermediate leg with first tarsal
segment emarginate on inner margin.

Location of Types: Holotype male, allotype female, 6 male and
3 female paratypes, vicinity of Rioja, Dept. San Martin, Peru,
Sept. 9 to Oct. 3, 1936, F. Woytkowski; other paratypes: 1 male
and 4 females, Boqueron del Padre Abad, Dept. Loreta, Peru,
Aug. 3-8, 1946, F. Woytkowski. The type series is in the Francis
Huntington Snow Entomological Collections, University of Kansas.

Data on Distribution: Known only from type series.

Buenoa salutis Kirkaldy

(PI. CIII, fig. 39; pi. CXV, fig. 74)

1904. Buenoa salutis Kirkaldy, G. W. Wiener Ent. Zeit., vol. XXIII, p. 124.
1909. Buenoa salutis, Kirkaldy, G. W. and Torre-Bueno, J. R. de la. Proc.

Ent. Soc. Washington, vol. X, p. 201 (catalogue).
1928. Buenoa mallochi Jaczewski, T. Ann. Musei Zoologici Poloniei, vol. VII,

pp. 129-130 (described from Brazil). New synonomy.

Size: Male, length 3.38 mm. to 3.70 mm., greatest body width
1.07 mm. to 1.17 mm.; female, length 3.70 mm. to 4.35 mm., greatest
body width 1.10 mm. to 1.30 mm.

Color: General facies sordid white to pale testaceous. This spe-
cies entirely sordid white to testaceous except abdomen mostly
brown to black. Metathoracic dorsum often with a light brown to
black, longitudinal stripe on each side. Hemelytra hyalin, often
with a light brown to black area at tip of corium.

Male Structural Characteristics: As viewed from above, outline
of head rounded with vertex protuberant; greatest width of head
approximately three and one half times the anterior width of vertex
and equal to or slightly less than humeral width of pronotum;
synthlipsis very narrow, less than one tenth the anterior width of

1470 The University Science Bulletin

vertex; along median longitudinal axis, head is slightly less than
the length of pronotum; notocephalon wide, sulcate dorsally; tylus
flat, not inflated; labrum short, with basal width more than twice
its median length and apex moderately to bluntly rounded; rostral
prong (pi. CXV, figs. 74b, 74c) variable, equal to or slightly longer
than third rostral segment, with base originating laterally near proxi-
mal end of third rostral segment, and with apex moderately to
bluntly rounded. Pronotum with its median length two fifths to one
half its humeral width; disk usually with two shallow, elongate de-
pressions toward the middle forming a faint median carina; lateral
margins slightly divergent; posterior margin slightly convex, medi-
anly concave. Scutellum large, with median length distinctly
greater than that of pronotum. Fore femur (pi. CXV, fig. 74a)
narrow and not thickened at apex; without stridulatory area. Fore
tibia (pi. CXV, fig. 74a) with stridulatory comb (pi. CXV, fig. 74d)
consisting of approximately eighteen to twenty teeth; teeth approxi-
mately same size and thickness. Chaetotaxy of male front leg as
shown on Plate CXV. Male genital claspers normal. Spine from
caudo-sinistral margin of seventh abdominal tergite normal, tapering
gradually from broad base to acuminate apex.

Female Structural Characteristics: As viewed from above, outline
of head rounded with vertex protuberant; greatest width of head
approximately three and one half times the anterior width of vertex
and slightly less than humeral width of pronotum; synthlipsis
very narrow, one ninth to one seventh the anterior width of vertex;
along median longitudinal axis, head is approximately equal to the
length of pronotum; notocephalon wide, usually sulcate dorsally;
tylus flat, not inflated. Pronotum with its median length approxi-
mately two fifths its humeral width; disk usually unimpressed, occa-
sionally with a faint median carina; lateral margins divergent; pos-
terior margin slightly convex, medianly concave. Scutellum large,
with median length distinctly greater than that of pronotum. Fe-
male ovipositor (pi. CIII, fig. 39) of normal shape with teeth ar-
ranged in three longitudinal rows; one inner row of large teeth, one
median row of normal teeth, and one outer row of smaller teeth; ap-
proximately four to seven small, lateral, toothlike setae near apex.

Macropterous forms are occasionally found. These specimens

A Revision of the Genus Buenoa 1471

have head distinctly narrower than humeral width of pronotum;
pronotum with lateral margins more divergent; scutellum much
larger; hemelytra with claval sutures present and large membranes;
fully developed flight wings.

Variation Within Species: As is indicated under measurements of
length, this species varies somewhat in size; there is also consider-
able variation in the proportional size of notocephalon, synthlipsis,
and pronotum. The greatest contrast is shown between a series
from Itaquaquecetupa, Brazil and one from San Esteban, Venezuela.
The former is a larger form, with a slightly wider synthlipsis, and
a less curved rostral prong.

Comparative Notes: This is our smallest Buenoa species. Super-
ficially it somewhat resembles B. amnigenus (White). Examina-
tion of the male, however, will show distinct differences. This spe-
cies differs from B. amnigenus in having a wider notocephalon, and
in the form of the rostral prong, fore femur, and tibial comb. Buenoa
salutis is much smaller than B. amnigenus.

Nomenclatorial Notes: Mr. C. O. Bare labeled a series of B. salu-
tis as types and paratypes, using a manuscript name based on the
state of origin, Amazonas, Brazil. As such paratypes may have
been widely distributed, it seems desirable to point out that the
name has not been, and should not be, validated by publication.

Examination of large series of B. salutis has led to the conclu-
sion that B. mallochi Jaczewski is a variant of this species and is
therefore placed in synonomy with B. salutis.

Location of Types: The type, a macropterous female, labeled
"I. du Salut, Mus. Paris, Gayane, Pingi 1882", is located at the Paris
Museum. A cotype, also a macropterous female and labeled as
above, is now in the Francis Huntington Snow Entomological Col-
lections, University of Kansas.

Data on Distribution: Recorded from Venezuela, British Guiana,
French Guiana, Brazil, Bolivia and Paraguay. Specimens from the
following localities have been examined:

VENEZUELA: San Esteban, Jan. 15, 1940, Pablo J. Anduze,
4 males and 5 females brachypterous, 1 nymph.

BRITISH GUIANA: Lamaha Conservancy, Demerara, July 23,
1932, S. Harris, 12 males and 12 females brachypterous, 22 males and
25 females macropterous, 5 nymphs; Georgetown Bot. Gardens,
Oct. 23, 1937, S. Harris, 5 females macropterous; Georgetown Bot.
Gardens, Nov. 6, 1937, S. Harris, 4 males macropterous.

FRENCH GUIANA: lies du Salut, 1882, Pingi, 1 female macrop-

63—6490

1472 The University Science Bulletin

terous; He Roy ale, April, 1913, Planchon, 1 male brachypterous, 1
male macropterous; Charvein, July, Coll. Le Moult, 1 male and 2
females macropterous (U.S.N.M.); Charvein, September, Coll. Le
Moult, 1 female macropterous (U.S.N.M.).

BRAZIL: Amazonas: Manacapuru, Manaos, March, 1928, S. M.
Klages, 2 males and 3 females brachypterous, 4 females macrop-
terous; Rio Negro, Manaos, Oct., 1935, A. M. Olalla, 9 males and 12
females brachypterous, 1 male and 3 females macropterous.

Para: Santarem, Dec. 11, 1909, No. 4043, 2 females macropterous
(Carnegie Mus.).

Ceard: Nr. Ico, artificial Lake, July 22, 1937, S. Wright, 1 male
brachypterous; Lagoa, Fortaleza, Aug. 3, 1937, S. Wright, 1 male
brachypterous.

Parahiba: Santa Luzia, No. 245 & 260, S. Wright, 3 males and
2 females brachypterous; Vic Joao Pessoa, River Jurua, July 10 and
Sept. 20, 1936, A. M. Olalla, 1 male and 5 females brachypterous.

Pemambuco: Triumfo, No. 5475, 11017, and 11395, S. Wright, 9
males and 2 females brachypterous, 1 male macropterous.

Scio Paulo: Sao Paulo, Aug. 7, 1927, E. D. Townsend, 4 females
brachypterous; Sao Paulo, Nov., 1928, E. D. Townsend, 3 females
brachypterous; Itaqiiaquecetupa, July, 1933, W. O. Townsend, 16
males and 46 females brachypterous; Itaqiiaquecetupa, E. Town-
send, 105 males and 130 females brachypterous, 4 females macrop-
terous, 7 nymphs; Ypirango, R. Spitz, 2 females brachypterous;
Pirassununga water reservoir, July 30, 1940, H. Kleerekoper, 7 males
and 4 females brachypterous.

Rio Grande do Suh Porto Alegre, July 1941, H. Kleerekoper, 13
males and 8 females brachypterous; Near Porto Alegre, Aug., 1941,
H. Kleerekoper, 4 males and 4 females brachypterous.

BOLIVIA: Prov. del Sara, Nov. 3, 1912, Steinbach; 1 female
macropterous; Rio Beni, Victoria, Oct., 1937, A. M. Olalla, 2 males
brachypterous, 1 male macropterous; Rio Beni, Consuelo, Jan.,
1938, A. M. Olalla, 2 females macropterous; Rio Beni, Las Pampas,
Mojos, Apr. & May, 1938, A. M. Olalla, 7 males and 4 females
brachypterous, 9 males and 2 females macropterous; Santa Ana del
Yacuma, Feb., 1938, A. M. Olalla, 3 males and 5 females brachyp-
terous; Santa Rosa del Yacuma, Feb., 1938, A. M. Olalla, 2 males
and 2 females brachypterous, 1 female macropterous; Puerto Suarez,
No. 3844, J. Steinbach, 3 males and 3 females macropterous (Car-
negie Mus.).

PARAGUAY: Villarrica, Dec, 1923, Fran. Schade, 1 male and 1
female brachypterous; Villarrica, Apr. 16, 1924, Fran. Schade, 1

A Revision of the Genus Buenoa 1473

male macropterous; Albovena Srojoguasi, Dec, 1926, Fran. Schade,
1 male and 4 females brachypterous, 1 nymph; Reimoser, Centurion,
3 females macropterous ( Berlin Mus. ) .

All specimens listed above are in the Francis Huntington Snow
Entomological Collections, University of Kansas, Lawrence, Kansas,
unless otherwise indicated.

Buenoa thomasi n. sp.
(PI. CIII, fig. 25; pi. CXV, fig. 76)

Size: Male, length 5.26 mm. to 5.72 mm., greatest body width
1.49 mm. to 1.69 mm.; female, length 5.46 mm. to 6.04 mm., greatest
body width 1.69 mm. to 1.82 mm.

Color: General facies sordid white to gray. Head, pronotum,
most of thoracic venter, and limbs sordid white to pale testaceous.
Scutellum usually pale testaceous, occasionally with base brown
to black; metathoracic dorsum pale testaceous to black. Abdomen
black except ventral keel and portions of connexivum and dorsum,
sordid white to testaceous. Some specimens entirely sordid white
to testaceous except portions of abdomen, black. This species varia-
ble in color.

Male Structural Characteristics: As viewed from above, outline
of head rounded with vertex usually indented at lateral margins;
greatest width of head approximately five to five and one half times
the anterior width of vertex and less than humeral width of pro-
notum; synthlipsis one third to one half the anterior width of vertex;
along median longitudinal axis, head is one half to two thirds the
length of pronotum; notocephalon usually sulcate dorsally; tylus
slightly inflated; labrum with basal width not quite twice its median
length and apex bluntly rounded; rostral prong (pi. CXV, fig. 76c)
equal to or longer than third rostral segment, with base originating
laterally midway of third rostral segment, and with apex bluntly
rounded. Pronotum with its median length approximately three
fifths its humeral width; disk with two elongate depressions toward
the middle and a large subtriangular depression on each side, thus
appearing tricarinate; lateral margins divergent; posterior margin
convex, slightly concave medianly. Scutellum with median length
equal to or greater than that of pronotum. Fore femur (pi. CXV,
fig. 76a) relatively narrow and not greatly thickened at apex; without
stridulatory area. Fore tibia (pi. CXV, fig. 76a) with stridulatory
comb (pi. CXV, fig. 76b) consisting of approximately twenty-
four to thirty-six teeth; apical teeth thicker than basal; ten to seven-
teen short, stout, club-shaped setae (pi. CXV, fig. 76d) on inner sur-

1474 The University Science Bulletin

face of tibia at apex. Chaetotaxy of male front leg as shown on
Plate CXV. Male genital claspers (pi. CIII, fig. 25) normal. Spine
from caudo-sinistral margin of seventh abdominal tergite tapering
gradually from base to strongly acuminate apex.

Female Structural Characteristics: As viewed from above, out-
line of head rounded with vertex usually indented at lateral margins;
greatest width of head five to five and one half times the anterior
width of vertex and less than humeral width of pronotum; synthlip-
sis one third to one half the anterior width of vertex; along median
longitudinal axis, head is three fifths to two thirds the length of pro-
notum; notocephalon sulcate dorsally; tylus slightly inflated. Pro-
notum with its median length approximately half its humeral width;
disk with two shallow, elongate depressions toward the middle and
a shallow, subtriangular depression on each side, thus appearing
faintly tricarinate, often with median carina only; lateral margins
divergent; posterior margin convex, medianly truncate to slightly
concave. Scutellum with median length greater than that of pro-
notum. Female ovipositor of normal shape with teeth arranged in
two longitudinal rows which merge in proximal half of ovipositor
valve; one inner row of large teeth and one outer row of smaller
teeth; approximately four to six small, lateral, toothlike setae near
apex.

Comparative Notes: Superficially this species resembles B. platy-
cnemis (Fieber). Examination of the male, however, will show
distinct differences. This species differs from B. platycnemis in
having an area of short, stout, club-shaped setae on inner surface
of fore tibia at apex, in lacking a femoral stridulatory area, and
in the form of the rostral prong and tibial comb.

Location of Types: Holotype male, allotype female, 18 male and
3 female paratypes, San Luis Babarocos (=Barbacoas ?), Chi-
huahua, Mexico, Dec. 30, 1934, H. S. Gentry; other paratypes: 7
males and 6 females, Carimechi, Rio Mayo, Chihuahua, Mexico,
Jan. 6, 1935, H. S. Gentry. The type series is in the Francis Hunt-
ington Snow Entomological Collections, University of Kansas.

Data on Distribution: Recorded from Mexico and known only
from type series.

Buenoa alterna n. sp.

(PI. CIII, fig. 35; pi. CXV, fig. 77)

Size: Male, length 5.65 mm. to 5.98 mm., greatest body width
1.56 mm. to 1.62 mm.; female, length 6.30 mm. to 6.76 mm.; greatest
body width 1.88 mm. to 2.08 mm.

A Revision of the Genus Buenoa 1475

Color: General facies sordid white to gray. Head, pronotum,
thoracic venter, and limbs sordid white to testaceous. Scutellum
with basal portion black, apical portion yellow to orange; meta-
thoracic dorsum usually black with broad, median, transverse
yellow to orange band, occasionally entirely black. Abdomen
black except ventral keel and portions of connexivum and dorsum,
sordid white to testaceous. Hemelytra colorless but with black
portions of thorax and abdomen shining through giving a gray
appearance to the specimen. Some specimens entirely sordid
white to testaceous except portions of abdomen, black.

Male Structural Characteristics: As viewed from above, outline
of head rounded with anterior margin of vertex continuous with
that of eyes, occasionally indented at lateral margins; greatest
width of head six to six and one half times the anterior width of
vertex and less than humeral width of pronotum; synthlipsis nar-
row, approximately half the anterior width of vertex; along median
longitudinal axis, head is approximately three fifths the length of
pronotum; notocephalon narrow, slightly sulcate dorsally; tylus not
inflated; labrum with basal width slightly more than twice its me-
dian length and apex bluntly rounded; rostral prong (pi. CXV,
fig. 77b) short, shorter than third rostral segment, with base origi-
nating laterally at proximal end of third rostral segment, and with
apex bluntly rounded. Pronotum with its median length slightly
more than half its humeral width; disk usually with two shallow,
elongate depressions toward the middle forming a faint median
carina, not tricarinate; lateral margins divergent; posterior margin
convex, medianly truncate to slightly concave. Scutellum large,
with median length greater than that of pronotum. Fore femur
(pi. CXV, fig. 77a) neither wide nor strongly thickened at apex;
without stridulatory area. Fore tibia (pi. CXV, fig. 77a) with
stridulatory comb (pi. CXV, fig. 77c) consisting of approximately
fifteen to nineteen teeth; apical teeth slightly thicker than basal.
Chaetotaxy of male front leg as shown on Plate CXV. Male genital
claspers normal. Spine from caudo-sinistral margin of seventh
abdominal tergite tapering gradually from base to strongly acumi-
nate apex.

Female Structural Characteristics: As viewed from above, out-
line of head rounded with anterior margin of vertex continuous
with that of eyes; greatest width of head five and one half to six
times the anterior width of vertex and less than humeral width of
pronotum; synthlipsis slightly less than half the anterior width of
vertex; along median longitudinal axis, head is one half to three

1476 The University Science Bulletin

fifths the length of pronotum; notocephalon slightly sulcate dor-
sally; tylus not inflated. Pronotum with its median length slightly
less than half its humeral width; disk usually unimpressed, occa-
sionally with faint median carina; lateral margins divergent; pos-
terior margin convex, medianly truncate to slightly concave. Scu-
tellum large, with median length distinctly greater than that of
pronotum. Female ovipositor (pi. Ill, fig. 35) of normal shape
with teeth arranged in three longitudinal rows which merge proxi-
mally; one inner row of large teeth, one short median row of normal
teeth, and one long outer row of small teeth; approximately five or
six small, lateral, toothlike setae near apex.

Comparative Notes: Superficially this species resembles B. muta-
bilis n. sp. and B. omani n. sp. Examination of the male, however,
will show distinct differences. This species differs from B. omani
in having tylus not inflated, frons just above tylus wider, fore femur
narrow at apex, and in lacking a femoral stridulatory area. Buenoa
alterna differs from B. mutabilis in having a distinctly different
rostral prong and in lacking a femoral stridulatory area.

Location of Types: Holotype male, allotype female, 6 male and
5 female paratypes, Puebla, Mexico, July 25, 1937, H. D. Thomas;
other paratypes: 4 males, Hda. La Libertad, Chiapas, Mexico,
Sept. 1, 1937, H. D. Thomas; 1 male and 4 females, Los Potosi,
Zacatecas, Mexico, Aug. 8, 1944, H. D. Thomas. The type series
is in the Francis Huntington Snow Entomological Collections, Uni-
versity of Kansas.

Data on Distribution: Known only from Mexico. Specimens
from the following localities have been examined:

MEXICO: Zacatecas: Los Potosi, Aug. 8, 1944, Henry Thomas,
4 males.

Hidalgo: Real del Monte, Sept. 23, 1938, H. D. Thomas, 1 male,
2 females.

Puebla: Puebla, July 25, 1937, H. D. Thomas, 7 males, 6 females.

Chiapas: Hda. La Libertad, Sept. 1, 1937, H. D. Thomas, 1 male,
4 females.

All specimens listed above are in the Francis Huntington Snow
Entomological Collections, University of Kansas, Lawrence, Kansas.

Buenoa unguis n. sp.

(PI. CXV, fig. 78)

Size: This species varies considerably in size within the same
population. Male, length 5.90 mm. to 7.07 mm., greatest body width
1.49 mm. to 1.75 mm.; female, length 5.85 mm. to 7.10 mm., greatest
body width 1.62 mm. to 1.88 mm.

A Revision of the Genus Buenoa 1477

Color: General facies sordid white to pale testaceous. Head,
thorax, and limbs sordid white to pale testaceous; scutellum occa-
sionally with two anterolateral brown to black areas. Abdominal
venter black except keel and portions of connexivum, yellowish
white; abdominal dorsum brown to black with anterior portion
usually yellowish white to pale testaceous.

Male Structural Characteristics: As viewed from above, outline
of head rounded with anterior margin of vertex slightly indented to
continuous with that of eyes; greatest width of head four and one
half to five times the anterior width of vertex and less than humeral
width of pronotum; synthlipsis narrow, one sixth to one fifth the
anterior width of vertex; along median longitudinal axis, head is
approximately three fifths the length of pronotum; notocephalon
sulcate; tylus flat, depressed longitudinally forming distinct lateral
carinae; labium with basal width more than twice its median
length; rostrum with a distinct, median, longitudinal carina; rostral
prong (pi. CXV, fig. 78c) short, distinctly shorter than third rostral
segment, with base originating laterally at proximal end of third
rostral segment, and with apex bluntly rounded. Pronotum with
its median length approximately four sevenths its humeral width;
disk with two elongate depressions toward the middle forming a
median carina, occasionally with a shallow, subtriangular depres-
sion on each side, appearing faintly tricarinate; lateral margins
divergent; posterior margin convex, slightly concave medianly.
Scutellum large, with median length greater than that of pronotum.
Fore femur (pi. CXV, fig. 78a) narrow, somewhat thickened at apex;
without stridulatory area. Fore tibia (pi. CXV, fig. 78a) with
stridulatory comb (pi. CXV, fig. 78b) consisting of approximately
nineteen to twenty teeth, with four or five long setae at apex; apical
teeth are slightly thicker than basal. Fore tarsus ( pi. CXV, fig. 78d )
robust with unusual tarsal claws. Chaetotaxy of male front leg as
shown on Plate CXV. Male genital claspers normal. Spine from
caudo-sinistral margin of seventh abdominal tergite normal, taper-
ing gradually from base to a curved, strongly acuminate, apex.

Female Structural Characteristics: As viewed from above, outline
of head rounded with anterior margin of vertex continuous with
that of eyes; greatest width of head four and one half to five times
the anterior width of vertex and less than humeral width of prono-
tum; synthlipsis narrow, one seventh to one fifth the anterior width
of vertex; along median longitudinal axis, head is two fifths to
three fifths the length of pronotum; notocephalon slightly sulcate;
tylus flat, depressed longitudinally forming faint lateral carinae.

1478 The University Science Bulletin

Pronotum with its median length slightly less than half its humeral
width; disk with two, shallow elongate depressions toward the
middle forming a faint median carina; lateral margins divergent;
posterior margin convex, medianly concave. Scutellum large, with
median length greater than that of pronotum. Female ovipositor of
normal shape with teeth arranged in two longitudinal rows; one
inner row of large teeth and one outer row of smaller teeth; approxi-
mately three or four small, lateral, toothlike setae near apex.

Comparative Notes: Superficially this species somewhat resem-
bles B. albida (Champion) and B. gracilis n. sp. Examination of
the male, however, will show distinct differences. This species
differs from B. albida and B. gracilis in having a distinctly shorter
rostral prong, rostrum with a median carina, fore femur without
stridulatory area, and distinct differences in the tibial comb, fore
tarsus and tarsal claws.

Location of Types: Holotype male, allotype female, 40 male and
40 female paratypes, Vicinity of Rioja, Dept. San Martin, Peru,
Sept. 9 to Oct. 3, 1936, F. Woytkowski. The type series is in the
Francis Huntington Snow Entomological Collections, University of
Kansas.

Data on Distribution: Known from Brazil, Peru, Bolivia, Para-
guay, and Argentina. In addition to type series, specimens from the
following localities have been examined:

BRAZIL: Paid: Lago Grande, Feb., 1939, A. M. Olalla, 7 males
5 females.

Ceard: Agua Verde, July 6, 1937, S. Wright, 16 males, 13 fe-
males; Russas, July 22, 1937, S. Wright, 2 males, 2 females; Sobral,
Oct. 26, 1937, S. Wright, 3 males, 12 females; Primavera, Oct. 28,
1937, S. Wright, 2 males, 18 females; Chere, Oct. 29, 1937, S. Wright,
2 females; Maranguape, Nov. 3, 1937, S. Wright, 2 males, 1 female;
Bom Acude Successo, Nov. 4, 1937, S. Wright, 3 males, 10 females;
Sobral, Nov. 26, 1937, S. Wright, 3 males, 19 females.

Rio Grande do Norte: Ouro Branco, No. 258, S. Wright, 2 males;
Caico, No. 327, S. Wright, 1 male.

Perahiba: Souza, No. 5530, S. Wright, 1 male, 1 female.

Pcrnambnco: Belem, No. 643, S. Wright, 1 male; Itapissuma,
Oct. 25, 1946, F. S. Barbosa, 2 males, 2 females.

Minas Geraes: Bello Horizonte, Apr., 1935, D. M. Cochran, 3
males, 1 female ( U.S.N. M.).

Rio de Janeiro: Sao Paulo Road, June 19, 1945, Wygodzinsky, 1
male, 1 female.

A Revision of the Genus Buenoa 1479

PERU: Vic. Rioja, Dept. San Martin, Sept. 9 to Oct. 3, 1936,
F. Woytkowski, 37 females.

BOLIVIA: Junction of Madre de Dios and Beni Rivers, Victoria,
Oct., 1937, A. M. Olalla, 32 males, 45 females; Santa Ana del Yacuma,
Feb., 1938, A. M. Olalla, 17 males, 15 females; Rio Beni, Las Pampas,
Mojos, April and May, 1938, A. M. Olalla, 8 males, 4 females.

PARAGUAY: Estancia Postilion, Puerto Max a. Rio Paraguay,
Mar. 9, 1905, Louis Des Arts, 2 males, 1 female (Hamburg Mus.).

ARGENTINA: Chaco, No. 7726, 1 male.

All specimens listed above are in the Francis Huntington Snow
Entomological Collections, University of Kansas, Lawrence, Kansas,
unless otherwise indicated.

Buenoa excavata n. sp.
(PL CII, fig. 12; pi. CXV, fig. 75)

Size: Male, length 4.55 mm., greatest body width 1.30 mm.;
known only from the male.

Color: General facies sordid white. Head, pronotum, most of
thoracic venter, and scutellum sordid white. Portions of thoracic
venter orange to brown; metathoracic dorsum colorless to sordid
white. First two pairs of legs testaceous to nigro-violaceous; hind
legs mostly yellowish white. Abdomen black with anterior portion
of connexivum orange. Hemelytron colorless except for nigro-
violaceous band covering humeral angle and extending along an-
terior margin of wing for approximately one third its length and a
large black area at tip of corium.

Male Structural Characteristics: As viewed from above, outline
of head rounded with anterior margin of vertex continuous with
that of eyes; eyes almost holoptic; greatest width of head approxi-
mately five and one half times the anterior width of vertex and
slightly less than humeral width of pronotum; synthlipsis extremely
narrow, approximately one fifteenth the anterior width of vertex;
a'ong median longitudinal axis, head is distinctly longer than pro-
notum; notocephalon indented at synthlipsis, sulcate anterodorsally;
tylus deeply excavate with a short antero-medial ridge; labrum
distinctly inflated, with basal width more than twice its median
length and apex truncate; rostral prong (pi. CXV, fig. 75b) as long
as third rostral segment, with base originating laterally near proximal
end of third rostral segment, and with apex moderately rounded.
Pronotum short, with its median length approximately two fifths
its humeral width; disk unimpressed, not carinate; lateral margins

1480 The University Science Bulletin

divergent; posterior margin convex, slightly concave medianly.
Scutellum large, with median length distinctly greater than that of
pronotum. Fore femur (pi. CXV, fig. 75a) neither wide nor greatly
thickened at apex; without stridulatory area. Fore tibia (pi. CXV,
fig. 75a) angulate anteriorly, with stridulatory comb (pi. CXV,
fig. 75c) consisting of approximately seventeen teeth; apical teeth
thicker than basal; a swollen area on inner surface of tibia at apex,
densely covered with fine setae. Chaetotaxy of male front leg as
shown on Plate CXV. Male genital claspers normal. Spine from
caudo-sinistral margin of seventh abdominal tergite (pi. CII, fig. 12)
unusually wide for most of length; apical one fourth narrow and
apex strongly acuminate.

Only the brachypterous male is known for this species.

Comparative Notes: This species is quite distinct and its rela-
tionship to the other species of the genus is obscure. The deeply
excavate tylus, the inflated labrum, and the form of the fore tibia
separates B. excavata from all other species.

Location of Types: Holotype male (brachypterous), Santa Elena
Boqueron Padre Abad, Dept. Loreto, Peru, Aug. 8, 1946, F. Woyt-
kowski. Holotype is in the Francis Huntington Snow Entomo-
logical Collections, University of Kansas.

Data on Distribution: Known only from type.

Buenoa macrophthalma (Fieber)
(PI. CII, fig. 3; pi. CIII, fig. 27; pi. CXVI, fig. 79)

1851. Anisops mocrophthahnus Fieber, F. X. Abhandlungen Kongl. Boh-

mischen Ceselschaft Wissensehaften, vol. VII, Folge 5, pp. 482-483.
1904. Buenoa macrophthalma, Kirkaldy, G. W. Wiener Ent. Zeit., vol. XXIII,

pp. 121-122, and 134 (taxonomic notes).
1930. Buenoa macrophthalma, Wolcott, G. N. Jr. Agriculture Univ. Puerto

Rieo, vol. XX, p. 149.
1939. Buenoa macrophthalma, Barber, H. G. New York Acad. Sci., vol. XIV,

p. 420.

Size: This species varies considerably in size due to the larger
macropterous forms. Male, length 8.51 mm. to 10.85 mm., greatest
body width 2.47 mm. to 3.18 mm.; female, length 9.10 mm. to 9.75
mm., greatest body width 2.73 mm. to 2.92 mm.

Color: The brachypterous and macropterous forms vary in color
as well as in other characteristics. The general facies of the former
is sordid white to testaceous while the latter is black. The body
of the brachypterous form is entirely sordid white to testaceous ex-
cept for abdominal venter, portions of connexivum, and laterally
above connexivum on dorsum of abdomen, black. The macropter-
ous form has head, anterior portions of pronotum, thoracic venter,

A Revision of the Genus Buenoa 1481

and limbs, mostly testaceous to light brown. Scutellum, metathor-
acic dorsum, and abdomen black except for ventral keel and mar-
gins of connexivum, testaceous. Hemelytron black except for an
area behind humeral angle and a wide band along the claval su-
ture, colorless.

Male Structural Characteristics: As viewed from above, outline
of head laterally rounded, anteriorly truncate with vertex indented
especially at its lateral margins; greatest width of head approxi-
mately five and one half times the anterior width of vertex and
usually slightly less but occasionally equal to or greater, than
humeral width of pronotum; synthlipsis less than one third the an-
terior width of vertex; along median longitudinal axis, head almost
as long as pronotum; notocephalon sulcate dorsally; tylus strongly
inflated with median depression forming two lateral protuberances;
labrum with basal width approximately two thirds greater than its
median length and apex bluntly rounded; rostral prong (pi. CXVI,
fig. 79a) extremely long, much longer than third rostral segment,
with base originating laterally at distal end of third rostral segment,
and with apex moderately rounded. Pronotum with its median
length slightly more than half its humeral width; disk usually with
two elongate depressions toward the middle and a shallow, sub-
triangular depression on each side, thus appearing faintly tricari-
nate; lateral margins divergent; posterior margin slightly convex,
medianly concave. Scutellum with median length distinctly greater
than that of pronotum. Fore femur (pi. CXVI, fig. 79b) wide and
thickened at apex; without stridulatory area. Fore tibia ( pi. CXVI,
fig. 79b) with stridulatory comb (pi. CXVI, fig. 79b) consisting of
approximately seventeen thick teeth of uniform size. Chaetotaxy
of male front leg as shown on Plate CXVI. Male genital claspers
(pi. C1II, fig. 27) abnormal in shape. Spine from caudo-sinistral
margin of seventh abdominal tergite (pi. II, fig. 3) relatively short
and thick, tapering gradually from broad base to acuminate apex.

Macropterous forms are occasionally found. These specimens are
black and larger in size than the brachypterous forms. They have
head distinctly narrower than humeral width of pronotum; prono-
tum with lateral margins more divergent; scutellum larger; hemely-
tra with claval sutures present and large membranes; fully de-
veloped flight wings.

Female Structural Characteristics: As viewed from above, outline
of head laterally rounded, anteriorly truncate with vertex indented
only at lateral margins; greatest width of head five to five and one
half times the anterior width of vertex and less than humeral width

1482 The University Science Bulletin

of pronotum; synthlipsis approximately one third the anterior width
of vertex; along median longitudinal axis, head almost as long as
pronotum; notocephalon sulcate dorsally; tylus inflated. Pronotum
with its median length less than half its humeral width; disk only
slightly impressed, usually not tricarinate; lateral margins divergent;
posterior margin slightly convex, medianly concave. Scutellum
with median length distinctly greater than that of pronotum. Fe-
male ovipositor of normal shape with teeth arranged in two longi-
tudinal rows; one very short inner row of approximately seven large
teeth and one long outer row of small teeth; approximately four or
five small, lateral, toothlike setae near apex.

Variation Within Species: As is indicated under measurements of
length, this species varies a great deal in size; there is also some
variation in the proportional size of the pronotum. The greatest
contrast is shown between males of series from Jamaica, B. W. I.,
and Cuba. The former is a small form, the males are seldom more
than 8.50 mm. in length, lack a short longitudinal carina on inner
surface of intermediate tibia at the base, and have normal and
similar claws on intermediate tarsus; the males of the latter form are
approximately 10 mm. in length, have a short longitudinal carina
on inner surface of intermediate tibia at the base, and have one long,
narrow, distorted claw and a short, broad, flat claw on intermediate
tarsus. Due to the fact that all important and constant characters
used in species determination are identical in these two forms, no
specific separation appears justified.

Comparative Notes: Superficially this species resembles B.
hungerfordi n. sp. Examination of the male, however, will show
distinct differences. The form of the rostral prong alone will serve
to distinguish this species from all others. It differs from B. hunger-
fordi not only in the form of the rostral prong, but also in having
a strongly inflated tylus, the fore femur greatly thickened at apex,
a distinct tibial stridulatory comb, and a difference in the form of the
tarsal claws on fore and intermediate legs.

Location of Types: The type specimen, a male, from Port au
Prince, Haiti, is located at the Berlin Museum. This specimen is
leucochromatic and I assume therefore, a brachypterous form.

A Revision of the Genus Buenoa 1483

Homotype male, labeled "Adjuntas, P. R., June 8-13, 1915", com-
pared with type by Dr. H. B. Himgerford, University of Kansas,
now located in the Francis Huntington Snow Entomological Collec-
tions, University of Kansas.

Data on Distribution: Recorded from West Indies — (Cuba,
Haiti, Jamaica, and Puerto Rico). Specimens from the following
localities have been examined:

WEST INDIES: Cuba: Cuba, Feb. 2, 1932, Ernesto Pujals y de
Quesada, 1 male and 2 females macropterous, 2 males and 1 female
brachypterous.

Jamaica: St. Andrew, Shooters Hill, Dec. 3, 1946, G. B. Thomp-
son, 1 female brachypterous; St. Andrew, Hermitage, Feb. 14, 1947,
G. B. Thompson, 1 male brachypterous.

Puerto Rico: Adjuntas, June 8-13, 1915, 1 male macropterous;
Maricao R., Stn. Maricao, Feb. 20, 1934, S. Hildebrand, 1 female
macropterous, 1 male brachypterous (U.S.N.M.); Ponce, Rio Finca,
July 12, 1934, R. G. Oakley, 1 female macropterous (U.S.N.M.);
Lares Guajataca R., Mar. 22, 1935, Julio Garcia Diaz, 1 female
macropterous, 1 male and 3 females brachypterous, 2 nymphs.

All specimens listed above are in the Francis Huntington Snow
Entomological Collections, University of Kansas, Lawrence, Kansas,
unless otherwise indicated.

Buenoa hungerfordi n. sp.
(PI. CII, figs. 14, 21; pi. Cm, figs. 23, 41; pi. CXVI, fig. 80)

Size: Male, length 10.01 mm. to 10.79 mm., greatest body width
2.99 mm. to 3.12 mm.; female, length 9.94 mm. to 10.20 mm., greatest
body width 3.12 mm. to 3.51 mm.

Color: General facies yellowish white to black. Head, most of
pronotum, thoracic venter, and limbs yellowish white to pale tes-
taceous. Pronotum usually with a median, subtriangular, light
brown to black area; scutellum usually black with posterolateral
margins yellowish white; metathoracic dorsum light brown to black.
Abdomen black except ventral keel and portions of connexivum and
dorsum, yellowish white. Hemelytra hyalin with posterior half
light brown to black. Some specimens entirely yellowish white to
pale testaceous except most of abdomen, black. This species varia-
ble in color.

Male Structural Characteristics: As viewed from above, outline
of head laterally rounded, anteriorly truncate with vertex protuber-
ant; greatest width of head four and one half times the anterior
width of vertex and less than humeral width of pronotum; synthlip-

1484 The University Science Bulletin

sis wide, approximately two thirds the anterior width of vertex;
along median longitudinal axis, head is slightly less than half the
length of pronotum; notocephalon wide, sulcate dorsally; tylus in-
flated, distinctly pilose; labrum pilose, with basal width not quite
twice its median length and apex bluntly rounded; rostral prong
(pi. CXVI, fig. 80b) short, distinctly shorter than third rostral seg-
ment, with base originating laterally near proximal end of third ros-
tral segment, and with apex moderately to bluntly rounded. Pro-
notum with its median length slightly less than half its humeral
width; disk with two shallow, elongate depressions toward the mid-
dle forming a faint median carina; lateral margins divergent; pos-
terior margin convex, medianly concave. Scutellum large, with me-
dian length greater than that of pronotum. Fore femur (pi. CXVI,
fig. 80a ) neither wide nor greatly thickened at apex; without stridu-
latory area. Fore tibia with stridulatory comb (pi. CXVI, fig. 80a)
consisting of approximately fifteen to seventeen thick teeth; all teeth
approximately same size and thickness. Tarsal claws of fore leg
dissimilar; one with narrow, acuminate apex and the other with
blunt, slightly bifurcate apex. Chaetotaxy of male front leg as
shown on Plate CXVI. Male genital claspers (pi. CIII, fig. 23) ab-
normal in shape. Spine from caudo-sinistral margin of seventh ab-
dominal tergite (pi. CII, fig. 14) broad, sickle-shaped, lying horizon-
tally rather than in usual vertical position.

Female Structural Characteristics: As viewed from above, out-
line of head laterally rounded, anteriorly truncate with vertex pro-
tuberant; greatest width of head three and one half times the
anterior width of vertex and distinctly less than humeral width of
pronotum; synthlipsis wide, approximately half the anterior width
of vertex; along median longitudinal axis, head is slightly more
than half the length of pronotum; notocephalon wide, sulcate dor-
sally; tylus inflated, not pilose. Pronotum with its median length
approximately two fifths its humeral width; disk with two shallow,
elongate depressions toward the middle forming a faint median
carina; lateral margins divergent; posterior margin convex, medianly
concave. Scutellum large, with median length distinctly greater
than that of pronotum. Female ovipositor ( pi. CIII, fig. 41 ) abnor-
mal in shape with teeth arranged in a long, irregular, curved row;
approximately two or three small, lateral, toothlike setae.

Variation Within Species: Occasionally specimens are found with
flight wings not fully developed. These specimens are pale in color
with pronotum narrower and lateral margins less divergent, scutel-
lum smaller, and hemelytral membranes smaller than the form with

A Revision of the Genus Buenoa 1485

fully developed flight wings. Both forms have claval sutures present
in the hemelytra.

Comparative Notes: Superficially this species resembles B. ma-
crophthalma (Fieber) and B. distincta n. sp. Examination of the
male, however, will show distinct differences. This species dif-
fers from B. macrophthalma in having the tylus less inflated, rostral
prong much shorter, fore femur narrower at apex, and differences
in the tibial comb and tarsal claws. Buenoa hungerfordi differs
from B. distincta in having the tylus distinctly more pilose, spine
from caudo-sinistral margin of seventh abdominal tergite sickle-
shaped, differences in the genital capsule as shown on Plate CII, and
in its larger size.

Location of Types: Holotype male, allotype female, 1 male and
2 female paratypes, Conejos, Dist. Alamos, Sonora, Mexico, Oct. 26,
1934, H. S. Gentry; other paratypes: 2 males and 3 females, Arroyo
S. Marcial, District Alamos, Sonora, Mexico, Oct. 28, 1934, H. S.
Gentry; 1 male, L. Tepancuapan, Chiapas, Mexico, Aug. 28, 1937,
H. D. Thomas; 1 male, Sabino Canyon, Arizona, U. S. A., July 12,
1932, R. H. Beamer. The type series is in the Francis Huntington
Snow Entomological Collections, University of Kansas.

Data on Distribution: Known only from United States and
Mexico. In addition to type series, specimens from the following
localities have been examined:

U.S.A.: Arizona: Catalina Mts., Sept. 29, 1941, Victor Potter,
2 males, 2 females ( U. of Mich. ) .

MEXICO: Chihuahua: Carimechi, Rio Mayo, Dec. 12, 1934, H.
S. Gentry, 1 female; San Luis Babarocos (= Barbacoas ?), Dec. 30,
1934, H. S. Gentry, 1 female.

All specimens listed above are in the Francis Huntington Snow
Entomological Collections, University of Kansas, Lawrence, Kansas,
unless otherwise indicated.

Buenoa distincta n. sp.
(PI. CII, figs. 13, 22; pi. CIII, fig. 24; pi. CXVII, fig. 81)

Size: Male, length 8.45 mm., greatest body width 2.27 mm.;
known only from the male.

Color: General facies gray. Head, pronotum, most of thoracic
venter, and limbs sordid white to pale testaceous. Scutellum black
with apex and lateral margins yellowish white; metathoracic dor-
sum black, appearing gray through hyalin hemelytra. Abdomen
black except ventral keel and portions of connexivum and dorsum,
yellowish white. Hemelytra hyalin with membrane light brown to
black.

1486 The University Science Bulletin

Male Structural Characteristics: As viewed from above, outline
of head laterally rounded, anteriorly truncate with vertex pro-
tuberant; greatest width of head approximately four and one half
times the anterior width of vertex and distinctly less than humeral
width of pronotum; synthlipsis slightly less than half the anterior
width of vertex; along median longitudinal axis, head is approxi-
mately four sevenths the length of pronotum; notocephalon wide,
sulcate dorsally; tylus slightly inflated; labrum with basal width not
quite twice its median length and apex moderately rounded; rostral
prong (pi. CXVII, fig. 81c) short, shorter than third rostral seg-
ment, with base originating laterally near proximal end of third
rostral segment, and with apex bluntly rounded. Pronotum with
its median length slightly less than half its humeral width; disk
with two elongate depressions toward the middle forming a median
carina; lateral margins divergent; posterior margin convex, medianly
concave. Scutellum large, with median length distinctly greater
than that of pronotum. Fore femur (pi. CXVII, fig. 81a) neither
wide nor greatly thickened at apex; without stridulatory area. Fore
tibia (pi. CXVII, fig. 81a) with stridulatory comb (pi. CXVII, fig.
81b) consisting of approximately fourteen thick teeth; all teeth
approximately same size and thickness. Tarsal claws of fore leg
dissimilar; one with narrow, acuminate apex and the other with
blunt, slightly bifurcate apex. Chaetotaxy of male front leg as
shown on Plate CXVII. Male genital claspers (pi. CIII, fig. 24)
abnormal in shape. Spine from caudo-sinistral margin of seventh
abdominal tergite (pi. CII, fig. 13) sword-shaped, lying horizontally
rather than in usual vertical position.

Comparative Notes: Superficially this species resembles B. hun-
gerfordi n. sp. Examination of the male, however, will show dis-
tinct differences. This species differs from B. hungerfordi in having
the tylus distinctly less pilose, spine from caudo-sinistral margin of
seventh abdominal tergite straight and sword-shaped, differences in
the genital capsule as shown on Plate II, and in its smaller size.

Location of Types: Holotype male, Acapulco, Gro., Mexico, July
12, 1937, H. D. Thomas. The holotype is in the Francis Huntington
Snow Entomological Collections, University of Kansas.

Data 07i Distribution: Known only from type.

Buenoa paranensis Jaczewski

(PI. CXVII, fig. 82)

1928. Buenoa paranensis Jaczewski, T. Ann. Musei Zoologici Polonici, vol.
VII, pp. 126-127.

This species was not present in the material at my disposal.

Dr. T. Jaczewski's types which were located at the Warsaw Museum,

A Revision of the Genus Buenoa 1487

Poland, have been destroyed. The following is a copy of the
original description and copies of Jaczewski's figures are included
among the illustrations.

"Colour yellowish white, eyes dark, abdomen partly black be-
neath and above.

Head with eyes distinctly wider than pronotum in front, as wide
as posterior width of the latter [Fig. 82f]. Synthlipsis about 3
times narrower than the greatest width of the notocephalon. Fron-
tal arch very feebly convex, not projecting between the eyes. Eyes
strongly convex and prominent in both sexes. Notocephalon in
front with two distinct longitudinal swellings and a shallow mediane
groove between them. Prongs of the third rostral joint of the
<J $ comparatively long, straight, gradually tapering towards their
end [Fig. 82e].

Pronotum about 1/2 times longer than the head, and about 1%
times as wide as long. Scutellum somewhat ( about Yq ) longer than
pronotum. Pronotum in both sexes evenly convex, without any
longitudinal keels. Claval orifice about % of the length of the
scutellum.

Relative length of the various parts of the legs, measured in per-
centages of the length of the corresponding femora, as follows:

Femur

Tibia

Tarsus 1 + 2

Tarsus 3

Front legs $ :
Front legs 2 :
Int. legs:
Hind legs:

100
100
100
100

116,7

126,5

79,5

84,1

57,4
58,5
34,2
31,1

31,1
32,1
27,2
30,6

The ratio of length of the femora of the three pairs of legs appears
as follows:

Fr. fern.: Int. fem.: Hind fern. = 100: 160,4: 246,3.

Front legs of the £ ^shaped as shown on fig. [82d]. Femora
with a triangular stridulatory area about the middle of their anterior
surface. Tibial prong with about 25 setaceous teeth.

Lateral (sinistral) spine of the seventh abdominal tergite of the
$ $ moderately long [Fig. 82c], thick in its basal portion, very
thin and strongly pointed towards the apex.

Gonapophyses of the $ $ shaped as shown on fig. [82].

Length 6-6,25 mm.

Rio da Areia, a fairly large pond in the forest; 27.111. 1922, 45 ad.,
9 larvae. A slide prepared of one of the $ $ is chosen as the type.

This species differs at once from the two preceding ones by its
smaller size, by the more slender body and by the sexual characters
of the $ $ r

64—6490

1488 The University Science Bulletin

Dr. Jaczewski refers in the above paragraph to B. crassipes
(Champion) and B. femoralis (Fieber).

LITERATURE CITED

( Additional references cited under genus and species )

Bare, C. O.

1925. A new species of Buenoa (Hemiptera, Notonectidae ) . Entomo-
logical News, vol. XXXVI, pp. 225-228.

1928. Haemoglobin cells and other studies of the genus Buenoa

( Hemiptera, Notonectidae ) . The University of Kansas Science
Bulletin, vol. XVIII, pp. 265-349.

1931. A Buenoa of southwest United States and Mexico (Hemiptera).

The Pan-Pacific Entomologist, vol. VII, pp. 115-118.

Berg, C.

1879. Hemiptera Argentina, pp. 18-316.

Champion, G. C.

1901. Rhynchota Heteroptera II. Biologia Centrali Americana, vol. II,
416 pp.

Fabricius, J. C.

1803. Systema Rhyngotorum, 314 pp.

Fieber, F. X.

1851. Rhynchotographieen. Abhandlungen Kongl. Bohmischen Gesell-
schaft Wissenschaften, vol. VII, pp. 469-486.

Hungerford, H. B.

1919. The biology and ecology of aquatic and semiaquatic Hemiptera.
The University of Kansas Science Bulletin, vol. XI, pp. 1-341.

1922. Oxyhemoglobin present in backswimmer Buenoa margaritacea

Bueno (Hemiptera). The Canadian Entomologist, vol. LIV, pp.
262-263.

1923. A new species of the genus Buenoa (Hemiptera, Notonectidae).

Entomological News, vol. XXXIV, pp. 149-152.

1924. Stridulation of Buenoa limnocastoris Hungerford and systematic

notes on the Buenoa of the Douglas Lake region of Michigan, with
the description of a new form. Annals of the Entomological So-
ciety of America, vol. XVII, pp. 223-227.

1933. The genus Notonecta of the World. The University of Kansas

Science Bulletin, vol. XXI, pp. 5-195.

1950. Two new generic names. The Journal of the Kansas Entomo-

logical Society, vol. XXIII, p. 73.

Jaczewski, T.

1928. Notonectidae from the state of Parana. Annales Musei Zoologici
Polonici, vol. VII, pp. 121-136.

KlRKALDY, G. W.

1899. On some aquatic Rhynchota from Jamaica. The Entomologist, vol.
XXXII, pp. 28-30.

A Revision of the Genus Buenoa 1489

1904. Uber Notonectiden. Wiener Entomologische Zeitung, vol.

XXIII, pp. 93-135.

Mayr, E.

1942. Systematics and die origin of species. New York, N. Y., Columbia
University Press, xiv + 334 pp.

Moore, R. C.

1949. Introduction to historical geology. New York, N. Y., McGraw-Hill
Book Company, Inc., ix -f- 582 pp.

Poisson, R.

1925. L'Anisops producta Fieber (Hemiptera, Notonectidae ) observations
sur son anatomie et sa biologic Archives de Zoologie Experi-
mental et Generate, vol. LXV, pp. 181-208.

TORRE-BUENO, J. R. DE LA.

1909. The notonectid genus Buenoa Kirkaldy. Journal of the New York
Entomological Society, vol. XVII, pp. 74-77.

Truxal, F. S.

1949. A study of the genus Martarega (Hemiptera, Notonectidae). The
Journal of the Kansas Entomological Society, vol. XXII, pp. 1-24.

1952. The comparative morphology of the male genitalia of the Noto-

nectidae (Hemiptera). The Journal of the Kansas Entomological
Society, vol. XXV, pp. 30-38.

White, F. B.

1879. List of the Hemiptera collected in the Amazons by Prof. J. W. H.
Trail, M. A., M. D., in the years 1873-1875, with descriptions of
the new species. Transactions of the Entomological Society of
London, Pt. IV, p. 271.

1490 The University Science Bulletin

INDEX

PAGE

absidata n. sp 1391

albida ( Champion ) 1412

alterna n. sp 1474

amnigenus ( White ) 1462

antigone ( Kirkaldy) 1374

antigone antigone ( Kirkaldy) 1376

antigone carinata ( Champion) 1379

arida n. sp 1435

arizonis Bare 1387

arfafrons n. sp 1444

communis n. sp 1442

confusa n. sp 1453

crassipes ( Champion ) 1385

distincta n. sp. 1485

excavata n. sp 1479

femoralis ( Fieber ) 1382

fuscipennis ( Berg ) 1460

gracilis n. sp 1439

hungerfordi n. sp 1483

ida Kirkaldy 1383

incompta n. sp 1466

limnocastoris Hungerford 1450

macrophthalma ( Fieber) 1480

macrotibialis Hungerford 1446

macrotrichia n. sp 1429

margaritacea Torre-Bueno 1397

mutabilis n. sp 1432

nitida n. sp 1430

oculata n. sp 1467

omani n. sp 1426

pallens ( Champion ) 1414

pallipes ( Fabricius ) 1418

paranensis Jaczewski I486

platycnemis ( Fieber) 1421

rostra n. sp 1395

salutis Kirkaldy 1469

scimitra Bare 1404

speciosa n. sp 1437

tarsalis n. sp 1392

thomasi n. sp 1473

uhleri n. sp 1409

unguis n. sp 1476

PLATE CI

Fig. 1. Male Buenoa — dorsal aspect.
Fig. 2. Male Buenoa — ventral aspect.

A Revision of the Genus Buenoa

1491

PLATE CI

o
a

Eo
^ °2

0 _

1 £

t JL

3
ffi

a>
o

3
V)

w
O

■D

1492 The University Science Bulletin

PLATE CII

Fig. 3. Buenoa macrophthalma (Fieber). Spine from caudo-sinistral mar-
gin of seventh abdominal tergite of male.

Fig. 4. Buenoa femoralis ( Fieber ) . Spine from caudo-sinistral margin of
seventh abdominal tergite of male.

Fig. 5. Buenoa pallens (Champion). Spine from caudo-sinistral margin
of seventh abdominal tergite of male.

Fig. 6. Buenoa scimitra Bare. Spine from caudo-sinistral margin of seventh
abdominal tergite of male.

Fig. 7. Buenoa fuscipennis (Berg). Spine from caudo-sinistral margin of
seventh abdominal tergite of male.

Fig. 8. Buenoa limnocastoris Hungerford. Spine from caudo-sinistral mar-
gin of seventh abdominal tergite of male.

Fig. 9. Buenoa crassipes (Champion). Spine from caudo-sinistral margin
of seventh abdominal tergite of male.

Fig. 10. Buenoa arizonis Bare. Spine from caudo-sinistral margin of seventh
abdominal tergite of male.

Fig. 11. Buenoa pallens (Champion). Spine from caudo-sinistral margin
of seventh abdominal tergite of male.

Fig. 12. Buenoa excavata n. sp. Spine from caudo-sinistral margin of
seventh abdominal tergite of male.

Fig. 13. Buenoa distincta n. sp. Spine from caudo-sinistral margin of
seventh abdominal tergite of male.

Fig. 14. Buenoa hungerfordi n. sp. Spine from caudo-sinistral margin of
seventh abdominal tergite of male.

Fig. 15. Cephalic aspect showing wide interocular cephalic space.

Fig. 16. Cephalic aspect showing narrow interocular cephalic space.

Fig. 17. Diagram showing method of measuring rostral prong.

Fig. 18. Diagram showing method of measuring width of head, vertex, and
synthlipsis.

Fig. 19. Diagram showing method of measuring length of fore femur and
width at apex.

Fig. 20. Buenoa confusa n. sp. Male genital capsule.

Fig. 21. Buenoa hungerfordi n. sp. Male genital capsule.

Fig. 22. Buenoa distincta n. sp. Male genital capsule.

A Revision of the Genus Buenoa

1493

PLATE CII

3. B mocrophtholmo 4 B. fomorolis 5. B. pollens 6 B. scimitro 7 B. (uscipennis 8 B. limnocostoris

21- B. hjngerford

22. B. distmcto

1494 The University Science Bulletin

PLATE CIII

Fig. 23. Buenoa hungerfordi n. sp.

23a. Left genital clasper.

23b. Right genital clasper.
Fig. 24. Buenoa distincta n. sp.

24a. Left genital clasper.

24b. Right genital clasper.
Fig. 25. Buenoa thomasi n. sp.

25a. Left genital clasper.

25b. Right genital clasper.
Fig. 26. Buenoa oculata n. sp.

26a. Left genital clasper.

26b. Right genital clasper.
Fig. 27. Buenoa macrophthalma (Fieber).

27a. Left genital clasper.

27b. Right genital clasper.
Fig. 28. Buenoa fuscipennis (Berg).

28a. Left genital clasper.

28b. Right genital clasper.
Fig. 29. Buenoa arizonis Bare.

29a. Left genital clasper.

29b. Right genital clasper.
Fig. 30. Buenoa antigone ( Kirkaldy ) . Laterodorsal view of left ovipositor
valve.

Fig. 31. Buenoa uhleri n. sp. Laterodorsal view of left ovipositor valve.
Fig. 32. Buenoa arizonis Bare. Laterodorsal view of left ovipositor valve.
Fig. 33. Buenoa margaritacea Torre-Bueno. Laterodorsal view of left ovi-
positor valve.

Fig. 34. Buenoa confusa n. sp. Laterodorsal view of left ovipositor valve.
Fig. 35. Buenoa alterna n. sp. Laterodorsal view of left ovipositor valve.
Fig. 36. Buenoa amnigenus (White). Laterodorsal view of left ovipositor
valve.

Fig. 37. Buenoa omani n. sp. Laterodorsal view of left ovipositor valve.
Fig. 38. Buenoa macrotibialis Hungerford. Laterodorsal view of left ovi-
positor valve.

Fig. 39. Buenoa salutis Kirkaldy. Laterodorsal view of left ovipositor valve.
Fig. 40. Buenoa limnocastoris Hungerford. Laterodorsal view of left ovi-
positor valve.

Fig. 41. Buenoa hungerfordi n. sp. Laterodorsal view of left ovipositor
valve.

A Revision of the Genus Buenoa

1495

PLATE CIII

25o. 25b.

25. 8. thomosl

26o. 26b

26. B. oculofo

30. B ontigonc 31 B uhleri

27b V ^ 29o.

27. B. macrophtholmo 29. B. orizonls

28o. 28b.

28. 6. fuscipennis

32 B arizonis 33 B morgaritaceo 34 B confusa 35 B olterno

36 B omnigenus 37 B omoni 38 B mocroliblolis 39 B solutis 40 B limnocaslorr's 41 B. hungerfordi

1496 The University Science Bulletin

PLATE CIV

Fig. 42. Buenoa antigone antigone ( Kirkaldy ) .

42a. Inner surface view of male left fore leg.

42b. Left lateral view of male rostrum and tylus.

42c. Enlarged view of left tibial stridulatory comb.

Fig. 43. Buenoa antigone carinata ( Champion ) .

43a. Inner surface view of male left fore leg.

43b. Left lateral view of male rostrum and tylus.

43c. Enlarged view of left tibial stridulatory comb.

A Revision of the Genus Buenoa

1497

PLATE CIV

42. B. ANTIGONE ANTIGONE

43 B ANTIGONE CARINATA

1498 The University Science Bulletin

PLATE CV

Fig. 44. Buenoa femoralis (Fieber).

44a. Inner surface view of male left fore leg.

44b. Left lateral view of male rostrum and tylus.

44c. Enlarged view of left tibial stridulatory comb.
Fig. 45. Buenoa ida Kirkaldy.

45a. Inner surface view of male left fore leg.

45b. Left lateral view of male rostrum and tylus.

45c. Enlarged view of left tibial stridulatory comb.

A Revision of the Genus Buenoa

1499

PLATE CV

45. B. IDA

1500 The University Science Bulletin

PLATE CVI

Fig. 46. Buenoa crassipes (Champion).

46a. Inner surface view of male left fore leg.

46b. Left lateral view of male rostrum and tylus.

46e. Enlarged view of left tibial stridulatory comb.

46d. Enlarged view of variable form of femoral stridulatory area.

46e. Enlarged view of variable form of tibial stridulatory comb.

Fig. 47. Buenoa arizonis Bare.

47a. Inner surface view of male left fore leg.

47b. Left lateral view of male rostrum and tylus.

47c. Enlarged view of left tibial stridulatory comb.

A Revision of the Genus Buenoa

1501

PLATE CVI

46o

46 B CRASSIPES

47. B ARIZONIS

1502 The University Science Bulletin

PLATE CVII

Fig. 48. Buenoa absidata n. sp.

48a. Inner surface view of male left fore leg.

48b. Left lateral view of male rostrum and tylus.

48c. Enlarged view of left tibial stridulatory comb.

Fig. 49. Buenoa tarsalis n. sp.

49a. Inner surface view of male left fore leg.

49b. Inner surface view of intermediate tarsus of male.

49c. Left lateral view of male rostrum and tylus.

49d. Enlarged view of left tibial stridulatory comb.

Fig. 50. Buenoa rostra n. sp.

50a. Inner surface view of male left fore leg.

50b. Left lateral view of male rostrum and tylus.

50c. Enlarged view of left tibial stridulatory comb.

A Revision of the Genus Buenoa

1503

PLATE CVII

48. B. ABSIDATA

50o

50. B. ROSTRA

65—6490

1504 The University Science Bulletin

PLATE CVIII

Fig. 51. Buenoa margaritacea Torre-Bueno.

51a. Inner surface view of male left fore leg.

51b. Enlarged view of left tibial stridulatory comb.

51c. Left lateral view of male rostrum and tylus.

Fig. 52. Buenoa scimitra Bare.

52a. Inner surface view of male left fore leg.

52b. Left lateral view of variable form of male rostral prong.

52c. Left lateral view of male rostrum and tylus.

52d. Enlarged view of left tibial stridulatory comb.

Fig. 53. Buenoa uhleri n. sp.

53a. Inner surface view of male left fore leg.

53b. Left lateral view of male rostrum and tylus.

53c. Enlarged view of left tibial stridulatory comb.

A Revision of the Genus Buenoa

1505

PLATE CVIII

53 B. UHLERI

1506 The University Science Bulletin

PLATE CIX

Fig. 54. Buenoa albida (Champion).

54a. Inner surface view of male left fore leg.

54b. Left lateral view of male rostrum and tylus.

54c. Enlarged view of left tibial stridulatory comb.
Fig. 55. Buenoa pollens (Champion).

55a. Inner surface view of male left fore leg.

55b. Left lateral view of male rostrum and tylus.

55c. Left lateral view of variable form of male rostral prong.

55d. Inner surface view of variable form of male fore femur.

55e. Enlarged view of left tibial stridulatory comb.

55f. Enlarged view of variable form of tibial stridulatory comb.
Fig. 56. Buenoa pallipes (Fabricius).

56a. Inner surface view of male left fore leg.

56b. Left lateral view of male rostrum and tylus.

56c. Enlarged view of left tibial stridulatory comb.

A Revision of the Genus Buenoa

1507

PLATE CIX

56. B. PALLIPES

1508 The University Science Bulletin

PLATE CX

Fig. 57. Buenoa platycnemis (Fieber).

57a. Inner surface view of male left fore leg.

57b. Left lateral view of male rostrum and tylus.

57e. Enlarged view of left tibial stridulatory comb.

Fig. 58. Buenoa omani n. sp.

58a. Inner surface view of male left fore leg.

58b. Left lateral view of male rostrum and tylus.

58c. Enlarged view of left tibial stridulatory comb.

Fig. 59. Buenoa macrotrichia n. sp.

59a. Inner surface view of male left fore leg.

59b. Left lateral view of male rostrum and tylus.

59c. Enlarged view of left tibial stridulatory comb.

A Revision of the Genus Buenoa
PLATE CX

1509

57. B. PLATYCNEMIS

58o

58. B. OMANI

59a

59. B. MACROTRICHIA

1510 The University Science Bulletin

PLATE CXI

Fig. 60. Buenoa nitida n. sp.

60a. Inner surface view of male left fore leg.

60b. Left lateral view of male rostrum and tylus.

60e. Enlarged view of left tibial stridulatory comb.
Fig. 61. Buenoa mutabilis n. sp.

61a. Inner surface view of male left fore leg.

61b. Left lateral view of male rostrum and tylus.

61c. Left lateral view of variable form of male rostral prong.

61d. Enlarged view of left tibial stridulatory comb.
Fig. 62. Buenoa arida n. sp.

62a. Inner surface view of male left fore leg.

62b. Left lateral view of male rostrum and tylus.

62c. Enlarged view of left tibial stridulatory comb.

A Revision of the Genus Buenoa
PLATE CXI

1511

61 a.

61. B. MUTABILIS

62 o.

62. B. ARIDA

1512 The University Science Bulletin

PLATE CXII

Fig. 63. Buenoa speciosa n. sp.

63a. Inner surface view of male left fore leg.

63b. Enlarged view of left tibial stridulatory comb.

63c. Left lateral view of male rostrum and tylus.

Fig. 64. Buenoa gracilis n. sp.

64a. Inner surface view of male left fore leg.

64b. Left lateral view of male rostrum and tylus.

64c. Enlarged view of left tibial stridulatory comb.

64d. Enlarged view of peg-like setae on inner surface of fore tibia.

Fig. 65. Buenoa communis n. sp.

65a. Inner surface view of male left fore leg.

65b. Left lateral view of male rostrum and tylus.

65c. Enlarged view of left tibial stridulatory comb.

A Revision of the Genus Buenoa
PLATE CXII

1513

65 o

65. 8. COMMUNIS

1514 The University Science Bulletin

PLATE CXIII

Fig. 66. Buenoa artafrons n. sp.

66a. Inner surface view of male left fore leg.

66b. Left lateral view of male rostrum and tylus.

66c. Enlarged view of left tibial stridulatory comb.

Fig. 67. Buenoa macrotibialis Hungerford.

67a. Inner surface view of male left fore leg.

67b. Left lateral view of male rostrum and tylus

67c. Enlarged view of left tibial stridulatory comb.

Fig. 68. Buenoa limnocastoris Hungerford.

68a. Inner surface view of male left fore leg.

68b. Left lateral view of male rostrum and tylus.

68c. Enlarged view of left tibial stridulatory comb.

A Revision of the Genus Buenoa
PLATE CXIII

1515

68. B. LIMNOCASTORIS

1516 The University Science Bulletin

PLATE CXIV

Fig. 69. Buenoa confusa n. sp.

69a. Inner surface view of male left fore leg.

69b. Left lateral view of male rostrum and tylus.

69c. Enlarged view of left tibial stridulatory comb.
Fig. 70. Buenoa amnigenus (White).

70a. Inner surface view of male left fore leg.

70b. Enlarged view of left tibial stridulatory comb.

70c. Left lateral view of male rostrum and tylus.
Fig. 71. Buenoa fuscipennis (Berg).

71a. Inner surface view of male left fore leg.

71b. Left lateral view of male rostrum and tylus.

71c. Enlarged view of left tibial stridulatory comb.
Fig. 72. Buenoa oculata n. sp.

72a. Inner surface view of male left fore leg.

72b. Left lateral view of male rostrum and tylus.

72c. Enlarged view of left tibial stridulatory comb.

72d. Inner surface view of intermediate tarsus of male.
Fig. 73. Buenoa incompta n. sp.

73a. Inner surface view of male left fore leg.

73b. Left lateral view of male rostrum and tylus.

73c. Enlarged view of left tibial stridulatory comb.

A Revision of the Genus Buenoa

1517

PLATE CXIV

73 B. INCOMPTA

1518 The University Science Bulletin

PLATE CXV

Fig. 74. Buenoa salutis Kirkaldy.

74a. Inner surface view of male left fore leg.

74b. Left lateral view of male rostrum and tylus.

74c. Left lateral view of variable form of male rostral prong.

74d. Enlarged view of left tibial stridulatory comb.
Fig. 75. Buenoa excavata n. sp.

75a. Inner surface view of male left fore leg.

75b. Left lateral view of male rostrum and tylus.

75c. Enlarged view of left tibial stridulatory comb.
Fig. 76. Buenoa thomasi n. sp.

76a. Inner surface view of male left fore leg.

76b. Enlarged view of left tibial stridulatory comb.

76c. Left lateral view of male rostrum and tylus.

76d. Enlarged view of club-like setae on inner surface of fore tibia.
Fig. 77. Buenoa alterna n. sp.

77a. Inner surface view of male left fore leg.

77b. Left lateral view of male rostrum and tylus.

77c. Enlarged view of left tibial stridulatory comb.
Fig. 78. Buenoa unguis n. sp.

78a. Inner surface view of male left fore leg.

78b. Enlarged view of left tibial stridulatory comb.

78c. Left lateral view of male rostrum and tylus.

78d. Enlarged view of male fore tarsus.

A Revision of the Genus Buenoa

1519

PLATE CXV

78. B UNGUIS

66—6490

1520 The University Science Bulletin

PLATE CXVI

Fig. 79. Buenoa macrophthalma (Fieber).

79a. Left lateral view of male rostrum and tylus.

79b. Inner surface view of femur and tibia of male fore leg.

79c. Inner surface view of male fore tarsus.

Fig. 80. Buenoa hungerfordi n. sp.

80a. Inner surface view of femur and tibia of male fore leg.

80b. Left lateral view of male rostrum and tylus.

80c. Inner surface view of male fore tarsus.

A Revision of the Genus Buenoa
PLATE CXVI

1521

1522 The University Science Bulletin

PLATE CXVII

Fig. 81. Buenoa distincta n. sp.

81a. Inner surface view of male right fore leg.

81b. Enlarged view of right tibial stridulatory comb.

81c. Left lateral view of male rostrum and tylus.

Fig. 82. Buenoa paranensis Jaczewski.

82a. Right genital clasper of male.

82b. Left genital clasper of male.

82c. Spine from caudo-sinistral margin of seventh abdominal tergite of
male.

82d. Inner surface view of male right fore leg.

82e. Frontal view of male third rostral segment.

82f. Dorsal view of head, pronotum, and scutellum.

A Revision of the Genus Buenoa

1523

PLATE CXVII

82 d

82 f.

62 c.

82 b.
82. B. PARANENSIS

82 a.

THE UNIVERSITY OP KANSAS

SGIENGE BULLETIN

Vol. XXXV, Pt. II] September 10, 1953 [No. 12

A Review of the Lizards of Ceylon

Edward H. Taylor

Abstract. — The paper deals with the lizard fauna of Ceylon exclusive of the
family Scincidae. The material studied is that in the United States National
Museum, and that in the Edward H. Taylor — Hobart M. Smith collection
(Lawrence, Kansas). Since the species of skinks were treated recently, keys,
but not descriptions are included for this Family.

In the total lizard fauna six families, twenty-four genera and sixty-two
species and subspecies are represented.

TABLE OF CONTENTS

PAGE

Introduction 1527

Distribution of Asiatic genera 1528

Taxonomic treatment 1530

Suborder Sauria 15S0

Key to Families of the Suborder Sauria 1530

Family Gekkonidae 1530

Key to genera of Gekkonidae 1531

Genus Gymnodactylus 1531

Key to Ceylonese species of Gymnodactijlus 1532

Gymnodactylus yakhuna Deraniyagala 1534

Gymnodactylus collegalensis Beddome 1532

Gymnodactylus triedrus Giinther 1535

Gymnodactylus frenatus Giinther 1536

Genus Cnemaspis 1537

Key to Ceylonese species of Cnemaspis 1538

Cnemaspis jerdoni (Theobald) 1539

Cnemaspis podihuna Deraniyagala 1538

Cnemaspis gracilis ( Beddome) 1539

Cnemaspis kandiana (Kelaart) 1539

Cnemaspis tropidogaster (Boulenger) 1541

Genus Hemidactylus 1542

Key to Ceylonese species of Hemidactylus 1542

Hemidactylus brookii Gray 1543

Hemidactylus depressus Gray 1544

Hemidactylus triedrus ( Daudin ) 1546

Hemidactylus leschenaulti Dumeril and Bibron 1547

(1525)

1526 The University Science Bulletin

PAGE

Hemidactijlus frenatus Schlegel 1549

Hemidactylus maculatus hunae Deraniyagala 1550

Genus Cosymbotus 1551

Cosymbotus platyurus (Schneider) 1551

Genus Peropus 1551

Peropus mutilatus (Wiegmann) 1551

Genus Hemiphyllodactylus 1552

Hemiphyllodactylus typus typus Bleeker 1552

Genus Lepidodacti/lus 1553

Lepidodactylus lugubris Dumeril and Bibron 1553

Genus Lophopholis 1554

Lophopholis scabriceps (Annandale) 1554

Family Agamidae 1554

Key to Ceylonese genera of the Agamidae 1555

Genus Sitana 1555

Sitana ponticeriana Cuvier 1555

Genus Otocryptis 1556

Otocryptis wiegmanni Wagler 1556

Genus Cophotis 1558

Cophotis ceylanica Peters 1558

Genus Ceratophora 1559

Key to the species of Ceratophora 1560

Ceratophorus stoddarti Gray 1561

Ceratophorus tennanti Giinther 1560

Ceratophora aspera Giinther 1561

Genus Lyriocephalus 1564

Lyriocephahis scutatus ( Linnaeus) 1564

Genus Calotes 1565

Key to Ceylonese species of Calotes 1565

Calotes versicolor (Daudin) 1566

Calotes calotes ( Linnaeus ) 1568

Calotes liocephalus Giinther 1569

Calotes nigrilabris Peters 1570

Calotes ceylonensis Miiller 1572

Calotes liolepsis Boulenger 1573

Family Scincidae 1574

Key to Ceylonese genera of Scincidae 1574

Genus Mabuya 1575

Key to the Ceylonese species of Mabuya 1575

Genus Riopa 1575

Key to Ceylonese species of Riopa 1576

Genus Dasia 1576

Genus Sphenomorphus 1576

Key to the Ceylonese species of Sphenomorphus 1576

Genus Chalcidoseps 1576

Genus Nessia 1576

Key to Ceylonese species of Nessia 1576

Family Lacertidae 1577

Genus Cabrita 1577

Cabrita leschenaulti Milne-Edwards 1578

Taylor: Lizards of Ceylon 1527

PAGE

Family Varanidae 1578

Genus Varanus 1578

Key to Ceylonese species of the Varanus 1578

Varanus bengalensis bengalensis (Daudin) 1579

Varanus salvator salvator (Laurenti) 1580

Suborder Rhiptoglossa 1580

Family Chamaleonidae 1580

Genus Chamaeleon 1580

Chamaleon zeylanicus (Laurenti) 1580

INTRODUCTION

This paper concludes my study of the Ceylonese Squamata *
contained in the United States National Museum and those in the
Edward H. Taylor-Hobart M. Smith collection. A study of the
amphibians is being pursued at the present time.

The material in these collections (approximately 1367 speci-
mens) has recently been augmented by a collection presented by
Mr. W. W. A. Phillips, Esq., noted mammalogist of Tonacombe
Estates, Namunukula, Ceylon. The lizards from the latter collec-
tion are reported in this paper. The snakes from the collection are
being reported under another title ** in this journal.

While this paper treats of the lizards, exclusive of the family
Scincidae, I have, however, listed the genera and species of the
seines, reprinting also keys published in a previous paper ( loc. cit. ) .

The recent papers of Mr. P. E. P. Deraniyagala and the recent
monograph of Malcolm Smith, ( The Fauna of British India, includ-
ing Ceylon and Burma. Reptilia and Amphibia vol. II, Sauria),
have been of great help in this study.

* Papers previously published are: Taylor, Comments on Ceylonese snakes of the genus
Typhlops, with descriptions of new species. Univ. Kansas Sci. Bull., vol. 31, pt. 2, Nov. 1,
1947, pp. 283-298, text-figs. 1-3. Tavlor, Ceylonese lizards of the family Scincidae. Univ.
Kansas Sci. Bull., vol. 33, pt. 2, March 20, 1950, pp. 481-518, text figs. 1-8.

Taylor, A brief review of Ceylonese snakes. Univ. Kansas Sci. Bull., vol. 33, pt. 2,
March 20, 1950, pp. 519-603, pis. 12-25, text-figs. 1-3.

** Report on a small collection of snakes from Uva Province, Ceylon.

1528

The University Science Bulletin

DISTRIBUTION OF ASIATIC LIZARD GENERA

Table of Lizard genera for South Asia.

Africa

West

Asia

India

Ceylon

East Asia

East Ind.
Arch.

Teratoscincus. .

X
X
X
X
X
X

X
X
X
X
X
X
X
X
X

Stenodaetylus

X
X
X

Alsophylax

Gymnodactylus

Agamura

x (Tibet)

x (Aust.)

Cnemaspis

Calodactylodes

Ptyodactylus

Phyllodactylus. . .

Dravidogecko

X
X
X

Hemidactylus

Cosymbotus

X
X
X

X
X
X
X
X

x (Aust.)

Peropus

Hemiphyllodactylus . .

X
X

Gekko

Lepidodactylus

Ptychozoon

Andaman Is.

Phelsuma ....

X
X
X
X
X
X
X

Lophopholis . .

X
X

Eublepharis

X
X

Draco

Sitana ....

Otocryptis. . .

Ptyctolaemus . .

Cophotis

X
X
X

Ceratophora

Lyriocephalus

Goniocephalus

Mictopholis. . .

X
X

Oriocalotes . .

Japalura . . .

Salea . ,

X
X
X
X
X

Calotes

Psammophilus

Agama .

X
X

Phrynocephalus

Physignathus . .

X
X

x (Papuasia)

Leiolepis . . .

x (Aust.)

Uromastix ....

X
X

X
X
X

Mabuya

X
X
X
X
X
X

Dasia

Lygosoma

Sphenomorphus

Ateuchosaurus .

Leiolopisma

X
X
X
X
X

Ablepharus . .

Riopa

X
X

Tropidophorus

Ristella

Taylor: Lizards of Ceylon

1529

Distribution of Asiatic Lizard Genera — Concluded

Africa

West
Asia

India

Ceylon

East Asia

East Ind.
Arch.

Ophioscincus

X
X

Eumeces

X
X
X
X

X
X
X

Ophiomorus

Barkudia

X
X

Sepsophis

Chalcidoseps

X
X

Nessia

Dibamus

X
X

Takydromus

Acanthodactylus

Cabrita

X
X
X
X
X
X
X

Ophisops

X
X
X
X
X

Ophisaurus

X
X

Chamaeleon

Of the 69 genera appearing in the South Asian list, some 23 have
been found to occur in Ceylon and approximately 41 are known to
occur in territory formerly regarded as India. However when one
considers that Lyriocephalus, Ceratophora and Chalcidoseps are
presumably endemic in Ceylon, and Peropus and Cophotes pre-
sumably do not occur in India, only 18 genera (less than one-half)
of the 41 known in India occur in Ceylon.

There is a large number of genera largely confined to Africa and
southwest Asia which enter India in the drier regions in the north-
west, and their further distribution eastward is seemingly limited
by their adaptation to relatively low rainfall. In this group are
Teratoscincus, Stenodactylus, Alsophylax, Agamura, Pristurus, Ptyo-
dactylus, Eublepharis, Agama, Phrynocephalus, Uromystax, Eu-
meces, Scincus, Ophiomorus, Chalcides, Acanthodactylus, Ophisops,
Ereinias and Chamaeleon.

A few genera seem to have entered the Indian territory from the
east, among which Calotes, Dasia, and Tropidophorus may be men-
tioned.

A few genera bespeak a considerable age, and such groups as
Varanus, Ophisaurus, Leiolopisma, and Mabuya, widespread as
they are, seemingly have obliterated the evidence of their route of
movement in Asia.

1530 The University Science Bulletin

TAXONOMIC TREATMENT

Key to Suborders of Reptilia (treated in this work)

Tongue terminally club-shaped, greatly extensile; skin covered
with flattened or rounded tubercles; fingers and toes webbed
together in groups of two and three, the groups opposed; eye
except pupil covered by a thick granular single lid, each eye
independently movable Rhiptoglossa

Tongue not club-shaped, fingers and toes maintain freedom of
individual movement; when present not webbed together into
groups; eye usually with an upper and lower lid Sauria

Of the groups of reptiles treated in this paper a total of six
families are represented. The Gekkonidae and Scincidae are wide-
spread cosmopolitan families; the Agamidae, Lacertidae and Varani-
dae are old world; the Chamaeleonidae occur in Africa and Mada-
gascar with one species entering southwestern Asia and Ceylon.

The Varanidae have formerly been represented in the new
world by the Genera Saniwa, Palaeosonkoa, and Parasaniwa. The
past history extends from the Upper Cretaceous to the Oligocene.
The Chamaeleonidae likewise appear to have been present formerly
in the new world based on the Genus Chamaeleo in the Upper
Cretaceous.

There is no evidence of the presence of the Lacertidae or the
Agamidae, the latter being replaced in the Western Hemisphere by
the Iguanidae.

Two other Asiatic families are the Anguidae and Dibamidae,
neither of which are known to enter Ceylon. The former is repre-
sented by the same genus in North America. The Dibamidae have
presumably never reached the Western Hemisphere.

SUBORDER SAURIA

Key to Ceylonese Families of Sauria

1. Scales on top of head large, arranged symmetrically 2

No large symmetrical scales on top of head 3

2. Scales on entire body cycloid, imbricate, smooth or keeled, with

osteodermal plates; no femoral or preanal pores; pleurodont;
tongue feebly nicked, covered with imbricate scalelike papilla,

Scincidae
Imbricating dorsal and lateral scales on body, strongly differ-
entiated from ventral scales; no osteodermal plates; pleuro-
dont; femoral pores; tongue forked, anteriorly covered with
imbricate papillae, or plicae Lacertidae

3. Body scales imbricate; tongue rather broad and short, smooth or

with villose papillae; eyes with movable lids; acrodont Agamidae
Body skin tubercular or granular, rarely with some imbricating
dorsal scales; eyes without or with movable lids; pleurodont . 4

Taylor: Lizards of Ceylon 1531

4. Tongue broad, short, covered with villose papillae, eyes without
movable lids; femoral and preanal pores usually present; scales

granular or tubercular, rarely imbricating Gekkonidae

Tongue long and slender, deeply forked, retractile into a sheath;
back with rounded scales, nonimbricate Varanidae

Family Gekkonidae

Eight genera of the Gekkonidae are represented in the fauna of
Ceylon. Some fourteen other genera occur in Asia, a few of which
are known in India. In most cases the genera known to occur in
Ceylon also occur in India and often the same species is found in
both areas. The genera Lepidodactylus and Peropus are excep-
tions, neither being known in India. I suspect that India received
Hemiphyllodactylus from the same outside source, as did Ceylon.

Key to the Ceylonese Genera of Gekkonidae

1. Digits bent at an angle, and not or but little dilated 2

Digits not bent at an angle, and more or less dilated 3

2. Pupil round; small adult lizards under 40 mm Cnemaspis

Pupil vertical; larger, adult lizards more than 40 mm. . . Gymnodactylus

3. Dorsal scales imbricate Lophopholis

Dorsal scales not imbricate 4

4. A lateral cutaneous flap or expansion on sides of body; claws on

all digits Cosymbotus

No lateral cutaneous flap; digits variable 5

5. Terminal phalanges of four outer digits joined to the dilated

terminal portion and not arising angularly Lepidodactylus

Terminal phalanges of four outer digits free, rising angularly from
the dilated portion 6

6. Inner digit well developed, with terminal phalanx clawed; lamel-

lae under toes distinctly divided Hemidactylus

Inner digit variable without or with only a very small claw 7

7. Inner digit well developed, lacking a free terminal phalanx, the

claw small and often concealed; lamellae divided Peropus

Inner digit represented by a vestige without a free terminal
phalanx, clawless or with a minute claw; lamellae, part di-
vided, part single Hemiphyllodactylus

Genus Gymnodactylus Spix

Gymnodactylus Spix, Specimen novum Lacerta Brasil., 1825, p. 17, pi. 17,
fig. 1 (type of genus Gymnodactylus geckoides).

This genus which has its greatest development in South Asia, the
East Indies and the Philippines has four representatives known in
Ceylon, two of which Gymnodactylus triedrus and Gymnodactylus
frenatus are endemic. The latter species is the largest of the genus
in Asia, perhaps in the entire range. Gymnodactylus collegalensis
on the other hand is one of the smallest species of the genus.

1532 The University Science Bulletin

Deraniyagala has recently described a species, Gymnodactylus
yakhuna from the Island.

Key to Ceylonese Species of Gymnodactylus

1. Body above with small equal granules; no femoral pores 2

Granules on back intermixed with numerous larger keeled or

rounded tubercles; femoral pores present 3

2. Smaller, 40 mm.; the dark bands on body equal to or shorter than

the light interspaces; dark spots subrectangular, in two trans-
verse rows on body; a large, curving, dark, occipital spot; black
dots abundant on light interspaces; a few feebly enlarged sub-

caudals yakhuna

Larger, adults, 50-53 mm.; head flecked or dimly spotted; a pair
of occipital spots; neck and body with seven pairs of black
spots, bordered by lighter brown, or tan; tail, dorsally with a
few dark, transverse flecks, ventrally with seven or eight black
bars, separated by cream spots collegalensis

3. Granules of back intermixed with larger, keeled, trihedral tuber-

cles; three or four femoral pores; length, snout to vent, to

65 mm triedrus

Granules of back intermixed with large, rounded tubercles; males
with four to six preanal pores frenatus

Gymnodactylus collegalensis Beddome

Gt/mnodactijlus collegalensis Beddome, Madras Month. Journ. Med. Sci., vol. 2,
'1870, p. 173 (type locality, Balarangams, near Yelandur, Mysore, India);
Smith The fauna of British India, including Ceylon and Burma; Reptilia
and Amphibia, vol. 2, Sauria, 1935, pp. 56-57, pi. 1, figs. Bl, B2
Gymnodactylus nebulosus ( part. ) Boulenger, the Fauna of British India, in-
' eluding Cevlon and Burma; Reptilia and Batrachia, 1895, pp. 64-65; Annan-
dale, Spolia Zeylanica, 1906, p. 189 (18 mi. N. Kandy) Deraniyagala,
Ceylon Journ. Sci., B., vol. 16, 1932, p. 293, pi. 59.

Diagnosis: A small species, snout-to-vent length 54 mm.; tail 36;
limbs short, the leg not reaching axilla; scales on dorsum small uni-
form except for a few on distal part of flanks and on back between
hind legs; no preanal or femoral pores; more than 50 rows of sub-
circular ventral scales, not or but scarcely imbricating; whorls on
tail scarcely discernible, each with six rows of small scales and with-
out larger spiny scales.

Description of species: (From E.H.T.-N.M.S. No. 30483, 12-16
mi. N Trincomalee, Ceylon.) Rostral 2.2 mm. wide, 1.5 mm. high;
nostril surrounded by rostral, internasal, and three or four post-
nasals, the first labial excluded; scales on snout rather irregular,
much larger than on occiput, distinctly larger than dorsal granules;
rostral bordered by four nearly equal-sized scales; the internasals
widely separated; ten supralabials, six to a point below middle of

Taylor: Lizards of Ceylon 1533

orbit; infralabials, 9, 8; forty granules across head between the
edges of eyelids; granules uniform on dorsum and sides except for
about 20 scattered, larger, conical tubercles on flanks and between
hind limbs; mental relatively small, its labial border equal or a little
less than the labial border of rostral; a median pair of postmentals,
their common suture longer than the suture of one with the mental,
each touching one labial; second pair of postmentals small, touch-'
ing first and second labials; two or three rows of scales nearest
labial series somewhat enlarged; scales on chin granular, subcorneal;
scales on venter granular, subcorneal, not imbricating, about 58
rows on ventral surface, not differentiated from adjoining lateral
scales; scales of preanal region larger, flattened, somewhat imbri-
cating, lacking trace of femoral or preanal pores; tail showing
whorls, each with about six transverse rows of flattened scales, all
more or less imbricating, their surfaces finely striated or corrugated;
on under surface of tail scales slightly larger, the median row not
distinguishable from adjoining scales; no larger tubercles on tail;
fingers slightly angular, the distal half scarcely narrower than
proximal; scales on basal portion enlarged at least one half to two
thirds of width of digit and extending onto palm; the formula of
scales from claw to base of fingers one to five is: 7, 11, 12, 12, 11;
one or more scales may be divided; scale formula for toes: 8, 12, 16,
14, 14, with certain ones divided; hind leg reaches to within one
millimeter of axilla.

Color: Rich brown, with head flecked or dimly spotted with
darker color and a pair of distinct spots on occiput; neck and body
with seven pairs of small irregular black spots, bordered with much
lighter brown; upper and lower labials cream, with blackish brown
spots; chin cream with a series of brown, vermiform flecks or spots;
belly dirty white, with some scattered pigment; tail with a few
dark, transverse flecks; under surface of tail with seven or eight
black bars separated by cream spots; sides nearly uniform brown.

Measurements in mm.: Snout to vent, 45; tail, 37; arm, 16; leg,
20.2; axilla to groin, 22.5; width of head, 10.6; length of head, 13.

Remarks: All specimens of this species in the collections were
found along the beach north of Trincomalee (12-16 mi.) under
masses of coral and about a plant, Zanzeveria zeylanica, growing
in sand and coral along the beach. Here they are very incon-
spicuous. They are seemingly confined to regions of low elevation
in Ceylon. The species is known to occur also in southern India.

1534

The University Science Bulletin

Data on Gymnodactylus Collegalensis, from Ceylon'

Number

Sex

Length

Tail

Head, long

Head, wide

Axilla to groin. .

Arm

Leg

Supralabials

Infralabials

30466

30484

30475

30454

30471

30470

30467

30482

30483

52.3

35.5

50
32.5

44
38
14.4

15.3

14.2

14.3

10.5

10.3

9.2

29.2

22.5

17.5

17.6

15.2

21.5

20.3

21.4

9-9

11-11

10-10

9-9

10-9

10-11

10-10

9-9

8-9

9-10

9-9

8-8

9-8

8-7

8-9

30470
&
41.5

18.5

11-12

9-9

° All specimens from 12 m. N Trincomalee, Ceylon, in E. H. Taylor-H. M. Smith collec-

tion.

Gymnodactylus yakhuna Deraniyagala

Gymnodactylus yakhuna Deraniyagala, Proa, 32dn. Indian Science Congress,
III, (Abstracts) 1945, p. 114; Spolia Zeylanica, vol. 24, part 2, Dec. 22,
1945, pp. 100-102 (type locality, Kalivila, Northwestern Province, Ceylon)
( "forma typica" ) .

PGymnodactylus yakhuna variety zonatus Deraniyagala, Spolia Zeylanica, vol.
24, pt. 2, Dec. 22, 1945, p. 101 (type locality, Manampitiya, Ceylon).

I am somewhat confused by the recent treatment of a small spe-
cies of Gymnodactylus by Deraniyagala. It is described as G.
yakhuna. Concerning it, Deraniyagala states: " — The Ceylon gecko
also exists as two color varieties, and it is possible that further
examination might reveal them to be subspecifically distinct. To
be in keeping with the procedure adapted for Gymnodactylus col-
legalensis [by Smith 1935, pp. 56-57, pi. 1, figs. Bl, B2] it is now
proposed to designate the spotted variety as the 'forma typica' and
confer the name of zonatus upon the banded one."

I presume that the form previously recorded as G. collegalensis
occurring in Ceylon is regarded as a synonym of G. yakhuna. Un-
fortunately I have not been able to examine specimens of this latter
species (or the color varieties) and cannot judge the merit of dis-
tinguishing these from collegalensis by name. Nevertheless, it
would appear that a species differing from yakhuna occurs on the
east coast, and this is recognized in this paper as G. collegalensis.

Diagnosis: Head rather heavy; snout slightly longer than distance
between eye and ear; body and tail subcylindrical; no lateral fold;
middle of tail as thick as neck; limb short; supralabials, 10-9, infra-
labials, 9-7; rostral subrectangular touching two supranasals that
are separated by a single median scale; dorsal granules rounded,

Taylor: Lizards of Ceylon 1535

sometimes two or three enlarged ones on each side near the hips;
mental subpentagonal followed by two enlarged postmentals that
are broadly in contact with one another mesially; ventrals subim-
bricate, some apt to be subhexagonal in shape; caudals and sub-
caudals imbricate, a few may be somewhat enlarged; subdigital
lamellae equal to half width of basal part of digit; head with an
arrow-shaped, dark, median band which is interrupted in many
specimens, and there are three or four supraocular dark blotches.
A dark band runs laterally from snout to nape, and is confluent with
its fellow; ventrally dusky gray; throat with a number of wavy dark
lines running from lips to the midgular region. (Data from
Deraniyagala. )

Remarks: That Gymnodactylus yakhuna differs materially from
specimens at hand that I have designated Gymnodactylus colle-
galensis, is obvious. The specimens of the latter are larger, fully-
grown adults, ranging from 45 to 53 mm., snout to vent, and none
in my series displays the type of coloration recorded for yakhuna.
Specimens referred by me to collegalensis are all from 12-16 miles
north of Trincomalee along the shore in Eastern Province, but at a
point probably as far north as the type locality of the variety
zonatus.

The differences in the two color varieties are: Forma typica —
there are two crossrows of dark blotches on the body, each row
consisting of two subrectangular markings. Zonatus — there are two
crossbands on the body, which are equal to or shorter than the
light interspaces.

Gymnodactylus triedrus Giinther

Gymnodactylus triedrus Giinther, Reptiles of British India, 1864, p. 113, (type
locality, Ceylon); Boulenger, Catalogue of the Lizards in the British Mu-
seum, vol. 1, 1885, p. 38; The Fauna of British India, including Ceylon and
Burma; Reptilia and Batrachia, 1890, pp. 67-68; Annandale, Jour. Asiat. Soc.
Bengal, vol. 73, 1904, Suppl. p. 13; Rec. Ind. Mus., vol. 9, 1913, p. 320;
Deraniyagala, Ceylon Journ. Sci., B, Vol. 16, 1932, p. 295-296, pi. 61;
Smith, The Fauna of British India, including Ceylon and Burma; Reptilia and
Amphibia, vol. 2, Sauria, 1935, pp. 55-56.

Geckoella punctata Gray, Proc. Zool. Soc. London, 1867, p. 99, pi. 9, (type
locality, Ceylon).

Diagnosis: A large Gymnodactylus, the snout-to-vent measure-
ment 62 mm.; tail 56 mm.; 12-13 supralabials, 10-11 infralabials;
venter with 35 rows of cycloid, imbricating scales; body with small
granular scales mixed with larger, rounded, keeled, or trihedral
tubercles; toes short, the lamella on basal part scalelike, half as wide
as the digit; tail with small rounded imbricate scales, those on under
surface somewhat larger; three or four femoral pores.

67—6490

1536 The University Science Bulletin

Brown on dorsum; numerous white spots with darker edges
usually present; light brown on venter.

I have not examined this species. It is endemic in Ceylon, and
has not been taken elsewhere.

Gymnodactylus frenatus Giinther

Gymnodactyhis frenatus Giinther, Reptiles of British India, 1864, p. 113, pi. 12,
fig. D. (type locality, Ceylon). Boulenger, Catalogue of the Lizards in the
British Museum, 2nd ed., vol. 1, 1885, p. 42; The Fauna of British India,
including Ceylon and Burma; Reptilia and Batrachia, 1890, p. 68; Annan-
dale, Spolia Zeylanica, vol. 3, Jan. 1906, p. 190 (Kandy); Henry, Ceylon
Journ. Sci., B, vol. 16, 1928, p. 339; Deraniyagala, Ceylon Journ. Sci., B,
1932, pp. 294-295, pi. 41; Smith, The Fauna of British India, including
Ceylon and Burma; Reptilia and Amphibia, vol. 2, Sauria, 1935, p. 49;
Bhatt, Animal Life (Colombo Ceylon) 1942, pp. 122.

Diagnosis: A large lizard (head-body length 100 mm., tail 120
mm.) with dorsal scales granular, intermixed with large rounded
tubercles; tail with flat scales, the basal whorls with four somewhat
enlarged rounded tubercles, quickly becoming smaller on succeed-
ing whorls; male with four to six preanal pores; 33-35 scales across
belly; four or five large dark bands whose anterior and posterior
borders form angles.

Description of species: ( From K.U.M.N.H. No. 31258, Tonacombe
Estate, Namunukula, Ceylon). A large species, the head (20 mm.)
nearly as wide as long (21.5 mm.); snout-to-vent length 85 mm.,
tail, broken from body, the tip regenerated, 87 mm.; rostral large,
its width (4.7 mm.) greater than its height (2.3 mm.), a median
entrant suture more than half height of scale, the upper edges
curving toward middle; internasals large, separated by a median
scale and followed by one somewhat enlarged; nostril surrounded
by rostral, first labial, two or three postnasals and the internasal,
the latter almost excluded; scales on snout variable in size in sev-
eral areas; fifty scales across snout between the fourth labials; a
depression somewhat behind nostrils and a well-defined median
depression beginning at about posterior level of orbits and extend-
ing onto frontal region, narrowing between orbits; approximately
48 scales between outer edges of eyelids at middle of orbit, small,
irregular with a few larger flat rounded tubercles intermixed; supra-
labials, 9-10; infralabials, 8-8; scales adjoining labials somewhat
enlarged, at least anteriorly; mental border on lip greater than
rostral border, the sides of mental being nearly straight, forming
approximately a right angle posteriorly. A pair of postmentals
forming a median suture as long as their sutures with mental,
bordering first labial for about half its length; on each side a small
outer postmental touching two labials; two or three scales follow-

Taylor: Lizards of Ceylon 1537

ing somewhat enlarged; chin scales small, circular, rather regular;
ear diameter less than half eye diameter; distance from ear to eye
a little greater than distance from eye to nostril; ventrolateral folds
more or less distinct; scales of occiput and dorsum small, with about
14 irregular, longitudinal rows of larger rounded, flat, rarely
slightly keeled, tubercles; the three or four basal whorls of tail may
have as many as six tubercles, but reduce posteriorly to scales that
are not or scarcely differentiated; 6 transverse rows of small scales
on each whorl; on ventral surface two widened scales to each
whorl; abdominal scale rows, 30-33, cycloid, imbricate; an enlarged
row of femoro-preanal scales the inner two of the series bearing
preanal pores in the male, separated by three scales mesially; males
with three enlarged flattened tubercular scales on each side of tail
base; bases of angular digits widened, bearing single, transverse
lamellae, 7 to 9 in number; on both hands and feet lamellae con-
tinue below metacarpals and metatarsals for some distance; all
digits clawed; hind leg long, reaching axilla when adpressed.

Color: Above gray, the top of head with small spots or marbling
of brown; a light-edged, brown band from behind each eye unites
with a dorsal brown spot on beginning of neck; three brown W-
shaped blotches on back of body which are separated by gray bands
with some indefinite brown flecks; arms and legs banded or mottled;
tail with regular broad bands of brown or black; grayish below on
chin and venter, everywhere peppered with black pigment.

Remarks: The single specimen at hand is from an elevation of
4000 ft. on the Tonacombe Estate, Namunukula, Ceylon (Uva
Hills), W. W. A. Philipps, collector.

The species is endemic in Ceylon and has not been taken else-
where.

Genus Cnemaspis Strauch

Cnemaspis Strauch, Mem. Acad. St. Petersburg, vol. 35 (7), 1887, p. 41 (type
of genus, Cnemaspis bonlcngeri).

This genus is represented by at least four forms in Ceylon. Cer-
tain of these are regarded as variable and Malcolm Smith (1930)
throws some doubt on the validity of certain forms recognized
by Deraniyagala and Boulenger. Probably the last word cannot be
said on the matter until rather large series are available from
various parts of Ceylon and south India. Only then can one prop-
erly evaluate the differences in populations. It seems likely that
size of the animals may be a specific character. Measurements
offered by Smith for kandiana are considerably greater than any
specimens in my series of thirteen. Whether the forms here con-

1538 The University Science Bulletin

sidered are to be regarded as species or subspecies will depend on
more material than has been collected. I have regarded them as of
specific rank, but without conviction.

Key to Forms of Cnemasfis in Ceylon

1. Femoral pores less than five (rarely five) median subcaudals

not strongly enlarged 3

Femoral pores five or more; median subcaudals much enlarged . . 2

2. Femoral pores eight to fifteen jerdoni

Femoral pores eight or less podihuna

3. No keeled scales on ventral neck region or abdomen; femoral

pores three or four; preanal pores two to four gracilis

Keeled scales on ventral neck region or on both neck and ab-
domen 4

4. Ventral nuchal scales keeled over a greater or lesser area; ab-

dominal and subcaudal scales smooth; femoral pores, three to

five; preanal pores, two to four kandiana

Keeled scales on entire nuchal, ventral and ventrolateral areas, as
well as under tail; femoral pores 3; preanal pores absent (in
specimen examined) tropidogaster

Cnemaspis podihuna Deraniyagala

Cnemaspis podihuna Deraniyagala, Journ. Royal Asiat. Soc. (Ceylon branch)

no. 97, 1944, pp. 226-227, 1 fig. (type locality).
? Gyninodactylus scalpensis Ferguson, Reptile Fauna of Ceylon, Colombo,

1877, pp. 1-30.

Diagnosis: Femoral pores five; preanal pores four; subcaudal
scales strongly enlarged; tail with small lateral spines; some enlarged
lateral spinelike scales on the sides; nine supralabials; six or seven
infralabials.

I have not examined specimens of this species. It appears to
be closest to Cnemaspis jerdoni. The type locality of the latter is
unknown. Ferguson has described a form from Ceylon (without
specific locality) which he named scalpensis. Unfortunately the
type is lost. There is a possibility that podihuna is a synonym of
scalpensis.

Cnemaspis jerdoni (Theobald)

Gyninodactylus jerdoni Theobald, Catalogue of the Reptiles in the Asiatic

Society Museum, 1868, p. 31 (type locality not stated, India by inference).
Gonatodes jerdoni Boulenger, Catalogue of the Lizards in the British Museum,

vol. 1, 1885, p. 71; The Fauna of British India, including Ceylon and Burma;

Reptilia and Batrachia, 1890, p. 78. Deraniyagala, Ceylon Journ. Sci. B,

vol. 16, pt. 3, 1932, pp. 298-299.
Cnemaspis jerdoni Smith, The Fauna of British India, including Ceylon and

Burma; Reptilia and Amphibia, vol. 2, Sauria, 1935, pp. 74-75.

Diagnosis: This species is similar to gracilis in general scale char-
acters: the postmentals two or three; tail cylindrical; median subcau-

Taylor: Lizards of Ceylon 1539

dal scales larger than others. A series of five to fifteen femoral pores
on each side; ventral scales cycloid, smooth. Gray brown clouded
with darker. Small lateral spines white, sometimes a black spot
on back of occiput; "Snout to vent, 40 mm., tail 44 mm."

Cnemaspis gracilis ( Beddome )

Gymnodactylus gracilis Beddome, Madras Month. Journ. Med. Sci., vol. 1, 1870,
p. 32 (type locality, Palghat Hills, Madras, India).

Gonatodes gracilis Boulenger, Catalogue of the Snakes of the British Museum,
vol. 1, 1885, p. 70, pi. 6, fig. 5; The Fauna of British India, including Ceylon
and Burma, 1890, p. 78; Hora, Rec. Indian Mus., vol. 28, part 3, 1926,
pp. 191-192, pi. 7, fig. 7, test fig. 3 a. (Indian Records only).

Gonatodes kandianus gracilis Deraniyagala, Ceylon Journ. Sci., B, vol. 16,
1932, p. 297.

Cnemaspis gracilis Smith, The Fauna of British India, including Ceylon and
Burma; Reptilia and Amphibia vol. 2, Sauria, 1935, p. 74. (Hills of Ceylon)
("Doubtfully distinct from kandianus perhaps only a varietal form not
correlated with geographical distribution" ) .

Diagnosis: This form differs from the species kandiana in having
two instead of three scales bordering the mental. Scales on the
neck are larger, smooth, lacking keels; scales under basal part of
foot smaller; grayish or brownish above, the back of occiput with
darker spots; sometimes a series of light vertebral spots present.
A forest form.

Cnemaspis kandiana (Kelaart)

Gymnodactylus Kandianus Kelaart, Prod. Fauna Zeylanicae 1852 (1853),
p. 186 (type locality, "Kandian-hills", [hills about Kandy, Ceylon]);
Gimther, Reptiles of British India, 1864, p. 114.

Gymnodactylus humei Theobald, Cat. of the Reptiles of British India, 1865,
p. 89, (type locality, Kandy, Ceylon).

Gonatodes kandianus Boulenger, Catalogue of the Lizards of the British Mu-
seum, vol. 1, 1885, p. 68; The Fauna of British India, including Ceylon and
Burma; Reptilia and Batrachia, 1890, pp. 77-78; Hora, Rec. Indian Mus.
vol. 28, pt. 3, 1926, pp. 191-193 (Kandy and Peradeniya, Ceylon). "G.
kandianus possesses a relatively longer and more pointed snout and three
chinshields behind the mental").

Diagnosis: Small geckoes, the maximum snout-to-vent length
approximately 40 mm.; tail, 45 mm.; fingers slender, the basal por-
tion of digits with a few enlarged, flat plates; femoral pores few,
widely separated from a small series of two or three preanal pores;
scales on back small, somewhat keeled granules, with some scat-
tered larger ones; on flanks, scales prominent, spinelike; belly
scales imbricate; those on neck with fine keels.

Description of species: (from E.H.T.-H.M.S. No. 30517). Ros-
tral (1.5 mm. x9 mm.) reaches above level of nostril, which is
surrounded by rostral, two postnasals and an internasal (the first
labial being excluded), and the internasal forming a very small
contact; latter scales separated mesially by a scale nearly as large

1540 The University Science Bulletin

as internasals but extending somewhat farther forward (in this
specimen, partially fused to rostral); seven or (eight) supralabials,
the fourth extending under front of orbit; between front part of
fourth labials across snout, 28 relatively large scales; scales bor-
dering labials not larger than snout scales; infralabials, 8-8; mental
very large, its labial border considerably greater than labial border
of rostral; a pair of chinshields separated mesially by a much
smaller scale; second chinshields smaller, separated by three inter-
vening scales all larger than other chin scales which are keeled on
side of jaws; scales on sides of neck larger, with keels, the keeled
extending on sides of neck to arm insertion; scales of venter some-
what cycloid, imbricate, smooth, in 24 rows; scales on sides keeled,
while on back they are smaller, almost trihedral, with a few larger
pointed scales often elevated to resemble small spines; tail with
small scales arranged in eight rows to a whorl, each whorl bearing
two small pairs of dorsal spines and one lateral pair; a very heavy
pyramidal scale at base of tail on each side; tail regenerated (ven-
trally, on original tails, three scales to a whorl, every third scale
being somewhat enlarged); four femoral pores placed ventropos-
teriorly; two preanal pores separated from femoral pores by seven
or eight scales; digits with a long slender distal part and a widened
basal part except on first inner finger and toe where the free basal
part is missing; on basal part of four outer fingers, there are four
or five widened flat scales; on toes five or six; a single large scale
at base of first finger and toe; ear about one third of eye diameter;
the distance between ear and eye more than a third less than dis-
tance from eye to nostril.

Color: Above dark, gray-brown with some blackish flecks tend-
ing to form a transverse band; pigment on ventral surface often
arranged in half-moon groupings on posterior edges of scales; chin
and throat dark (at least in males); tail indistinctly barred (when
complete, the dark marks narrow).

Measurements in mm.: Snout-to-vent length, 31; tail regenerated;
width of head, 5.45; length of head, 9.

Remarks: The following specimens have been examined: E.H.T.-
H.M.S. Nos. 30514 Kandy; 30515-30518 Badulla; K.U.M.N.H. Nos.
31265-31275 Tonacombe Estate, Namunukula, 4000 ft. elevation, in
Uva Hills.

The largest of this series of specimens measures only 31 mm.
snout to vent; tails for the most part are incomplete or reproduced.
No. 30514 measures 25 mm., the tail being 32 mm.

One of the specimens has only a single preanal pore, and four

Taylor: Lizards of Ceylon 1541

femoral pores. Those from Tonacombe Estate have a curving, light
gray line from eye around occiput to eye, bordered anteriorly by a
darker mark; mesially on occiput there is a lighter spot; behind this
there is a curving darker spot; some specimens show eight or nine
transverse bars on body, the dark bars being somewhat angulate or
W-shaped.

Cnemaspis tropidogaster (Boulenger)

Gonatadcs kandiamus tropidogaster Boulenger, Catalogue of the lizards in the
British Museum, vol. 1, 1885, p. 70 (type locality, Ceylon); The Fauna of
British India including Ceylon and Burma; Beptilia and Batrachia, 1890,
p. 78.

Diagnosis: Entire ventral surface of throat, venter and under side
of tail strongly keeled; ventral scales under tail all relatively small,
more or less equal in size; three femoral pores; no preanal pores (?).

Description of species: (From E.H.T.-H.M.S., No. 30246 Galle,
Ceylon. ) The form agrees in certain characters with kandiana but
differs in the character of the squamation; 24 mm. snout-to-vent
(tail broken and posterior part missing); nostril surrounded by
rostral, an internasal and two postnasals; pair of internasals sepa-
rated mesially by one scale; 29 scales across snout between fourth
labials; supralabials, 7-7; infralabials, 7-8; approximately 42 scales
across head between edges of eyelids at middle of orbit; scales on
snout larger than on occiput, rather equal and regular; granules
much finer on occiput; on body, scales nearly as large as those on
snout, but keeled or trihedral with a few larger, scattered, irregu-
larly placed tubercles; scales on arms and legs, as well as on sides
and ventral surfaces, keeled, except for a few in preanal region;
median scales under tail pointed, keeled and subequal, but little
larger than other subcaudals, which are all strongly keeled; scales
above arranged in whorls, each whorl with eight or nine rows of
keeled scales and a transverse series of large pointed spines, six at
least on basal whorls; a very heavy, spiny tubercle on each side at
base of tail; a few lateral spines present.

Distal part of fingers compressed, the basal part wider, bearing
three or four flat scales on four outer fingers; one at base of the
compressed part of first finger; four or five scales under four outer
toes, one under first toe; hind leg brought forward, the toes touch
axilla (skin damaged so no count of abdominal scales can be made);
three femoral pores; I cannot discern preanal pores.

Color: Brown or gray-brown above; a pair of irregular occipital
dark spots with a lighter cream spot between them; some darker
clouding on back and sides, the median region a little lighter.

Remarks: Most records of Cnemaspis tropidogaster are from low-

1542 The University Science Bulletin

land localities, while kandiana is from the mountainous regions or
highlands. Boulenger, loc. cit., regarded tropidogaster a variety
(subspecies) of kandiana. Smith throws it into synomomy of
kandiana. Deraniyagala reports it from Peradeniya however.

It seems to me that a recognition of the form is essential as a
species or a subspecies. I prefer to use the former relationship
since the characters seem to warrant such usage.

Genus Hemidactylus Oken

Hemidactylus Oken, Isis, 1817, p. 1183 (type of the genus, Gecko tuberculosa).

This very large, cosmopolitan genus is represented in Ceylon by
six species. The two smaller species, Hemidactylus frenatus, and
Hemidactylus brookii, the most common of the domestic geckoes,
are widespread forms, the first of the two having become estab-
lished in Mexico. Four other species are tree-dwelling forest
geckoes, although rarely they may be found in human habitations.
These are: Hemidactylus triedrus and leschenaulti, which occur
also in India, Hemidactylus depressus, an endemic Ceylonese form
unknown elsewhere, and Hemidactylus maculatus hunae, a Cey-
lonese representative of an Indian species.

Key to the Species of Hemidactylus in Ceylon

1. Dorsal tubercles usually trihedral, strongly developed, in 16-20

rows 2

Dorsal tubercles few, rounded, not or but slightly keeled, usually
confined to posterior part of back 5

2. Small forms usually less than 52 mm. snout to vent length;

femoro-preanal pores, 11-15, separated by three poreless scales

mesially; dorsal tubercles in 18-20 rows brookii

Larger forms, 80-115 mm. in snout-to-vent length 3

3. Femoral pores only, 19-25 on each side; 5 transverse bands on

body with median lighter spots; very large, snout to vent

115 mm maculatus hunae

Femoro-preanal pores; markings varied; smaller, probably not ex-
ceeding 90 mm 4

4. Digits webbed at base; 16-19 femoro-preanal pores on each side,

separated mesially; tubercles smaller depressus

Digits free at base; preanal pores 6-9, separated mesially by one
to three scales; tubercles larger triedrus

5. Dorsal tubercles reduced in number (rarely absent), smooth or

faintly keeled; head and body broad; snout-to-vent length,
83 mm.; male with 10-17 femoro-preanal pores on each side,

separated mesially by a distinct interval leschenaulti

Dorsal tubercles few, usually two or three smooth rows on pos-
terior part of back; male with a continuous series of femoro-
preanal pores, 13 to 18 on each side; snout to vent, 60 mm.,

frenatus

Taylor: Lizards of Ceylon 1543

Hemidactylus brookii Gray

Hemidactylus brookii Gray, Catalogue of the Specimens of Lizards in the
collection of the British Museum, 1845, p. 153 (type locality, Borneo);
Boulenger, Catalogue of the Lizards of the British Museum, vol. 1, 1885,
p. 128; Deraniyagala, Ceylon Journ. Sci., B, 16, 1932, p. 300.

Hemidactylus gleadovii Boulenger, The Fauna of British India, including Ceylon
and Burma; Reptilia and Batrachia, 1890, pp. 86, 87, fig. 27.

Hemidactylus maculatus? Kelaart, Prod. Fauna Zeylanica, 1852 (1853) p. 158
( Ceylon, widely distributed ) ( not of Dumeril and Bibron ) .

Diagnosis: A small, widespread species with a maximum snout-
to-vent length of 60 mm., the tail approximately 78 mm.; femoro-
preanal pores, 11-15; scales of tail in whorls, each bearing eight or
six sharp, spiny tubercles; back with 18-22 rows of trihedral tuber-
cles; scales across abdomen cycloid, imbricate, 36-40 in number.

Description of species: (From E.H.T.-H.M.S. No. 30417.) A
rather small species, the head-width in head-length, 1.5 times;
diameter of ear distinctly less than half of eye-opening, its distance
from eye a little less than distance between eye and nostril; width
of rostral, 2.1 mm., height, 1.45 mm.; a median, entrant suture less
than half height of scale; a pair of internasals separated by a pair
of median scales; nostril surrounded by rostral, first labial, three
postnasals and an internasal; scales on snout regular, with some
variation in size, all scales larger than elsewhere on head and dor-
sum (exclusive of tubercles); 37 scales across snout between the
fourth supralabials; approximately 43 scales between edges of upper
eyelids; scales on head minute, the small intermixed tubercles
reaching forward to anterior level of eye; scales of sides and dorsum
small, juxtaposed, intermixed with trihedral tubercles, forming
approximately 20 irregular, longitudinal rows, but diagonally the
rows are relatively regular; nine or ten rows between hind legs;
scales on tail arranged in whorls, seven or eight scale-rows basally
to a whorl, with a transverse series of eight large spiny scales (at
base) and reduced to six and later to four farther back; ventral
scales on the base of the tail divided, becoming single on the fifth
whorl, two scales to a whorl; the regenerated tail has very wide,
ventral scales, while the scales of side and dorsal part of tail are
nearly uniform and more or less keeled; a number of large tubercles
on dorsal surface and hind limb; supralabials, 11-10, infralabials,
9-9; mental with a wider labial border than rostral, its sides straight,
forming an acute angle behind; two pairs of chinshields, the inner,
larger pair with a suture much less than half that bordering the
mental, the scales touching two labial scales; outer pair smaller,
touching only one labial; chin scales small, regular, cycloid, a little
larger on sides; venter with larger cycloid, imbricate scales, in 38

1544 The University Science Bulletin

rows at widest part of belly; a rather large tubercular scale on each
side of base of tail; preano-femoral pores, 13-13, separated mesially
by three scales; ventrolateral folds scarcely discernible. All digits
with claws, and lamellae under basal part; terminal lamellae, fore
and aft, are single, others double, arranged slightly diagonally, seven
or eight on outer four toes, five on inner toe; six or seven on four
outer fingers, five on inner finger, the proximal lamellae on all
somewhat scalelike.

Color: Above fawn with a series of brown flecks or spots on head
and a curving mark from back of eye around to back of eye on
opposite side; a dark line directly back from eye to behind ear;
dorsum with three moderately distinct series of spots, one series
medial, the others dorsolateral; venter yellowish white; tail indis-
tinctly barred.

Remarks: A series of specimens of this species (E.H.T.-H.M.S.
Nos 30401-30417) was acquired at a point 12 to 14 miles north of
Trincomalee. Most of these were collected in human habitations.
One other specimen, No. 30486, E.H.T.-H.M.S. from "Ceylon," is in
the collection.

There is considerable variation in color and markings, dependent
on the amount of pigment. Specimens taken and preserved at
night are light in color.

Hemidactylus depressus Gray

Hemidactylus depressus Gray, Zoological Miscellany, 1842, p. 58 (type lo-
cality unknown: here restricted to Ceylon); Catalogue of the Specimens of
Lizards in the collection of the British Museum, 1845, p. 153; Giinther,
Zoology of the Erebus and Terror, vol. 2, 1874-75, p. 16, pi. 15, fig. 1
(figure of type) [not seen by me]; Boulenger, Catalogue of the Lizards in
the British Museum, 2nd ed. vol. 1, 1885, p. 134; Theobald, Descriptive
Catalogue of the Reptiles of British India, 1876, p. 76; Boulenger, The Fauna
of British India, including Ceylon and India; Reptilia and Batrachia, 1890,
p. 90-91; Haly, First Report on the Collection of Lizards in the Colombo
Museum, 1886, p. 4, (Colombo, Ceylon); Mehelv, Tennes Fiizet, vol. 20,
1897, p. 57; Deraniyagala, Ceylon Journ. Sci., B, vol. 16, 1932, p. 302, pi. 58.

Nubilia argentii Gray, Catalogue of the specimens of Lizards in the collection
of the British Museum, 1845, p. 273 ( type locality "Singapore", ex errore,
corrected to and restricted to Ceylon ) .

Hemidactylus pieresii Kelaart, Prod. Fauna Zeylanica, vol. 1, 1852 (1853),
p. 159 (type locality, Kandy, Ceylon ["The Kandian Gecko"]).

Diagnosis: Head moderately wide the maximum snout-vent
length reaching 80 mm., the tail 90; scales across belly, 35; 15 to 17
femoral pores on each side, almost continuous; back with fine gran-
ules and 16 (14-18) irregular rows of trihedral tubercles; scales of
tail in whorls, each whorl bearing 8 spinelike tubercles; a cream line
from eye to nostril.

Taylor: Lizards of Ceylon 1545

Description of species: (from E.H.T.-H.M.S. No. 30399). Head
one and one-half times as long as wide; rostral nearly twice as wide
as high, reaching distinctly higher than upper level of nostril; a pair
of internasals separated mesially by two small scales (tandem);
two, small, indistinct postnasals, one supralabial, the rostral and
internasal complete the border of nostril; 38 small granular scales
from eyelid edge across to eyelid edge at middle of eye; 48 scales
from fifth labial across snout to fifth labial; scales on snout larger
on interorbital area and occiput, but in latter area larger scattered
scales are present; ear large, diagonal, its greatest diameter a little
more than half eye diameter; ear a little closer to eye than nostril;
supralabials 13-13, infralabials 10-10; scales bordering labials
scarcely larger than other scales; width of mental about equal to
that of rostral, the sides of mental forming an angle; first pair of post-
mentals touch two labials, forming a common suture as long as their
sutures with the mental, the scales nearly pointed behind; outer pair
of postmentals touch one labial and are very narrowly separated
posteriorly, the third scale on each side being separated from the
third labial by another small scale; scales on chin and throat small,
nearly uniform; on belly, scales cycloid, imbricate, about thirty-six
rows between indistinct dorsolateral folds; femoro-preanal pores,
17-17, separated mesially by three scales; terminal (anterior and
posterior) lamellae single; other lamellae paired, arranged slightly
diagonally; outer fingers with eight or nine lamellae; inner finger
with six; four outer toes with nine or ten lamellae, the inner toes
with six lamellae; tail with numerous whorls, bearing at base eight
spiny tubercles and seven or eight rows of tiny scales; ventrally each
whorl with two enlarged scales.

Color: Above light gray or brownish gray; a silvery or cream line
from eye to nostril, dark bordered above and below; a black stripe
from eye to near point of arm insertion and a short upward projec-
tion from it behind ear; back gray-brown with four narrow trans-
verse markings.

Measurements in mm.: Snout-to-vent length, 69; tail length, 73
(tip regenerated); width of head, 15; length of head, 21; axilla to
groin, 36; arm, 19; leg, 26.

Remarks: A series of eleven specimens have been examined:
E.H.T.-H.M.S. Nos. 30393-30400 from 12 miles north of Trincomalee
Ceylon; Nos. 31255-31257, 31287 Tonacombe Estate, Uva Hills,
Namunukula Ceylon, W. W. A. Phillips, collector.

1546 The University Science Bulletin

Hemidactyhis triedrus (Daudin)

Gecko triedrus Daudin, Histoire Naturelle generale et particuliere des Reptiles,
vol. 4, year X (1802) pp. 155-157 (unknown locality; here fixed and re-
stricted to Trincomalee, Ceylon).

Hemidactyhis triedrus Kelaart, Prod. Faun. Zeylanica, 1852, p. 157; Giinther,
Reptiles of British India, p. 107; Boulenger, Catalogue of the Lizards of the
British Museum (Natural History, 2nd ed., vol. 1, 1885, p. 133; Fauna of
British India, including Ceylon and Burma; Reptilia and Batrachia, 1890,
pp. 89-90; Mehely, Termes Fiizet., vol. 20, 1897, p. 57. ( Kala-Wewa; tuber-
cular scales in 18 rows; femoral pores 7-7); Deraniyagala, Ceylon Joum.
Sci., B, vol. 16, 1932, p. 303, pi. 44; Smith, The Fauna of British India,
including Ceylon and Burma; Reptilia and Amphibia, vol. 2, Sauria, 1935,
pp. 88-89.

Hemidactyhis trihcdrus Kelaart, Prod. Faun. Zeylanicae, 1852 (1853), p. 157,
( rare in Ceylon; found 'in ant hills at Trincomalee; it lays 3 to 6 eggs. )

Diagnosis: Large, snout-to-vent length, 80 mm., tail 90 mm.;
trihedral tubercles arranged in 16 to 18 longitudinal rows; femoral
pores, 8-8, separated by a single scale, mesially.

Description of species: (from E.H.T.-H.M.S. No. 30418.) Head
moderate, its length, 1.6 times the width; height of rostral, reaching
upper level of nostril slightly more than half its transverse width;
internasals separated mesially; nostril surrounded by rostral, inter-
nasal, first labial and three postnasals; supralabials, 10-10; infra-
labials, 8-8; 37 scales across snout between the fourth supralabials;
scales between edges of eyelids, 47, extremely minute but with a
few larger tubercles scattered in this region and on occiput; scales
on dorsum and sides distinctly larger than those on occiput, but
smaller than those on snout; trihedral tubercles especially large,
measuring 1.5 mm. in length and width, and .5 mm. high, these
separated from each other by from one or two rows of small scales;
approximately 24 longitudinal rows of scales on venter, the ventro-
lateral folds being nearly obsolete; scales on chin subequal, very
small; mental large its labial border much larger than labial border
of rostral; first pair of postmentals touching one labial, forming a
median suture much shorter than their suture with mental; outer
( second ) pair of postmentals small, widely separated, touching two
labials; third outer postmentals touching second and third labials;
auricular opening distinctly more than half length of eyes; preanal
pores, 8-8, separated mesially by a single scale, and scarcely extend-
ing onto femora. All fingers with claws; second joint of fingers and
toes with lamella, the distal and sometimes proximal lamellae of
series single, others double, arranged diagonally; four outer fingers
with eight, inner finger with six lamellae; four outer toes with eight,
inner toe with seven lamellae; terminal joint arising back from edge
of lamella, the joint long; scales of tail in whorls, bearing either six
(on basal whorls) or four, large trihedral tubercles; about nine rows

Taylor: Lizards of Ceylon 1547

of small scales to each whorl; a pair of broad scales on ventral sur-
face of each whorl.

Color: Above rather light, brownish-fawn, with six or seven very
indistinct, transverse, darker spots, the posterior ones scarcely dis-
cernible; on edges of these spots several cream-colored trihedral
tubercles; behind eye a dark stripe, bordered by three or four cream
tubercles above, and below, behind this, a single row of cream tuber-
cles extending along neck and on side, with others on back; tail
nearly uniform brown ( banded in young ) .

Remarks: 1 have three specimens, E.H.T.-H.M.S. Nos. 30418-
30420 taken 12-15 miles north of Trincomalee, in forest. The largest
specimen measures 72 mm. snout to vent; tail, 85 mm. Two speci-
mens in the U.S.N.M., Nos. 120311 $ to 120313 <? are from Rattota,
Matale District.

The two latter specimens differ but little in the characters re-
corded for the preceding. The male pores are 7-8, separated
mesially by two scales.

Hemidactylus leschenaulti Dumeril and Bibron

Hemidactylus leschenaulti Dumeril and Bibron, Erpetologie generale vol. 3,
1836, p. 364 (type locality, Ceylon); Boulenger, Catalogue of the Lizards
in the British Museum, 2nd ed. 1884, pp. 136-137; Fauna British India in-
eluding Ceylon and Burma, Reptilia and Batrachia, 1890, pp. 91-92; Der-
aniyagala, Ceylon Journ. Sci., B, vol. 16, 1932, p. 301, pi. 43; Smith, the
Fauna of British India. . . . Reptilia and Amphibia, vol. 2, Sauna,
1935, pp. 97-98.

Hemidactylus coctaei Kelaart, Prod. Faun, Zeylanica, 1853, p. 160, (nee Du-
meril and Bibron).

Hemidactylus pustulosus Lichtenstein and von Martens, Nomenclator Rep-
tilium et Amphibiorum musei zoologici Berolinensis, 1856, p. 5. (type lo-
cality, Ceylon).

Hemidactylus kelaartii Theobald, Journ. Asiat. Soc. Bengal, extra number 88,
1868, p. 29 (type locality, Ceylon).

This large Indian gecko was found to be common in the lowland
forests eastward and northward of Trincomalee. One smooth-
barked tree species common in this vicinity, was almost certain to
have a family of these lizards, the mottled gray bark rendering
the lizard almost invisible. I have found no individuals of this
species in human habitations.

Diagnosis: Head and body relatively broad for the length; 50
granules across head between eyelids; whorls on tail with 12-14
rows of small scales and two pairs of small flat tubercles; granules
on back small with flat rounded inconspicuous tubercles; 17-18
femoral pores.

Description of species: (from E.H.T.-H.M.S. No. 30430). A
large-headed, large-bodied species, reaching 92 mm. snout-vent

1548 The University Science Bulletin

length; tail length, 86 mm. ( which is, I believe, the largest recorded
specimen); rostral much wider than high, reaching up to level of
middle of nostrils; a short groove enters rostral from above; inter-
nasals (separated by a minute scale, or often by two or three), a
pair of postnasals and one labial helping form nostril rim; upper
labials, 9-9, followed by three or four tiny scales; infralabials, 9-8;
rows of scales bordering the labials both upper and lower, some-
what larger than other head scales; granules between edges of
upper eyelids approximately 50; ear opening large, its diameter
about 2M times in diameter of eye; its distance from eye a little less
than distance from eye to nostril; head, dorsum and sides covered
with small, nearly uniform granules although those on front of
head larger, irregular; median scales on back somewhat smaller
than those on sides, the granules juxtaposed rather than imbricating;
a few (two or four), large, rounded, flat tubercles on back between
hind limbs; venter covered with cycloid, imbricating scales, approxi-
mately 50 between edges of the two slight ventrolateral folds;
mental large, its labial border distinctly larger than that of rostral;
a pair of median postmentals touching first labials and forming a
median suture a little more than half their length; second, outer
pair, smaller, touching first and second labials; femoral pores 17-18,
separated mesially by six scale rows; tail with whorls, each bearing
12-14 rows of fine, juxtaposed or partially imbricating granules, and
dorsally with two pairs of flat rounded tubercles, and dorsolateral^
with one flattened spinelike scale; ventral scales widened, two to
each whorl; posteriorly the flat rounded tubercles become more
pointed and scalelike (tail partly regenerated, this without whorls
or tubercles ) ; toes widened with a double series of slightly diagonal
lamellae; four outer fingers with eight or nine lamella, first toe with
six, the terminal joint arising back from anterior lamella; all fingers
with claws; four outer toes with eight or nine paired lamella (the
terminal one always single on all digits ) ; inner toe with six or seven
lamellae.

Color: Grayish or grayish black, with dark indistinct markings
forming wavy, crossbars, each lighter anteriorly, darker posteriorly
and mesially; tail dark with lighter bands of cream or yellow-white;
under side of tail, pigmented; a dark line from eye more or less
traceable to groin on the side.

Measurements in mm.: Length, snout to vent, 92 mm.; tail length,
86; width of head, 21; length of head, 26; axilla to groin, 39; arm,
29; leg, 36.

Taylor: Lizards of Ceylon 1549

Hemidactylus frenatus Schlegel

Hemidactylus frenatus Schlegel, in Dumeril and Bibron Erpetologie Generale
. . . vol. 3, 1836, p. 366 (type locality, "Java," here restricted to
Batavia, Java); Kelaart, Prod. Faun. Zeylanicae, 1853, p. 161 ("found in all
parts of the Island except in Newera Ellia"); Deraniyagala, Ceylon Journ.
Sci., B, vol. 16, 1932, p. 299, pi. 42; Smith, The Fauna of British India,
including Ceylon and Burma; Beptilia and Amphibia, vol. 2, Sauria, 1935,
pp. 95-96, fig. 29.

Diagnosis: A relatively small species, snout-to-vent length 62
mm.; tail, 70 mm.; preano-femoral pores in a continuous series of
29-33 on each side; two postmentals, the second outer pair nearly
as large as inner pair; body with few, scattered, rounded or feebly
keeled tubercles chiefly confined to posterior part of body. Tail
with whorls, each bearing a series of spinelike scales.

Description of species: Rostral nearly a half wider than high;
nostril surrounded by rostral, first labial, an internasal and two
postnasals; internasals separated mesially by a single scale; approxi-
mately 41 scales across head between edges of eyelids; occiput,
neck and body covered with minute granules and with one or two
scattered rows of rounded tubercles beginning on shoulders or
perhaps not extending farther forward than middle of body; some-
times tubercles trihedral.

Tail with 10-12 rows of small scales arranged in whorls, each
bearing a series of six larger spinelike scales, the outermost ventro-
lateral in position, and on ventral surface bearing two enlarged
scales, the anterior the larger on each whorl; supralabials, 12-12,
the last four very small; infralabials, 9-9; chin scales small, regular,
a little larger on the sides; ventral scales cycloid, about 38 to 40
between the ventrolateral folds; however those along sides of venter
nearly a half smaller than those in middle; digits with claws; inner
digits very small, less than half length of the adjoining digit;
lamella, eight to ten on four outer toes, six to eight on four outer
fingers, five on inner finger and toe.

Color: Grayish or fawn to blackish brown, uniform or with some
indistinct marking; usually a dark, light-bordered line on head;
yellow or yellow-white below.

Remarks: Some specimens may have a pinkish tinge above. The
color is changeable and while at night they may be nearly fawn,
in daylight they may become blackish.

The species is widespread in South Asia and Indonesia about
human habitations. I have not found individuals in forest, away
from the proximity of such habitations.

The following Ceylonese specimens are at hand: E.H.T.-H.M.S.,
Nos. 30433-30463, from 12-14 mi. N of Trincomalee, Ceylon; K.U.

1550 The University Science Bulletin

M.N.H., Nos. 31259-31264, Tonacombe Estates, Namunukula, Cey-
lon. The pore-counts vary from 29 to 33, usually in a continuous
series ( one exception has the series separated by one scale ) .

Hemidactylus maculatus hunae Deraniyagala

Hemidactylus maculatus hunae Deraniyagala, Ceylon Jonrn. Sei. B. vol. 20,
pt. 2, Apr. 15, 1937, pp. 185-189, fig. 1 (type locality, Okanda, Eastern
Province, Ceylon).

Diagnosis: A large species 105 mm. snout to vent; nineteen to
twenty-five femoral pores on each side, separated mesially.

Description: Snout obtuse, as long as or longer than the distance
from eye to ear; supralabials ten to twelve, the infralabials nine or
ten; mental subtriangular, wider than the rostral; and usually as
long as wide; postmentals two pairs, the inner enlarged; gular scales
fine, granular; rostral subquadrangular, broader than high; nostril
between rostral, first labial and several small scales; a pair of inter-
nasals; snout with convex scales, the occiput with granular scales
and larger conical tubercles; back covered with small scales and
about fifteen to twenty longitudinal rows of tubercles, which are
smallest along the middorsal line and feebly trihedral; ventrals
smooth, imbricate, rounded scales which grade imperceptibly into
the laterals posterodorsally, immediately anterior to the hind legs;
digits free, moderately dilated with almost straight, transverse
lamellae, nine or ten under inner toe, eleven to thirteen under
fourth toe; tail feebly depressed, with about sixty wide subcaudals
and dorsally with six or four longitudinal series of tubercles which
are usually low and subcorneal, rarely trihedral; males with nine-
teen to twenty-five femoral pores on each side, separated by from
two to sixteen scales.

Color: Dorsally silvery gray with five broad transverse olive-
brown bands from neck to rump; each band possesses a black mar-
gin and forms a rhomboid vertebral enlargement with a light center
containing a black ring. Tail with about six dark rings a little
wider than the interspaces; limbs with five or six irregular rings on
each; two lateral bands on each side of head; ventrally white, dusted
with brown especially on tail; young with dorsal crossbands without
any light areas.

The type is from Okanda, Eastern Province, Ceylon. It was
taken in a rock cave.

Measurements: Snout to ear, 28 mm.; snout to vent, 105 mm.;
tail length, 115 mm.

Data from the type description.

Taylor: Lizards of Ceylon 1551

Malcolm Smith has not included this form in his "Fauna," nor
docs he list H. maculatus from Ceylon.

Genus Cosymbotus Oken

Cosymbotus Fitzinger, Syst. Rept. 1843, pp. 19, 104 (type of genus Stellio
platyurus Schneider); Myers, Copeia, 1943, No. 3, Oct. 15, p. 192.

Two species are recognized. The species, which occurs in Ceylon,
is widely distributed in India, Indo-China, and the East Indian
Archipelago.

Cosymbotus platyurus (Schneider)

Stellio platyurus Schneider, Amphib. Physiol., vol. 2, 1792, p. 30 (type locality
not given [fide M. Smith. I have not seen this paper] ).

Hemidactylus platyurus Boulenger, Catalogue of the Lizards in the British Mu-
seum, vol. 1, 1885, p. 143; The Fauna of British India, including Ceylon
and Burma; Reptilia and Batrachia, 1890, p. 95.

Cosymbotus platyurus Deraniyagala, Ceylon Journ. Sci., B, vol. 15, 1932, p. 306.

Platyurus platyurus Smith, The Fauna of British India including Ceylon and
Burma; Reptilia and Amphibia, vol. 2, Sauna, 1935, pp. 102-103.

Diagnosis: A medium sized gecko, the snout-vent length reaching
60 mm.; tail 65; digits webbed, broadened, with the lamellae divided
below by a deep groove; all digits clawed; a narrow lateral web
or fringe on each side of body; males with continuous preanal and
femoral pores, 13-20 on each side; nostril between rostral, first labial,
an internasal and two postnasals; back with uniform small granules;
venter with smooth, cycloid scales; gray or gray-brown above,
marbled or with spots, sometimes with large distinct dorsal blotches;
a bar from eye onto side.

I have seen no Ceylonese specimens of this species.

Genus Peropus Wiegmann

Peropus Wiegmann, Nova Acta Acad. Leop. Carol., vol. 17, 1835, p. 238 (type
of genus [Peropus mutilatus]).

A single species of the genus occurs in Ceylon.
Peropus mutilatus Wiegmann

Hemidactylus (Peropus) mutilatus Wiegmann, Nova Acta Acad. Leop. Carol.,

vol. 17', 1835, p. 238 (type locality, Manila, P. I.).
Gehyra mutilata Boulenger, Catalogue of the Lizards of the British Museum,

vol. 1, 1885, p. 48; The Fauna of British India, including Ceylon and Burma;

Reptilia and Batrachia, 1890, p. 96; Smith, The Fauna of British India,

including Ceylon and Burma; Reptilia and Amphibia, vol. 2, Sauria, 1935,

pp. 105-106, fig. 30.

Diagnosis: A specimen (No. 41 $ ) from Tonacombe Estate,
Namunukula, Ceylon, is used as the basis of the diagnosis and de-
scription.

Three pairs of chinshields, the outer small, extending but slightly
farther back than inner; digits webbed at base; seven to nine strongly

68—6490

1552 The University Science Bulletin

oblique lamellae under fourth toe; labial border of mental equals
that of rostral; head with small granules; on snout granules a little
larger; dorsum with very small granular scales; scales cycloid,
imbricate on venter; skin-fold on flank and on posterior margin of
leg; male with 25-41 preano-femoral scales, meeting mesially.

Description: (from No. 41 Tonacombe Estate, Namunukula,
Ceylon, a small species, snout-to-vent length, 51 mm., tail 50 mm.)
Rostral not twice as wide as high; large internasals separated
mesially; nostril surrounded by rostral, first labial, internasal and
two postnasals; 47 scales across snout between fourth supralabials;
42 granules between edges of eyelids; scales on back distinct,
rounded, imbricate or subimbricate, larger on sides than on middle
of dorsum, and merging into scales of venter; no distinct ventro-
lateral fold; approximately 48 scales across venter, cycloid, imbri-
cating; supralabials, 10-10, infralabials, 9-10, the last of both series
small; labial border of mental smaller than that of rostral; two very
large postmentals touching each other, their common suture twice
that with the mental; each postmental touching a single labial; outer
pair of postmentals slender, not extending behind first pair, touch-
ing two labials; a few enlarged scales follow second pair; ear nearly
quadrangular, less than half diameter of eye. Digits with a narrow
angular joint and a basal part with well-defined, paired, diagonally
placed lamellae distally, the proximal lamellae being almost flat,
scalelike, 10-13 on four outer fingers and toes, nine or ten on inner;
claws present on all digits; skinfold or web on back of leg ample;
tail flattened below, with fine scales arranged in indefinite whorls,
about ten rows to each whorl.

Color: Above lavender-gray with a more or less definite series
of rounded light spots on each side near middle of back and some
scattered light spots on sides; scattered spots on occiput; yellowish
white below.

Remarks: I have examined one other specimen from Ceylon,
U.S.N.M. No. 59014 § without exact locality. This specimen has
a well-developed third pair of postmentals separated from labials
by small granular scales.

HemiplujUodactyhis Bleeker

Hemiphyllodactylus Bleeker, Nat. Tii'dschr. Ned. Ind., vol. 20, 1860, p. 327
(type of the genus, Hemiphyllodactylus typus.) A single representative
of this genus is known in Ceylon.

Hemiphyllodactylus typus typus Bleeker

Hemiphyllodactylus typus Bleeker, Nat. Tijdschr. Ned. Ind., vol. 20, 1860,
p. 327 (type locality, Gunong Paring, Java); Deraniyagala, Ceylon Journ.

Taylor: Lizards of Ceylon 1553

Sci., B, vol. 16, 1932, p. 308; Smith, The Fauna of British India, including
Ceylon and Burma; Beptilia and Amphibia, vol. 2, Sauria, 1935, pp. 107-108,
(part).
Lepidodactylus ceylonensis Boulenger, Cat. Lizards of the British Museum, vol.
1, 1885, p. 164, pi. 13, fig. 3, Fauna of British India, including Burma and
Cevlon, 1890, p. 98; Annandale, Journ. Asiat. Soc. Bengal, N. S., vol. 1,
1950, and Spolia Zeylanica, vol. 8, 1915, p. 239 (Ganipola near Kandy,
1,600-1,700 ft. Peradeniya.)

Diagnosis: A small, slender lizard reaching a snout-vent length
of 60 mm.; tail 60 mm.; digits free, subcylindrical at base; four toes
with lamellae of which the terminal portion is actually or partly
divided, the terminal joint rising angularly from within the widened
basal part; inner finger and toe without free distal phalanx but
sometimes bearing a claw; an angular series of 10-11 preanal pores,
separated from a series of 8-10 femoral pores; supralabials, 10-12;
infralabials in like number; back and top of head covered with uni-
form small granules, without tubercles; no postmentals, but sev-
eral polygonal scales present behind the mental.

I have no specimens at hand from Ceylon. I am not wholly
certain that the Ceylon specimens are referable to the typical form.

Genus Lepidodactylus Fitzinger

Lepidodactylus Fitzinger, Systema Reptilium, 1843, pp. 19, 98 (type of genus,

Platydactyhis lugubris ) .
Amydosaurus Gray, Catalogue of the Specimens of Lizards in the collection of

the British Museum, 1845, p. 162 (type, Platydactyhis lugubris).

Whether the form of this genus occurring on Ceylon is subspecifi-
cally referable to the widespread oceanic form of lugubris may be
doubted. I have not seen a specimen.

Lepidodactylus lugubris Dumeril and Bibron

Platydactyhis lugubris Dumeril and Bibron, Erpetologie Generale, vol. 3, 1836,
p. 304 (type locality, Tahiti); Deraniyagala, Ceylon Journ. Sci., B, vol. 15,
1929, p. 157, pi. 33; ibid vol. 16, 1932, p. 307, fig; Smith, The Fauna of
British India, including Ceylon and Burma; Reptilia and Amphibia, vol. 2,
Sauria, 1935, pp. 115-116, fig. 33 (part).

Not having examined the Ceylonese form of this genus, I follow
Deraniyagala in placing it in the species lugubris and take data
from Smith for the diagnosis.

Diagnosis: A diminutive species (snout to vent 42 mm.; tail 40
mm.), with digits webbed at base, strongly dilated, bearing trans-
verse (or distally) strongly oblique lamellae; supralabials 11-13;
mental smaller than adjacent labials; no distinct postmentals, the
space occupied by some rows of polygonal scales; snout granules
somewhat larger than those on head and dorsum; venter with flat,
cycloid imbricate scales; 12-14 lamellae under longest toe; hind leg

1554 The University Science Bulletin

reaches two thirds of the distance to axilla; tail swollen at base, with
a sharp denticulated edge, covered above with small subimbricate
scales and below with larger imbricate scales; male with a con-
tinuous series of 25-30 preano-femoral pores which form a slight
angle mesially.

Light, pinkish gray or brownish above generally with a vertebral
series of paired spots, or pigment may be arranged as a sinuous
marking; a dark streak from tip of snout passing through eye to ear;
young may have one or more dark stripes; lower surfaces light or
speckled with brown.

Genus Lophopholis Smith and Deraniyagala

Lophopholis Smith and Deraniyagala, Ceylon Journ. Sci., B, vol. 18, 1934,
p. 235 ( type of genus, Teratolepis scabriceps Annandale ) .

This monotypic genus is known only from southern India and
Ceylon.

Lophopholis scabriceps (Annandale)

Teratolepis scabriceps Annandale, Mem. Asiat. Soc. Bengal, vol. 1, 1906, p. 187,
pi. 9, fig. 1, a-c (type locality, Bamnad, Madura District, south India).

Lophopholis scabriceps Smith and Deraniyagala, Ceylon Journ. Sci., B, vol. 18,
1934, pp. 235-236 ( Mariccukatti, Northern Province, Ceylon); Smith, The
fauna of British India, including Cevlon and Burma; Beptilia and Amphibia,
vol. 2, Sauria, Feb. 7, 1935, pp. 124-125, fig. 37.

Diagnosis: A small gecko (snout to vent 45 mm., tail 50 mm.), the
dorsum covered with imbricate scales a little larger than ventral
scales; caudal scales uniform with body scales; three preanal pores
on each side, separated mesially by a scale; ear subcircular, its
diameter about one third of the eye diameter; supralabials, seven
or eight, infralabials, six or seven; two pairs of postmentals, the inner
pair largest; head with small granules; body scales slightly elongated,
striated, feebly keeled; digits short; hind limb reaches about half
way to axilla; tail round, tapering to a point, covered with uniform,
imbricate scales.

Gray-brown with brown marking above arranged as transverse
bars; below dirty whitish; known in southern India and Ceylon (one
locality ) . ( Data from Smith, 1935. I have seen no specimen. )

Family Agamidae

Six genera occur of which Lyrioccphalus and Ceratophora are
endemic. Cophotis is excluded from India but a species occurs in
Java and Sumatra. Sitana, Otocryptis, and Calotes occur both in
India and Ceylon. Twelve genera occurring in South Asia are un-
known in Ceylon.

Taylor: Lizards of Ceylon 1555

Key to the Ceylonese Genera of Agamidae

1. Four digits on feet, five on hands Sitana

Five digits on hand and feet 2

2. Fifth toe short, not longer than first Otocryptis

Fifth toe longer than first toe 3

3. A rostral appendage 4

No rostral appendage 5

4. A strong globular protuberance on tip of snout Lyriocephalus

A narrow pointed rostral protuberance, scaled or naked ( reduced

in female ) Ceratophora

5. Dorsal scales enlarged, irregular, unequal; tail prehensile .... Cophotis
Dorsal scales more or less equal-sized, regularly arranged; tail

greatly elongated, not prehensile Calotes

Genus Sitana Cuvier

Sitana Cuvier, Regne Animal, 2nd ed., vol. 2, 1829, p. 43 (type of genus,
Sitana ponticeriana ) .

Only a single species of this genus is recognized.

Sitana ponticeriana Cuvier

Sitana ponticeriana Cuvier, Regne Animal, 2nd ed., vol. 2, 1829, p. 43 (type
locality Pondicherry, India); Kelaart, Prodr. Faun. Zeylanicae, 1852-1853,
p. 164; Roulenger, Catalogue of the Lizards in the British Museum, 2nd
ed., vol. 1, 1885, pp. 270-271; The Fauna of British India, including Ceylon
and Burma; Reptilia and Batrachia, 1890, pp. 114-115; Deraniyagala,
Ceylon Journ. Sci. B, vol. 16, 1931, pp. 141-142; Smith, The Fauna of
British India including Ceylon and Burma; Reptilia and Amphibia, vol. 2,
Sauria, 1935, pp. 144-146.

Sitana minor Gunther, Reptiles of British India, 1864, pp. 135-136, pi. 14,
fig. A, (type locality, Madras, India) ("it is more probable that the Cey-
lonese Sitana is identical with the Madras species" [i. e. S. minor]).

Diagnosis: A small terrestrial agamid, with five fingers and four
toes; body compressed, covered with regular keeled scales; no
crest on back or neck; a strong gular appendage on throat, extend-
ing to abdomen; no femoral or preanal pores; tympanum present,
visible.

Description of species: (E.H.T.-H.M.S. No. 30645 12 mi. N Trin-
comalee. ) Rostral small, bordered laterally by a labial, and behind
by five small scales; nasal large, elevated where the nostril is pierced,
separated from rostral by a single scale; an elongate supranasal
contiguous or imbricating with canthal scales; two median scales
on anterior frontal region elevated and with a more or less con-
tinuous keel; on each side two very slight ridges may be traced
back between the orbits; supraocular scales enlarged, irregular as
are the occipital scales; a very fine nuchal ridge or crest; a few
enlarged scales on the middle edge of upper eyelid; supralabials,
8-9; infralabials, 8-8; mental much narrower than the rostral; an

1556 The University Science Bulletin

enlarged gular appendage extending onto anterior part of abdo-
men; no femoral or preanal pores; leg long, the fourth toe reaching
beyond snout; fifth toe lost; five fingers; scales on dorsum enlarged,
in ten or eleven rows; adjoining scales on sides much smaller, but
with a few scattered enlarged scales; scales on gular appendage
enlarged, distinctly larger than ventral scales; approximately 20
rows of large ventral scales; tail rounded, the scales keeled but not
arranged in distinct whorls.

Color: Above brownish, with a very indistinct lighter border on
the outer row of enlarged dorsal scales; six pairs of black spots on
each side, divided by the median line; sides darker with fine, light,
ill-defined dots; a metallic cream spot on tympanum; venter lighter,
with a black line from tip of chin back and onto the front of the
gular appendage; tail very indefinitely barred.

Measurements in mm.: Length snout to vent, 51; tail, 114; head
length, 15; head width, 10; axilla to groin, 25; arm, 25; leg, 50.

Remarks: The following specimens were examined: E.H.T.-
H.M.S. 30628-30640, 30642, 30645, 30647-30655; Nos. 30660-30670,
30671, 30672 "Ceylon"; U.S.N.M. 58508 "Ceylon."

Genus Otocryptis Wagler

Otocryptis Wagler, Natiirliches System der Amphibien, 1830, p. 150, (type of
genus, Otocryptis wiegmanni); Wiegmann, Isis, 1831, p. 291 (type of genus,
Otocryptis bivitatta ) .

This genus may be readily recognized by the greatly reduced
outer toe which is smaller than the first toe. The species are terres-
trial. Two species are known, one of which occurs in Ceylon, the
other in southern India.

Hallowell (Proc. Acad. Nat. Sci. Philadelphia, 1860, p. 491), and
Stejneger (U. S. Nat. Mus. Bull. 58, 1907) consider the possibility of
a member of this genus occurring in the Riukiu Islands.

Otocryptis wiegmanni Wagler

Otocryptis wiegmanni Wagler, Natiirliches System der Amphibia 1830, p. 150
(type locality, America, [ex errore]. Here corrected to Ceylon.

Otocryptis bivittata Wiegmann, Isis, 1831, p. 291. (type locality unknown);
Peters, Monatsb. Akad. Berlin, 1860, 184 ( Trincomalee, Hinida, Ratnapura,
Adam's Peak); Giinther, Reptiles of British India 1864, pp. 127-128; Bou-
lenger, Catalogue of the Lizards in the British Museum, 2nd ed., vol. 1,
1885, p. 271; Haly, First Report on the collection of Lizards in the Colombo
Museum, 1886, p. 6 ( Ramboda and the Western Province); Theobald,
Descriptive catalogue of the Reptiles of British India, 1876, p. 98. Bou-
lenger, The Fauna of British India, including Ceylon and Burma; Reptilia
and Batraehia, 1890, pp. 115-116; Green, Journ. Bombay Nat. Hist. Soc,
vol. 15, 1903, p. 817. Deraniyagala, Ceylon lourn. Sci., B, vol. 16, 1931,
pp. 142-143; Smith, The Fauna of British India, including Ceylon and Burma;
Reptilia and Amphibia, vol. 2, Sauria, 1935, pp. 146-147. ("common at
Peradeniya.")

Taylor: Lizards of Ceylon 1557

Diagnosis: A very small outer toe; male with a very extensive
gular flap extending to center of abdomen. Hind leg reaches far
beyond tip of snout.

Description of species: (from E.H.T.-H.M.S. No. 30613).
Length, snout to vent, 64 mm.; the length of head (19 mm.) much
less than twice the width (12 mm.); rostral rather large, bordered
by two labials and five scales, separated from nasal by a single pre-
nasal; nasals large forming a mound with crater, the rim the highest
part; a large supranasal, nearly as large as nasal; four canthals con-
tinuous with the elongate, broadly imbricating superciliary series;
scales on snout rough, unequal, keeled, two median ones forming a
short ridge beginning at posterior level of nasals; from here two
smaller series of scales diverge that are continuous with the enlarged
keeled scales bordering inner edge of orbits, and which continue
around behind eye, the two series separated by from three to five
scales; supralabials, 12-12; infralabials, 10-10; mental narrow, longer
than wide, much smaller than rostral; scales around middle of body,
approximately 78; all dorsal and lateral scales keeled and pointing
backward and upwards; the ventral scales strongly keeled, some-
what mucronate; limbs long, the terminal joints of fingers reach be-
yond snout; leg brought forward, the entire free part of fourth toe
extends beyond snout; 28 keeled scales under free part of fourth
toe; tail cylindrical not compressed, longer than head and body. A
broad "dewlap" in males beginning on chin and extending more than
half length of abdomen, covered with larger keeled scales (absent
in females and very young).

Color: Body dull brown to blackish brown; head blackish brown,
the dewlap black on the anterior part; limbs light brown; tail in-
distinctly banded with broader dark bands, and narrow, dull,
light bands; sides lighter, with some dull whitish marks on each
side scarcely discernible unless specimen is submerged.

Measurements in mm.: Snout to vent, 64; tail (broken); width
of head, 12; length of head, 19; axilla to groin, 27; arm, 30; leg, 75.5.
Length of dewlap, 40; depth of dewlap, 16.

Remarks: The series of 18 specimens available in the E.H.T.-
H.M.S. collection, Nos. 30610-30627, are all from 12 miles north of
Trincomalee, Ceylon. Here they were found most frequently in the
forest bordering on the shore. They are terrestrial, the reduced size
of the outer toe presumably being a handicap in climbing.

Females are lighter brown than the males and occasionally speci-
mens show a series of four deep-black, dorsal spots somewhat
chevron-shaped; the legs and feet may be spotted black; in these
the head is darker than the body; usually whitish below.

1558 The University Science Bulletin

A young specimen (E.H.T.-H.M.S. 30622) has a distinct dark bar
across the frontal region of head with some spots on neck.

Otocryptis sp.?

A specimen, U.S.N.M., No. 120328 from Nandana Estate, Pera-
deniya, Kandy District, differs considerably from the species de-
scribed under the name wiegmanni. The differences, as seen by
comparing this highland specimen with the lowland series, would
seem to warrant a specific separation. The more striking differences
are as follows: scales on nuchal region distinctly larger, as are the
scales along the dorsum; a distinct dorsal ridge along the middle of
back with crest; scales on lower half of sides directed backward and
downward; gular dewlap smaller ( length, 24 mm., depth, 12 mm. ) ;
head and gular fold not blackish.

I strongly suspect that two species are involved but until the
characteristics of the type are ascertained one is uncertain to which
of the forms the name wiegmanni must apply. It is presumed that
wiegmanni and bivitatta have the same type.

Genus Cophotis Peters

Cophotis Peters, Monatsb. Akad. Wiss. Berlin, 1861, p. 1103, (type of genus,
Cophotis ceylonica) .

Cophotis is a genus containing only two species, one of which
occurs in Ceylon and this chiefly in the higher regions. The other
occurs in Java and Sumatra, without any known form occupying the
Indian and southeastern Asiatic regions.

The Ceylon species is ovoviviparous. It may be recognized from
other species by the large, scattered, flattened, dorsal scales on a
strongly compressed body, by absence of femoral and preanal pores,
in males, the absence of a tympanum, the presence of a short nuchal
crest, a dorsal crest, and the presence of a small gular pouch.

Cophotis ceylanica Peters

Cophotis ceylanica Peters, Monatsb. Akad. Wiss. Berlin, 1861, p. 1103 (type
locality, Ceylon); Giinther, Beptiles of British India, 1864, p. 132, pi. 13,
fig. H; Theobald, Descriptive Catalogue of the Reptiles of British India,
1876, pp. 100-101; Boulenger, Catalogue of the Lizards in the British Mu-
seum, vol. 1, 1885, p. 275; Haly, First Report on the collection of Lizards
in the Colombo Museum, 1886, p. 6 (Ramboda; Le Vallon Estate); Bou-
lenger, The Fauna of British India including Ceylon and Burma; Reptilia and
Batrachia, 1890, p. 118; Willey, Spolia Zeylanicae, vol. 3, pt. 12, Apr. 1906,
pp. 235-237, pi. and fig. in text; Deraniyagala, Ceylon Journ. Sci., B, vol.
16, 1931, pp. 148-149, pi. 34; Hora, Records of the Indian Museum, vol. 28,
pt. 4, 1926, pp. 216-217 (Nuwara Eliya); Smith, The Fauna of British India,
including Ceylon and Burma; Reptilia and Amphibia, vol. 2, Sauria, pp. 150-
151, fig. 46.

Diagnosis: A small lizard ( snout-to- vent length approximately

Taylor: Lizards of Ceylon 1559

60 mm.; the tail, 85 mm.); body compressed; dorsal scales relatively
large; about 33 scales around body; head slender; a discontinuous
nuchal and dorsal crest; a small gular sac; five fingers and five toes;
tail prehensile, curving down; tympanum hidden; no preanal or
femoral pores.

Description of species: (from E.H.T-H.M.S. No. 30500, "Cey-
lon"). Length of head twice as long as width; snout longer than
eye; head narrowed, rather pointed, the three postrostrals and three
adjoining scales form an indistinct "boss or nose"; nostril in a large
scale, separated from the rostral by three scales, touching the sec-
ond and third labials; scales in frontal region large, irregularly
elevated, with a depression between the orbits; a somewhat ele-
vated median scale in depression; five or six enlarged supraoculars,
all very irregular; a pair of enlarged tubercular scales on parietal
region, behind which are two distinctly elevated tubercular scales;
supralabials, 10; infralabials, 8 or 9; mental smaller than rostral; a
series of irregular keeled scales extend back from lower edge of
eye in temporal region; scales under chin, smooth; a small gular sac
present; approximately eight lateral rows of enlarged scales point-
ing backward and downward, strongly imbricating, smooth or
slightly keeled; lower part of sides and venter with strongly mucro-
nate, smaller, keeled scales in about 16 rows; a nuchal crest con-
sisting of four or five spines, separated from the dorsal crest, which
consists of soft, spiny scales about fifteen in number, separated by
short intervals; tail compressed, prehensile; hind leg reaching axilla;
arm reaching beyond tip of snout; third and fourth toes practically
of equal length.

Color: Olive green above with some lighter markings and some
scattered darker markings; a light brown mark along upper lip,
covering labials and adjoining scale row and extending onto
shoulder; a lighter spot on back of occiput; lower labials dark
brown, the throat and chin cream with scattered dark flecks, or
lines; venter dirty gray-white; tail banded darker and lighter gray;
some trace of darker bands low on sides.

Measurements in mm.: Snout-to-vent length, 61; tail, 79; head
length, 18; head width, 9; arm, 29; leg, 30; axilla to groin, 30.

Remarks: The species is ovoviviparous. It is a highland form,
known up to elevations of 7000 feet.

Ceratophora Gray

Ceratophora Gray, Illustrations of Indian Zoology, vol. 2, 1835, pi. 68, fig. 2,

(type of genus: Ceratophora stoddartii).
Lyriocephalus (part) Theobald, Descriptive Catalogue of the Reptiles of British

India, 1876, p. 99.

1560 The University Science Bulletin

This genus, endemic in Ceylon, is represented by three species, all
seemingly clearly differentiated, occupying territory at an elevation
in excess of 3500 ft. All have a characteristic soft appendage pro-
truding from the tip of the snout.

Key to the Species of Ceratophora

1. Gular scales smaller than ventral; rostral appendage scaly,

pointed aspera

Gular scales larger than ventrals 2

2. Gular scales smooth or feebly keeled; lateral scales large very

unequal; rostral appendage smooth pointed stoddartii

Gular scales strongly keeled; lateral scales large, nearly equal;
rostral appendage, compressed, suboval, covered with scales or
granules tennentii

Ceratophorus tennentii Giinther

Ceratophoras tennentii Giinther, in Tennent, Natural History of Ceylon, 1861,
p. 281, fig. (type locality, Ceylon); Reptiles of British India, 1864, pp. ISO-
LSI (more complete description); Boulenger, Catalogue of the Lizards of
the British Museum, 2nd ed., vol. 1, 1885, p. 278; Haly, First Report of
the Collection of Lizards in the Colombo Museum, 1886, p. 6; Boulenger,
Fauna British India, including Ceylon and Burma; Reptilia and Batrachia,
1890, p. 120; Deraniyagala, Ceylon Journal Sci., B, vol. 16, 1931, p. 145-146.

Liiriocephalus tennentii Theobald, Descriptive Catalogue of the Reptiles of
British India, 1876, pp. 99-100.

Diagnosis: Head scales small, irregular; a ridge on each side of
occiput; rostral appendage large in both sexes, fleshy, compressed,
suboval, covered with scales and granules; supralabials, 9; infra-
labials, 9 or 10; gular scales feebly keeled, large, quadrangular,
forming regular longitudinal series; a low toothed nuchal crest;
scales on dorsum irregular and unequal in size, the larger ones
feebly keeled; lateral scales equal, large, strongly imbricating, point-
ing upward and backward; smooth or feebly keeled; ventral scales
smaller; leg reaches to eye or a little beyond; tail slightly com-
pressed, scales keeled.

Olive above, irregularly marbled with brownish, young with an
angular crossband between eyes; sometimes with white longi-
tudinal lines; a more or less distinct, white line along hinder side
of thighs. Length, 260 mm.; tail, 170; arm, 48; leg, 76. (After
Boulenger, 1884. )

Tennent, in writing of the form, anticipates Giinther by publishing the name and brief
description as follows: "Among the specimens sent from Ceylon by Dr. Kelaart, and now in
the British Museum, there is one which so remarkably differs from C. Stoddartii, that it
attracted my attention by the peculiar form of this rostral appendage. Dr. Giinther has
pronounced it to be a new species; and Dr. Gray concurring in this opinion, they have done
me the honour to call it Ceratophora Tennentii". From this statement one might suppose
that the name should be attributed to Giinther and Gray. However Giinther refers to the
Tennent reference as "Giinther, in Tennent" so it would appear that Gray had no part in
the naming.

Taylor: Lizards of Ceylon 1561

Ceratophora aspera Giinther

Ceratophora asjjera Giinther, Reptiles of British India, 1864, p. 131, pi. 13,
figs. G. and G' ( type locality, Ceylon ) ; Boulenger, Catalogue of the Lizards
in the British Museum, 2nd ed., vol. 1, p. 278 (Ceylon, "south Ceylon");
The Fauna of British India, including Ceylon and Burma; Reptilia and
Batrachia, 1890, p. 120; Deraniyagala, Ceylon Journ. Sci., B, vol. 16, 1931,
pp. 146-147; Smith, The Fauna of British India, including Ceylon and Burma;
Reptilia and Amphibia, vol. 2, Sauria, 1930, p. 154.

Li/riocephalus asper Theobald, Descriptive Catalogue of the Reptiles of British
India, 1876, p. 100.

Diagnosis: A diminutive form, total length, 82 mm.; snout-to-
vent length, 37 mm.; head scales small irregular, each of which
forms a small tubercle; a larger tubercle behind superciliary edge;
back of head with a pair of low ridges convergent anteriorly; rostral
projection slender, covered with small imbricating scales, all keeled,
the projection equaling head length in males, rudimentary in fe-
males; throat with small strongly keeled scales; scales on the dor-
sum and sides small with larger, irregular, keeled scales intercalated;
no dorsal or nuchal crests, but some of the larger scales form series
across the vertebral line, with their angles pointing posteriorly;
ventral scales keeled; preanal region with small scales; tail not com-
pressed, all scales keeled; hind leg reaches to, or near to orbit;
brownish, marbled with darker; a rhombic light spot on rump;
brown spots on arm, edged with white in male.

Ceratophora stoddartii Gray

Ceratophora stoddartii Gray, Illustrations of Indian Zoology, vol. 2, 1834,
pi. 168, fig. 2 (type locality, Ceylon, Stoddart, coll.); Catalogue of the speci-
mens of lizards in the collection of the British Museum, 1845, p. 237; Kelaart,
Prod. Faunae Zeylanicae, 1852 (1853), p. 165; Giinther, the Reptiles of
British India, 1864, p. 129, pi. 13, fig. F, F', F"; Boulenger, Catalogue of
the Lizards in the British Museum, 2nd ed., vol. 1, 1885, p. 277; Haly, First
Report on the Collection of Lizards in the Colombo Museum, 1886, p. 6,
( "numerous specimens from the hill districts". ) ; Boulenger, The Fauna of
British India, including Ceylon and Burma; Reptilia and Batrachia, 1890,
p. 119; Boettger, Ber. Offenb. Ver. Nat., vol. 29-32, 1892, p. 69; Willey,
Spolia Zeylanica, vol. 3, 1906, p. 236; Deraniyagala, Ceylon Journ. Sci.,
vol. 16, B, 1931, p. 144; Smith, The Fauna of British India, including Ceylon
and Burma; Reptilia and Amphibia, vol. 2, Sauria, 1935, pp. 152-153.

Lyriocephalus stoddartii Theobald, Descriptive Catalogue of the Reptiles of
British India, 1876, p. 99.

There are six specimens in the collections available here and all
seem to belong to Ceratophora stoddartii. However, there is con-
siderable variation in this group of specimens as to character of the
enlarged scales, the rostral appendage, keeling of dorsal and ventral
scales, length of hind limb and coloration. Unfortunately the exact
provenance of these specimens is unknown. None of these variations
can be attributed to sex, since all are males. I suspect that there
may be distinct geographical races represented among them.

1562 The University Science Bulletin

Diagnosis: Five fingers and toes; body compressed, the scales
on the sides irregular in size; in the male, a smooth, compressed or
pryamidal appendage on tip of snout without any scales; tympanum
hidden; no preanal or femoral pores; gular pouch absent.

Description of species: (from E.H.T.-H.M.S. 30495 "Ceylon").
Rostral appendage conical, sharply pointed, its base surrounded by
a median pair of supralabials (paired rostral?) and five other small
scales; nasal scale somewhat craterlike, separated from appendage
by three scales, separated from its fellow by six scales; four can thai
scales, the canthus rather rounded; a pair of low pyramidal scales
in frontal region; supraoculars irregular, the inner row largest sepa-
rated from its fellow by three small scale-rows; last four scales of
inner row pass behind eye, covering an angular elevation; a pair of
knobs or bosses in parietal region, covered by a few large scales
and separated from each other by approximately seven scales; a
transverse row of slightly elevated scales across head in front of
these bosses; two large, prominent, elevated scales in temporal
region above tympanum; behind eye, a row of three, large, elevated
contiguous scales; supralabials, 14-14; infralabials, 13-13; a group
of larger scales on edge of upper eyelid, the largest scales (3 or 4)
bordering edge; mental small, separating first postmentals; scales
on chin quadrangular, forming definite rows, smaller mesially, the
rows nearest the labials larger than ventral scales; all ventral scales
on chin and body smooth; scales on limbs very unequal with a few
keeled scales; a few scales on the posterior border of arm and leg
that bear small tubercles; scales on venter small, more or less regu-
lar, smooth; scales on body very irregular; a small nuchal crest
beginning behind occipital bosses continues to near level of arm
insertion; no dorsal crest; scales above arm largest, forming a
curving row; crossing the back are series of scales somewhat larger
than intervening scales; no femoral or preanal pores; fourth toe
much longer than third reaching to a point near back of orbit; tail
somewhat compressed, the whorls indistinct and bordered behind
by larger irregular scales; the scales above and below keeled.

Color: ( In preservative. ) Blackish with a series of six, dim, gray
lines crossing back on the larger scales; arms and legs barred with
black and ultramarine; tail dimly barred with olive and gray; labials,
chin, throat, and area about angles of jaws, whitish; below, olive
with some scattered gray flecks; larger scales on sides ultramarine.

Measurements in mm.: Snout to vent, 80; tail, 162; length of head,
25; width of head, 15; length of rostral appendage, 105; arm, 42;
leg, 64.

Taylor: Lizards of Ceylon 1563

Remarks: There are certain variations which may be arranged
in the form of a key.

1. Rostral appendage laterally compressed; scales on top of head,

gular region, breast, under tail, on arms and legs, to some
extent on sides and ventral surfaces, keeled; appearing be-
tween the occipital bosses is a short, denticulate, nuchal crest
rising from an elevated fold about four scales high, reaching
posteriorly to back level of arm insertion; a row of six, en-
larged, lateral, white scales, their position marking lower edge
of dim light transverse bands that follow series of slightly en-
larged scales; a slight diagonal series below eye, each scale
with a whitish elevated center "Ceylon" E.H.T.-H.M.S. No. 30494
Rostral appendage not laterally compressed; scales on head, tail,
and at least lower part of arms and legs with keels; venter,
chin and neck smooth; a white lateral line present or absent;
crest variable in height, length and origin 2

2. A pair of dorsolateral whitish or cream lines extending onto tail;

lips and posterior jaw-angle whitish; chin and throat cream;
sides darker, some of the large lateral scales lavender; arms
and legs banded lavender and brown; skin on neck black; a
crest begins a short distance behind occipital bosses, with ap-
proximately 10 denticulations; ventral scales smooth. "Cey-
lon" U.S.N.M. 192165

No dorsolateral lines; keeling variable, origin and length of crest
variable 3

3. Scales on venter and breast, and median scales on chin, keeled; a

low crest arises between occipital bosses, which are rather
closely approximated; denticulated scales of crest compressed,
widened at base, the ridge four scales high at highest point; the
denticulations continued beyond crest; there are 14 scales in
the nuchal crest and approximately 10 in dorsal crest, con-
tiguous or more rarely narrowly separated,

K.U.M.N.H. Nos. 20131-20132
Ventral scales smooth; dorsals smooth; dorsal head scales, tail and
some scales on limbs, keeled; nuchal crest rather elongate,
consisting of 15 denticulate scales, with a few, low, incon-
spicuous median dorsal scales following; first two scales of
crest paired between bosses in one, but not so in the second
specimen E.H.T.-H.M.S. 30495; U.S.N.M. 19217

The single copy available to me of Gray's, Illustrations of Indian
Zoology has the plate 168 missing and I cannot judge which of the
above specimens show closest resemblance to the type figure. Since
no specific localities are available with these specimens, it is im-
possible for me to determine whether these differences represent
geographic variations or not. Aside from the scale differences it is
possible that the living specimens may have shown considerable
color and pattern differences. Collections of series of this species

1564 The University Science Bulletin

from very numerous localities is essential for an understanding of
the meaning of this variation, and no new form should be named
until adequate series are available, and compared with the type of
C. stoddartii.

Genus Lyriocephalus Merrem

Lyriocephalus Merrem, Tentamen Systematis Amphibiorum, 1820, p. 49, (type
of genus, Lyriocephalus margaritaceus) .

A single species is known. It is endemic in the highlands of
Ceylon.

Lyriocephalus scutatus Linnaeus

Lucerta scutata Linnaeus, Systema Naturae, ed. 10, 1758, p. 201, (Based on a
figure in Seba, Thesaurus, vol. 1, p. 173, pi. 109, fig. 3 [type locality,
"Amboyna" ex errore. Restricted to Ceylon.]).

Luriocephalus margaritaceus Merrem, Tentamen Systematis Amphibiorum,
' 1820, p. 49 (Based on Seba).

Lyriocephalus scutatus Kelaart, Prod. Faun. Zeylanieae, 1852 (1853), p. 166;
Gunther, Reptiles of British India, 1864, p. 128; Theobald, Descriptive
Catalogue of the Reptiles of British India, 1876, p. 99; Boulenger, Catalogue
of the Lizards in the British Museum, vol. 1, 1885, pp. 281-282 (synonymy);
Boulenger, The Fauna of British India including Ceylon and Burma; Rep-
tilia and Batrachia, 1895, pp. 121-122; Deraniyagala, Ceylon Journ. Sci.,
B, vol. 16, 1931, p. 147-148, pi. 33, and ibid. vol. 17, 1932, p. 46.

L'iriocephalus macgrcgorii Gray, Illustrations of Indian Zoology, vol. 2, pi. 68,
' fig- 1.

Diagnosis: A pair of sharp high bony crests on sides of snout
following the superciliary edge, terminating in a bony point behind
orbit; a rounded protuberance on tip of snout.

Description of species: (from K.U.M.N.H. No. 19534). Head
ridges above flare out anteriorly, the distance between them greater
than posteriorly at their termination; scales on snout protuberance
large, about 20 in number; scales on head variable in size; an irregu-
lar larger scale series outlines the inner dorsal part of orbit, separated
from opposite series by five scale rows; three or four enlarged
supraoculars; crests bordered by enlarged compressed scales, ter-
minal one largest; rostral and two adjoining labials form part of
the globular protuberance on snout; supralabials 16, infralabials
about 18, the posterior scales not well differentiated; tympanum
wanting; a transverse series of four compressed spines on occiput,
behind which are a pair of high compressed soft spines; nostril
directed outward and slightly downward; a row of 26 suborbital
scales from nostril, the three posterior largest, terminating in the
temporal region; a crest present, composed of soft, dorsal, serrate
scales; posteriorly, the scales of the crest are separated; body cov-
ered with small, flat, imbricating scales, directed upward, with three
series of larger scales on sides of neck, shoulders, and sides; below
these, on sides, numerous scattered trihedral or heavily keeled

Taylor: Lizards of Ceylon 1565

scales; approximately 28 keeled ventral scale series; a strongly
defined, gular pouch or fold; tail rather short, compressed, with
four rows of keeled scales below; no femoral or preanal pores; five
fingers and five toes, the limbs strong, well developed; hind limb
reaching mouth angle; 22 scales under free part of fourth toe.

Color: Leaf green in life; gular sac yellow, the enlarged scales
green; belly sometimes bluish; globular protrusion on snout, light
brownish or cream.

Measurements in mm.: (from K.U.M.N.H. Nos. 19534, 19535,
19536 Ceylon respectively.) Snout to vent length, 125, 132, 154;
tail, 105, 127, 152; arm, 63, 70, 77; leg, 81, 85, 105; width of head,
24, 25, 27; length of head, 34, 34, 40.

Remarks: This species has a superficial resemblance to a species
of Chamaeleon found in Ceylon and it is perhaps on this general
resemblance of the skulls that some are inclined to reduce the
chamaeleon group to the status of a family, Chamaeleonidae.

Variation: A female specimen, U.S.N.M. No. 58489, from Pera-
deniya, Ceylon, is gray, the upper free edges of the scales being
edged with brown. The rows of enlarged scales and the scattered
individual scales or groups of larger keeled scales are blue. The
dorsal crests are low. The enlarged scales on the tail and limbs
are bluish.

A pair of sharp incisors, above and below, are present in this
species.

Genus Calotes Cuvier

Calotes Rafinesque, Anal. Nat., 1815, p. 75, nomen nudum.

Calotes Cuvier, Regne Animal, vol. 2, 1817, p. 35 (type of genus, Lacerta
calotes ) .

The genus Calotes extends through southern Asia, throughout the
Philippines and most of the East Indian Archipelago. Six clearly
differentiated species occur in Ceylon.

Key to the Species of Calotes in Ceylon

1. Scales on sides point backwards and upwards 2

Scales on sides pointing backwards or backwards and downwards, 4

2. Two separated spines above tympanum versicolor

A row of 8 or 9 compressed spines above tympanum calotes

4. No spines on head liocephalus

Spines present on head 5

5. A row of spines above and somewhat behind tympanum; ventrals

larger than dorsals nigrilabris

At least two separated spines above tympanum 6

6. Lateral scales pointing backwards but not downward; no dorsal

crest ceylonensis

Lateral scales pointing backwards and downwards; a dorsal crest,

liolepis

1566 The University Science Bulletin

No less than six species of the genus Calotes are known to occur
on the island of Ceylon, and of these four, C. ceylonensis, nigrilabris,
liolepis and liocephalns, are endemic; two others occur in India
also, one, C. calotes being confined to Southern India and the
Nicobar Islands; the other, C. versicolor, has a very wide distribu-
tion in southern and southeastern Asia, and parts of Sumatra.

These species are arboreal, some (perhaps all) coming to the
ground to deposit their eggs. I have recently described the process
of nestbuilding in Calotes calotes*

Calotes versicolor (Daudin)

Agama versicolor Daudin, Histoire Naturelle des Reptiles, vol. 3, 1802, p. 395,
pi. 44 (type locality, "India") restricted to Pondicherry, India by Smith
["terra typica"].

Calotes versicolor Kelaart, Prod. Fauna Zeylanicae, 1852 (1853), p. 170; idem,
vol. 2, 1853 (1854), p. 7; Boulenger, Catalogue of the Lizards in the British
Museum, 2nd ed. vol. 1, 1885, p. 321; Haly, First Report on the Collection
of Lizards in the Colombo Museum, 1886, pp. 2, 7; Boulenger, The Rep-
tiles of British India, . . . Reptilia and Amphibia, 1890, pp. 135, 136,
fig. 42; Deraniyagala, Ceylon Journ. Sci., B, vol. 16, 1931, p. 150-151;
Smith, The Fauna of British India . . ., Reptilia and Amphibia, vol. 2,
Sauria, 1935, pp. 189-193.

Diagnosis: A Calotes with a strongly compressed body; a distinct
gular sac, but lacking a pit or depression on front of shoulder; a
swelling at base of tail in males that tapers regularly, the basal
scales, especially dorsal ones, greatly thickened; scales on sides
pointing backwards and upwards; two separated spines above
tympanum, sometimes flanked by flattened, erect scales.

Description of species: (E.H.T.-H.M.S., No. 30594, 12 mi. N.
Trincomalee, Ceylon ) . Body compressed with a rather high nuchal
crest continuous with the dorsal crest; in males, cheeks strongly
swollen, the forehead not concave; snout short, its length from orbit
equal to distance between tympanum and eye; rostral relatively very
small, less than one fifth of mental; nasal scale more than twice size
of nostril, separated from the rostral by two scales; supralabials, 11-
11, infralabials, 9-10; dorsal head scales smooth, irregular; mental
large, sharply pointed behind; the labial border more than three
times that of rostral; scale rows delimiting supraocular area scarcely
distinguishable from other scales; five superciliaries; a pair of high,
soft spines above tympanum, each flanked by erect, smaller, soft
spines or scales; nuchal crest of flattened, somewhat curved, sharply
pointed soft spines reaching a height of eleven millimeters; the
dorsal crest lower, continuous, gradually diminishing in height, the
scales contiguous, terminating abruptly eighteen millimeters be-

* Taylor, Herpetologica, vol. 7, 1951, pp. 59-60.

Taylor: Lizards of Ceylon 1567

hind level of hind leg; scales on sides of body larger than ventrals,
the scales pointing backwards and upwards, all keeled; gular pouch
inconspicuous, or absent, in preserved specimens; gular scales as
large as ventrals, mucronate; no pit or groove on shoulder; ventrals
and caudals keeled; tail rather rounded, much swollen at base; ap-
proximately 42 scale rows around body; eighteen scales about widest
part of tail base in males, the dorsal scales greatly thickened with a
median rounded ridge and depression on each side; scales bordering
crest directed nearly vertically, tending to hold the crest erect; hind
leg brought forward, the toe reaches the middle of eye; third and
fourth toes nearly equal; fourth toe much longer than third.

Color: Above olive brown or grayish in males, often with scat-
tered distinguishable flecks of bluish white; tail very dimly barred
with darker spots, some scarcely discernible; venter gray-white;
limbs with some darker markings.

Measurements in mm.: Length, snout to vent, 128; tail, broken;
head length, 40; head width, 31; axilla to groin, 56; arm, 57; leg, 85.

Remarks: The specimens available are as follows: E.H.T.-H.M.S.
Nos. F. 984, F. 989, 30592-30609, 30641, 30646, 30656, 30659, 12
miles north Trincomalee, Ceylon. U.S.N. M. No. 38283 Kandy,
Ceylon; 102317-102320, 102322-120324, Clodagh Estate, Rattota,
Matale District, Ceylon; K.U.M.N.H. No. 19531-19532 "Colombo,"
Ceylon.

Variation: The local population differs very greatly in coloration,
the females often being of a dark reddish-tan color with seven,
rather broad, transverse bands across back, the bands interrupted on
each side by a more or less distinct cream line running from shoulder
onto tail; tail barred with dark bands at least on basal half; younger,
half-grown males may have similar banding on a grayish background
but lacking the light line ( save in very young males ) ; chin may have
some black, converging lines, and some scattered black flecks on
the under side of limbs and venter.

The known distribution of this species is from Afghanistan to
Hong Kong, south in India and Ceylon, occurring also in Malaya
and Sumatra. There is considerable variation in this very wide-
spread species, especially in the size of the lateral scales and the
number of rows around the body, the smallest numbers being found
in Indian and Ceylonese specimens. It varies too in its vertical
distribution, occurring in lowlands and at elevations up to 7000 feet.
Subsequent study may prove that subspecific differences exist.

69—6490

1568 The University Science Bulletin

Calotes calotes (Linnaeus)

Lacerta calotes Linnaeus, Systema Naturae, 10th ed. 1758, p. 207 (type lo-
cality, Ceylon ) .

Agama ophiomachus Merrem, Tentamen Systematis Amphibiorum, 1820,. p. 51.

Calotes ophiomachus Kelaart, Prodr. Faun. Zeylanieae, 1852 (1853) p. 169;
idetn, vol. 2, 1854, p. 7; Boulenger, Catalogue of the Lizards in the British
Museum, vol. 1, 1885, p. 327; Haly, First Report on the Collection of
Lizards in the Colombo Museum, 1886, p. 8 ( Rattota; Western and Eastern
Provinces). The Fauna of British India, including Ceylon and Burma;
Reptilia and Batrachia, 1890, p. 140; Deraniyagala, Ceylon Journ. Sci., B,
vol. 15, 1931, p. 153.

Calotes calotes Smith, The Fauna of British India, including Ceylon and Burma;
Reptilia and Amphibia, vol. 2, Sauria, pp. 201-202; Taylor, Herpetologica,
1951, p. 59 (egg laying habits).

Diagnosis: A very long-tailed species, green, with a series of
narrow, transverse, bluish-white lines crossing dorsum; dorsal scales
of body not larger than the ventral scales; an irregular row of com-
pressed spines above tympanum; tail extremely long, usually at
least three and one-third times snout-vent length; only 30-35 scale
rows around body; a pit or depression present in advance of
shoulder.

Description of species: (From E.H.T.-H.M.S. No. 30580, 12 mi.
N Trincomalee, Ceylon.) Head rather short, the forehead very
slightly concave; no gular fold; body compressed, rather triangular
in cross section; rostral small, flanked by two labials and bordered
behind by three scales, its labial border less than that of mental;
nasal scale larger than rostral, at least three times area of nostril,
separated from the rostral by two scales; scales on snout and head
not keeled but the edges bordered by minute pores or tubercles
giving them a roughened appearance; irregular scale-rows de-
limiting the supraocular areas, separated by three scale rows; supra-
ocular scales, larger, irregular, about 10 or 11 in number; four
superciliaries; supralabials, 10-10, the infralabials, 10-10; tympanum
large, its diameter (3 mm.) slightly less than eye opening (3.2
mm. ) ; a small pit at beginning of shoulder, covered with granular
scales on black skin; scale rows on sides directed backwards and
upwards, faintly keeled, not or scarcely larger than ventrals which
are distinctly keeled; a pair of elongate, soft spines above tympanum
each flanked by two or three, flattened, soft, erect scales, sometimes
forming a continuous row above and extending somewhat anterior
to tympanum level; nuchal crest composed of soft flattened spines
and continuous with a short dorsal crest that rapidly diminishes in
height; upper scale-row composed of mucronate, erect scales seem-
ingly holding the crest erect; (tail in males not strongly inflated at
base, the dorsal scales not especially modified); tail in females

Taylor: Lizards of Ceylon 1569

rather slender, the length in both sexes 3.32 to 3.36 times head-body
length; * leg reaches to nostril.

Color: Bright green on head, body and tail, with five narrow,
bluish-white lines crossing the body transversely, and one or two
near base of tail; the lines as wide as a scale-row; tail dimly banded
with darker and lighter in preserved specimens; males in life may
show a reddish area on neck; belly light green or greenish white.

Measurements in mm.: Length, snout to vent, 91; tail, broken;
length of head, 25; width of head, 17; length of snout, 8; length of
orbit, 7.6; axilla to groin, 43; arm, 50; leg, 83.

Remarks: Specimens of this species available are as follows:
E.H.T.-H.M.S. Nos. 30577-30591, 12 mi. N Trincomalee, Ceylon;
U.S.N.M. Nos. 120316, 120321 Clodagh Estate, Rattota, Matale Dis-
trict, Ceylon; K.U.M.N.H. Nos. 20094 "Colombo," Ceylon; 31288,
Tonacumbe Estates, Namunukula, Ceylon.

This species seemingly has the longest tail of any species in the
genus, this appendage reaching a length equivalent to 3.84 times
the snout-vent length. The white marks are usually visible, but
during egg-laying activity all trace of this marking is lost and the
color may become a dirty, blackish olive, or greenish brown.

After the eggs have been buried, the head is used for pounding
the loose soil to make it firm above the nest.

CaJotes liocephalus Giinther

Calotes liocephalus Giinther, Ann. Mag. Nat. Hist., ser. 4, vol. 9, 1872, p. 86
(type locality, Peradeniya district Ceylon); Theobald, Descriptive Catalogue
of the Reptiles of British India, 1876, p. 109; Boulenger, Catalogue of the
Lizards in the Eritish Museum, 2nd ed., 1885, p. 329, pi. 26; Haly, First
Report on the Collection of Lizards in the Colombo Museum, 1886, p. 8
( Agrapatanas ) ; Boulenger, The Fauna of British India . . .; Reptilia
and Batrachia, 1890, pp. 141-142; Miiller, Verb. Nat. Gesellsch. Basel, vol.
8, 1889, p. 698; Deraniyagala, Ceylon Joum. Sci., B. vol. 16, 1931, p. 154-
155; Smith, The Fauna of British India, including Ceylon and Burma; Rep-
tilia and Amphibia, vol. 2, Sauria, 1935, p. 204 (Gammaduwa, Agrapatnas,
Punduluoya, in the Central Provinces).

Diagnosis: This species may be diagnosed by the absence of a
pair of erect spinelike scales above and anterior to tympanum; as
well as by the absence of a row of compressed scales above and
somewhat behind tympanum; dorsal scales smaller than liolepis and
feebly keeled, about as large as ventral scales; gular scales as large
or a little larger than ventrals; 43-50 scale-rows around middle of
body; hind limb reaches to eye or nearly as far; in male head large
and base of tail swollen.

* Malcolm Smith, loc. cit. gives the measurement of Calotes calotes as: snout to vent
130 mm.; tail 500, which is 3.84 times head body length.

1570 The University Science Bulletin

Male uniform green, bluish green or olivaceous above, with five
or six angular, reddish-brown crossbars; upper lips and cheeks with
a brown streak, or spotted with brown; sometimes dark crossbars
on top of head; base of tail light olive brown, the rest of it banded
light and dark; below greenish white.

Female may be uniform green except for some black markings
on snout and flanks. Snout to vent, 90 mm.; tail, 250. The species
is known from the Central provinces (Gammaduwa, Agrapatnas,
Punduluoya ) .

Data from Malcom Smith (1935, p. 204).

Calotes nigrilabris Peters

Calotes rouxi Blyth, Journ. Asiat. Soc. Bengal, vol. 22, 1853, p. 647 (not of
Dumeril and Bibron).

Calotes (Bronchocele) nigrilabris Peters, Monatsb. Akad. Wiss. Berlin, 1860,
p. 183 (type locality, "Newerelia" = PNuwara Eliya).

Calotes nigrilabris Giinther, Reptiles of British India, 1864, pp. 143-144, pi. 40,
fig. D. (Giinther distinguishes two varieties, Alpha and Beta.); Boulenger,
Catalogue of the Lizards of the British Museum; vol. 1, 1885, pp. 328-329
(Ceylon); Miiller, Verh. Nat. Gesellsch. Basel, vol. 8, 1889, p. 698; Annan-
dale, Spolia Zeylanica, vol. 8, pt. 30, June 1912, pp. 135-136 (Nuwara
Eliya; Pattipola, alt. 6000 ft.); Deraniyagala, Ceylon Journ. Sci., B, vol. 16,
1931, p. 153-154; Smith, The Fauna of British India, including Ceylon and
Burma; Reptilia and Amphibia, vol. 2, Sauria, 1935, p. 206 (Ceylon, in
hills at high altitudes).

Diagnosis: A row of four or five flattened spines arising at a point
above tympanum and extending behind its posterior level; a nuchal
crest, and in males at least, a low, serrate, dorsal crest; scales of
dorsum directed backwards and downwards; scales keeled; leg ex-
tends beyond orbit; coloration variable; an area of small tuber-
cular scales on shoulder forming a "pit."

Description of species: Head large, areas about jaw angles rather
inflated; rostral small, bordered by two labials and five postrostral
scales; nasal rather large, the nostril occupying less than half its
area, separated from rostral by a single scale; an elongate supra-
nasal; four scales in canthal row, the canthal edge rather short; three
superciliaries; dorsal head scales smooth, unequal; supraorbital area
bordered by a row of larger, unequal scales, separated from its
fellow by two scale rows; supraoculars, 15 to 17, unequal; nuchal
crest rather high (beginning between a pair of slightly elongate,
more or less elevated ridges on occiput); scales of crest flattened,
compressed, highest at back level of head and continuous with a
low serrate crest on dorsum; lateral scales distinctly smaller than
ventral scales, more or less keeled, the scales pointing backward and
downward; scales on side of neck pointing up or up and back; ven-
tral scales, scales on limbs and tail keeled, mucronate, the mucrone

Taylor: Lizards of Ceylon

1571

heavy, distinct; throat with a suggestion of a pouch, the median
scales slender with strong mucrones; mental with a labial border
equal to that of rostral; supralabials, 9-9, infralabials, 8-8; when leg
is brought forward the toes reach halfway between eye and nostril;
a rather large area of fine granular scales in front of arm insertion
on side of neck; base of tail swollen, surrounded by 15 rows of
scales; approximately 43 scale-rows around middle of body; tym-
panum large.

Color: Olive or greenish in life, becoming dirty grayish blue on
head and on body and limbs; back part of head and neck region
gray olive; tail distinctly greenish olive; infralabials bluish; a black
mark along supralabials, leaving a series of tiny bluish spots on
labials; a light line below eye to tympanum bordered above by
light.

Measurements in Millimeters of Calotes nigrilabris Peters

Museum

EHT-HMS

USNM

EHT-HMS

Number

30502

58666

38281

38282

30503

Locality

Nuwara
Eliya

"Ceylon"

Nuwara
Eliya

Sex

d1
86

9
83

Snout-to-vent length

Tail length

293
32.2

264
30.5

197 + ?
29.5

238
26

205

Head length

Head width

22.3

19.2

14.6

Axilla to groin

36?

Arm

51.5
84

49
71

46.5
64

Leg

63.5

Remarks: The coloration varies in life but usually it is uniform
green with a broad black band along the lips to behind the tym-
panum; tail brownish with darker-bordered light band or spots.
Sometimes the lips are green as is the remainder of head and body.

A female specimen in the collection differs in being olive brown
above with two chevronlike bands across shoulders pointing back-
wards. The remainder of the dorsum has small white spots, the tail
a series of cream, brown-edged spots. The nuchal crest of the
female is low, and the dorsal crest is scarcely discernible. In this

1572 The University Science Bulletin

specimen the toe reaches to the orbit. There are two eggs in each
oviduct.

Calotes ceijlonensis Miiller

Calotes mystaceus ceylonensis F. Miiller, Verh. Naturf. Gesellsch. Basel, vol. 8,
1887, p. 292, pi. 3 (type locality, Kumbukan-aar, southwest Cevlon).

Calotes Haly, Taprobanian, vol. 2, pt. 5, Oct. 1887, p. 133 (described, but not
named ) .

Calotes kclaartii Nevill, Taprobanian, vol. 2, 1887, p. 134 (type locality, by
inference, North Western Province, Ceylon — species named but not de-
scribed. The description is that of Haly. )

Calotes ceylonensis Boulenger, The Fauna of British India, including Ceylon
and Burma; Reptilia and Batrachia, 1890, pp. 139-140; Deraniyagala, Ceylon
Journ. Sci., B, vol. 16, 1931, p. 151-152; Smith, The Fauna of British India
including Ceylon and Burma; Reptilia and Amphibia, vol. 2, Sauria, 1935,
pp. 202-203.

Calotes saleoides Werner, Verh. Zool-bot Ges. Wien, vol. 46, 1896, p. 7 (type
locality, Ceylon ) .

Diagnosis: A diagonal, pitlike depression in front of shoulder,
its surface covered by small tubercular scales; rows of scales on sides
of dorsum directed backwards and downwards; two separated
spines on occiput above and somewhat forward of the tympanum;
a nuchal crest but no dorsal crest; scales about body, 54 to 60; none
of the scale rows directed downward.

Description of species: (E.H.T.-H.M.S., No. 30576, $ , "Ceylon").
Rostral small, flanked by two labials, bordered behind by four
scales, its width a little greater than its height; nasal separated
from rostral by two scales; supranasals larger than other scales on
dorsal part of snout, equaling or nearly equaling the nasal; supra-
labials, 10-10; infralabials, 10-9; inner row of scales bordering the
supralabials not strongly defined, separated by three scale-rows;
supraoculars about 20, irregular; five superciliaries, canthal series of
four scales partly interrupted; tympanum more than half length of
eye opening; a pair of erect, soft spines above the tympanum; a
low nuchal crest (female) beginning on a level with the more
anterior of the two spines; depression or pit on shoulder distinct, its
surface covered with small imbricating scales; scales on snout di-
rected forward or outward, irregularly ridged; scales on nuchal
region with small tubercles; on dorsum scales larger than those on
ventral surface, the two rows bordering the nuchal spines largest;
upper lateral rows directed back and upwards; others directed
straight back, all more or less keeled; all scales on ventral regions
keeled; leg reaches to tympanum; approximately 54 scales around
middle of body; no gular sac.

Color: Head dark brown above; a light stripe from nostril back
across area below eye to tympanum; dorsum light brown, barred
with bands of varying width, those more posterior may be six scales

Taylor: Lizards of Ceylon 1573

long; sides with some brown blotches and on flanks the lighter
ventral color encroaches; tail dimly barred brown and light brown;
some brown flecks on chin.

Measurements in mm.: Length, snout to vent, 72; tail, 155; length
of head, 21; width of head, 13; axilla to groin, 37; arm, 31; leg, 50.

Remarks: Males differ considerably in having the cheeks inflated
or swollen and presumably the head scales are smoother. The
nuchal crest is much higher and stronger and the base of the tail is
swollen and covered with large thickened scales, the median form-
ing a serrated edge.

The color of the male is olivaceous, the back of head and front
of the dorsum being more or less pinkish or reddish in life. Some
transverse darker bars may be present and a pale stripe passing
from below the eye to the end of the jaw is usually discernible.

This species occurs chiefly in lowland forest in the northern half
of the Island.

Calotes Holepis Boulenger

Calotes nemoricola Giinther, Proc. Zool. Soc. London, 1869, p. 507 (not of
Jerdon ) .

Calotes liolepis Boulenger, Catalogue of the Lizards in the British Museum,
vol. 1, 1885, pp. 326-327 plate 25, fig. 2 (type locality, Ceylon); Nevill,
Taprobanian, vol. 2, pt. 5, Oct. 1887, pp. 133-134 (Kandy, Kotmali);
Boulenger, The Fauna of British India, including Ceylon and Burma; Bep-
tilia and Batrachia, 1890, p. 140; Deraniyagala, Ceylon Journ. Sci., B,
vol. 16, 1931, p. 152-153; Smith, The Fauna of British India, including
Ceylon and Burma; Beptilia and Amphibia, vol. 2, Sauria, 1935, p. 203
( Punduluoya, Kandy, Gammaduwa ) .

Diagnosis: A species having a pair of soft, spinelike scales above
and forward of tympanum; a nuchal crest narrowly separated from
a short dorsal crest; scales on sides of neck pointing upward; those
on sides of body pointing backward and downward; scales on chin,
throat and venter with minute mucrones, and sometimes additional,
minute, hairlike, posterior projections; scales about base of tail
largest; leg reaches to tympanum; an area of small tubercular scales
on shoulder, tending to form a pit.

Description of species: (From K.U.M.N.H. No. 19533, "Cey-
lon.") Rostral small bordered by two labials and three postrostrals;
nostrils occupying half surface of nasal scales; nasals separated from
rostral by two scales; a large supranasal; five scales on canthal edge
behind nasal; scales on head as far back as occiput smooth, unequal;
rows of large scales outline orbits, separated from each other by
three scale-rows; three elongate superciliaries; supraorbitals very
large, irregular; scales on occiput and in anterior nuchal region each
bearing an elevated tubercle; approximately 32 scales around middle

1574 The University Science Bulletin

of body; nuchal crest with eight, soft, compressed spines, the largest
five to eight millimeters in length; two soft spines, one directly
above, one anterior to level of naked tympanum; area about jaw
angles swollen; a short dorsal crest consisting of eight or ten, soft,
small, posteriorly directed spines; scales on base of tail larger than
body scales; scales on sides pointing back and downward, not
keeled. Third and fourth toes nearly of equal length; toes reaching
to tympanum.

Color: In preservative, head light reddish brown, the color ex-
tending to tympanum but the swollen areas at jaw angles blackish;
nuchal crest and occipital spines black; dorsum blackish brown with
three indistinct transverse lighter bands on back; throat and neck
blackish, the anterior part of chin lighter; venter lighter than sides.

Measurements in mm.: Snout to vent, 82; tail, 212; length of head,
30.5; width of head, 19; arm, 40.5; leg, 59.

Family Scincidae Gray

The Ceylonese species of this family have recently been reviewed

by me, and specimens in the collections reported on. Since this

paper is readily available,* I am not repeating this data here other

than to repeat the keys, since workers may find it convenient to

have an entire listing of the species under a single cover. The

following species are known:

Mabuya bibronii Sphenomorphus taprobanense

Mabuya macularia Sphenomorphus fallax

Mabuya beddomii Sphenomorphus rufogulus

Mabuya flowed Chaleidoseps thwaitesi

Mabuya carinata Nessia burtonii

Mabuya madaraszi Nessia didactyla

Riopa punctata Nessia monodactyla

Riopa singha Nessia bipes

Dasia haliana Nessia sarasinorum

Sphenomorphus dussumieri Nessia hikanala

Sphenomorphus megalops Nessia layardi

Sphenomorphus deignani Nessia deraniyagalai
Sphenomorphus striatopunctatus

Key to the Genera of Scincidae in Ceylon

1. Palatine bones in contact or overlapping along medial palatal line, 2
Palatine bones not in contact along medial palatal line 6

2. Supranasals present 3

Supranasals absent 5

* Ceylonrsc Lizards of the Family Scincidae, Univ. of Kansas Sci. Bull., vol. 33, pt. 2,
Mar. 20,' 1950, pp. 481-518.

Taylor: Lizards of Ceylon 1575

3. Scales keeled with 2 to 7 keels 4

Scales smooth; body somewhat elongated; limbs pentadactyl,

but short, widely separated when adpressed; scales smooth;
lower eyelid with a semitransparent disk Riopa

4. Pterygoid bones not in contact, the palatal notch extending for-

ward to level of centers of eyes; limbs pentadactyl, well de-
veloped, distinctly overlapping when adpressed; no distinct

transverse bands Mabuya

Pterygoid bones in contact anteriorly, the palatal notch not reach-
ing forward to level of centers of eyes. Limbs pentadactyl,
failing to touch or barely overlapping when adpressed; color
pattern of transverse bands Dasia

5. Limbs pentadactyl, meeting, overlapping or failing to meet when

adpressed; lower eyelid scaly; frontoparietal single or double;
scales smooth or at most, with slight suggestion of keels or

striations Sphenomorphus

Limbs not pentadactyl, greatly reduced or absent; body angui-
form; the nostril in rostral 6

6. Nostril in anterior part of rostral, connected to the posterior edge

of scale by a groove; limbs variable, never tetradactyl Nessia

Nostril near the posterior edge of rostral; limbs short, tetradactyl,

Calcidoseps

Genus Maruya Rafinesque

Six species occurring in Ceylon may be differentiated by the
following key:

Key to Species of Mabuya in Ceylon

1. A transparent disc or lower eyelid bibronii

No transparent disc, but several larger transparent scales on

lower eyelid 2

2. A postnasal macularia

No postnasal 3

3. Scales almost smooth or with three to five very feeble keels . beddomii
Scales not smooth, each bearing 3, 5 or 7 strong keels 4

4. Scales tricarinate ( or sometimes also with a small tubercle on

outer edge of scale); dorsum with a paired series of black

markings floweri

Scales with five or seven keels (except very young which may
have three); no paired series of black markings on dorsum. ... 5

5. Larger; snout to vent, 123 mm.; adpressed hind limb to wrist or

elbow carinata

Smaller; snout to vent, 77 mm.; adpressed hind limb to axilla or
farther madaraszi

Genus Riopa Gray

Two species occurring in Ceylon may be differentiated by the
following key:

1576 The University Science Bulletin

Key to the Species of Riopa in Ceylon

Scales 24-26 rows, each dorsal and lateral scale with a dark spot.

In young, spots forming 6 dark lines separated by light lines;

tail uniform red; dorsolateral light lines from rostral,

Riopa punctata
Scales in 28 rows; four very narrow dark lines on median scale

rows; dorsolateral lines from supraoculars; vertical rows of

white spots on neck and scattered white spots on the sides;

a dorsolateral line from nuchal Riopa singha

Genus Dasia Gray

A single endemic species, Dasia haliana, occurs in Ceylon. The
characters being those of the genus.

Genus Sphenomorphus Fitzinger

Seven species from Ceylon are referred to this genus. They may
be differentiated by the following key:

Key to the Species of Sphenomorphus in Ceylon

1. Frontoparietal divided 2

Frontoparietal single 6

2. Less than thirty scale rows about body 3

Forty scale rows about body dussumieri

3. Parietals enclosing interparietal 4

Parietals separated by interparietal megalops

4. Dorsal scales striated deignani

Dorsal scales unstriated 5

5. Adpressed limbs barely overlap; prefrontals forming a common

suture taprobanense

Adpressed limbs separated by seven scales; prefrontals usually
separated striatopunctatus

6. Males with sides of head and throat blue-black, each scale with

a whitish spot fallax

Males with throat bright, rosy red, lacking black color and white
spots rufogulus

Genus Chalcldoseps Boulenger

A single species Chalcldoseps thwaitesi Giinther is known in
Ceylon. The characters are those of the genus.

Genus Nessia Gray

The following key will differentiate the eight forms recognized
under specific names:

Key to the Species of Nessia

1. Two or four limbs present; interparietal broader than frontal

(except sarasinorum) ; ear opening present 2

Limbs absent 6

Taylor: Lizards of Ceylon 1577

2. Limbs bearing clawed digits 3

Limbs budlike, lacking clawed digits 4

3. Four limbs present, tridactyl; 24 scales at midbody burtonii

Four limbs present, didactyl; 24 scales at midbody didactyla

4. Four limbs present; 24-26 scales about midbody monodactyla

Two limbs present; scales variable 5

5. Scale rows about midbody, 28 bipes

Scale rows about midbody, 22; interparietal narrower than

frontal sarasinorum

6. Snout flattened below, projecting sharklike; frontonasal one third

width of rostral; ear-opening present hickanala

Snout not especially flat, not sharklike; frontonasal more than
half length of rostral 7

7. One large elongate loreal; preoculars small; frontonasal nearly

as long as rostral laijardi

Two loreals, the posterior lower than anterior; frontonasal a little
more than half of rostral length deraniijagalai.

Sphenomorphus fallax Peters

Lijgosoma fcdlax Peters, Monatsb. Akad. Berlin, 1860, p. 184, (Ratnapura,
Trincomalee, Ceylon); Taylor, Univ. Kansas Sci. Bull., vol. 33, pt. 2, no. 13,
Mar. 20, 1950, pp. 501-504, fig. 4, A and B.

I have recently received a series of specimens from Tonacombe
Estate, Namunukula, Ceylon, 4000 ft. elev. These were not avail-
able when the lizard report was made.

The specimens are K. U. M. N. H. Nos. 31276-31286. The fe-
males of the species show a fairly well-defined pair of dorsolateral
lines, covering halves of two scale rows ( third and fourth from mid-
dle of dorsum); this bordered laterally by a broad, dark stripe be-
ginning at nostril, passing through eye to side of tail, one whole
scale-row and two half scale-rows wide; the adjoining four rows
with a narrow lighter line through their middle, the four median
dorsal rows each may have a narrow indistinct, lighter line. The
light lines continue on the tail; venter white.

Males with sides of head and throat blue, each scale with a milk-
white or bluish white spot.

The lineation on the body is lost in older males and no trace is
evident. The males seemingly are a little larger than females, with
a somewhat larger head proportionally.

Family Lacertidae Gray

Genus Cabrita Gray

Cabrita Gray, Ann. Mag. Nat. Hist., vol. 1, 1838, p. 282, (type of genus,
Cabrita brunnea ) .

This genus, which ranges in southern India is represented by
a single species in Ceylon. It is presumably confined to the lower,
dryer parts of northern Ceylon.

1578 The University Science Bulletin

Cabrita leschenaultii Milne-Edwards

Lacerta leschenaulti Milne-Edwards, Ann. Sci. Nat. Paris, vol. 16, 1829, pp. 80,
86; pi. 6, fig. 9, (type locality, Coromandel Coast, India).

Cabrita leschenaulti Boulenger, Catalogue of the Lizards of the British Mu-
seum, 2nd ed., vol. 3, l887, p. 70; The Fauna of British India, including
Ceylon and Burma; Beptilia and Batrachia, 1890, p. 172; Monograph of the
Lacertidae, vol. 2, 1921, p. 194; Deraniyagala, Ceylon Journ. Sci., B, vol. 16,
1931, p. 157, pi; Smith, The Fauna of British India including Ceylon and
Burma; Beptilia and Amphibia, vol. 2, Sauria, 1935, pp. 374-375.

Diagnosis: (from Smith, loc. cit.) Anterior labials ridged, form-
ing a projecting margin; upper head scales strongly keeled and
finely striated; canthus rostralis sharp; a single frontonasal; nostril
between two swollen nasals; one or two postnasals; prefrontals in
contact mesially; frontal long and narrow touching first three supra-
oculars; interparietal small touching a smaller occipital; four supra-
oculars; dorsal scales subequal, smaller than caudals; ventrals in six
longitudinal rows, in 24-28 transverse series; 42 to 50 scales around
body at middle; a large preanal plate; 12-16 femoral pores on each
side.

Brownish or golden above; a light stripe, edged above with black
behind superciliary edge, passes along body to tail; a second light
line borders upper lip, and passes along to flank; a black stripe be-
tween light lines (or green, spotted black); greenish white on
venter; snout to vent, 50 mm.; tail 100 mm.

Family Varanidae
Genus Varanus Merrem

Varanus Merrem, Tentamen Systematis Amphibiorum, 1820, p. 58 (type of
genus, Lacerta varia Shaw).

This genus comprises the largest living lizards, one species reach-
ing a length of 10 or more feet. Two species are known from
Ceylon, a smaller Varanus begalensis begalensis Daudin, which
reaches a length of nearly 6 feet, and Varanus salvator Laurenti,
even larger, reaching a length of eight feet, four inches. These
animals run rapidly, often climbing trees and often actually living
in hollow trees.

Key to Ceylonese Species of Varanus

Nostril round or oval, nearer tip of snout than to orbit; abdominal
scales weakly keeled, in 80 to 90 transverse rows . . salvator salvator

Nostril a rather narrow slit directed backwards, nearer orbit than
tip of snout; abdominal scales smooth, in 90-110 transverse
rows bengalensis bengalensis

Taylor: Lizards of Ceylon 1579

Varanus bengalensis bengalensis (Daudin)

Tupinambis bengalensis Daudin, Histoire Naturelle des Reptiles, vol. 3, p. 67
(type locality, Bengal).

Lacerta monitor Linnaeus, Systema Naturae, ed. 10, 1758, p. 201 (type lo-
cality, India).

Monitor dracaena Kelaart, Prod. Faun. Zeylanicae, 1853, p. .

Varanus bengalensis Boulenger, Catalogue of the Lizards in the British Mu-
seum, 2nd ed., vol. 2, 1885, p. 310 (Ceylon); The Fauna of British India,
including Ceylon and Burma; Reptilia and Batrachia, 1895, pp. 164-165
(Ceylon, "Whole of India"); Deraniyagala, Ceylon Journ. Sci., B, vol. 16,
1931, p. 161; Bhatt, Animal Life (Colombo Ceylon) 1942, pp. 118, 120
( native name onlv ) .

Varanus (Indovaranus) bengalensis bengalensis Mertens, Abh. Senckenb.
Naturf. Gesellsch. No. 462, 1942, pi. 3, figs. 9, 10; pi. 11, fig. 49; pi. 13,
figs. 64, 67; ibid. No. 465, 1942, pp. 182-184, pi. 22, figs. 160-161; pi. 25,
figs. 194-196; pi. 29, figs. 232-234; pi. 32, figs. 258-259; pi. 34, figs. 285,
287; ibid. No. 466, pp. 334-338.

Diagnosis: A large species reaching six feet in total length; ven-
tral scales smooth; no widened scales in supraorbital series; nostril
a narrow slit, nearer orbit than to tip of snout.

Description of species: Rostral as high as wide, bordered laterally
by first labials, bordered behind by a pair of postrostrals; largest
scales on head form three median rows between the orbits, the ad-
joining rows somewhat smaller; a median groove begins two scale-
lengths behind rostral and extends behind anterior level of nostril;
supraorbital scales somewhat wider than other scales, one or two
of which may be widened; canthus rather obtuse; supralabials, 31;
approximately 34 infralabials; mental larger than rostral, followed
by four somewhat enlarged scales separated by a groove which
continues back for some distance; ear-opening subtriangular, the
distance between eye and ear distinctly greater than distance from
eye to anterior end of nostril; 32 scales between rostral and parietal;
latter bearing pineal or parietal eye; between occipital and a point
above the vent, 158 scales; between nuchal fold and line joining
front level of legs, 104 smooth scale rows; leg when adpressed fails
to reach axilla.

Digits below with transverse series of small scales, at least 30
series under the fourth toe, one row on inner edge enlarged on basal
region; 20 series under first toe.

Color of young: Olive to brown with short black lines or spots,
those on head longitudinal, those on body tending to form narrow
transverse bands; sides, and to a lesser extent back, with small dark
spots having yellow centers; a distinct black stripe behind eye; chin
and neck more or less transversely marked with black; belly indis-
tinctly marbled, with irregular transverse lines somewhat in evi-
dence; toward distal part of tail there is a broad cream band with

1580 The University Science Bulletin

some darker lines or marbling (one or two specimens also have tip
of the tail cream color).

Adults usually dull, blackish brown, or, recently after shedding,
some yellowish may be in evidence.

Varanus salvator salvator (Laurenti)

Stcllio salvator Laurenti, Specimen medicum exhibens Synopsin Reptilium
emendatum, 1868, p. 56 ( based on plate 88, fig. 2 in vol. 2, Seba, Thesaurus,
a species of unknown provenance).

Varanus salvator Boulenger, Catalogue of the Lizards in the British Museum,
2nd ed., vol. 2, 1885, p. 314; The Fauna of British India, including Ceylon
and Burma; Reptilia and Batrachia, 1890, p. 166. Deraniyagala, Ceylon
Journ. Sci., B, vol. 16, 1931, p. 159; Smith, The Fauna of British India,
including Ceylon and Burma; Reptilia and Amphibia, vol. 2; Sauria, 1935,
pp. 406-407; Deraniyagala, Spolia Zeylanica, vol. 24, pt. 1, 1944, pp. 59-62.

Varanus salvator salvator Mertens, Abh. Senckenb. Natur. Gesellsch. no. 466,
1942, pp. 245-253; (ibid. Abh. 462, 1942, pi. 2, fig. 5; pi. 5, figs. 19-21;
pi. 15, fig. 85; pi. 16, fig. 103.)

Diagnosis: Snout depressed at end, its length three times its
height; canthus rostralis distinct but somewhat rounding; rostral
small, as wide as high bordered by a pair of postrostrals; nostril
round or oval in shape, much closer to tip of snout than to the eye;
scales on crown of head larger than nuchal scales; four to eight
median supraoculars transversely enlarged; scales of dorsal surface
oval, more or less keeled; abdominal scales keeled in 85-95 rows;
tail strongly compressed laterally, with a crest composed of paired
scales ( "doubletoothed" ) ; tongue very long, divided, snakelike.
Total length more than eight feet (2500 mm. fide Malcolm Smith).

Color: Adults dark brownish olive, usually indistinctly marked
with transverse series of yellow spots.

Suborder— RHIPTOGLOSSA

Family Chamaeleonidae Gray

Genus Chamaeleon Gronovius

Chamaeleon Gronovius, Zooph. Anim., vol. 1, 1763, p. 12 (type of genus,
Lacerta charnaeleon Linnaeus).

A single species occurs in Ceylon.

Cha?nacleon zeijlanicus Laurenti

Chamaeleo zeylanicus Laurenti, Specimen medicum exhibens Synopsin Rep-
tilium emendatum, 1768, p. 46 (based on a figure in Seba, Thesaurus, vol. 1,
pi. 82, fig. 3).

Chamaeleon calcaratus ( part. ) Boulenger, Catalogue of the Lizards in the
British Museum 2nd ed., vol. 3, 1887, p. 445, pi. 39, fig. 2; The Fauna of
British India, including Ceylon and Burma; Reptilia and Batrachia, 1890,
p. 232, text figs. 66, 67; Werner, Zool. Tahrb., vol. 15, 1902, p. 332; Der-
aniyagala, Ceylon Journ. Sci., B, vol. 15, 1931, p. 156.

Taylor: Lizards of Ceylon 1581

Chamaeleon zeylonicus Jerdon, Journ. Asiat. Soc. Bengal, vol. 22, 1853, p. 466;
Smith, The Fauna of British India, including Ceylon and Burma; Reptilia
and Amphibia, vol. 2, Sauria, Feb. 7, 1935, pp. 251-253.

Diagnosis: Foot with digits in groups of two and three opposed
to each other; tongue club-shaped and extremely extensive, (nearly
as long as head and body); head casque elevated posteriorly with
a parietal crest, a serrated dorsal crest and a median ventral crest
on chin, throat and venter.

Description of species: Head large, decorated with two elevated
lateral crests beginning on the snout, following canthus rostralis to
above eye, and temporal region, where they become lost on side
of a high, parietal crest; parietal crest begins between eyes at their
back level, and rises rapidly to the median highest point of the
bony casque; latter sharply truncate on neck; width of casque at
widest part slightly less than width of head at jaw angle; eyelids
with a narrow median opening, closely applied to the somewhat
conical eyeball; ear completely covered, but a slight depression
suggests its position; diameter of bony orbit of eye equal to its dis-
tance from tip of snout; supralabials, 25-25; infralabials, 27, the
last four not differentiated from other head scales; rostral and
mental not differentiated from labials.

Scales on head unequal, relatively larger than scales on body
and chin; body strongly compressed laterally leaving no ventral
surface except a crest; dorsum likewise merely a high crest some-
what serrate, supported by neural spines of vertebrae covered with
scales that are much larger than those on sides and limbs; feet with
the upper three toes grouped together, directly opposed to lower
two, all strongly clawed, and bound together, only the tips free;
tail slender, compressed, higher than wide, decurving, prehensile.

Color: Green in life; in preservative nearly uniform blackish on
body, the median dorsal crest and head somewhat grayish, the
median ventral crest white or cream; palms and soles cream or
yellowish, without pigment; under part of tail lighter than its sides.

Measurements in mm.: Length, snout to vent, 191; tail, 245, total
length, 436; width of head, 34; head length to back level of casque,
51; back of jaw to snout tip, 40; arm, 84; leg, 84.

Remarks: A specimen (K.U.M.N.H. No. 24137, from Puttalam,
Ceylon, W. C. Osman-Hill collector) has furnished the preceding
description. The species is probably confined to the lower, dryer
forests in the northern half of the island.

1582 The University Science Bulletin

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1944. Four new races of the kabaragoya lizard Varanus salvator. Spolia
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1945. A new gymnodactylid gecko from Ceylon. Spolia Zeylanica, vol.
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Drieberg, C.

1913. Kabaragoya raiding crow's nest. Spolia Zeylanica, vol. 8, pt. 32,
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Essex, R.

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1877. Reptile fauna of Ceylon, 1877 (Colombo), pp. 1-30.

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1845. Catalogue of specimens of lizards in the British Museum, 1845,
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1867. On a new geckoid lizard from Ceylon. Proc. Zool. Soc. London
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Green, E. Ernest

1903. Bipedal locomotion of a Ceylonese lizard. Journ. Bombay Nat. Hist.
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1908. The bite of the "Brahminy Lizard," Spolia Zeylanica, 1908, vol. 5,
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70—6490

1584 The University Science Bulletin

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1886. First Report on the collection of lizards in the Colombo Museum
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1928. Notes on the Gecko, Gymnodactylus fremitus. Ceylon Journ. Sci.,
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1929. Some Scincidae from S. Africa, Madagascar, and Ceylon. Ann.
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1926. Notes on lizards in the Indian Museum. II. On the unnamed
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1860. Verzeichniss der von Schmarda aus Ceylon gesammelten Amphibien,
und Beschreib. neue. Arten. Monatsb. Akad. Wiss. Berlin 1860,

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THE UNIVERSITY OF KANSAS

SCIENCE BULLETIN

Vol. XXXV, Pt. II] September 10, 1953 [No. 13

Fourth Contribution to the Herpetology of San Luis Potosi

Edward H. Taylor

Abstract. — A report is made on a collection of reptiles and amphibians,
containing approximately 608 specimens from San Luis Potosi, property of
Louisiana State University Museum of Natural History. The collection con-
tains a total of 71 species distributed as follows: one salamander, seventeen
species of Salientia, twenty-one species of lizards, thirty-one species and sub-
species of serpents, and one crocodile. The following are new records for the
State: Bufo occidentalis Camerano (previously reported as Bufo simus [er-
roneously]); Hemidactylus turcicus turcicus (Linnaeus); Sceloporus dugesi
intermedins Duges; Ophisaurus sp.; Abronia taeniata (Wiegmann); Rhinocheilus
lecontei tessellatus Garman; and Crocodijlus moreleti Dumeril and Bibron. Two
other species, Sceloporus serrifer cariniceps Martin, and Leiolopisma cauda-
equina (Smith) have not appeared in the author's previous lists.

Elaplie triaspis intermedia (Boettger) was formerly reported erroneously as
Elaphe chlorosoma (Gunther). Thamnopliis cyrtopsis crytopsis was formerly
reported erronously as Thamnophis eques eques (Reuss). Thamnopliis eques
eques (Reuss) was formerly reported as Thamnopliis subcarinata subcarinata
Gray.

INTRODUCTION

Three recent herpetological collections, made for the Louisiana
State Museum of Natural History and submitted to me for identifica-
tion and study by Dr. George Lowery, form the basis for this pub-
lication.* These collections were made by personnel from the mu-
seum:

Mr. and Mrs. Richard Graber spent nearly a year in San Luis
Potosi and visited several isolated mountain ranges from which no
previous herpetological collections have been reported. Their chief
interest was ornithology.

* Three other papers based on previous collections have been published as follows:

A preliminary account of the herpetology of the State of San Luis Potosi, Mexico, Univ.
Kansas Sci. Bull., vol. 33, pt. 1, Apr. 20, 1949, pp. 169-215.

Second contribution to the herpetology of San Luis Potosi, Univ. Kansas Sci. Bull., vol.
33, pt. 2, Mar. 1950, pp. 441-457, pi. 4-9, map.

Third contribution to th> herpetology of San Luis Potosi, Univ. Kansas Sci. Bull., vol. 34,
pt. 2, Feb. 15, 1952, pp. 793-815, map.

(1587)

1588 The University Science Bulletin

Mr. Richard T. Gregg was in the country pursuing limnological
studies in the summer of 1951 and obtained more than a hundred
herpetological specimens.

The third collection was made by Mr. Charles Fugler, largely in
1952, while engaged in ornithological collecting.

A total of 608 specimens f is included in the material sent.
When one realizes that the primary interests of the collectors were
in other fields, and the herpetological specimens merely the result
of chance collecting, the number is indeed large.

The materials have proved especially interesting and the number
of species and subspecies represented — seventy — is especially high.
A number of forms are reported from the State for the first time.

The following species are treated in this paper:

Caudata
Ambystoma velasci Duges

Salientia
Scaphiopus couchii Baird
Scaphiopus multiplicatus Cope
**Bufo occidentalis Camerano
Bufo valliceps Wiegmann
Bufo horribilis Wiegmann
Bufo punctatus Baird and Girard
Leptodactylus labialis ( Cope )
Syrrhophus cystignathoides (Cope)
Tomodactylus macrotympanum Taylor
Smrfisca baudinii (Dumeril and Bibron)
Acrodytes spilomma ( Cope )
Hyla arenicolor Cope
Hijla eximia Baird
Hyla miotympanum Cope
Rana pipiens Schreber
Rana montezumae Baird

Sauria

Hemidactylus turcicus turcicus (Linnaeus)
Anolis sallaei Gunther

Phrynosoma orbiculare orbicidare (Linnaeus)
Holbrookia maculata approximans Baird
Sceloporus parvus parvus Smith
Sceloporus grammicus disparilis Stejneger
Sceloporus spinosus spinosus Wiegmann
Sceloporus variabUis variabilis Wiegmann
Sceloporus jarrovi minor Cope
^Sceloporus dugesii intermedins Duges

t Not including specimens being studied by Dr. Hobart M. Smith.
** Change in nomenclature of a previously reported form.
* First state record.

Herpetology of San Luis Potosi 1589

***Sceloporus serrifer cariniceps Martin

Sceloporus torquatus melanogaster Cope

Eumeces hjnxe lynxe Wiegmann
***Leiolopisma caudaequinae (Smith)

Ameiva undulata podarga Smith and Laufe

Cnemidophorus sacki gularis Baird and Girard
*Ophisaurus sp.

Gerrhonotus liocephalus infernalis Baird

Gerrhonotus liocephalus loweryi Tihen
*Abronia taeniata taeniata (Wiegmann)

Barisia imbricata ciliaris (Smith)

Serpentes

Leptotyphlops myopicus myopicus ( Garman )

Constrictor constrictor imperator ( Daudin )

Ficimia streckeri Taylor

Geophis mutitorques Cope

Lampropeltis triangulum polyzona Cope

Rhadinaea crassa Smith
*Rhinocheilus lecontei tessellatus Garman

Salvadora lineata Schmidt
**Elaphe triaspis intermedia (Boettger)

Elaphe flavirufa flavirufa ( Cope )

Masticophis flagellum testaceus ( Say )

Masticophis taeniatus ruthveni Ortenburger

Pituophis catenifer affinis Hallowell

Pituophis deppei jani (Cope)

Drymarchon corais erebennus (Cope)

Dryadophis melanolomus veraecrucis (Stuart)

Spilotes pullatus mexicanus (Laurenti)

Thalerophis mexicanus mexicanus ( Dumeril, Bibron, and Dumeril )
*'*Thamnophis cijrtopsis cyrtopsis (Kennicott)
**Thamnophis eques eques (Reuss)

Thamnophis melanogaster canescens Smith

Thamnophis phenax Cope
*'*Thamnophis halophilus Taylor

Thamnophis sirtalis proximus (Say)

Leptodeira annulata septentrionalis (Kennicott)

Leptodeira annulata taylori Smith

Leptodeira maculata ( Hallowell )

Coniophanes miperialis imperialis (Kennicott)

Coniophanes fissidens proterops Cope

Crotalus triseriatus triseriatus (Wagler)

Crotalus atrox Baird and Girard

LORICATA

Crocodylus moreletii Dumeril and Bibron

* First state record.

** Change in nomenclature of a previously reported form.

*** Recently described forms.

1590 The University Science Bulletin

REPORT ON SPECIMENS

Caudata
A single salamander is present in the collection.

Ambystoma velasci Duges

Amblystoma velasci Duges, La Naturaleza, ser. 2, vol. 1, 1888, p. 142 (type
locality, Laguna Santa Isabel, near Guadalupe Hidalgo, D. F. Mexico).

One specimen, No. 4294, was taken at Presa Gonzales Santos,
San Luis Potosi. The specimen is somewhat hardened so that
accurate comparison with other forms is somewhat difficult. I have
referred the species to velasci with some hesitation. The specimen
is black with a pair of small yellow spots on the beginning of the
neck and another pair near the angles of the mouth. The sides of
the neck and body together have six to eight small canary-yellow
spots, irregularly placed. The tail is strongly compressed with a
series of five or six spots on each side on the upper part of the tail
and a few scattered spots about tail-base. Some indistinct light
flecks are present on the limbs. The venter is blackish with whitish
or yellowish markings, the underside of the limbs dark The
vomerine teeth are in two, somewhat diagonal series meeting
mesially and anterior to the level of the choanae, forming a broad
chevron. About 25 teeth are present on the left side, and 22 on the
right, where there is a short hiatus near the choana. The tongue is
well developed. The phalangeal formula is: 2, 2, 3, 2; 2, 2, 3, 4, 2.
The webbing on the hand does not extend to the ends of the meta-
carpals. On the foot the webbing extends slightly beyond the
metacarpals and on each digit continues as a diminishing fringe to
near the end of the digits. Snout to front of vent, 70 mm.; tail,
54 mm.; basal depth of tail, 8 mm.; arm, 25 mm.; leg, 27 mm.

Salientia
The collection of Salientia contains 239 specimens, and 16 species.

Scaphiopus couchii Baird

Scaphiopus couchii Baird, Proc. Acad. Nat. Sci. Philadelphia, vol. 7, 1854,
p. 62 (type locality, Matamoros, Tamaulipas [restricted]).

The following specimens are in the collection: Nos. 5464, 5465
from Ebano; No. 4980, from 12 mi. W of Ebano, 100 ft. elev.; No.
4289, from 11 mi. W of Ebano; Nos. 4298, 4299, recently transformed
specimens, that seemingly belong to this species.

Herpetology of San Luis PoTOsi 1591

Scaphiopus multiplicatus Cope

S(caphiopus) multiplicatus Cope, Proc. Acad. Nat. Sci. Philadelphia, vol. 15,
1863, p. 52 (type locality, Coyoacan, D. F. Mexico, [restricted]).

A series of the northern variant of this species (Nos. 5458-5463,
5789 ) was taken at Luna Media.

Btifo occidentalis Camerano

Bufo occidentalis Camerano, Atti Accad. Sci. Torino, vol. 14, 1879, p. 887 (type
locality, Guanajuato, Guanajuato [restricted]).

One adult specimen, No. 4982, from Pozo del Carmen, 6500 ft.
elev., is in the collection. The tympanum is distinct.

This species was formerly reported from San Luis Potosi, as Bufo
simus.

Bufo valliceps Wiegmann

Bufo valliceps Wiegmann, Isis von Oken, vol. 26, 1833, pp. 657-659 (type lo-
cality, Veracruz, Veracruz, [restricted] ) .

The following numerous specimens of this species are in the col-
lection: Nos. 4315-4318, 5432-5437, Tamazunchale; 4319-4328, 4990-
4991, 5438-5441, Nacimiento del Coy; 4329-4332, Rio Valles, at
Valles; 4985-4986, Aqua Sonadora, 4 km. N Tanchachin; 4987, 2 mi.
N Valles, 300 ft. elev.; 4988, 5455, Valles, 300 ft.; 4995-4996, 6 mi. W
Ahuacatlan; 5442-5450, Ebano; 5414-5431, El Salto; 5457, Rio Axtla
(ferry on Xilitla Road); 5451, Luna Media; 5456, 38 km. SW Rio
Verde.

In a previous publication * I noted certain toads that differed
from typical valliceps in having a narrower head. A specimen,
No. 4996, from 6 mi. W of Ahuacatlan, likewise differs from the
typical valliceps. This specimen is a large male. The head crests
are thicker and the dark parotoid gland extends farther, reaching
laterally to a level of the middle of the tympanum. The hands and
feet are broader and the toes are slightly more webbed. Seem-
ingly a smaller part of the femur is included in body skin. The
vocal sacs open on both sides.

Bufo horribilis Wiegmann

Bufo horribilis Wiegmann, Isis von Oken, vol. 26, 1833, pp. 654-655 (type lo-
cality, Veracruz, Veracruz, [restricted] ) .

Nos. 4333, 4994, and 4998 are from El Nacimiento del Coy, 300
ft. elev.; Nos. 4334-4347, 4350, 4351, 4353, 4356, from Rio Valles,
at Valles; Nos. 4348, 4352, 4354, 4355, from El Sol Courts, Tamazun-
chale; No. 4357, from El Salto. Males may usually be distinguished
from females by their rougher skin.

* Univ. Kansas Sci. Bull., vol. 34, pt. 2 Feb. 15, 1952.

1592 The University Science Bulletin

Bnfo punctatus Baird and Girard

Bufo punctatus Baird and Girard, Proc. Acad. Nat. Sci. Philadelphia, vol. 6,
1852, p. 173 (type locality, Rio San Pedro [now Devil's River], Val Verde
Co., Texas).

Six specimens are in the collection: Nos. 4311-4314, from Presa
San Jose, 4 mi. SW of San Luis Potosi; Nos. 4290-4291, from 5 miles
SW Bledos, E. Cabrera, coll.; No. 4981, Pozo del Carmen, 5600 ft.
elev., J. Graber, coll.; No. 4983, Sierra San Miguel, 7500 ft. elev.,
R. Graber, the specimen "gray with red-brown warts"; No. 4984, 2
mi. W of Amoles, 5600 ft., R. Graber, coll.

Leptodactylus labialis (Cope)

Cystignathus labialis Cope, Proc. Amer. Philos. Soc, vol. 17, 1877, p. 90 (type
locality, Potrero Viejo, Veracruz [restricted]).

A small, recently transformed specimen (No. 4961), from El
Nacimiento del Coy, 300 ft. elev., collected by Jean Graber, seem-
ingly belongs to this species although too young to make positive
identification. The vomerine teeth are well developed. The color
is "dark gray with darker markings."

Syrrhophus cystignathoides (Cope)

Phyllohates cystignathoides Cope, Proc. Amer. Philos. Soc, vol. 17, 1877,
pp. 89-90 (type locality Potrero, near Cordoba; Veracruz, Mexico).

Two specimens of this species in the collection are from La Joya,

5 km. W of Aquismon, elev. 2500 ft., Nov. 22, collected by R. Graber.
No. 4962 is "olive with black spots; black band through eye and on
side of head." No. 4963 is "gray with darker spots; black on sides
of head and around eye."

Tomodactylus macrotympanum Taylor

Tomodactylus macrotympanum Taylor, Univ. Kansas Sci. Bull., vol. 26, 1939
(1940) pp. 496-499, pi. 55, figs. 2, 2a, 2b, (type locality, La Placita, south
or Jacala, Hidalgo, Mexico).

Three specimens, Nos. 4968, 4969, and 4970 were taken at a point

6 miles west of Ahuacatlan, at an elevation of 5200 ft. by R. Graber,
July 30 and 31, 1951. "Sits in bushes about three to four feet above
the ground." "Gray, almost transparent, with darker spots."

Smilisca haudinii (Dumeril and Bibron)

Hyla haudinii Dumeril and Bibron, Erpetologie generale, vol. 8, 1941, pp. 564-
565 (type locality, Cordoba, Veracruz [restricted]).

Three specimens are in the collection: Nos. 4274, El Nacimiento
del Coy, L. Guerrero, collector; Nos. 4300, 4301, El Sol Courts,
Tamazunchale, R. T. Gregg, collector.

Herpetology of San Luis Potosi 1593

Acrodytes spilomma ( Cope )

Hyla spilomma Cope, Proc. Amer. Philos. Soc, vol. 17, 1877, p. 86 (type lo-
cality, Cosomaloapam, Veracruz, Mexico).

A specimen, No. 5786, is from El Salto. The spotting on the limbs
and sides of the body is typically nigropunctate.

Hijla arenicolor Cope

Hyla arenicolor Cope, Jour. Acad. Nat. Sci. Philadelphia, ser. 2, vol. 6, 1866,
p. 84 ( substitute name for Hyla affinis Baird preoccupied; type locality,
Santa Rita Mts. [restricted]).

The northeastern specimens are on the whole smaller than speci-
mens from the southern part of the range. Nos. 4966, 4967 are
from 12 mi. E of Santiago, 8200 ft. elev., taken Sept. 24, 1951; Nos.
4974, 4975 are from Sierra San Miguel, 7500 ft. elev., taken Oct. 20,
1951; and No. 5785 is from 38 km. SW of Rio Verde de los Pinos.

Hyla eximia Baird

Hyla eximia Baird, Proc. Acad. Nat. Sci. Philadelphia, vol. 7, 1854, p. 61 (type
locality, Coyoacan, Distrito Federal, Mexico [restricted]).

Four specimens in the collection are from the following localities:
No. 4964, Bledos, 6200 ft. elev.; Nos. 4965, 4971, 4972, are from 10
mi. E Pozo del Carmen, 6500 ft. elev. The adults are typical.

Hyla miotympanum Cope

Hyla miotympanum Cope, Proc. Acad. Nat. Sci. Philadelphia, vol. 15, 1863,
p. 49 (type locality, Jalapa, Veracruz [restricted]).

Two specimens, Nos. 5780, 5781 from Xilitla, are in the collection.

Rana pipiens Schreber

Rana pipiens Schreber, Der Naturforscher, Halle, vol. 18, 1872, p. 185, pi. 4
(Raccoon, Gloucester Co., N. J.).

The following specimens are in the collection: Nos. 4278-4284,
Bledos; Nos. 4286, 5 mi. SW Bledos; No. 4288, El Nacimiento del
Coy; No. 4960, Agua Sonadoro; No. 4976, Villa Reyes, Laguna de la
Rusias; Nos. 4977-4979, Pozo del Carmen, 5600 ft. elev. No attempt
is made to determine the subspecific relationship of these specimens.

Rana montezumae Baird

Rana montezumae Baird, Proc. Acad. Nat. Sci. Philadelphia, vol. 7, 1854, p. 61
(type locality, Mexico, Distrito Federal, Mexico).

The following specimens are in the collection: Nos. 4302-4306,
Presa Gonzales Santos, 10 mi. SW San Luis Potosi; 4307-4310, 4390,
Presa San Jose, 4 mi. SW San Luis Potosi; 4275-4277, Bledos; 4285,

1594 The University Science Bulletin

4287, 5 mi. SW Bledos in pools in mountain canyon, 7000 ft. elev.;
4973, Sierra San Miguel, 7500 ft.; 5654, Laguna de las Rusias; 5668,
5699-5671 Presa Prudentia.

Sauria

The lizard collection contains considerably more than three hun-
dred specimens, but all are not recorded here since certain ones
are being studied by Dr. Hobart M. Smith. His report will appear
elsewhere. There are 21 species represented here.

Hemidactylus turcicus turcicus ( Linnaeus )

Lacerta turcicus Linnaeus, Systema Naturae, ed. 10, 1758, p. 202 (type lo-
cality, Cairo, Egypt [restricted]).

The presence of this species in the State is attested by a series of
specimens from Tamazunchale, collected by Fugler (Nos. 5798-5806).
They were captured in the El Sol Courts and suggest a recent in-
troduction possib'y by tourists. Previous collecting in Tama-
zunchale has yielded no specimens. Specimens were previously
known from 12 mi. E of Llera in Tamaulipas. These are the first
records for San Luis Potosi.

Anolis sallaei Giinther

Anolis sallaei Giinther, Proc. Zool. Soc. London, 1859, p. 421 (type locality,
Jalapa, Veracruz, Mexico [restricted]).

This species was formerly reported from the State under the
name Anolis sericens (Hallowell) which I now believe refers to a
different species.* The present collection contains two specimens,
Nos. 4209 and 5S06, from Nacimiento del Coy. The latter is a
female, having a white throat, which bears a purplish-red spot. The
body is generally gray and lacks the broad, dorsal, cream stripe
that is present in the type ( also a female ) .

Phrynosoma orbicalare orbiculare (Linnaeus)

(Plate CXVIII)

Lacerta orbicularis Linnaeus, Systema Naturae, ed. 12, 1789, p. 1062 ( part. )
(type locality, Mexico, D. F. Mexico [restricted]).

A specimen, No. 4944, was taken on the Sierra San Miguel, at an
elevation of 7200 ft., Oct. 21, 1951, by R. Graber. The color is:
"Dorsally brown with dark brown and cream colored markings;
spines olive green; occipital spines pink. Ventrally blotched with
blue-gray; legs yellowish."

* Taylor, Univ. Kansas Sci. Bull., vol. 34, pt. 2, Feb. 15, 1952, p. 805.

Herpetology of San Luis Potosi 1595

Plate CXVIII. Phrynosoma orbiculare orbiculare L.S.U. No. 4944. Sierra

San Miguel, S.L.P.

1596 The University Science Bulletin

Holbrookia maculata approximans Baird

Holbrookia approximans Baird, Proc. Acad. Nat. Sci. Philadelphia, 1858, p. 253
type locality, Tamaulipas, Tamaulipas, Mexico [restricted]).

A mutilated female specimen, No. 5796, from Presa Gonzales
Santos, is referred to this species.

Sceloporus parvus parvus Smith

Sceloporus parvus Smith, Trans. Kansas Acad. Sci., vol. 37, 1934, pp. 263-267,
pi. 8, figs. 1, 3, pi. 10, fig. 10 (type locality, 5 mi. W Sabinas Hidalgo, Nuevo
Leon, Mexico).

Six specimens are referred to the typical subspecies: Nos. 4924,
? mi. W Amoles; 4925, 4926, Sierra San Miguel, 7500-8000 ft.
elev.; 4927, 12 mi. SE Santiago, 8000 ft.; 5628, 5629, approximately
1 mi. W Ciudad del Maiz.

These specimens have 14 or 15 femoral pores on each side, the
series separated mesially by four scales. The scales in a dorsal row
from the occipital to above vent are 63-68 and the enlarged dorsal
scale rows on neck are 17 or 18. Since the specimens are discolored
by formalin, the dorsolateral light line is scarcely distinguishable.
The chin has light spots on a blackish ground color.

An area under the thighs in the anal region and under the tail is
whitish and without pigment. In younger specimens the tail is
white to the tip on the under side while the dorsal and lateral
coloring of the tail is gray, much lighter than the dorsal body color.

Sceloporus grammicus disparilis Stejneger

Sceloporus disparilis Stejneger, Proc. Biol. Soc. Washington, vol. 29, 1916,
pp. 227-230 (type locality, Lomita Ranch 6 mi. N Hidalgo, Texas).

A considerable series of this species is present in the collection.
The following localities are represented: Nos. 4248-4254, Cerro
Conejo; 4242," Canada Grande; 4227, Bledos; 4912, 4915-4916, 6 mi.
W Ahuacatlan, 5400-6500 ft. elev.; 4917, 4918, 4920-4922, Sierra
Azul, 10 mi. NE Bledos, 900 ft. elev.

Sceloporus spinosus spinosus Wiegmann

Sceloporus spinosus Wiegmann, Isis von Oken, vol. 21, 1828, p. 370 (type lo-
cality, Puebla, Puebla [restricted]).

The following specimens are in the collection: Nos. 4260-4262,
4365, 4934, 4935, Laguna de las Rusias; 4237-4240, Bledos; 4241,
5 mi. SW Bledos; 4242, Canada Grande; 4936, 10 mi. E of Pozo del
Carmen; 4937, 3.5 mi. W Pozo del Carmen; 4272 (young), Villar;
5490-5499, 5539, 5794, Presa Prudentia; 5500-5511, 5529, 5532-5537,
Luna Media; 5516-5521, 5523-5528, 5530, 5531, 5540, Presa Gonzales

Herpetology of San Luis Potosi 1597

Santos; 5515, 1% mi. W Leoncito; 5522, Presa San Jose; 5512-5514,
5538-5539, Ciudad del Maiz.

SceJoporus variabilis variabilis Wiegmann

Sceloporus variabilis Wiegmann, Herpetologia Mexicana, 1834, p. 51 (type
locality, Veracruz, Veracruz, Mexico [restricted]).

The following specimens are in the collection: Nos. 4236, Axtla;
4271, 2 mi. W Tamuin; 4257-4259, 5563-5573, 5600-5612, El Salto;
4267-4268, Platanito; 4931-4933, 3 mi. W Platanito, 4000 to 4600 ft.
elev.; 4269, Puente de Dios; 4264, 5590, Tamazunchale; 4243, 4244,
4246, Canada Grande; 4256, 10M mi. W Ebano; 4263-4266, 4361-
4363, 4928-4930, 5574-5585, 5616-5619, Nacimiento del Coy; 5553-
5562, 5587, 5596-5599, 3 mi. W Xilitla; 5588, 5589, 5613-5615, Luna
Media; 5591, 5620-5625, Valles; 5590, 5626, 5627, Cerro de la Paz;
5586, 5587, 5592-5595, approx. 24 mi. SW Rio Verde, Los Pifios.

Sceloporus jarrovi minor Cope

Sceloporus torquatus minor Cope, Proc. Amer. Philos. Soc, vol. 22, 1885, p. 402
(type locality, Valparaiso Mts. Zacatecas, Mexico [restricted]).

Specimens Nos. 4366, 4905, 4919, 4923 were collected 10 miles
northeast of Bledos in the Sierra Azul, 9000 ft. elev.; No. 4366 at
Presa San Jose, 4 mi. SW of San Luis Potosi.

Sceloporus dugesii intermedius Duges

Sceloporus intermedius Duges, La Naturaleza, vol. 4, 1877, pp. 29-34, pi. 1,
figs. 21-32 (type locality, La Noria, near Zamora, hda. P. Epifanio Jimenez
in Miehoacan).

A series of specimens taken in southern San Luis Potosi is referred
to the above species. These are: Nos. 4232, 4908, 4911, 4913, 4914,
6 mi. W Ahuacatlan, 5400 ft. elev.; 4906, Sierra Azul, 10 mi. N
Bledos, 9000 ft. elev.

The latter specimen, a male, has a white chin, throat, and venter;
the underside of the tail is a shade of orange.

Sceloporus serrifer cariniceps Martin

Sceloporus serrifer cariniceps Martin, Occ. Pap. Mus. Zool. Univ. Michigan,
no. 543, Oct. 22, 1952, pp. 1-7 (type locality, Rancho Pano Ayuctle, along
Rio Sabanas, 5 mi. NE of Gomez Farias, Tamaulipas, Mexico.

No. 5550, from 6 km. E of Tamazunchale, is a typical example of
this recently described species. It has been reported previously
from Ebano.

Sceloporus torquatus melanogaster Cope

Sceloporus torquatus melanogaster Cope, Proc. Amer. Philos. Soc, vol. 22, 1885,
pp. 400-401 (type locality, Tupataro, Guanajuato [restricted]).

The species is represented by Nos. 4231, 4234 from 6 mi. W
Ahuacatlan; 5541, 5548, Presa Prudentia; 5549, Presa San Jose.

1598 The University Science Bulletin

Eumeces lynxe lynxe Wiegmann

Eumeces lynxe Wiegmann, Herpetologia Mexicana, 1834, pp. 36-37 (type lo-
cality El Chico, Hidalgo).

The following specimens are in the collection: Nos. 4210, 4211,
4213, Cerro Conejo at an elevation of 7000 ft.; No. 4212, Canada
Grande; No. 4942, 6 mi. W Ahuacatlan, 5400 ft. elev.; J. Graber coll.
This latter specimen has "yeh"ow stripe on dorsum, the back bronze;
throat somewhat orange in color; sides black; tail bright blue; belly
bluish gray."

Leiolopisma caudaequinae (Smith)

Scincella caudaequina Smith, in Smith and Taylor, U. S. Nat. Mus. Bull. 199,

1950, p. 158 (type locality, Salto Cola de Caballo, 25 mi. S of Monterrey,
Nuevo Leon, Mexico).*

Leiolopisma caudaequinae Smith, Univ. Kansas Sci. Bull., vol. 34, pt. 1, Oct. 1,

1951, pp. 195-200 (complete description).

Four specimens are in the collection: Nos. 4940 and 4941 from
La Joya, 3.1 mi. W of Aquismon, 2500 ft. elev.; No. 4943, from 4 mi.
W of Pendencia, 4600 ft. elev.; No. 5793, 3 mi. W of Xilitla.

The scale rows around the middle of the body are respectively,
28, 30, 28, 30; the lamellae under the fourth toe are, 18, 19, 19, 19.
The last specimen is a female, distended with eggs, in which the
limbs touch but do not overlap. There are 69 dorsal scales in a row
from parietals to above vent. This species has been taken pre-
viously in the State from Naranjo.

Ameiva undulata podarga Smith and Laufe

Ameiva undulata podarga Smith and Laufe, Univ. of Kansas Sci. Bull., vol. 31,
pt. 1, May 1, 1946, pp. 40-43, figs. Id, 2a (type locality, 7 mi. W Victoria,
Tamaulipas ) .

The following are in the collection: Nos. 4218, Puente de Dios,
Rio Santa Maria; 4228, 5700, 5701, 5769, and 3 unnumbered speci-
mens, El Salto; 4358, 5714, 5715, 5746, 5764, 5765, Tamazunchale;
5768, Rio Elera, 3.7 mi. E Tamazunchale; 5717, Nacimiento del Coy.

Cnemidophorus sacki gularis Baird and Girard

Cnemidophorus gularis Baird and Girard, Proc. Acad. Nat. Sci. Philadelphia,
1852, p. 128 (type locality, mouth of Devil's Biver, Texas [restricted]).

Nos. 4945-4947, Pozo del Carmen, 5600 ft. elev.; 4948-4950, from
2 mi. W of Valles, 300 ft. elev.; 5693, 5694, 5719-5722, 5750-5758,
Luna Media; 5695, 5696, 5702-5704, 5723-5725, 5761, 5772, Ciudad
del Maiz; 5726-5731, 5760, and four unnumbered specimens El Salto;

* The data in the key serve as a description and prevent this name from being nomen
nudum in this publication.

Herpetology of San Luis Potosi 1599

5742-5744, 5774, 5775, Presa Prudentia; 5723, 5748, 5749, Valles;
5705-5708, 5732-5738, 5766, 5767, 23.75 mi. SE Rio Verde (Los
Pifios); 5712, 5713, Presa San Jose; 5697-5699, 5763, Laguna de las
Rusias; 5709-5711, and 2 specimens without numbers, Presa Gon-
zales Santos; 5747, 5771 Cerro de la Paz; 5718, 5739, 5740, 5745,
5762, one-half mi. W of Leoncito.

Ophisaurus sp.

A single specimen, No. 4886 J , captured seven miles south of
Valles, San Luis Potosi by C. O. Peterson, Nov. 15, 1951, represents
the first record for the State and one of two for Mexico. The nearest
point in the United States where the genus is known to occur is
Cameron and Hidalgo Counties in extreme southern Texas.

It was reported from Jalapa by Yarrow,* a record that Smith
and Taylor ** believe to be incorrect. However, the finding of a
specimen near Valles proves the presence of the genus in Mexico.

Diagnosis: Ground color gray, with a series of black marks form-
ing lines on each scale-row above the lateral groove, the lines vary-
ing in width, the widest and most complete being that of the fourth
from the groove, which is not wider than one half scale-row; no
pigmentation below groove, but a faint peppering on edges of
groove; venter pure white; ten ventral scale-rows; 14 rows between
the grooves; the scales around neck behind ear, 34; scales between
end of groove and ear, 17; upper labials, 12-12.

Description of species: Rostral wider than high, bordered by
four internasal scales lying between the first labials, the median
scale rather unequally divided; nasal small, divided, the posterior
section very narrow, scarcely more than a partial rim around nostril;
a well-defined supranasal; a pair of posterior internasals broadly
in contact, touching the upper postnasal and the supranasal later-
ally; frontonasal large, more or less rounded in front, with nearly
parallel sides; posterior border curving from each side to a short
mesial point, barely separated from a large frontal by a rather
large pair of prefrontals; frontal bordered by three of the five inner
supraoculars; five small outer supraoculars; six small superciliaries;
interparietal broadly in contact with the frontal, pentagonal, but
generally triangular in shape, narrowing posteriorly to a point, in
contact with the interoccipital; a pair of parietals border the inter-
parietal, but are separated by three scales from the supraoculars,

* Yarrow. U. S. Nat. Mus. Bull., 24, 1884, p. 46.

Smith and Taylor. U. S. Nat. Mus. Bull. 199, 1950, p. 194.

* *

71_6490

1600 The University Science Bulletin

and touch three lateral, temporal scales; upper labials, 11-11; lower
labials approximately nine; at least nine scales in the presub- and
postocular series. One postnasal, two or three canthals; loreals? *
a single postmental, separated from mental by inner submentals;
three large paired chinshields; an inner and an outer row of sub-
mentals.

Ear small, separated from the last labial by three scales, from the
beginning of the lateral groove, by 15 scales; scales in a dorsal row
from parietal to above vent, 114-116; 14 dorsal scale rows, the outer,
bordering lateral groove, more than half size of adjoining row; ten
ventral scales, bordering lateral groove more than half size of ad-
joining rows; vent bordered by at least 11 scales only slightly differ-
entiated; scales on venter, from last enlarged paired postmental to
preanals, 119.

Measurements: Width of head, 15 mm.; head length to ear, 23
mm.; snout to vent, 231 mm. (tail broken and regeneration begun).

Colors: Above and on sides as far as the lateral groove, grayish
to grayish white, each scale of a given row with equal-sized darker
areas which together form rows of dots or discontinuous lines; the
two median lines darker and more nearly continuous than adjoining
rows; on each side of these two rows, are two rows of grayish-black
dots; the next row on each side is the darkest and practically con-
tinuous; width of the black equaling more than half width of
scale-row, while borders of this and succeeding rows whiter than
those dorsally; next two rows broad but discontinuous, while the
outermost bordering lateral groove is smallest, the individual dots
most widely separated; just posterior to level of vent third and
fourth rows of each side unite to form only a single strong stripe on
each side; labials and temporals with black spots; a black spot on
interoccipital, and a few small blackish flecks elsewhere on head.
Below lateral groove, belly uniform, dull white.

Remarks: The genus Ophisaiirus has recently been re-examined
by Mr. Edward McKonkey who has a paper in press dealing with
the genus at this writing. He suggests that I refer to this specimen
as I have done, until his paper appears.

Gerrhonotus liocephalus infernalis Baird

Gerrhonotus infernalis Baird, Proc. Acad. Nat. Sci. Philadelphia, 1858, p. 255
(type locality, Devil's River, Texas).

One specimen of this species, No. 4958, was acquired at Birmania,
3 mi. S of Valles, 300 ft. elev., by C. A. Peterson, November 10, 1951.

* The front of the head has been injured and certain characters are obscured.

Herpetology of San Luis Potosi 1601

The specimen has the following characters: snout to vent, 137
mm., tail, 291 mm., partly regenerated; total length, 428 mm.; ad-
pressed limbs separated by 10-11 scales; four scales bordering rostral
across snout between first labials; a distinct axillary pit, the lateral
nuchal pit being almost obsolete; nine transverse markings of tan,
dark brown and whitish; white mark from eye to ear bordered by
black above; dark marks crossing the fine scales of the lateral groove.
This specimen differs from the typical infernalis in having two
labials and four other scales bordering the rostral, and the number
of scale rows approaches the number in loweryi.

Gerrhonotus liocephalus loweryi Tihen

Gerrhonotus liocephalus loweryi Tihen, Trans. Kansas Acad. Sci., vol. 51, 1948,
pp. 302-305 ( type locality, Xilitla, San Luis Potosi ) .

A series of five specimens, Nos. 4953-4957, are from six miles W
Ahuacatlan, elevation approx. 5400 ft., Oct. 4 to 6, 1951, R. Graber,
collector. The median body scale-counts are, 53, 55, 52, 54, 53
respectively. There are three scales bordering rostral across snout
between the first labials. No. 4954 has a snout-vent length of 192
mm., the tail, 338 mm. The caudal scales of this specimen number
161. The head is widened in the males. The legs when adpressed
are separated by 2-3 scales in males; by 4-4/2 in females.

Abronia taeniata taeniata ( Wiegmann)

(Plate CXIX)

Gerrhonotus taeniatus Wiegmann, Isis von Oken, vol. 21, 1828, p. 379 (type
locality El Chico, Hidalgo [restricted] ).

A single specimen, No. 4208, is from Llano Conejo on Cerro
Conejo, collected by E. Esquibel. The specimen is somewhat dis-
colored but when compared with the Wiegmannian figure of the
type * it agrees very well. However, this specimen has somewhat
less light marking on the head and labials. This is a new record
for San Luis Potosi.

Barisia imbricata ciliaris ( Smith )

Gerrhonotus levicollis ciliaris Smith, Proc. U. S. Nat. Mus., vol. 92, 1942,
pp. 365-367 (type locality, Sierra Guadelupe, Coahuila, Mexico).

One specimen, No. 4207, 7000 feet, Cerro Conejo, measures 138
mm. snout to vent. The tail is regenerated. The color of the pre-
served specimen is nearly uniform olive above without markings,
the head, especially the sides of head, somewhat lighter.

Two specimens, Nos. 4951, 4952, are from 10 mi. NE of Bledos

* Wiegmann, Herpetologia Mexicana, pi. 9.

1602

The University Science Bulletin

Plate CXIX. Abronia taeniata taeniata L. S. U. No.
4208 Cerro Conejo S. L. P.

Herpetology of San Luis Potosi 1603

in the Sierra Azul, 9000 ft. elev., collectors, Jean Graber and R.
Graber. The former is "dorsally brown-olive drab, ventrally pale
orange; especially orange on under side of tail." The latter speci-
men is "olive-drab dorsally; whitish green ventrally except for tail,
which is orange on under surface."

These two specimens differ somewhat from the Cerro Conejo
specimen. The heads are larger, the bodies shorter. No. 4207 has
the adpressed limbs separated by ten scales, the other two by five
scales.

Serpentes

The collection contains 68 specimens of snakes. For the size of
the collection, the number of species is large, no less than 31 species
and subspecies being represented. Three are the first records for
the State.

Leptotyphlops myopicus myopicus (Garman)

Stenostoma mijopicum Garman, Mem. Mus. Comp. Zool., vol. 8, 1883, pp. 6,
130, 131 (type locality, Savineto, near Tampico, Tamaulipas, Mexico).

Three specimens of this species are in the collection: Nos. 5411,
Nacimiento del Coy; 5412, Cerro de la Paz; 5413, Luna Media,
Fugler collector.

Constrictor constrictor imperator (Daudin)

Boa imperator Daudin, Histoire Naturelle . . . reptiles, vol. 5, 1803,
pp. 150-152 (type locality, Cordoba, Veracruz, Mexico [restricted]).

A skin containing a complete head (No. 4871) is from near Cor-
rones, 15 mi. W of Ebano, altitude 100 ft., R. Graber, collector.

Ficimia streckeri Taylor

Ficimia streckeri Taylor, Copeia, 1931, no. 1, pp. 5-7 (type locality, 3 mi. E
Rio Grande City, Texas).

A single specimen ( No. 5391 ) is from 3 mi. W of Xilitla.
Geophis mutitorques * Cope

Geophis mutitorques Cope, Proc. Amer. Philos. Soc, vol. 22, 1885, p. 384 (type
locality, Zacualtipan, Hidalgo, Mexico).

A small specimen, No. 4202, from Cerro Conejo (7700 ft.), is in
the collection. It is black above, while below it is black with more
or less quadrangular white spots, alternating or more rarely fused.
There is a nuchal collar, almost interrupted medially, widening
laterally and connecting with the large whitish area on chin.

* In Taylor, Univ. Kan. Sci. Bull., vol. 33, pt. 1, Apr. 20, 1949, pp. 171, 194, this
specific name is incorrectly spelled maltitorques.

1604 The University Science Bulletin

Lampropeltis triangulum polyzona Cope

Lampwpeltis polyzona Cope, Proc. Acad. Nat. Sci. Philadelphia, vol. 12, 1860,
p. 258, ( type locality, Cuatupe, near Jalapa, Veracruz ) .

Two specimens, No. 4370, $ , from El Sol Courts at Tamazun-
chale, and No. 4203, $ , from 11.2 km. S of Valles, are in the col-
lection. The black-white-black bands, 18 or 19 in number, are not
or scarcely wider dorsally than laterally in No. 4203, while in the
smaller (No. 4370), the bands are distinctly wider dorsally than
laterally. The lengths of the two specimens are respectively, 339
mm. and 687 mm., tails, 48 mm., 88 mm.

Rhadinaea crassa Smith

Rhadinaea crassa Smith, Proc. Biol. Soc. Washington, vol. 55, 1942, pp. 190-
191, figs. 4-5 (type locality, Durango, Hidalgo).

A specimen (No. 4206, R. Graber, collector), typical in practi-
cally all details, is from Cerro Conejo.

Rliinocheihis lecontei tessellatus Garman

Rhinocheilus lecontei tessellatus Garman, Mem. Mus. Comp. Zool., vol. 8,
1883, p. 74, 159 (type locality, Monclova, Coahuila, Mexico).

The first State record for this species is No. 4875, from 3.5 mi. W
of Pozo del Carmen, 5600 ft. elev. — the collector, Emilio Esquibel.

The following characters obtain: snout turned up and elevated
above other snout scales; internasals angular, nearly three fourths
the size of prefrontals; part of rostral seen from above, equal in
area to an internasal; one preocular, two postoculars; upper labials,
8-8; lower labials, 8-9; temporals, 2 + 3; first pair of chinshields
longer and wider than posterior pair (on one side the second right
chinshield is broken transversely); second pair of chinshields sepa-
rated by two scales anteriorly, by four scales posteriorly; ventrals,
194M; anal single; subcaudals, 49 + 1 ( 1st divided; 2nd to 43rd
single, 44th divided, 45th single, 46th to 49th divided, last terminal
scale single); scale formula: 23, 23, 23, 19, 19; scales with a single
apical pit. Total length, 769 mm.; tail, 92 mm. Above, black, red
and yellow; 25 broad, black, dorsal blotches, separated by rosy red;
9 black spots on tail separated by rosy red; ventral surfaces pale
yellow; every third or fourth ventral with black spots on outer edges.

Salvadora lineata Schmidt

Salvadora lineata Schmidt, Publ. Field Mus. Nat. Hist., Zool. Ser., vol. 24,
1940, pp. 148-150, fig. 15 (type locality Kingsville, Kleburg County, Texas).

Two specimens are in the collection (No. 4197, Ventilla, May 16,
1951, and No. 4880, 12 mi. SE of Santiago, Sept. 24, 1951, R. Graber,

Herpetology of San Luis Potosi 1605

collector). The former has 194 ventrals and 88 subcaudals; the
latter has 192 ventrals; the tip of the tail is missing; the scale formula
of the latter is: 19(20), 17, 17, 13(15). The color is: "Ventrally
pale yellow-white; dorsally, median stripe yellow; lateral stripes
brown; sides gray-brown." The total length is 841 mm., the tail,
217 mm.
A third specimen, No. 5390, is from 1.5 mi. W Ciudad del Maiz.

Elaplie triaspis intermedia (Boettger)*

Pityophis intermedins Boettger, Ber. Offenbacher Ver. f. Naturk. Bd. 22, 1883,
pp. 147-152 (type locality, Mexico).

Coluber chlorosoma Giinthcr, Biologia Centrali-Americana, Beptilia and Ba-
trachia, 1894, pp. 115-116, pi. 41 (type locality San Ramon, Jalisco [re-
stricted] ).

A specimen, No. 5387 from 3 mi. W of Xilitla was taken June 18,
1952.

Elaplie flavirufa flavirufa (Cope)

Coluber flavirufus Cope, Proc. Acad. Nat. Sci. Philadelphia, vol. 18, 1866
(1867) p. 319 (type locality, Campcche, Campeche, Mexico [restricted]).

A specimen, No. 4874, from Rancho Sabanal, collected by Jean
Graber, Nov. 12, 1951, has the following characters: one very large
preocular, reaching frontal; two postoculars; temporals, 3 + 4; up-
per labials, 9-9; lower labials, 12-13; ventrals, 253; subcaudals,
109 + 1; scale formula: 25, 27, 29, 23, 19. "Tan above with red-
brown, black-edged spots." Head scales more or less outlined in
black. A second specimen, No. 5381, is larger than the preceding
and darker. It was taken at Nacimiento del Coy.

Masticophis flagcUum testaceus (Say)

Coluber testaceus Say, in Long's Expedition to the Rocky Mountains, vol. 2,
1823, p. 48 (type locality, Pueblo, Pueblo County, Colorado [restricted]).

Two specimens, one a young, road-killed specimen, No. 5797,
from Cerro de la Paz and a larger specimen ( road-killed ) , No. 5393,
from 15 mi. W of Ebano, are referred to this species.

Masticophis taeniatus ruthveni Ortenburger

Masticophis ruthveni Ortenburger, Occ. Papers, Mus. Zool. Univ. Michigan,
No. 139, 1923, pp. 3-8, pis. 1-3, (part) (type locality, Brownsville, Texas).

In specimen No. 4204, from Bledos, San Luis Potosi, J. Graber,
collector, there are seven upper labials, seemingly the second rep-
resenting a fusion of the normal second and third. There are two
preoculars, two postoculars, and eight lower labials present. The

* For use of this name fide Dowling, Occ. P. Mus. Zool. U. Mich. no. 541, Oct. 10, 1951.
p. 8.

1606 The University Science Bulletin

anterior part of the preocular is partially divided. The scale for-
mula is, 15, 15, 15 (14), 13; the ventrals, 199; the subcaudals,
137 + 1; and the anal divided. The specimen has a total length of
1300 mm., the tail, 290 mm.

A second specimen, No. 5396 from 38 km. southwest of Rio Verde,
shows the dorsal scales with white edges throughout the anterior
half of the body. The outer fourth of each ventral is actually lateral
and colored like the sides but it is separated from the other lateral
color by narrow white dashes, distinct and forming a white line
on the anterior fourth of the body but becoming obsolescent or ob-
solete more posteriorly. The ventral area of the ventrals is heavily
peppered with gray pigment. The ventrals are 199, the subcaudals,
99 + 1, the anal divided.

Pittiophis catenifer affinis Hallowell

Pityopkis affinis Hallowell, Proc. Acad. Nat. Sci. Philadelphia, 1852, p. 181
(type locality [restricted] Zuni, New Mexico).

One specimen ( No. 4369, $ , 6 mi. E of El Huisacha ) , is present
in the collection. The head is rather reddish tan without markings.
The dark blotches are not of solid color and the intervening lighter
spaces have almost every scale marked with black. Posteriorly on
body the darker markings become brown while the intervening
scales are lighter brown. On the extreme posterior part of the body
and tail the blotches are brownish red, the scales flecked with black
while the intervening scales are immaculate with a slight pinkish
edging. There are 23 dark blotches on body and ten on tail.

Some short discontinuous lines are more or less evident on the
scales of the anterior part of the body. The ventrals number 235,
the subcaudals 56 -f- 1, while the scale formula is, 29, 25, 22. The
total length is 1800 mm., the tail, 193 mm.

Pituophis deppei jani (Cope)

Arizona jani Cope, Proc. Acad. Nat. Sci. Philadelphia, vol. 12. 1860 (1861),
p. 369 (type locality, Buena Vista, Coahuila, Mexico).

A specimen (No. 4876 from Amoles, San Luis Potosi, 5600 ft.
elev., R. Graber, collector) has the head a light, reddish tan, the
anterior half of the body, tan with black spots. The posterior half
of the body is gray with red-brown spots. The ventrals are 220,
the subcaudals 59+1. There are 24 dorsal body blotches, with
seven on the tail.

No. 5383, 2 3 Ciudad del Maiz, has the head light fawn, the light
intervening blotches usually less than five scales long. The middle

Herpetology of San Luis Potosi 1607

and terminal dark blotches are reddish brown, those on the posterior
parts being edged with black. There are 30 dark body blotches.

No. 5382, g , 3 mi. north of Bledos, has the light anterior bands
five scales long or longer, the median dark body spots red-tan be-
coming black posteriorly. There are 33 body blotches.

No. 5384 is a young specimen from Presa Prudentia. It seem-
ingly agrees with the preceding specimen, save that there are 31
body blotches and certain differences due to youth.

It seems quite probable that there is some intergradation of char-
acters of the two subspecies, deppei deppei and deppei jani since
they show characters of both forms as delineated by Stull.*

Drymarchon corais erebennus (Cope)

Spilotcs erebennus Cope, Proc. Acad. Nat. Sci. Philadelphia, vol. 12, I860,
p. 342 (type locality, Eagle Pass, Texas).

One specimen, No. 5392, is from Xilitla, collected by Fugler.

Dryadophis melanolomus veraecrucis (Stuart)

Eudryas boddaerti mexicanus Stuart, Occ. Papers Mus. Zool. Univ. Michigan,
no. 254, 1933, pp. 8-9 (type locality, Zacuapan, Veracruz).

Two typical specimens, Nos. 5394 from 3 miles W of Xilitla, and
5395 from El Salto, are in the collection.

Spilotes pullatus mexicanus (Laurenti)

Spilotes mexicanus Laurenti, Specimen medicum exhibens synopsin reptilium,
1768, p. 83 (Mirador, Veracruz, [restricted]).

A single specimen, No. 4191, is from El Nacimiento del Coy,
L. Guerrero, collector. The ventrals are 205, the anal single, the
subcaudals 130 -4- 1- There are 8-8 supralabials, 8-9 infralabials
with the fifth and seventh labials touching above the sixth. The
scale formula is: 18(17), 18, 18, 12. The total length is 1900 mm.,
the tail, 730 mm.

Thalerophis mexicanus mexicanus (Dumeril, Bibron, and Dumeril)

Leptophis mexicanus Dumeril, Bibron, and Dumeril, vol. 7, pt. 1, 1854, pp. 536-
537 (type locality, Potrero Viejo, Veracruz, Mexico [restricted]).

A specimen, No. 3200, g , taken at Nacimiento del Coy, has 166/2
ventrals, the anal divided and the subcaudals, 88 -f- 1. The scale
formula is: 15, 11, 11, the scales having single terminal pits.

A second specimen, No. 5808, was taken 1 mile W of San Felipe,
July 27, 1952.

* U. S. Nat. Mus. Bull., 175, 1940, pp. 25-47.

1608 The University Science Bulletin

Thamnophis cyrtopsis cyrtopsis (Kennicott)

Eutaenia cyrtopsis Kennicott, Proc. Acad. Nat. Sci. Philadelphia, vol. 12, 1860,
pp. 333-334 (type locality, Rinconada, Coahuila); Smith, Copeia, 1951,
No. 2, June 8, p. 138-140.

No. 4376 is from Presa Gonzalo Santos, 10 mi. W of San Luis
Potosi, R. T. Gregg, collector, and No. 4878, is from 4.5 mi. N of
Jesus Maria, 5900 feet elevation, R. Graber, collector. In this latter
specimen, the ventrals are 171, the subcaudals 75 -f- 1. The scale
formula is: 22, 21, 21, 19, 19. The coloration of this species is:
"Head blue-gray, the median stripe orange-yellow, lateral stripe
white. Dorsally olive-drab with yellow spots; gray with black spots
ventrolaterally; gray-white ventrally."

This species was formerly recognized under the name Tham-
nophis eques eques (Reuss).

Thamnophis eques eques (Reuss)

Coluber eques Reuss, Abh. Senck. Mus., 1834, pp. 152-155, pi. 8, fig. 2 (type

locality, El Limon, Totalco, Veracruz [restricted]); Smith, Copeia, 1951,

no. 2, June 8, pp. 138-140, pi. 1.
Thamnophis subcarinata subcarinata Smith, Herpetologica, vol. 5, 1949, p. 64

(type locality, Guadalajara, Jalisco).
ThamnojiJiis macrostemma macrostemma Smith and Taylor, U. S. Nat. Mus.

Bull., 187, 1945, p. 163.

The unfortunate mix-up in names used for this species has been
the result of a misinterpretation of the identity of the snake de-
scribed by Reuss. Many authors have confused it with the snake
here called Thamnophis cyrtopsis cyrtopsis (Kennicott).

The following specimens are in the collection: Nos. 4196, 4374,
4375, 5403, 5406, Laguna de las Rusias; No. 4870, 4 miles west of
Bledos (young); No. 4879, Bledos, 6200 ft. elev.; No. 5408, Presa
Prudentia.

The underside of the tail is much lighter than the ventral colora-
tion of belly, having in life a distinctly pink color.

Thamnophis melanogaster canescens Smith

Thamnophis melanogaster canescens Smith, Zoologiea, vol. 27, 1942, pp. 117-
120 (type locality, Chapala, Laguna de Chapala, Jalisco, Mexico).

The following specimens are in the collection: Nos. 4377, 5397,
5402, 5404, 5405, Laguna de las Rusias; No. 4378, Presa de San Jose;
No. 4379, Presa Gonzales Santos, 10 mi. SW of San Luis Potosi;
No. 4877, Villa de Reyes, Laguna de las Rusias, 6000 ft. elev.; Nos.
5407, 5409, Presa Prudentia.

In No. 4377, $ , small whitish dots or dashes are evident in rows
on the upper edges of the third and fourth rows of scales. These

Herpetology of San Luis Potosi 1609

may be largely on the skin between the scales, but a portion of the
white covers the edge of the scale. On the sixth and seventh rows,
small lighter dots are evident when the skin is stretched. The
stomach of this specimen contained 12 small fish. The scale-row
formula is 23, 17, 17. The ventrals are 149, the subcaudals 61. The
specimen has a total length of 828 mm., the tail, 144 mm.

Nos. 4378 and 4379 are young. Each has a well-defined light line
on the second scale-row. The first scale-row is light with two black
spots on each scale. In these the loreal enters the eye below the
single preocular.

Thamnophis plienax Cope

Eutaenia phenax Cope, Proc. Acad. Nat. Sei. Philadelphia, vol. 20, 1868, p. 134
(type locality, Cordoba, Veracruz).

A specimen of this species (No. 4198^ ), was taken on Cerro
Conejo, June 16, 1951. The specimen, while much discolored by
formalin, shows clearly the black edges on the 50 dorsal blotches
separated by a narrow lighter area one scale-length wide. The
ventrals are 163, the subcaudals 74 + 1. There were but 36 blotches
in the type of the species.

Thamnophis sirtalis proximns (Say)

Coluber proximns Say, in Long's Expedition to the Rocky Mountains, vol. 1,
1823, p. 187 (type locality, 3 mi. above mouth of Boyer's River, Washington
County, Nebraska).

A specimen, No. 4205, was taken near Valles. The median dorsal
stripe is obsolete. The scale rows are 19, 19, 17, the ventrals 152M
and the subcaudals 92. These counts place the specimen very close
to the subspecific form chalceus and a series might demonstrate that
the two forms actually intergrade in eastern San Luis Potosi.

Tha?nnophis halophilus Taylor

Thamnophis halophilus Taylor, Herpetologica, vol. 1, 1940, pp. 183-187, pi. 19,
text fig. 1 (type locality, seven mi. N of Zacualtipan, Hidalgo).

A specimen, No. 4199, from Cerro Conejo, Llano Chico, 7600
ft., is referred to this species.

A series of lateral spots, and a series of median spots are present.
Farther back, at the beginning of the second fourth of the body,
are two paired series of spots on each side. Sometimes those of the
upper series are contiguous or fused (more frequently not). The
skin between spots is white in life. The chin is light and there are
light areas on the labials below the eyes. Ventrals number 158, the
subcaudals 65. The scale-row formula is: 23, 19, 19, 19, 17. One

1610 The University Science Bulletin

preocular and three postoculars are present. There are 8-8 supra-
labials, and 9-9 infralabials.

The finding of Thamnophis phenax Cope and Thamnophis halo-
philus on the Cerro Conejo suggests that these forms do not bear
subspecific relationship to each other as is presumed in the Smith-
Taylor Check^st of Snakes. Consequently I revert to the binomial
forms for these names.

Leptodeira annulata septentrionalis (Kennicott)

Dipsas septentrionalis Kennicott, in Baird, Rep. U. S. Mex. Bound. Surv., vol. 2,
1859, R?pt., p. 16, pi. 8, fig. 1 (type locality, Brownsville, Texas [re-
stricted ] ) .

A specimen, No. 5380, from Luna Media, has 29 quadrangular
bands covering some scales of the first row. There are 15 tail bands.
Anteriorly the ventral pigmentation is confined to the outer edges
of the ventrals, but on the posterior fourth of the body it pushes
more toward the middle of the scales. The subcaudals are sprinkled
rather thickly with gray pigment. The light bands between black
bands dorsally, rarely are as wide as two whole scale lengths. These
bands widen a little laterally, and have but little pigment. The
nuchal band has some pigment and a median, narrow, elongate,
black spot.

Leptodeira annulata taylori Smith

Leptodeira annulata taylori Smith, Proc. Biol. Soc. Washington, vol. 54, 1941,
p. 115 (type locality, Orizaba, Veracruz).

One specimen of this form (No. 4872) was taken at La Joya, 5
km. W of Aquismon, 2500 ft., J. Graber, collector. The specimen
gives the following scale data: ventrals, 194; anal divided, sub-
caudals, 84 + 1; two preoculars, two postoculars; upper labials, 8-8;
lower labials 10-10; temporals, 1 + 3 (1 + 2); scale formula, 21, 21,
23, 17, 16. Thirty-four dark spots on body, 21 on tail; bands on body
not reaching to ventrals; no light lines bordering head scales; venter
with some scattered pigment usually bordering posterior edge of
each ventral; heavily pigmented under tail.

Leptodeira maculata (Hallowell)

Me°alops maculatus Hallowell, Proc. Acad. Nat. Sci. Philadelphia, vol. 12,
1860 (1861), p. 468, (type locality, 5 mi. E Jalapa, [restricted]).

One specimen ( No. 4201 ) is from Nacimiento del Coy, collected
April 27, 1951, by L. Guerrero.

The ground color is dark brown, not strongly contrasted with
the black blotches except on the median dorsal line, where one or
more scales between spots may be entirely lacking in pigment.

Herpetology of San Luis PotosI 1611

There are approximately 27 dorsal blotches, none reaching the
ventrals laterally, and 12 on the tail. Some of the blotches are
confluent dorsally, with a complete fusion of two on one side and
a separation on the other. The gray-white borders of the posterior
head scales (parietals, supraoculars and frontal) are evident. The
venter is immaculate. A second specimen (No. 4873 2 ) taken at
Rancho Sabinal, 200 ft. elev, Nov. 10, 1951, by R. Graber, has the
following characters: upper labials, 8-8; lower labials, 10-10; ven-
trals, 174; anal divided; subcaudals, 54 -f- 1; scale formula, 23, 23,
23, 19, 19. Color above dark brown with 26 black body bands and
about 8 caudal bands, separated by orange-tan areas; ventrally
orange-white on body and tail; some pigment spots on outer edges
of certain subcaudals. A few of the blotches are confluent. Head
scales partly edged with white, outer edge of parietals strongly so.
Dark blotches reach to the first scale row.

A specimen, No. 5384, from El Salto, has 25 bands on body and
eleven on the tail; the light neck-band is narrow, two to three scales
wide, the borders parallel. The venter is immaculate. The head
scales are faintly outlined with lighter color.

Coniophanes imperialis imperialis (Kennicott)

Taeniophis imperialis Kennicott, in Baird, Rep. U. S. Mex. Bound. Surv., vol. 2,

1859, Rept., p. 23, pi. 19, fig. 1 (type locality, Tamaulipas).

No. 5386, from Nacimiento del Coy was collected July 16, 1952.

The characteristics of the subspecies imperialis are clearly de-
fined, especially in the details of color and markings. The ventrals
are 135. The tail is missing. The subspecies is known from Texas,
southward, along the eastern lowlands, to Jalapa, where it seems to
intergrade with C. imperialis clavatus (Peters).

Coniophanes fissidens proterops Cope

C[oniophanes] proterops Cope, Proc. Acad. Nat. Sci. Philadelphia, vol. 12,

1860, p. 249 (type locality, vicinity of Jalapa, Veracruz).

A specimen from El Salto (No. 5385), collected by Fugler is
somewhat atypical. The head is dark gray, the labials light bor-
dered above by a narrow cream line which in turn is bordered
with black above, that extends to back level of the head. A gray
median stripe covers three whole scale rows and edges of adjoining
scales. The median row has a series of black dots visible to near
end of body. The scales on sides of body are lighter, without dark
or light stripes. Supra- and infralabials finely punctate with black,
a distinct row of diminutive dots on the outer edges of ventrals,
with other equally large flecks scattered sparsely on venter, but

1612 The University Science Bulletin

growing less, posteriorly. There is some trace of a light lateral
caudal line darker bordered above and below. The subcaudal area
is immaculate. The supralabials, 7; infralabials, 8; ventrals, 129;
subcaudals, 74 -j- 1.

Crotalus triseriatus triseriatus (Wagler)

Urosophis triseriatus Wagler, Natiirliches System der Amphibien, 1830, p. 176,
(type locality, Alvarez, San Luis Potosi [restricted]).

The following characters obtain in No. 4882, Ahuacatlan, 5800 ft.
elev.: ventrals 155, subcaudals 29 (1st divided, 2nd-19th single,
20th-27th divided, 28th single, 29th divided). Scale formula: 26,
23, 23, 19, 17; upper labials, 12-12, two touching the subocular;
lower labials, 11-11. Anterior nasal near twice size of second;
apical pits discernible on many scales. "Olive with red-brown
markings; belly orange; iris yellow-orange"; no line present from
eye to angle of mouth.

Crotalus triseriatus (melanistic form)

Two specimens are in the collection. The first, No. 4881, from
6 mi. W of Ahuacatlan, Aug. 3, 1951, has the following characters:
ventrals 199, subcaudals 24 ( 1st divided, 2nd to 20th single, 21st to
24th divided), scale formula: 26(27), 23, 23, 19, 19; supralabials
12-12; infralabials 11-12; anterior nasal only about one and one-
half times posterior; four rattles and a button present. Dorsally
blackish with brown spots; orange near tip of tail; ventrally almost
uniformly steel gray; white markings near head.

The second specimen, No. 4884, 6 mi. W Ahuacatlan, October 5,
1951, 5400 ft. elev., R. Graber, coll., has the following characters:
154 ventrals, 22 subcaudals (1st divided, 2nd to 13th undivided,
14th to 21st divided); scale formula: 27, 23, 23, 19, 17(18).

Both snakes contained partly digested specimens of Sceloporus
( not identified ) .

Crotalus atrox Baird and Girard

Crotalus atrox Baird and Girard, Catalogue of North American Reptiles, 1853,
pp. 5-6 (type locality, Indianola, Calhoun Co., Texas).

This species is represented by a single small specimen, No. 4883,
taken at Pozo del Carmen, 5600 ft. elevation, Aug. 30, 1951, by Jean
Graber. The specimen has 181 ventrals and 20 subcaudals, the
terminal one divided. A button represents the rattle. The scale
formula is: 23, 23, 23, 23. Blackish bands on body number 32.
There are four on the tail.

Herpetology of San Luis Potosi

1613

LORICATA

Crocodylus moreletii Dumeril and Bibron

Crocodilus moreletii Dumeril and Bibron, Catalogue methodique de la collec-
tion des reptiles, 1851, p. 28 (type locality, Lake Peten, Guatemala).

Three specimens, Nos. 4371-4373 from Tanchachin, near Aquis-
mon, R. T. Gregg, collector, are in the collection. The three speci-
mens measure from 482-515 mm. in total length; snout-to-vent length
( greatest ) 245 mm. This is the first published record for the State.

COLLECTING LOCALITIES

The map below gives most of the collecting localities mentioned
in the text.

102 101 100 99

/ \

SAN L0IS POTOSI

S Mat

ehuala

• 1

/""

;

• Huisa(

Amoles • ■.

"N -•' — v

Villar

\.^J >

• PI

atanlto * \

*».-^

.'''^

# Poso del (

armen

i Prese

Gonzales Santos

• M

COS »

Santiago •

Jesus Maria

Tanchachin • •

Puente de Dlos

Bledos *a

0 10 25 50 TOO

Ventllla

Rancho Sablnal •

• El Naclmlent
• del Co:

1 — 1 1 1 1

Aqulsmon «

\. < Vv

• AxtlaY.

V-

> ./

vir

102 101 100 99

Fig. 1. Map of San Luis Potosi showing collection localities.

Amoles

22° 49' N;

100° 20' W

Platanito

22° 30' N;

99° 26'

Aquismon

21° 38' N;

99° 00' W

Poso del Carmen

22° 19' N;

100° 37'

Axtla

21° 27' N;

98° 52' W

Presa Gonzales

Bledos

21° 49' N;

101° 07' W

Santos

22° 06' N;

101° 06'

Canada Grande

21° 58' N;

100° 23' W

Sabinal

21° 45' N;

99° 06'

El Nacimiento del

Santiago

21° 53' N;

101° 15'

Coy

21° 47' N;

98° 58' W

Sierra San Miguel .

22° 00' N;

101° 05'

Huisache

22° 55' N;

100° 23' W

Tanchachin

21° 50' N;

99° 09'

Jesus Maria

21° 56' N;

100° 54' W

Ventilla

. 21°47'N;

101° 02'

Matehuala

Micos

23° 39' N;

22° 08' N;

100° 38' W
99° 10' W

Villar

•

22°32'N;

100° 28'

1614 The University Science Bulletin

The following localities visited by Mr. Charles Fugler are not on

the map.

Presa Prudentia. A reservoir about 4 miles N of Bledos on the road to Bledos.
La Luna Media or Media Luna. A large, spring-fed area of deep lagunes, with

intertwining creeks, approximately 8 miles SE of Rio Verde.
38 Km. SW of Rio Verde. This locality is in the mountains to the SW of Rio

Verde. The distance is approximate. While in the area, we stayed at a

logging camp. I can best characterize the region as one of magnolias and

pines.
El Abra. A small village on the Valles-Tampico highway.
Rio Claro. 6 Ions. E of Tamazunchale.
San Felipe. 1 mile W. A small village on the Valles-Tampico highway.

THE UNIVEKSITY OF KANSAS

SCIENCE BULLETIN

Vol. XXXV, Pt. II] September 10, 1953 [No. 14

A Report on a Collection of Ceylonese Serpents

Edward H. Taylor

Abstract. — This is a study of a collection of serpents from Numunukula,
Ceylon made by Mr. W. W. A. Phillips. While several presumed new forms
are pointed out none are described as new since the author has not been able
to examine described types of related forms.

A well-preserved collection of reptiles and amphibians, recently
received as a gift from Mr. W. W. A. Phillips, Esq., the well-known
mammalogist of Namunukula, Ceylon, contains numerous interesting
species.

The specimens were taken by Mr. Phillips on the Tonacombe Es-
tates in the Uva Hills, Uva Province, Ceylon, at elevations between
3000 and 4500 feet. Altogether there are 61 specimens, distributed
as follows: 22 snakes, 34 lizards, and 5 amphibians.

In this paper I am reporting on the snakes of the collection. The
lizards are being treated elsewhere in a paper dealing with the Cey-
lonese lizard fauna, and the amphibians in a similar paper dealing
with the total amphibian fauna.

I take this opportunity of expressing my gratitude to Mr. Phillips.

Pseudotyphlops philippimis (Cuvier)

Uropeltis philippinus Cuvier, Regne animal, distribue d'apres son organization,
2nd ed., vol. 2, 1829, p. 76 (type locality, "Philippines"). Wagler, Natiirliches
System der Amphibien, 1831, p. 194-195; Griffith, The Animal Kingdom
arranged in conformity with its organization by the Baron Cuvier, vol. 9,
p. 251 (footnote); Miiller, Tiedemann und Treviranus Zeitschr. fur physiol.
Heidelberg, 1832, vol. 4, pp. 248-252, pi. 22, figs. 2, 3; pi. 21, figs. 4, 5;
Gervais, Guerin, Magasin Zool. 1837, Cl. 3, pi. 13; Schinz, Naturgesch.
Abbildung, Rept., 1833, p. 132; Dumeril and Bibron, Erpetologie Generale,
vol. 7, pt. 1, 1854, pp. 160-165, pi. 59, fig. 2, head, and 2A, tail (this con-
tains a detailed description of the type specimen). Peters, De Serpentum
Familia Uropeltaceorum, Berlin, 1861, p. 20 (places in synonymy Uropeltis
Saffragamus, U. pardalis, and U. grandis, three species described by
Kelaart, Prodromus Fauna Zeylanicae, vol. 2, pt. 1, 1853 (1854), pp. 15-16.)

Pseudo-typhlops phUippinus Cuvier, Abbildungen neuer . . . Amphibien
1838, p. 44; Gervais, Voyage de la corvette, Favorite, vol. 5, p. 66, pi. 26;
Tennent, Sketches of the Natural History of Ceylon . . . London,
1861, pp. 302-303, unnumbered figure.

(1615)
72—6490

1616 The University Science Bulletin

Uropeltis grandis Kelaart, Prodr. Faunae Zeylanicae, 1854, vol. 2, p. 15;
Giinther, The Reptiles of British India, 1864, p. 188; Beddome, Ann. Mag.
Nat. Hist., ser. 5, vol. 17, 1886, p. 9; Boulenger, The Fauna of British India,
Including Ceylon and Burma; Reptilia and Batrachia, 1890, p. 254; Cata-
logue of the Snakes in the British Museum (Natural History), vol. 1, 1893,
p. 139; Green, Spolia Zeylaniea 1906, p. 220; Wall, Ophidia Taprobanica,
or the Snakes of Ceylon, 1921, p. 26; Journ. Bombay Nat. Hist. Soc., vol. 29,
1923, p. 354.

Uropeltis saffragamus Kelaart, Prodr. Faunae Zeylanicae 1853 (1854), p. 15
( type locality Ratnapoora near Adam's peak, Ceylon ) .

Uropeltis pardalis Kelaart, Prodr. Faunae Zeylanicae 1853 (1854), p. 15 (type
locality, Matura, Ceylon); Gray, Proc. Zool. Soc. London 1858, pp. 263-264.

Pseudotyphlops philippinus Smith, The Fauna of British India, including
Ceylon and Burma; Reptilia and Amphibia, vol. 3, Serpentes 1943, pp. 93-
94, fig. 27.

The genus Pseudotyphlops has been considered by most writers
as monotypic. Kelaart, loc. cit., described three forms, that since
1861 have been placed in the synonymy of philippinus; or the three
have been regarded as a single species under the name grandis.
In 1861, Tennent ( loc. cit. ) suggested that Uropeltis grandis Kelaart
(including pardalis and saffragamus) and Uropeltis philippinus
Cuvier were synonyms, but this was disregarded for the most part
until Smith ( 1943, loc. cit. ) placed them under the older name.
Smith makes no comments on the Kelaart species save to place them
in synonymy.

Of the three forms described by Kelaart, the types of two, U.
pardalis and U. grandis are in the British Museum. The type of
U. saffragamus is presumably lost.

I regard it as probable that more than a single species (sub-
species?) occurs in Ceylon and an examination of the described
forms becomes necessary to ascertain which name must apply to
which form. Since the descriptions are brief I am repeating them
here. The material available to me does not permit a solution of
the problem.

Uropeltis saffragamus

Head dark olive brown, the rest of the upper surface of a blackish brown
color with bluish bronze reflections. Beneath white. A pale white spot on
each side of the neck near the head. Tail deeply truncated and nearly covered
with a large, flat, circular, blackish, granular shield, white and rounded be-
neath, and with the lower part covered with five series of small scales; the
central series broader than the lateral ones, vent shields, 1-2. The neck and
forepart of the body thicker. Length 9 inches.

District of Saffragam, near Adam's peak. The only specimen of this species
which I have seen, is one sent by Mr. Barnes de Zilva from Ratnapoora.

Uropeltis pardalis

Head small dark olive, upper parts black with beautiful bronze reflections
irregularly spotted white. Beneath yellowish white marked with large and
sAall black spots, variously shaped; some spots pale eyed. Tail very short

Collection of Ceylonese Serpents 1617

obliquely truncated, with a large flat orbicular granular shield. Length 6%
inches. Circumference % inches. Habitat, Matura.

I am indebted to the Rev. Mr. Ondaatje for the only specimen I have ex-
amined of this species. The black spots on the lower parts occupy more than
one scale, generally two or three contiguous scales, and they are placed without
any order in various directions. The chin and throat are immaculate.

Uropeltis grandis

Above dark brown with a bluish metallic lustre; anterior part of each scale
with a blackish spot. Beneath a pale yellow color, spotted brown on the
anterior part of scale. Head of a light olive brown color. Tail short, abruptly
truncated, the truncated surface entirely covered with a larger circular granular
shield. Vent scales 1-2.

Total length above 1 foot 7 in.; length below 1 ft. 8 in. Tail shield nearly
the size of a shilling piece. Head, 8/10 inch in length; greatest circumference
near neck, 2% in. Habitat Southern Province.

The only specimen I have seen of this very large rough tail, is one procured
by Mr. Balkhuysen of the Colonial Medical Service, from Kerinday, near
Matura, Ceylon.

The description given for Uropeltis partialis seems to approach
more closely to the description of the type specimen of philippinus
(as presented by Dumeril and Bibron, Erpet. Generale vol. 7,
pt. 1, pp. 160-165).

While the dorsal head scales seem to agree in their proportions
and relationships there are differences in the labials and the scales
on the chin.

Flower, loc. cit., p. 24, fig. 7, gives a figure showing the squama-
tion of the chin; there is no undivided postmental and the first
chinshields are regular both touching the mental.

In the specimens at hand three have the first pair of chinshields
present, only the one on the left, in contact with the mental. In
No. 31248, there is an undivided postmental. The subcaudal scales
of males are more numerous and there is some difference in the
character of scales about the vent.

No. 31250 has 5 lower labials, No. 31251 has five on one side,
four on the other. In Nos. 31248 and 31249 the labials are 4, the
second one being normally much larger than in Nos. 31250 and
31251.

The squamation of the head is remarkably similar in the two
color forms. The following characters obtain:

Rostral forming a terminal cap on snout, dorsally extending back
between nasals and partially dividing them; no internasals; suture
of nasals somewhat longer than that between prefrontals; frontal
six-sided, its length equal to dorsal length of rostral or length of
parietal, not equal to its distance from end of snout; common
parietal suture about equal to the prefrontal suture; no anterior

1618 The University Science Bulletin

temporal; ocular scale as high as long; the diameter of eye a little
less than half length of ocular; nostril in lower anterior corner of
nasal; no undivided postmental except in No. 17, a young speci-
men; ventral length of rostral less than dorsal length; rostral ex-
tending 2.7 mm. beyond mouth.

There is rather little variation in the character of the head scales.
In No. 31249 the snout region of the head is somewhat broader
than in Nos. 31250 and 31251.

Two of these Nos. 31248 and 31249 conform to the color descrip-
tion of U. philippinus and U. pardalis Kelaart (See Dumeril and
Bibron loc. cit. and fig.) and two Nos. 31250 and 31251 belong to
the form described as U. grandis.

The description of these follow:

In No. 31249 2 , the general color is deep, iridescent lavender
with a darker area on each dorsal and lateral scale. The three
ventral scale-rows together with the median ventral series are dis-
tinctly lighter than the dorsum, each scale, save on chin and neck,
with an indefinite darker area. A paired series of darker spots
(occasionally alternating or fused together) are on the ventral sur-
face. A prominent, yellow, curving spot borders the lower part of
the terminal plate border, and the labials, at least the lower part of
the labials, are light colored. Length 345 mm.

A young specimen, No. 31248 $ , measuring only 148 mm. is
black above with a bluish iridescence. There is a scattering of
very numerous yellow dots, no larger than a single scale, usually
single but occasionally on sides suggesting narrow, irregular, trans-
verse lines. The ventrals and three adjoining scale-rows are greenish
white with numerous black spots scattered over the venter. There
is no tendency to form transverse black bands, and there is no sharp
line of demarkation between the dorsal dark ground and the ventral
light coloration; chin and throat immaculate; labials whitish.

Nos. 31250 5 and 31251 § measure 318 mm. and 360 mm., re-
spectively. They are brownish above with dark markings on all
scales above and below, the ventral ground color is lighter than the
dorsal but the lighter areas are very indefinite and the spotting
such as occurs in the preceding two specimens is absent.

Data on Pseudotyphlops philippinus

Number
and
sex

31248 $

31249 9

31250 9

31251 9

Elev.

Ventrals

Length

from

Subcaudal

Scale

ft.

mental

pairs

rows

mm.

3000

140

148

2000

149

345

2000

144

318

2000

146 anal 2

360

Collection of Ceylonese Serpents 1619

Since the four specimens here listed are from the same general
locality, the variation cannot be regarded as geographic. Whether
two forms actually are represented cannot be determined from
the material at hand.

Lycodon aulicus ( Linnaeus ) ( var. )

The genus Lycodon is known to be represented by at least three
species in Ceylon. These are Lycodon aulicus (Linnaeus), Lycodon
osmanhilli Taylor, and Lycodon striatus (Shaw)* The first two
species are characterized by having angular ventral scales and nine
upper labials; the third species lacks the angular scales and has
only eight upper labials.

From the large synonymy of Lycodon aulicus presented by
Boulenger f and Smith } one suspects that certain of the names
probably represent forms worthy of subspecific (or perhaps even
specific) designation, more especially since certain of these dif-
ferences presumed to be individual variation occur throughout a
wide range in India and Ceylon. Series of specimens from the
same locality are of course necessary to ascertain the degree of
differentiation. In the collection at hand there are two specimens
of Lycodon from southern Ceylon having a different general appear-
ance from those at hand taken in the region north of Trincomalee §
in the lowlands. With adequate material it may be possible to
recognize a named form for southern Ceylon. Whether this is new,
or is an Indian form extending into Ceylon cannot be stated here.

No. 31232, g : The following scale and color characters obtain:
the ventrals, 194; anal single, (partly divided anteriorly, undivided
posteriorly); subcaudals, 29 -f- 1; scales smooth, with a single apical
pit, the outer row largest; scale formula: 17, 17, 17, 15, 15; supra-
labials, 9-9, the first not in contact with loreal; infralabials, 10-10,
five touching first chinshields, which are scarcely as large as first
lower labial; third, fourth, and fifth upper labials enter eye; pre-
ocular touches frontal; one preocular and two postoculars; frontal
length less than its distance from middle of internasals; prefrontals
angular laterally; internasals rounded laterally; diameter of eye,
2 mm.; distance from eye to nostril, 3.65 mm.; eye to level of tip of
snout, 5 mm.; above violet-brown banded with cream becoming
lighter laterally. A nuchal band separated from the first light body

* In Univ. Kan. Sci. Bull. vol. 33, pt. 2, 1950, p. 562. The scale data given under
Lycodon striatus applies to the young paratype of Lycodon osmanhilli and not to this species,
t Boulenger, Cat. Snakes Brit. Mus. vol. 1, 1893, p. 325.

t Smith, Fauna Brit. India. Rept. and Amph. vol. 3, Serpentes 1943, p. 263-264.
§ Taylor, Univ. Kans. Sci. Bull., vol. 33, pt. 2, 1950, pp. 560-562, pi. 19, fig. 2.

1620 The University Science Bulletin

band by a broad darker band covering 36 transverse scale rows;
first transverse light band 3 scales wide dorsally to 5/2 scales wide
where it borders the ventrals; between the first and second light
bands a dark band covering 36 scale-lengths; between second and
third light bands (barely indicated laterally) a darker band covering
31 scale-lengths; venter white, the pigment encroaching on the ven-
trals slightly; scales of white bands each with some pigment.

No. 31233 $ : This specimen has the following characters: ven-
trals, 196; anal divided; snbcandals, 60 -f 1; scale formula: 17, 17,
17, 15, 15 (14); supralabials, 9-9; infralabials, 10-10; head scales as
in specimen above; the preocnlar touches frontal. The color is
similar to the above but it is slightly more brownish. The dark band
between the neck band and first light body band, is 30 scale-lengths
wide; that between the first and second light bands, 23 scale-lengths;
that between the second and third light bands, 26 scale-lengths; that
between third and fourth light bands, 19 scale-lengths (latter
scarcely evident ) ; there is no band of white around the snout on the
upper labials, the labials being slightly lighter than scales on top of
head.

Oligodon sublineatus Dumeril, Bibron and Dumeril

Oligodon sublineatus Dumeril, Bibron and Dumeril, Erpetologie Generate,
vol. 7, pp. 57-58 (type locality, Ceylon).

One small specimen (No. 31242), bearing the typical coloration
and markings, has 142 ventrals, the anal divided, and the sub-
caudals 32 + 1. On the left side, the sixth labial is excluded from
the labial border. There are seven upper and seven lower labials
present. The scales are smooth, a few scattered ones having single
apical pits. The scale formula is 17-15-15.

The narrow diagonal lines from the fourth and fifth labials to the
prefrontals are narrowly separated mesially. An elongate sym-
metrical light-edged mark extends from the posterior third of the
frontal to a point two scale-lengths behind parietals. A pair of
lateral, nuchal spots are present. Anteriorly the dark blotches on
each side alternate with their fellows, and there are three rows of
ventral dots or dashes, the outer ones being more nearly continuous
than the inner.

Aspidura brack y orrhus * (Boie)

Scytale brachyorrhus Boie, Isis von Oken, 1827, p. 517, (type locality, Ceylon).
A female specimen, No. 3123S, from Tonacombe has the following
characters :

* This name is incorrectly spelled brachyorrhos by Taylor (1950).

Collection of Ceylonese Serpents 1621

Ventrals, 149; subcaudals, 28 + 1, all divided; anal single; scale
formula: 17, 17, 17, all rows smooth without keels; supralabials,
6-6; infralabials, 6-6; one preocular, two postoculars, both touching
the parietal; temporals, 1 + 2; frontal hexagonal, distinctly longer
than its length from tip of snout; parietals longer than their distance
from tip of snout; rostral small, only slightly visible above, the head
much narrowed at tip; first chinshields two and one-half times
larger than the second chinshields; upper secondary temporals
large; median scale following parietals also enlarged; no loreal, the
prefrontals touching the second and third labials, the fourth labial
alone entering orbital ring; eye small, its diameter into distance
between eye and nostril, 2.2 times. Dorsally rather light fawn, the
pigment varied so that a lighter stripe, covering much of the fourth
and fifth rows, is discernible from the neck band to end of tail; the
three outer scale-rows with scattered black pigment, darkest along
the edge of the fawn stripe; median dorsal row with a little less
pigment than three adjoining rows, and bearing a black spot on
each fourth or fifth scale, the series extending to end of tail; a few
irregular black spots on sides of neck and anterior fifth of body.
Head darker anteriorly, growing light on outer posterior part of
parietals and secondary upper temporal; a pair of black, nuchal
spots bordered front and back by lighter color behind angles of the
jaws; a median, spear-shaped, black mark from parietals extending
back six scale-lengths, the three spots nearly contiguous; labials
largely cream with black areas; ventrals nearly immaculate, with
only an occasional fleck of pigment on their outer edges.

There is a very young specimen (No. 31238a) of this species in
the collection, that shows no significant differences from the one
described.

Haplocercus ceylonensis Giinther

Haplocercus ceylonensis Giinther, Catalogue of the Colubrine Snakes in the
British Museum, 1858 (Feb. 12, 1859), p. 15 (type locality, Ceylon).

Two specimens are in the collection of which the first No. 31236
has the following characters: one preocular, two postoculars; scale
formula, 17, 17, 17, the scales (except on the anterior fourth of
body, which is smooth) strongly keeled to tip of tail; supralabials,
6-6; infralabials, 6-7; second pair of chinshields less than one half
(near one third) of first pair; three or four lower labials, touching
the first chinshields; ventrals, 167, the penultimate divided; anal
single; subcaudals, 47 -f 1, single. Head brownish followed by a
somewhat irregular, darker collar; usually a series of dark dots,
separated by three scale-lengths, are discernible on each side of

1622 The University Science Bulletin

the body throughout its length; posteriorly they are discernible
only with difficulty since the body color is also dark; anterior part
of ventral surfaces cream while gradually merging into pink or
salmon-pink at the beginning of the second fourth of the body;
subcaudal region magenta.

No. 31237. In the second specimen, taken at 4500 ft, 25 May,
1951, there is a distinct lighter band preceding the dark nuchal
band; labials and first temporals cream, the sutures edged with
dark; two or three light spots on sides of neck. The posterior two
thirds of body salmon-pink. The row of dark spots along the sides
of the body can be discerned with difficulty.

Dryophis nasutus (Lacepede)

Coluber nasutus Lacepede, Histoire Naturelle des Serpents, vol. 1, p. 100, vol.
2, p. 277, plate 4, fig. (type locality, Ceylon [restricted]).

One typical specimen No. 31239 from 4000 feet elevation, is
present in the collection. The ventrals are 182, the anal divided,
and the subcaudals are 154. The color is uniform dark green above,
while below it is yellow green with a pair of cream lines on the
outer sides of ventrals.

Natrix stolata stolata (Linnaeus)

Coluber stolatus Linnaeus, Systema Naturae, 1758, 10th ed. p. 219, (type lo-
cality, Asia).

A specimen in the collection, No. 31252, has the following char-
acters: scale formula, 19, 19, 17; ventrals, 120; subcaudals, 72; anal
divided; supralabials, 8-8; infralabials, 10-10; preocular, 1, not reach-
ing frontal; postoculars, 3; temporals, 1 4- 2 -|- 3; 3rd to 5th labials
enter orbit; four lower labials touch first chinshields, which are
(presumably abnormally) transversely divided; eye large, its diam-
eter equal to its distance from nostril; frontal longer than its distance
from tip of snout; shorter than the parietals.

Head scales largely edged with black; two yellow bars, one in
front, one behind eye on side of head, each bordered by black bars;
venter and under side of tail, chin and throat immaculate.

Boiga ceylonensis (Giinther)

DipsadomorpJuis cei/lonensis Giinther, Catalogue of the Colubrine Snakes in
the collection of the British Museum, Feb. 12, 1858 (1859), p. 176 (type
locality, Ceylon); Reptiles of British India, 1864, p. 314, pi. 23, fig. B.

Two female specimens, No. 31240 and No. 31241 are in the col-
lection. These specimens have the following characters: Inter-
nasals as long as broad or a little longer; prefrontals nearly a half

Collection of Ceylonese Serpents 1623

broader than long, and longer than internasals; frontal one-fifth or
one-sixth longer than broad, its length greater than its distance from
the tip of the snout; loreal quadrangular, a little higher than long;
preocular single, high, narrowly separated from frontal (or touch-
ing); two postoculars; temporals (respectively) 2 + 4 + 4; 2 +
3 + 4; upper labials, 8-8, 8-8; lower labials, 11-12, 11-11, five or six
lower labials touching first chinshields; scale formula, 19, 19, 19, 15;
19, 19, 18, 13 (in this latter specimen, the median dorsal scales
becoming veiy large where the rows reduce to 13); ventrals, 224,
222; anal, 1, 1; subcaudals 107 + 1, 98 + 1.

There are some indefinite flecks or spots on the interorbital area
and on snout; a pair of symmetrical spots with somewhat lighter,
brownish centers on parietals; a black line, beginning behind eye,
extends back, narrowing on the eighth labial, then widening a little
and terminating behind jaw angle; a median series of dark blotches,
58(59) from occiput to vent; some twenty indefinite spots discerni-
ble on tail; on sides an equal number of rather indefinite dark marks
alternating with the median series and below these is another series
alternating with the preceding and opposite the median series that
extends onto edges of ventrals. Venter dirty white with an in-
definite row of small spots or flecks on each side, and with finer
flecks or peppering over the ventrals, least dense on neck region,
most dense under tail. One of the specimens No. 31241 had the
remnants of a green Calotes in its stomach but I could not identify
the species with certainty.

Boiga trigonata (Schneider)

Coluber trigonatus Schneider in Beehstein (Lacepede, Histoire Naturelle des
Serpents, vol. 4, 1802, p. 256, pi. 40, fig. 1 (type locality, Vizagapatam,
India (Based on Russel).

This specimen, (No. 31234), has the following characteristics:
Scale formula: 23, 21, 21, 17, 17; ventrals, 237; anal, 1; subcaudals,
85 + 1; preoculars, 2; postoculars, 2; labials, 8-8; lower labials, 11-11;
temporals, 3 + 3 + 4; scales of median row nearly a half larger
than adjoining scales; upper head triangular with a pair of gray-
ish lines beginning on supraoculars and extending from supra-
ocular back to angle of mouth; bordering these on their inner edge,
is a pair of grayish brown, darker-edged stripes that run forward
and join the gray-brownish color of the snout and area in front of
eyes; these stripes are separated by a narrow gray stripe running
from the posterior level of eyes to the ends of the parietals where
the line forks and runs back behind the mouth angles inclosing a

1624 The University Science Bulletin

somewhat arrow-shaped, brownish, darker-edged spot on the nape.
Body generally gray-brown with approximately 59 narrow, trans-
verse, light gray bands edged with black. The black becomes more
important and may cover nearly a whole scale row as does the
lighter color; spots resulting come to alternate and the bands may
join a median light line that extends the greater part of length of
body; on the lower body scales and outer edges of ventrals there
are numerous small blackish dots rarely appearing near the mid-
ventral part of venter.

Bungarus ceylonicus Giinther

Bungarus ceylonicus Giinther, Reptiles of British India, 1864, p. 344 (type lo-
cality, Ceylon).

A specimen (No. 31230) has the head somewhat mutilated and
certain scale characters are obscured. The ventrals are 132, the
anal single, the subcaudals 33 4- 1. The color is black above with
18 whitish bands on the body and four on the tail. These bands are
narrowed dorsally to a width of 2 to 2M scales, but widen on the
sides and on the venter to a width of four, more rarely five, ventral
scales. Ventrally the intervening black areas are usually six scales
wide, the ventrals involved in most cases still having some white
color remaining. The black ventral markings are not present in the
young, but they become increasingly important in older specimens.
The first white band is separated from the head by approximately
20 black scale-rows. The first two white bands involve lOM and 8/2
ventrals respectively.

Agkistrodon hypnale (Merrem)

Cophias hypnale Merrem, Syst. Amph., 1820, p. 155 (type locality, Ceylon).

The five specimens in the collection, Nos. 31343-31347, are from
Tonacombe. The ventral and subcaudal scale-counts of these are,

spectively:

31343 3

155 ventrals

43 subcaudals

31344 $

153

31345 $

151

31346 5

149

31347 5

148

THE UNIVEKSITY OF KANSAS

SGIENGE BULLETIN

Vol. XXXV, Pt. II] September 10, 1953 [No. l5

Frogs of the Family Centrolenidae from Brasil

Edward H. Taylor * and Doris Cochran **

Abstract. — This paper treats of the frogs of the genus Cochranella, family
Centrolenidae, that are known to occur in Brasil. Twelve species are recog-
nized, of which nine are regarded as new and described under the names,
Cochranella surda, C. petropolitana, C. lutzorum, C. delicatissima, C. boker-
manni, C. divaricans, C. dubia, C. vanzolinii, and C. albotunica. The Brazilian
species, Hylella parvula Boulenger, Hyla (Hylella) eurygnatha A. Lutz, and
Hyla ( Hylella ) uranoscopa L. Miiller are referred to the genus Cochranella and
the family Centrolenidae since all have the bones of the tarsus fused into a
single bony shaft, similar to the tibio-fibula of all Salientia.

INTRODUCTION

For several years, as time permitted, the junior author has been
preparing a report on the frogs of Brasil, but in this work she has
omitted discussion of a group of small species occurring in Brasil,
that in the past have been regarded as having an uncertain family
relationship. Since the frogs of this group have been of especial
interest to the senior author she has proposed that they be treated
and published in a separate paper by the two of us.

Probably least known in the Brazilian amphibian fauna is this
group of tiny frogs (28 mm. maximum snout-vent length in known
Brazilian species), arboreal in habit, generally encountered cling-
ing close to leaves of herbs and trees usually in the immediate
vicinity of running water. Eggs in some, if not all, species are
placed on leaves above water, the young hatching and falling into
the water to continue their development.

In life most of the known species of the group are green above,
the transparent flesh on the ventral surfaces permitting a fairly
clear view of part of the viscera. Occasionally some yellow is

* Department of Zoology, University of Kansas, Lawrence, Kan.
** United States National Museum, Washington, D. C.

(1625)

1626 The University Science Bulletin

present. On fixation, in alcohol or formalin, the green disappears
rapidly and such pigment as is present is visible in chromatophores,
that may be equally distributed over head and dorsum, with a strip
of pigmentation on the dorsal surface of the limbs (or sometimes
absent on upper arm). Sometimes the chromatophores may be
segregated into spots and reticulations. The color of the pigment
under a lens is lavender or purplish.

The characters and habitus of these species are for the most part
those of the tree frogs of the Hylidae and Rhacophoridae, and like
them, there is a cartilage intercalated between the ultimate and the
penultimate phalanges. These frogs differ however from typical
hylids in that the terminal phalanges bear a pair of projections,
commonly described as T-shaped. Because of this particular char-
acter certain authors have regarded them as belonging to the Family
Leptodactylidae — believing that the greater weight should be placed
on the shape of the terminal phalanges than on other more signifi-
cant characters of the animal. Others have placed them with the
Hylidae. The discovery by the senior author of the fusion of the
astragalus and calcaneum in this group of frogs, together with
other significant characters, has caused us to treat them in a family
of their own.

The Family has a range extending through eastern and southern
Mexico from central Veracruz and Guerrero south to and including
most if not all of Brasil and a part of the various states bordering
northern and western Brasil. It comprises at least three known
genera and perhaps others now unknown or unrecognized. We
believe that a better understanding of the group is possible by
considering it of family rank, as proposed by Taylor, using for it
the name Centrolenidae.*

One will, of course, expect a large fauna of species and perhaps
genera in Brasil when competent exploration for these diminutive
forms has been made. In the limited exploration of Costa Rica no
less than eight species are now known, representing three genera.

In United States National Museum specimens of Centrolenidae,
collected in the Brazilian states of Rio de Janerio, Sao Paulo, Minas
Geraes, and the Districto Federal, and two specimens from the
zoological collections of Sao Paulo, we recognize some ten species.
Four of these belong to a spatulate-headed section; and six to the
short blunt-headed section, all seemingly referable to the genus
Cochranella. None of these species has a humeral hook and pre-

* Taylor, Proc. Biol. Soc. Washington, vol. 64, 1951, p. 36.

Family Centrolenidae from Brasil 1627

sumably none has an exposed pollical spine, although one species
has a pollical remnant. The vomerine teeth are absent in all of the
various Brazilian species here considered.

Certain external characteristics which these species have in com-
mon are: reduced webbing on the inner fingers; the presence of
an oval or somewhat circular (biscuit-shaped) palmar tubercle on
the posterior part of the palm; absence of a distinct, external meta-
tarsal tubercle; males with a vocal sac, and short or long vocal slits
that are somewhat diagonally placed, not tending to parallel the
line of the lower jaw, and the astragulus and calcaneum fused into a
single slender shaft of bone.

Characters used to show specific differences are those of the
tongue, the palatal groove, and the size of the vocal slit; size of
choanae and their relative distance from each other; character of
exposed tympanum or its covering, and its position with relation to
the eye; size of eye, its lid, and the covering of the hidden parts of
the eyeball ( tunic ) ; shape and extension of snout; head proportions;
position of nostril; constriction of body behind head; size (length)
of the suprascapula; anal "decoration" ( i. e. presence of specialized
granules, folds, or tubercles in the region lateral and posterior to
vent); relative leg length compared to body length; webbing and
character of the digits, and the character of the skin of venter
(smooth, granulate or striate).

To the students of amphibians who have not had the fortune to
examine these small frogs, the number of species recognized by us
may seem large. We are aware that some of this material leaves
much to be desired. Our experience, however, has shown that for
the most part in species of this family where a series is available
there is rather close conformity in characters such as body propor-
tions, general pigmentation, anal decoration, character of webbing,
etc.

It is not impossible that the eye tunic may change after long
preservation and become darker. However we have no definite
evidence that this occurs. In some of the species the posterior edge
of the pelvic girdle seemingly extends posterior to the level of the
thighs, when these are at right angles to the body; in others this is
not so. We find no evidence that the method of preservation is re-
sponsible for this appearance.

In the summer of 1951, the junior author made a journey to
Europe for the purpose of visiting some of the museums of Europe,
and while there examined Brazilian species in London and Munich.

1628 The University Science Bulletin

A few specimens which we and others had suspected of belonging

to this group were X-rayed or dissected and the findings were as

follows :

Hylella buckleyi Boulenger = Cochranella buckleyi (Boulenger)

Hylella parvula Boulenger = Cochranella parvula (Boulenger)

HijlcUa parabambae Boulenger = Centrolene (?) parabambae (Boulenger)

Hylella puncticrus Boulenger r= Centrolene prosoblepon (Boettger)

Hylella ocellata Boulenger = Cochranella ocellata (Boulenger)

There is no possibility that the generic name Hylella could be
used for any small frog of this family since the type of the genus
Hylella is Hylella tenera which the junior author has recently ex-
amined and which she regards as Hyla bipunctata. The only other
species placed in the genus by Reinhardt and Lutken was Hylella
punctatissima and this too is a hylid frog and not a member of this
family.

Three species of Centrolenid frogs have already been described
from Brasil, in each case under the genus Hyla or Hylella of the
Family Hylidae. These are Hylella parvula Boulenger,* Hyla
(Hylella) uranoscopa L. Mxiller,** and Hyla (Hylella) eurygnatha
A. Lutz.***

The junior author has examined cotypes of H. eurygnatha in
Brasil and there are several specimens, collected by Dr. A. Lutz at
the type locality, available in the collections of the United States
National Museum. These unquestionably are Centrolenidae and
are referable to the genus Cochranella. The tarsal segment has a
single bone (astragalo-calcaneum), and they conform in other
generic characters.

The type of uranoscopa was likewise examined and by dissection
the presence of a single bone in the tarsal joint was verified. This
species, the southernmost known is from the state of Santa Catha-
rina and is distinctly different from new forms herein described.

An X-ray photograph of the type of parvula, made by the au-
thorities of the British Museum, clearly demonstrates the presence
of a single bone in the tarsus of this species. This type likewise
was studied by the junior author. This species seemingly cannot
be identified with any of the forms here described as new.

We wish to offer our thanks to Dr. George Myers, Sr. Antenor
Leitao de Carvalho, and Mr. Jay Savage for having read the manu-
script and for offering helpful suggestions. Sr. Carvalho has assisted
in the preparation of the map.

* Proc. Zool. Soc. London 1894, p. 646, pi. 40, fig. 3.

** Zool. Anz., Bd. 59, 1924, pp. 234-235.

*** Compt. Rend. Mem. Soc. Biol. Paris, vol. 93, no. 21, 1925, pp. 137-138.

Family Centrolenidae from Brasil 1629

DESCRIPTIONS OF SPECIES

Synopsis of Described Forms of Cochranella in Southeastern Brash. *

1 . Tympanum completely concealed 2

Tympanum partially or entirely exposed 4

2. Two outer fingers one-third to one-fourth webbed 3

Two outer fingers with only a vestigial trace of web; a pair of

relatively large postanal pads or tubercles, with some vertical
wrinkles flanking pads laterally; eye tunic (covering concealed
parts of eyeball) blackish; heel reaching to middle of eye;
head rather narrow, not spatulate; chromatophores small,
equally scattered on dorsum; 24 mm surda sp. nov.

3. A pair of flattened postanal pads; eye tunic white; head short not

spatulate; heel reaching to nostril; chromatophores equally
distributed on dorsum; head wider than long; no pollex rudi-
ment; 22 mm eurygnatha

Probably no postanal decoration (none mentioned); eye tunic
white ("skin of eyelid like body"); heel reaching to tip of
snout; head as long as wide; skin with small, pinkish dots; a
distinct pollex rudiment; 25 mm uranoscopa

4. Venter completely or largely striate 5

Venter granular 6

5. Venter entirely covered with longitudinal striae; no special "dec-

oration" in region lateral to, behind, or below vent; tympanum
large, distinct, directed outward; head wide, somewhat spatu-
late; first and second fingers equal, or first slightly longer; heel
reaching beyond tip of snout; eye tunic dark; pigment equally

scattered; 23.5 mm. petropolitana sp. nov.

Venter longitudinally striate save on posterior part where striae
form irregular elongate granules; a pair of vertical ridges
curving across thighs to ventral surface behind vent; tym-
panum covered with skin, but outline clearly visible, directed
nearly vertically upward; head broad, spatulate, the front
sloping obliquely to lip in front of nostril; heel to a point
2 mm. beyond tip of snout; pigment chromatophores large, un-
evenly scattered on dorsum; eye with white tunic,

lutzorum sp. nov.

6. Tympanum directed laterad 7

Tympanum directed wholly (or largely) upward 8

7. Tunic of eye black; tympanum covered with skin but outline

clearly visible; heel to a point beyond tip of snout; no dec-
oration in region of vent; skull not transparent; outer fingers

only one-fourth webbed; 20.8 mm delicatissima sp. nov.

Eye with a brown or black tunic; tympanum covered with skin,
its outline visible, directed outward; heel to a point between
eye and nostril; a pair of swellings behind vent, widely sepa-
rated; webbing between outer fingers vestigial; pigment rather

* Since so many significant characters of Cochranella parvula are not known to us, we
are omitting this species from the synopsis. Its relationship may be with delicatissima.

1630 The University Science Bulletin

equally distributed in small ehromatophores; skull transparent,
showing outline of brain (not impossibly a result of preserva-
tion ) ; 20.4 mm. bokermanni sp. nov.

8. Eye with a dark tunic; no anal decoration; tympanum partly

covered, directed nearly vertically upward (and slightly back-
ward); heel to front edge of eye; head short, nonspatulate;
venter smooth ( ? ) ; dorsum smooth no granules evident on
under side of thigh (?); color nearly uniform lavender, the
individual ehromatophores not visible to eye; 20 mm.,

divaricans sp. nov.
Eye with a white or cream tunic; ehromatophores large not
equally distributed but tending to segregate; venter and thigh
distinctly granulate 9

9. Interorbital area elevated and sloping rather abruptly to base

of snout; nostril nearer eye than to mid-point on lip; anal dec-
oration lacking; heel reaching to tip of snout or slightly be-
yond; ehromatophores segregated into an indefinite pattern
of spots or groups; 23.5 mm dubia sp. nov.

Interorbital area not elevated 10

10. Smaller, a yellow-cream, spot almost covering tympanum; upper
eyelid heavily pigmented with purple; lip bordered by a row
of cream tubercles; upper arm very slender; forearm greatly
thickened, permanently flexed (?); heel to tip of snout; very
short anal flap; a pair of enlarged postanal granules; supra -
scapulae not especially broadened; 20 mm vanzolinii sp. nov.

Larger, no yellow or cream spot on tympanum; spatulate-headed;
tympanum small at least partly distinct, directed outward and
upward; heel reaching to a point much beyond snout; no row
of tubercles on lip; forearm not especially thickened; supra-
scapulae very broad, prominent; 27 mm albotunica sp. nov.

Cochranella stirda sp. nov.

(Fig. 1)

Type: U. S. National Museum No. 96916, Passo Quatro, Minas
Geraes, Brasil; Zikan, coll., November 1920.

Paratype: U.S.N.M. No. 96917. Same locality and collector.

Diagnosis: A short-headed species, the eyeball blackish, the con-
cealed parts surrounded by a dark tunic; pigment where present
equally distributed, not reticulated; tympanum hidden under skin,
the tympanic region nearly vertical; distance between orbits equals
width of an eyelid; nostrils nearer to eyes than to median point in
upper lip; dorsal skin finely corrugated, belly strongly granular;
outer fingers with a web vestige ( one-fifth webbed or less ) ; terminal
discs oval, the sides and tips curving, not truncate; first and second
finger of approximately even length; a pair of prominent anal pads;
a strong inner tarsal fold; a distinct heel ridge; heel to anterior edge
of eye.

Family Centrolenidae from Brasil

1631

Description of type: Head width (8 mm.) greater than head
length (7.1 mm.); region about nostrils swollen but no depression
evident on snout or on top of head; can thai region short, the canthus
rostralis absent, the loreal region not or but slightly concave, sloping
oblique1)7 to edge of lip; tip of snout, seen in profile, sloping some-

Fig. 1. Cochranella surda sp. nov. Type, U.S.N.M. No. 96916.
Passo Quatro, Minas Geraes, Brasil.

what, the nostrils not terminal; tympanum hidden under skin with
no trace of its outline; a fold above tympanic region faintly indi-
cated; eye moderate, prominent, directed somewhat forward, its
length (2.6 mm.) greater than length of snout (2.3 mm.); nostril
nearer the eye than median point on upper lip, which is not
notched; width of an eyelid (2 mm.) equals interorbital width;
region back of eyes not or but very slightly constricted.

Choanae longer than wide; width of one (.4 mm.) in distance
between them (1.8 mm.) 4/2 times; a strongly defined transverse

73—6490

1632 The University Science Bulletin

groove for openings of palatal glands close to anterior edge of
choanae; no trace of vomerine teeth; tongue broader than long, not
or but slightly free behind, slightly emarginate posteriorly; open-
ings of vocal sacs short, diagonal.

Wrist extending beyond tip of snout; a very slight axillary web;
first finger distinctly smaller and shorter than second; well-developed
pads or discs on fingers, the tips distinctly wider than digits, the
tips rounding rather than truncate, the sides rounded rather than
angular; the discs transversely oval rather than subtriangular; no
trace of web between first three fingers; a trace between fourth and
third with a slight ridge following outer edge of third digit; a
large broad metacarpal tubercle, and a well-defined, oval, palmar
tubercle.

Heel to middle of eye; toes with terminal discs smaller than
fingers, the first two fingers Vs webbed; the second and third, %; the
third and fourth, %; the fourth and fifth, % webbed; an ill-defined
inner tarsal fold; a small inner metatarsal tubercle; no distinct outer
tubercle.

Skin of dorsum, seen under a lens, minutely corrugated; chin and
breast smooth; venter strongly granulate, under surface of thigh
granular; anal opening high; a short anal flap with a broad free
edge; a pair of large postanal pads or tubercles, with some vertical
wrinkling flanking them laterally.

Color: Probably green in life. In preservative transparent flesh
with dorsal surface of body, head, and upper surfaces of limbs
covered with nearly equally distributed, fine, purplish-lavender
chromatophores; edge of lip lacking pigment; ventral and concealed
surfaces transparent, whitish-flesh; eyelids surrounded by a dark
purplish tunic; iris purplish.

Measurements in mm. of type and paratype

Number 96916 96917

Snout to vent 21.5 24

Width of head 8 8.5

Length of head 7.1 7.5

Length of eye 2.6 2.7

Length of snout 2.3 2.3

Arm 17 16

Hand 8.8 7.8

Leg 33.2 37.4

Tibia 11 10.4

Foot and tarsus 14 18

Family Centrolenidae from Brasil

1633

CochraneUa eurygnatha (A. Lutz)

(Fig. 2)

Hyla (HyleUa) eurygnatha A. Lutz, Compt. Rend. Mem. Soe. Biol., Paris, vol. 93,
no. 21, June 19, 1925, pp. 137-138 (type locality, Serra da Bocaina, Brasil);
Trabalho Inst. Oswaldo Cruz, Mar. 10, 1926, pp. 5, 12.

Centrolenella eurygnatha B. Lutz, Copeia, no. 4, 1947, p. 243.

Diagnosis: Snout short, not spatulate, the nostrils nearly terminal-
eyeball surrounded by a whitish tunic; tympanum concealed under
skin; belly strongly granular; outer fingers not more than one-
fourth webbed, the discs much widened, curved rather than trun-
cate; all digits with terminal grooves on hand and foot; heel to nos-
tril; anal flap rather broad, notched mesially, flanked behind by
indefinite, flattened, anal pads, with some few enlarged granules on
ventral surface; no outer tarsal tubercle; a deep groove across palate
for openings of palatal glands.

Fig. 2. CochraneUa eurygnatha (A. Lutz). No. 7926 Coll.
Departamento de Zoologia, Secretaria de Agrieultura Sao Paulo
Brasil; Itatiaia, Rio de Janeiro, Brasil. Actual length, snout to vent,
approximately 22 mm.

1634 The University Science Bulletin

Description of species: (From U.S.N.M. No. 96656 g from Bonito,
Serra da Bocaina, State of Rio de Janeiro, near the Sao Paulo bound-
ary, late February 1928, Dr. A. Lutz, B. Lutz, and J. Venancio,
colls.). Head width (8 mm.) greater than head length (7.1 mm.);
snout very short, truncate, the nostrils nearly terminal; canthus
rostralis somewhat swollen but only a meager depression between
canthi; tympanic area lateral, the tympanum concealed under skin;
the supratympanic fold not or only indistinctly indicated; length
of eye opening (2.4 mm.) about as long as the snout (2.5 mm.);
the eye directed somewhat forward, and prominently elevated; dis-
tance between eye and nostril (1.8 mm.) nearly equal to distance
between nostril and median notch on lip (1.82 mm.); interorbital
interval (2.1 mm.) about equal to width of eyelid (2.2 mm.); dis-
tance between nostrils about equal to their distance from eye.

Choanae large, the distance between them 2.7 mm.; diameter of
one choana in distance between, 3.5 times; a distinct deep trans-
verse palatal groove crosses palate a short distance in front of
choanae, almost reaching choanae on outer ends; the edges of the
groove may be thickened and the skin in front seems loose; tongue
broader than long, slightly emarginate behind, and free behind for
approximately a fourth of its length; vocal sac indicated externally
by longitudinal folding of the skin on chin and neck; vocal slits run
somewhat diagonally from near jaw forward and across to tongue.

Arm with wrist reaching considerably beyond tip of snout; first
finger shorter than second, but digits of nearly same width; fingers
bearing broad terminal pads with terminal grooves, the discs dis-
tinctly wider than the digits, and delimited; the tips moderately
rounded rather than sharply truncate; about one-fourth webbed
between outer fingers; web absent between other fingers; a large,
rather broad, inner metacarpal tubercle; a prominent rounded
median palmar tubercle; subarticular tubercles distinct; super-
numerary tubercles on palm and sole; heel reaches to nostril; when
legs are folded at right angles to body, heels overlap 2.5 mm.; when
laid along body, heel and elbow overlap; the presence or absence
of an inner tarsal fold cannot be determined; inner metatarsal
tubercle present, outer absent.

Skin everywhere smooth on dorsal surfaces; chin with surface of
vocal sac longitudinally folded; breast smooth; venter covered with
rather large granules; median part of ventral surface of thigh with
a few granules, some distinctly larger than others; vent high, cov-
ered with a short, rather wide flap, notched mesially, flanked be-

Family Centrolenidae from Brasil 1635

hind by two, irregular, large, flattened but relatively distinct anal
pads.

Color: Flesh color, with lavender-purple pigment in small chro-
matophores, their distribution on dorsal surfaces of body nearly
equal, not segregated into spots or a reticulum; chromatophores
sparse on sides of head; venter and concealed parts of limbs flesh-
color, probably transparent in life; tunic surrounding concealed
parts of eyeball, cream-white, the iris white with purple streaking
or clouding; anal flap pigmented. The species is probably green
on exposed surfaces in life.

Measurements in mm.: Snout to vent, 22; width of head, 8; length
of head, 7.1; length of eye, 2.4; interorbital width, 2.1; arm, 16;
hand, 8; leg, 35.5; tibia, 11; foot and tarsus, 17.

Remarks: All the specimens of the topotypic series (U.S.N.M.
nos. 96652-96661 ) are of the same species, all males except for a
single small female (No. 96662). Tadpoles associated under the
number 96663 seem to belong to more than a single species some of
which may belong to this species. (See discussion of these else-
where. )

The original description of this species is as follows: "Hyla
(Hylclla) eurygnatha; L. 17 mm. — Le dessus est vert, passant au
jaunatre au museau, aux doigts, aux orteils et sur une ligne mar-
ginale. Tete courte, elargie derriere les yeux, ce qui est du a la
largeur de la mandible. Iris or mat; lobule superieur et inferieur
au bord de la pupille. Pointille noiratre plus accentue apres la
mort. Membrane court entre les doigts, plus longue entre les
orteils. Un seul exemplaire provenant de la Serra da Bocaina."

The drawing is of a specimen recently forwarded by Mr. Werner
Bokermann (No. 7926) from Itatiaia, Brasil. The specimens re-
cently preserved show more pigment, especially on eyelids, and
there are some indistinct lighter spots, which may have been more
evident in life. The fingers may be a trifle less than one-fourth
webbed.

Cochranella uranoscopa L. Miiller

Hyla (HyleJla) uranoscopa L. Miiller, Zool. Anz., Bd. 59, 1924, pp. 234-235
(type locality, "Humboldt [Flussgebeit des Rio Novo], Santa Catharina",
S. E. Brasil; Bavarian States Herpetological Collection No. 81/1921, W.
Ehrhardt, coll. Nov. 1919).

Diagnosis: Tympanum hidden; a large angular pollex-rudiment.

Description of the type: The frog appears very strongly de-
pressed, at the same time proportionally slender; width of body
is contained two and one-half times in the snout-vent length; head

1636 The University Science Bulletin

rather large (contained two and three-fourths times in the snout-
vent length), as long as broad and very flat; head broadest in region
of eye and narrows itself from eye to region back of tympanum, so
that it appears to be set off from body by a slight necklike constric-
tion. The limbs are rather long and slender; choanae moderately
large; vomerine teeth lacking; the tongue oval, completely mar-
ginate, only free on edge; snout longer than eye, broadly rounded
anteriorly; canthus rostralis not present and loreal region slopes
gently out and down; eyes not lateral but are obliquely directed up-
ward; nostrils directed upward; the interorbital space equals the
length (diameter) of the eye; distance of nostrils from tip of snout
equals two thirds its distance from eye; tympanum hidden under the
skin; fingers slender, slightly flattened, with moderately large an-
teriorly truncate finger discs; first finger somewhat shorter than the
second; on its basal part a moderately large angularly protruding
flat spreading pollex-rudiment is in evidence; between third and
fourth fingers one-third webbed; between the second and third
somewhat less than one-third webbed; between the first and second
fingers the web is vestigial.

Toes moderately long, likewise flattened, half webbed; subarticu-
lar tubercles on fingers and toes weakly developed; a distinct, elon-
gate, strongly compressed metatarsal tubercle present; hind leg
pressed against body, the tibiotarsal articulation reaches tip of snout;
tibia somewhat longer than femur, measures one and two thirds the
head length.

Skin of the dorsum finely granular; that of the ventral side strongly
granular. A fold goes from back end of eye, over tympanum to in-
sertion of upper arm.

Color: Color of the head and dorsum pale, characteristically trans-
parent, bluish green which laterally becomes more yellowish; the
limbs yellowish; entire upper side, with the exception of upper arm
and parts of hind leg that are concealed when crouched, is covered
with thickly distributed brownish-gray spots consisting of thickly
distributed punctations; underside bright olive-yellow, transparent
on the belly.

Measurements in mm: Head-body length, 25; head length, 9;
head width, 9; length of hind leg, 46; length of tibia, 14; tibiotarsal
articulation to tip of longest toe, 19.
(Data from type description).

Family Centrolenidae from Brasil 1637

Cochranella petropoJitana sp. nov.

Type: U. S. National Museum No. 101135, Petropolis, Rio de
Janeiro, Brazil; A. Lutz, Bertha Luth and Doris Cochran, colls.,
May, 1935.

Diagnosis: Head rather broad, somewhat spatulate, body de-
pressed; tympanum large, very distinct, lateral, relatively close to
jaw, directed outward; no specialization of fold, grooves or tuber-
cles in anal region; the vent high, the anal flap short and wide; pig-
ment where present scattered nearly equally on dorsal surfaces, with
no tendency to segregate and form spots or reticulations; none, or
only an ill-defined fold above tympanum; eye tunic dark.

Description of type: (The specimen is indifferently preserved
and in the region of the tip of the snout and on hand and foot the
characters are somewhat obscured.) Small frog, snout-to-vent
length 23.5 mm.; head rather spatulate, widest part being opposite
eye, narrowing a little near jaw angle; head depressed, the canthus
not, or barely indicated, rounded, sloping obliquely to lip, with a
shallow depression behind nostril in loreal region; area about nostrils
slightly swollen, however no depression is indicated between swell-
ings; tympanum distinct, not covered with skin or pigmented; its
vertical diameter (1.1 mm.) a little greater than its length (1 mm.),
separated from eye by a distance equaling the greatest diameter;
length of eye (estimated 2.4 mm.) less than length of snout (esti-
mated 3.5 mm.); tongue somewhat wider than long; vocal slits
present in mouth, but no external evidence of vocal sac; no vomerine
teeth; choanae large, length equal to half distance between them.

Arm with wrist extending considerably beyond tip of snout; first
finger as long as second or ( on right hand ) a little longer than sec-
ond; tips of fingers widened into truncate discs, distinctly wider than
digit; inner fingers without web; two outer fingers about one-third
webbed; inner metacarpal tubercle moderately distinct; median
palmar tubercle subcircular, distinct.

Leg slender, tibiotarsal articulation extending two millimeters
beyond tip of snout; when legs are flexed at right angles to body,
heels overlap; when flexed on side of body, knee and elbow over-
lap; toes three-fourths to four-fifths webbed; the terminal discs
truncate, smaller than those on fingers; inner metatarsal tubercle
distinct, outer absent or obsolete.

Skin smooth on the dorsal and concealed surfaces; chin and
breast smooth; venter longitudinally striate; no granules evident on
ventral face of thighs; no distinct fold above tympanum.

1638 The University Science Bulletin

Color: Probably green or olive in life, the venter transparent; in
preservative flesh white with a thick scattering of lavender or pur-
plish pigment, the chromatophores of nearly equal size; on head,
pigment less dense on lips and loreal region; there is a narrow strip
of chromatophores on surface of upper arm and upper surface of
thigh, wider areas of chromatophores on forearm and tibia, the pig-
ment extending onto outer finger and toe; a small area below vent
pigmented; eyeball above, showing dark pigment through upper
eyelid, the iris likewise being dark.

Measurements in mm. of type: Snout to vent, 23.5; greatest width
of head, 8.6; width at jaw angle, 7.3; length of head, 7.6; arm, 16;
hand, 8; leg, 41; tibia, 14; foot and tarsus 17.5.

Remarks: The specimen is not well preserved and the character
of the subarticular tubercles, the eyelid and the length and shape
of the snout in front of the nostrils cannot be determined satis-
factorily. It is interesting that the specimen was taken at the same
place as the type of Cocliranella lutzorum. Both are rather similar
in general appearance, but the following specific differences are in
evidence:

1. Tympanum distinct, low, lateral, 1. Tympanum partially concealed

directed outward. under skin, and directed upward.

-. No anal folds or tubercles. 2. A pair of vertical folds extend

across the posterior face of thigh
behind vent to ventral surface.
3. Pigment rather equally distributed 3. Pigment forming a reticulum over
on dorsum; pigment on upper arm. dorsum, absent from upper arm.

Cocliranella lutzorum sp. nov.

Type: U. S. National Museum No. 101134, Petropolis, Rio de
Janeiro; colls., A. Lutz, Bertha Lutz, and Doris Cochran; May,
1935.

Diagnosis: A moderately large species of the genus with a broad
spatulate head, the front of the snout sloping obliquely to lip in
front of nostrils; tympanum covered with skin; the outline partly
concealed; a pair of vertical folds behind vent; no pigment on
upper arm.

Description of type: Area about nostrils swollen with a depres-
sion between continued on down to edge of lip and bordered by
two distinct elongate elevations; nostrils separated from eye by a
distance of 3 mm., from median edge of lip by 2.6 mm.; canthus
rostralis absent; a longitudinal depression midway of the loreal
region, the lower part sloping very obliquely to lip; tympanum
covered with thin skin but its entire outline distinct, directed up-

Family Centrolenidae from Brasil 1639'

ward, separated from the eye by a distance equal to VA times di-
ameter of tympanum; length of eye, 2.65 mm., a little more than 2M
times greatest diameter of tympanum, but much less than snout
length (5 mm. ); upper lip indistinctly notched; no trace of vomerine
teeth; choanae moderately large, longer than wide, the distance
between them (2.2 mm.) contains width of one (.5 mm.) a little
more than four times; tongue large, much wider than long, not of
but only a narrow fringe free behind; (vocal sacs present in males?).

Arms slender, the wrist reaching distinctly beyond mouth; the
digits moderately slender, elongate, with widened truncate terminal
discs; web lacking between first three fingers; about one-fourth
webbed between the two outer fingers; a flattened protruding inner
metacarpal tubercle; a prominent median rounded palmar tubercle;
subarticular tubercles moderately distinct. Leg long, the heel ex-
tending 2 mm. beyond tip of snout; discs on toes smaller than those
on fingers, truncate; the first toe with web reaching half its length;
on outer side of second three-fourths webbed, third and fifth toes
two-thirds webbed, the fourth toe with two and a half joints free;
subarticular tubercles small but distinct; inner metatarsal tubercle
small, slightly projecting; outer not distinctly indicated; the char-
acter of the tarsal fold cannot be determined but seemingly present;
a small fold or ridge on heel.

Skin smooth on dorsal surfaces; chin and breast smooth; anterior
part of venter finely striate longitudinally, the posterior part broken
up into somewhat elongate granules; on under side of thigh large
round granules indicated on posteroventral face; vent high, covered
with a short pigmented flap; a pair of high vertical ridges curve
across the thighs to ventral surface. Scapulae large, broad, ele-
vated, their inner edges forming two irregular elevations which
seemingly are discernible in life since the edge marks an elongate
spot following the elevated portion.

Color: (In preservative.) Flesh white with lavender pigmenta-
tion unequally distributed; tip of snout and lips with sparse un-
equal pigmentation; top of head and dorsum with some segregation
of chromatophores leaving small irregular pigmented areas; eye
with whitish tunic; upper arm lacking pigment or at most a
few chromatophores; forearm pigmented, the pigment continuing
sparsely on outer finger to near tip; a narrow line of chromatophores
on upper face of thigh, wider on tibia continuing sparsely on tarsus
and two outer toes. Entire ventral surface flesh white (probably
transparent in life ) .

1640 The University Science Bulletin

Measurements in mm.: Snout to vent, 28; width of head, greatest,
10.3; width at jaw angle, 9.3; length of head, 8; diameter of tym-
panum, 1.05; length of eye, 2.6; length of snout, 5; arm, 18; hand, 8;
leg, 46; tibia, 16; foot and tarsus, 21.

Remarks: The species is dedicated to Dr. Bertha Lutz and her
illustrious father, Dr. A. Lutz.

Cochranella delicatissima sp. nov.

Type: U. S. National Museum No. 96481, Angra dos Reis, Rio
de Janeiro; A. Lutz, coll.

Diagnosis: Snout short not spatulate the nostrils nearly terminal;
flesh of body very white with a very fine scattering of lavender pig-
ment; tympanum pigmented, visible, directed laterad; interorbital
space wider than an eyelid; anal decoration probably absent, the
pelvis extending a little behind level of thighs?; heel to beyond tip
of snout; outer fingers about one-fourth webbed.

Description of type: (where characters are somewhat obscured
the word "probably" is used. ) Width of head, ( 7.8 mm. ) greater
than length (6.5 mm.); the canthus barely indicated; the tip of
snout in lateral profile curving slightly to lip, the nostrils not quite
terminal; loreal region slightly concave near eye, sloping obliquely to
lip; tympanum distinct, pigmented, directed laterally outward and
perhaps a trifle upward; its distance from eye double its greatest
diameter; eye large, its length ( 3 mm. ) distinctly greater than snout
length (2.1 mm.); canthus rostralis barely indicated; the medial
loreal region slightly concave near eye, sloping obliquely to lip; area
about nostrils slightly swollen with a slight depression between,
more or less evident down to lip; nostril as near to eye as to median
point on upper lip; width of upper eyelid less than interorbital
interval.

Choanae large, the greatest diameter of one, contained in dis-
tance between them about two times; groove for palatal glands
absent or obsolete; openings of vocal slits elongate, diagonally
placed; tongue wider than long, narrowed posteriorly, not or but
slightly emarginate, free only on posterior edge; no trace of vomerine
teeth.

Arm long, more than half of forearm extends beyond tip of snout;
a small inner metacarpal tubercle; a rounded palmar tubercle; outer
fingers one-fifth webbed; other fingers without or with only a trace
of web; discs present (the tissues of the fingers have partially
sloughed off so details of the discs and webs are not wholly clear);
first finger shorter than second; heel to slightly beyond tip of snout;

Family Centrolenidae from Brasil 1641

probably no inner tarsal fold; small metatarsal tubercle, outer tuber-
cle absent; toes probably three-fifths webbed.

Skin smooth on dorsal surfaces; venter with distinct fine granula-
tions; probably no anal decoration but the sacrum extends slightly
behind level of thighs on median line.

Color: Everywhere flesh white with a thin regular scattering of
lavender in tiny chromatophores (in many places the skin has been
rubbed and the color is seemingly absent); tunic about concealed
parts of eye at least for the most part whitish. Iris light with some
light lavender; pigment confined to dorsal surfaces but extending
narrowly on limbs to outer fingers and toes.

Measurements in mm.: Snout to vent, 20.8; width of head, 7.8;
length of head, 6.5; length of eye, 3; length of snout, 2.1; arm, 15.5;
hand, 7.2; leg, 35.5; tibia, 11.4; foot and tarsus, 15.7.

Cochranella bokermanni sp. nov.

Type: Departamento de Zoologia, Secretaria de Agricultura, Sao
Paulo, Brasil, No. 328, Itatiaia, Rio de Janeiro, May, 1906, Leuder-
waldt, coll.

Diagnosis: A diminutive species, the head not spatulate, the
dorsal skin glassy smooth; the chin smooth, the venter and under
face of thigh granular, the granules large and irregular on thigh
and posterior part of venter; a pair of slight swellings behind vent,
widely separated; tympanum covered, its outline faintly visible;
neck not constricted; heel reaching to between eye and nostril; web
vestige between fingers two and three, a larger vestige between
three and four. A slight fringe on outer edge of third; inner toes
about one-half, or less webbed; outer toes three-fifths to three-
fourths webbed; first finger much shorter than second.

Description of type: Head short the width (7 mm.) greater than
length (6.1 mm.); canthus absent, the interorbital area rather flat
and transparent showing the complete outline of the brain; snout
somewhat oval rather than rounded, seen from above; nostrils nearly
terminal, minute, equidistant from eye and the median point on
upper lip; nostril not noticeably swollen and no depression occurs
between; eyes large, directed somewhat forward, the length (2.45
mm.) greater than length of snout (2 mm.); tympanum covered
with skin its outline partly visible, the loreal region sloping obliquely
to lip, not or scarcely concave; snout extending about .75 mm. be-
yond mouth; width of an eyelid (1.8 mm.) less than interorbital
interval (2.1 mm. ) ; choanae moderate, the diameter of one ( .7 mm. )
in distance between them approximately 2.2 times. Eye with a

1642 The University Science Bulletin

brown tunic ( the eye possibly red in life ) ; palatal glands open into
a nearly straight groove, nearer the choanae than front of palate;
tongue rather thick, wider anteriorly than posteriorly, free for less
than one fifth of its length, unemarginate behind; opening of Eu-
stachian tubes equal to choana; (specimen a female lacking vocal
slits ) .

Wrist reaching beyond snout tip; first finger shorter than second,
with no trace of a web between them; between second and third
fingers a distinct vestige and a larger vestige between two outer
fingers ( less than one-fifth webbed ) ; tips of digits widened, sharply
truncate, the subterminal "discs" subtriangular; subarticular tuber-
cles single, moderately distinct; a small inner metacarpal tubercle,
the palmar tubercle irregular, large; an indistinct fold on outer finger
continued along the outer, under edge of arm.

Legs slender the heel reaching to nostril; inner toes less than half
webbed, the outer nearly three-fourths webbed, subarticular tuber-
cles indicated but there are no distinct supernumerary tubercles on
sole or palm; a small, inner metatarsal tubercle and an indistinct
outer tubercle; a distinct inner tarsal fold. Outer fold, if present,
very indistinct.

Skin shining smooth above on head and dorsum; chin and breast
smooth; venter with smaller granules anteriorly, larger more irregu-
lar ones posteriorly; under side of thigh with still larger irregular
granules; presumably a pair of small pustular swellings behind
vent. Ana' Rao short, wide, the vent opening at upper thigh level.

Color: ( Faded in preservative ) . Whitish flesh above and whiter
below; a small pigmented area visible on right side, the brownish
lavender pigment scattered; eyelid appearing dark, taking color
from the tunic of eyeball.

Measurements in mm.: Snout to vent, 20.4; width of head, 7;
length of head, 6; length of eye, 2.45; length of snout, 2; arm, 18.4;
hand, 7; leg, 32; tibia, 11; foot and tarsus, 15.

Remarks: The exposed brain is clearly visible, the posterior part
being much widened, a little more than double that of the cerebral
hemispheres and equal to the brain length. (The width is 3.5 mm.
the greatest length 3.5 mm. )

The relationship of this species is probably closest to Cochranella
surda, but seemingly differs in the greater width of an eyelid in
relation to interorbital interval, in having the skin glassy smooth
instead of being finely corrugated, and in having the first and sec-
ond finger of nearly equal length, as well as other less obvious
differences. ••

Family Centrolenidae from Brasil 1643

The species is dedicated to Mr. Werner Bokermann of the Mu-
seum of the Departamento do Zoologia who has given us the privi-
lege of studying this form and Cochranella vanzolinii.

Cochranella divaricans sp. nov.

Type: U. S. National Museum No. 96562 $ , Serra da Bocaina;
A. Lutz coll.; Jan. 2-19, 1930.

Diagnosis: A small frog with a short head; lavender pigment in
large spreading chromatophores that are contiguous or nearly so;
eyeball with a dark tunic; the interorbital interval probably ap-
proximately equal to width of an eyelid; heel to front edge of eye;
tympanum covered (outline partly visible); moderate, necklike
constriction behind head; outer fingers one-fourth webbed; toes
three-fifths to three-fourths webbed, the digital disks sharply trun-
cate and angular, the pads subtriangular in shape.

Description of type: A small species, the type being 20 mm. in
snout-vent length; width of head ( 7.2 mm. ) greater than length of
head (6.3 mm.); tympanum covered with pigmented skin but on
one side its outline is dimly evident, showing it directed largely
upward and slightly backward; the distance of tympanum from
eye at least once and a half the diameter of the tympanum; eye
large, directed partly forward, its length (2.5 mm.) greater than
snout length (2 mm.); nostrils slightly injured (presumably by
insects ) , probably swollen somewhat, although there is no evidence
of a depression between them or on snout, and probably equidistant
between median point on lip and eye; canthus rostralis barely indi-
cated, rounded; loreal region not concave, but sloping rather ab-
ruptly to lip; eyelid probably nearly equal to interorbital distance;
a necklike constriction behind head and partly including back part
of head.

Choanae large, irregular, their greatest diameter contained in
distance between them about two and one-third times; no trace of
vomerine teeth; a pair of vocal slits somewhat diagonally placed;
tongue (distorted) wider than long, free for at least one fourth of
its length (membranes of the palatal region have sloughed so that
the character of the palatal groove is not discernible, and the bony
rim of the choanae is exposed ) .

Arm brought forward the wrist reaching a little beyond tip of
snout; first finger shorter than second; digits with sharply truncate,
angulate discs, considerably wider than the digits; the pads on the
disc subtriangular in shape; web vestigial between first and second
fingers; approximately one-fifth webbed between second and third,

1644 The University Science Bulletin

and one-fourth webbed between third and fourth; a well-deveJoped
inner metacarpal tubercle; without trace of a pollical spine; a
rounded palmar tubercle.

Heel reaching to a point immediately in front of eye; leg slender
when folded at right angles to body, the heels overlapping 2 mm.;
webbing between the toes reaches three fifths to three fourths of
the length of digits except on the fourth; discs at tip much wider
than the digits, and only a little smaller than those on fingers; a
small, flat, inner metatarsal tubercle (two indistinct tubercles on
posterior median part of sole ) ; no outer metatarsal tubercle.

Skin above smooth; venter seemingly smooth, definitely not striate
but some granules may have been present in life; no distinct anal
decoration, and no granules evident on the under side of the thigh.
Subarticular tubercles not strongly marked.

Color: Above on body uniformly lavender, the pigment spread,
rather than segregated in small dots, although on dorsal surfaces
of limbs the pigment is in rather discrete dots; very little pigment
on upper arm; venter and concealed surfaces of the limbs and body
transparent flesh; under a lens one sees numerous minute lighter
(probably white in life) areas scattered on dorsal surface of body
and head; on limbs pigment extends to tips of outer digits.

Measurements in mm.: Snout to vent, 20; length of head, 6.3;
width of head, 7.2; length of snout, 2; length of eye, 2.5; arm, 13;
hand, 6.3; leg, 31; tibia, 10.5; foot and tarsus, 13.

Cochranella dubia sp. nov.

Type: U. S. National Museum No. 96722, Serra da Bocaina,
boundary of Rio de Janerio and Sao Paulo, By A. Lutz; Dec, 1930-
Feb., 1931.

Diagnosis: A spatulate-headed form with the interorbital area
elevated, and sloping rather abruptly to base of snout; nostril a
little closer to eye than median point on upper lip; tympanum dis-
tinct, directed upward; concealed part of eyeball with a white
tunic; eye large, its length equals snout length; eyelid distinctly
less than interorbital distance; probably no especial anal decora-
tion, but the pelvic bones push back forming a decided posterior
median protrusion behind level of thighs; heel reaches tip of snout
or minutely beyond; discs on hand and feet nearly same size, rela-
tively smaller than those of other spatulate-headed forms, their
shape subtriangular rather than transversely oval; venter probably
granular ( ? ) ; purplish pigment in large chromatophores segregated
into groups forming indefinite spots.

Family Centrolenidae from Brasil 1645

Description of type: Head somewhat spatulate, moderately wide,
somewhat narrowed at jaw angles, the width (7.8 mm.) greater
than length (7.3 mm.); canthus rostralis indistinct or absent, the
snout sloping obliquely to lip, the loreal region not or but slightly
concave; tympanum distinct, round, directed upward, its distance
from eye greater than its diameter; eye very large, somewhat di-
rected forward, its length (3 mm.) equal to length of snout (3
mm. ) ; areas about nostrils swollen, with a slight depression between
them; snout sloping forward obliquely in front of nostrils; width of
an eyelid distinctly less than interorbital width, the area between
the orbits and occiput elevated, sloping abruptly down to snout at
anterior level of eyes; choanae moderate, their longitudinal di-
ameter in distance between them, approximately two times; upper
rim of choanae followed outward terminates in a lobulate projec-
tion of bone; a strongly defined transverse groove in palate for
openings of palatal glands, much closer to choanae than to front
of palate; tongue thick, subcircular, posteriorly emarginate, free
for one fourth of its length and attached (seemingly) by a series of
elongate lamellate, muscular fibers; large diagonal vocal slits; no
trace of vomerine teeth.

Arm long, nearly half of forearm extends beyond tip of snout;
digits with relatively narrow terminal discs the shape of pad sub-
triangular rather than transversely oval; first and second fingers
equal, or second a trifle longer; digits slender, with a vestige of
web between first two fingers, about one-fifth (or less) webbed
between second and third; between outer fingers one-fourth
webbed; inner metacarpal tubercle relatively small; palmar tubercle
large, distinct, somewhat rounded; leg with heel reaching to or
minutely beyond tip of snout; discs on toes small most of them not
or scarcely wider than digit; probably an inner tarsal fold; inner
metatarsal tubercle small, outer absent; foot about three-fourths
webbed throughout; when limbs are folded at right angles the
heels touch but do not overlap; knee and elbow overlap.

Skin of body is smooth; probably a short distinct skinfold above
tympanum; suprascapulae not forming ridges; venter indistinctly
granular but granules are not regular; indication of some granules
under the femora; pelvis somewhat protuberant behind level of
thighs.

Color: Deep purplish, on body and head, and dorsal surfaces of
limbs, on a ground of flesh, the large chromatophores segregated
into irregular groups sometimes suggesting reticulation; (probably

1646 The University Science Bulletin

no pigment on upper arm); below flesh; (a few brownish areas on
venter and under side of limbs are probably due to discoloration).

Measurements in mm.: Snout to vent, 23.5; width of head, 7.8;
length of head, 7.3; length of eye, 3; length of snout, 3; arm, 17;
hand, 8; leg, 40; tibia, 13; foot and tarsus, 17.

Remarks: This species was taken in the Serra da Bocaina by
Dr. A. Lutz together with the species described as Cochranella albo-
tunica sp. nov.

Cochranella vanzolinii sp. nov.

(Fig. 3)

Type: Collection of the Departamento de Zoologia, Secretaria
da Agricultura ( Ipiranga, Sao Paulo, Brasil ) No. 2952, from Boracea,
Sao Paulo. Collected December 12-19, 1947 by P. Vanzolini and
W. Bokermann.

Diagnosis: A diminutive species; pigment not equally distributed;
head slightly spatulate; venter granular; tympanum distinct, di-
rected only slightly upwards and covered by a cream-yellow spot
size of tympanum; choanae small, diameter of one in distance be-
tween choanae, 4.5 times; upper arm very slender; forearm greatly
thickened; fingers flattened, truncate at tips, the tips only slightly
wider than digits, distinctly wider than toes.

Description of species: Snout-to-vent measurement, $ , 20 mm.;
width of head (7.7 mm.) less than its length (7 mm.); canthus
rostralis present but rounded somewhat, the loreal region shallowly
concave, sloping to upper edge of upper jaw, the remainder nearly
vertical to tip; area about nostrils a little swollen with a slight de-
pression between them, the nostrils not terminal but the snout
sloping obliquely to lip; tympanum small, its diameter ( .8 mm. ) con-
tained in length of eye (2.7 mm.) approximately 3.4 times; pupil of
eye horizontal the upper edge with a tiny rounded median projec-
tion above giving the pupil a dumbbell shape when contracted;
length of eye greater than length of snout ( 2.4 mm. ) ; nostril almost
equidistant between eye and median point on upper lip; eye directed
somewhat forward; covering of the concealed parts of the eyeball
white but eyelid itself heavily pigmented with dark purple; extent
of upper eyelid not clearly defined but definitely less than the inter-
orbital distance; choanae small, the greatest ( diagonal ) diameter of
one choana in distance between them, 4.5 times; openings of the
palatal glands form a sinuous groove close to anterior level of
choanae; tongue broadest anteriorly, narrower posteriorly, unemar-
ginate, free for less than a fourth of its length; vocal slits large;
vocal sac evident on chin, displaying ample folds.

Family Centrolenidae from Brasil 1647

Upper arm very slender, about one third thickness of forearm;
(the arm appears to be permanently flexed at elbow); fingers flat-
tened, nearly truncate the subterminal discs somewhat triangular
in shape; second finger distinctly longer than first; a web vestige be-
tween first and second, and second and third fingers; approximately
two-fifths webbed between the third and fourth; subarticular tuber-

V-Ir"

Fig. 3. Cochranella vanzolinii sp. nov. No. 2952. Type. De-
partamento de Zoologia, Seeretaria da Agricultura, Sao Paulo, Brasil.
Boracea, Sao Paulo. Actual length, snout to vent, 20 mm.

cles distinct and single; supernumerary tubercles present on digits
and palm; a well-defined inner metacarpal tubercle; a well-defined
single palmar tubercle; fringe on outer finger continued along arm
as a distinct fold and bearing a number of elongate white tubercles;
heel reaching to tip of snout; a distinct outer tarsal fold continuous
with the fringe on outer toe and bearing occasional elongate white
tubercles, terminating in a slight heel-fold; subarticular tubercles
distinct; a small inner metatarsal tubercle; and a small but very

74—6490

1648 The University Science Bulletin

indistinct outer tubercle; inner metatarsal fold indistinct. When
legs are folded the heels overlap 2.5 mm.; no axillary web; toes
nearly four-fifths webbed.

Skin generally smooth on dorsum; side of head and dorsolateral
region with a few scattered low tubercles, whitish or cream in color;
other similar small tubercles scattered on dorsal or exposed surfaces
of limbs and in region below and about vent; the chin, breast, and
venter closely granular, this extending to the pigmented area on
sides; some granulation on under surface of thighs; skin on each
side of vent seemingly swollen; a pair of strong, somewhat conical
tubercles or granules on ventral surface of thigh below vent; anal
flap short and broad, the vent opening near upper thigh level.

Color: In preservative, whitish with a pigmentation of lavender
or purple in numerous chromatophores, unequally distributed leav-
ing lighter reticulation and spottings most distinct dorsolaterally;
concealed part of eye with a white tunic, the eyelid heavily pig-
mented with purple; tympanum almost entirely cream-yellow; bor-
der of nostril cream, this bordered by somewhat heavier pigment.
Lip bordered by a series of fine elongate white or cream tubercles;
limbs colored like back with all tubercles cream-white; ventral sur-
face, sides, and concealed parts of limbs lacking pigment, possibly
transparent or colorless in life, now dull, dirty, yellowish white.

Measurements in mm.: Snout to vent, 20; width of head, 7.7;
length of head, 7; length of eye, 2.7; length of snout, 2.4; tympanum,
.8; arm, 12; hand, 7; leg, 37; tibia, 12; foot and tarsus, 17.

Remarks: Due to the flexed condition of the limb at the elbow
and knee joints it is impossible to straighten the limbs completely
without injuring the specimen. In consequence the limb measure-
ments are partially an estimation. The specimen however is in a
perfect state of preservation. The relationship is thought to be
closer to Cochranelhi albotunica and Cochranella dubia than to the
other forms in southeastern Brasil. Neither of these species how-
ever have the cream-white tuberculation, the cream-colored spot
covering tympanum, or the outer tarsal fold. Many other differ-
ences likewise are in evidence.

The species is named for Dr. P. Vanzolini of the Departamento
de Zoologia, Sao Paulo, Brasil, one of the collectors.

This is one of the most distinctive forms of the genus.

Cochranella albotunica sp. nov.

Type: U. S. National Museum No. 96559, Serra da Bocaina,
boundary of Rio de Janeiro and Sao Paulo, Brasil, A. Lutz, collector;
Jan. 2-19, 1930.

Family Centrolenidae from Brasil 1649

Paratypes: U.S.N.M. No. 96557, same locality, data and collector;
U.S.N.M. No. 96723, same locality and collector; Mar., 1927.

Diagnosis: Small frog, snout to vent 27 mm. Head broad, tym-
panum small, at least partly distinct, directed outward and some-
what upward; pigment segregated forming spots and reticulations;
very prominent, somewhat protruding anal pads below and partly
lateral to vent; suprascapulae very broad more or less distinct; head
somewhat spatulate the snout sloping forward from nostrils, which
are a little closer to eye than to median point of lip, nostrils large;
canthus rostralis not or scarcely indicated; outer fingers one-fifth
webbed; choanae large; no vomerine teeth; tongue one-fourth free;
venter granular; heel to much beyond snout.

Description of type: Head rather spatulate, its width 8.4 mm.
greater than length ( 7.3 mm. ) , narrowed in tympanic region;
canthus rostralis not or but vaguely indicated, the loreal region not
or but slightly concave, sloping very obliquely to lip; area about
nostrils swollen, with a distinct depression between them; nostrils
moderately large, the snout sloping obliquely to lip; tympanum
small, directed almost upward, its greatest diameter about .9 mm.;
length of eye, 2.8 mm. greater than its distance from nostril, but
shorter than the length of snout (3.2 mm.); eyeball surrounded
largely by a white tunic (seen easily on inside of mouth); greatest
width of an eyelid ( 1.8 mm. ) distinctly less than interorbital interval
(2.15 mm.); (probably a distinct fold above and partly covering
upper edge of tympanum); outline of suprascapulae more or less
visible, their greatest (longitudinal) width 4.1 mm. Choanae large
the greatest diameter of one in distance between the choanae about
2.2 times; no vomerine teeth; tongue a little wider than long, free
behind for about one fourth of its length; vocal slits present, some-
what diagonal.

Arm rather long the wrist extending beyond tip of snout; first
finger smaller and shorter than second; all fingers with terminal
discs distinctly wider than digits, the tips slightly rounded and not
or but little angular on sides; none or but a slight vestige of web
between first two fingers; between second and third one-fourth,
between third and fourth one-third webbed; a thick ridge follows
outer side of fingers to disc; a rather large metacarpal tubercle; a
rounded, prominent palmar tubercle.

Leg very long the heel extending 3 mm. beyond tip of snout;
discs on toes smaller than those on fingers; webbing of foot as
follows: between first and second, one-third webbed; between
other toes, approximately three-fourths webbed; a small inner meta-

1650 The University Science Bulletin

tarsal tubercle, no outer tubercle visible; probably no inner tarsal
fold; when legs are folded at right angles to body the heels do not
overlap; when pressed on side of body, elbow and knee overlap.

Skin on dorsum generally smooth; venter with small granules;
under surface of femur with some granulation; vent high followed
by a pair of relatively large postanal pads or tubercles.

Color: (In preservative.) Everywhere transparent flesh. Dor-
sum with purplish or lavender pigment, the chromatophores segre-
gated to form irregular small spots and reticulations; dorsal surface
of limbs, with pigment extending onto outer finger and onto two
outer toes; upper arm devoid of pigment.

Measurements in mm.

96559 96557 96723*

Snout to vent 27 24 22**

Head width 8.4 9.1

Head length 7,3 7.4

Length of snout 3.2 3.3

Length of eye 2.8 2.6 2.5

Arm 17 16 16,3

Hand 8.8 8.9 7.2

Leg 42 41 39

Tibia 14.2 15 13.3

Foot and tarsus 16,3 16.5 15

Remarks: The three specimens are indifferently preserved, and
one, No. 96723, has the front part of the face missing. All show the
same general distribution of the pigment, and all show the postanal
pads well developed. The character of the granular belly, spatulate
head, position of the nostril, and segregation of the chromatophores
in groups forming "spots" and reticulations are characteristic of this
form.

Cochranella parvula Boulenger

Hylella parvula Boulenger Proc. Zool. Soc. London, 1894, p. 646, pi. 40, fig. 3
( Lages and Theresopolis, Santa Catharina, Brasil. )

Diagnosis: A diminutive species (17 mm. snout-vent length);
tympanum distinct belly granular; heel to a point a little beyond
snout; upper eyelid very narrow, the interorbital space broad and
convex; head as long as broad.

Description of type: (From Boulenger).

"Tongue circular, entire. Head as long as broad; snout short,
rounded; no canthus rostralis; eye large and very prominent; upper
eyelid very narrow; interorbital space broad and convex; tympanum
distinct, hardly one third the diameter of the eye. Fingers dis-

* Tip of snout missing.
** Estimated.

Family Centrolenidae from Brasil 1651

tinctly webbed at the base, first slightly shorter than second; toes
two-thirds webbed; disks moderate. Tibio-tarsal articulation reach-
ing a little beyond the tip of the snout. Skin smooth above; belly
and lower surface of thighs granulate. Grayish or pale brown
above, speckled with white; lower parts white.

"From snout to vent 17 millimeters. Two specimens. One
from Lages, Santa Catharina, collected by Hr. Michaelis; The other
from Theresopolis, presented by Dr. Goldi.

"This species appears to be most nearly allied to H. carnea, Cope,
but the tympanum is perfectly distinct and the coloration is entirely
different."

Several pertinent characters are omitted from the description.
However it does not seem to be synonymous with any of the forms
described here as new.

DESCRIPTION OF TADPOLES

Associated with one lot of U.S.N.M. specimens (Nos. 96652-
96662) is a series of four tadpoles bearing the number 96663 from
the same locality. The tadpoles differ in size and there is no assur-
ance that they belong to this species, or that all the tadpoles are
conspecific. There is a very strong probability that they are
Cochranella tadpoles. To differentiate the four tadpoles they are
referred to as: A (37 mm.), B (30 mm.), C (24 mm.), and D
( 23 mm. ) .

The specimens are indifferently preserved, and that all the exter-
nal tooth rows are present cannot be affirmed.

Specimen A: The largest specimen has a somewhat elongate oval
body; the eyes dorsal and rather close together, their greatest diam-
eter 1.3 mm.; their distance from the tip of the snout .35 mm.; dis-
tance between nostrils 2.1 mm.; the distance from eye to nostril
(1.25 mm.) distinctly less than their distance to end of head (2.3
mm.). Spiracle sinistral its distance from anterior head level, 9.4
mm.; length of body from head to end of anal tube 13 mm.; from
vent to tip of tail, 24; mouth located very close to front level of head
on ventral surface ( median point on upper horny beak .5 mm. from
tip of snout); a large free frill surrounds lateral and posterior rim
of mouth, 1.1 mm. wide mesially, its edge denticulate; just posterior
to mouth the frill has certain fleshy ridges which may have borne
horny teeth (the posteriormost has some minute fleshy denticula-
tions); horny upper beak broadly curving, bearing on its edge a
series of fine denticulations (approximately 38); lower beak much

1652

The University Science Bulletin

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Fig. 4. Tadpole presumed to be that of Cochranella. Specimen A enlarged.

Family Centrolenidae from Brasil

1653

narrower with approximately the same number of denticulations
but of smaller size, those near the outer ends a litttle larger. Hind
limbs approximately 3 mm. long, the feet showing no development
of the toes. Dorsal caudal fin begins at a point about 2 mm. in front
of level of developing legs; anal opening definitely mesial; greatest
width of the caudal expansion 4.2 mm., the musculature, continued
to extreme tip, strongly attenuated.

Fig. 5. Teeth of tadpole presumed
to belong to Cochranella. Specimen
A enlarged.

Specimen B: This specimen (30 mm. total length) is younger,
presumably, since there is no external trace of developing hind legs;
the body is somewhat elongate-spatulate, and the eyes are very small
comparatively ( about .35 mm. in greatest diameter ) , while the dis-
tance between them is approximately 1.1 mm.; eye to tip of snout,
3.05 mm.; eye to nostril, 1.55 mm.; spiracle sinistral, its distance
from anterior head level, 7 mm.; snout to end of anal tube, 11 mm.;
snout to base of tail, 10 mm.; vent to tip of tail, 19 mm.; greatest
width of head, 6 mm.; body width at spiracle, 4.3 mm.; greatest
elevation of fin, 2 mm.; greatest height of tail, 4 mm.; width between
nostril, 1.9 mm.

1654

The University Science Bulletin

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Fig. 6. Tadpole, presumed to be that of a species of Cochranella.

Specimen B enlarged.

Family Centrolenidae from Brasil

1655

Mouth is surrounded on lateral and posterior parts with a free
frill. Mouth ventral, 1 mm., back from the end of the head. The
front upper part of the beak is slightly curved, the denticulations
strongly marked, small at outer edges becoming gradually larger,
then diminishing in size in the mesial region. The eight median
teeth smaller, all of approximately the same size. There is a total

Fig. 7. Teeth of tadpole presumed to be
a species of Cochranella. Enlarged.

of 32 denticulations. The beak on lower jaw is deep within the
mouth and likewise denticulate, but more strongly curved, the row
consisting of about 30 denticulations suddenly increasing in size
near the outer ends. Within the mouth and invisible when mouth
is closed are two rows of very fine denticulations the more posterior
nearly on a level but following the curve of the jaw, the more
anterior is slightly arched but likewise following the curve of the
mandible back of the beak.

The frill is denticulate on its edge, mesially, a millimeter wide
but narrowing a little laterally; a row of small papillae border edge
of lip and there are no fleshy ridges on the surface of the frill. At
no point does the width of the tail and its fin exceed 4 mm. eleva-
tion.* No pigmentation whatever is visible.

Specimen C and D have lost much of the buccal frill but while
one seems to agree with B in the character of the denticulation, the

* This specimen was accidentally destroyed before the characters of the ventral caudal
fin were ascertained.

1656

The University Science Bulletin

other suggests still a third form, as regards the denticulations on
the beak. Here the teeth are fewer, large, and not reduced in the
middle part of the series.

Map of southern Brasil showing type localities

□

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UNIVERSITY OF KANSAS

SCIENCE BULLETIN

UNIVERSITY OF KANSAS PUBLICATIONS
University of Kansas Science Bulletin - Vol. XXXV - Part HI

November 20, 1953

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Editor Edward H. Taylor

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UNIVERSITY OF KANSAS

Science Bulletin

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Volume XXXV, Part III

University of Kansas Publications

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24-8663

Contents of Volume XXXV, Part III

No.

PAGE

16. The Biology of a Leafcutter Bee (Megachile brevis) and

its Associates Charles D. Michener, 1659

MIS. GOMP. ZOBL

JAN 4 1954
KNIVES

THE UNIVERSITY OP KANSAS

SGIENGE BULLETIN

Vol. XXXV, Pt. Ill] November 20, 1953 [No. 16

The Biology of a Leafcutter Bee
(Megachile brevis) and Its Associates1

Charles D. Michener -

ABSTRACT

This paper presents a detailed account of the habits of the commonest leaf-
cutter bee in Kansas. It is shown that the bee increases greatly in abundance
during the warmer part of each year, there being two to four generations per
year. This species is very mobile, populations regularly dispersing from certain
habitats and new assemblages concentrating at other places as their food
sources (flowers) change during the year. Dispersal distances up to several
miles are thought to be common. Places of concentration are of two kinds,
( 1 ) nesting habitats where the sexes are equally abundant and ( 2 ) nectar
habitats which hold principally males although females may be found there.

The kinds of flowers used as pollen sources are systematically diverse. Most
of them are blue, purple, or whitish in color. No flower constancy was detected
in bees merely sucking nectar, but in those gathering pollen there is a strong
tendency for only a single kind of pollen to be gathered on any one pollen col-
lecting trip. Exceptions were noted, however.

Nesting sites are always in pre-existing hollows, but these may be almost
anywhere. Most are in weed stalks but some are in the ground or even in very
dense foliage. Cell structure is also highly variable, both petals and leaves
being used in most cells. The processes of cell construction, provisioning,
and egg laying are described in detail. Growth and development are also de-
scribed. In summer the period from egg laying to adult emergence is about
a month. The species overwinters as mature larvae.

The life history of the cuckoo bee, Coelioxys octodentata, the principal
parasite of Megachile brevis, is described.

A section on the possible importance of this leafcutter as a pollinator of alfalfa
is included and to a considerable extent serves as a summary.

1. Acknowledgment is made to the General Research Funds of the University of
Kansas for aid which made this study possible.

2. Contribution number 841 from the Department of Entomology, University of Kansas.

(1659)

TABLE OF CONTENTS

PAGE

Abstract 1659

Introduction 1662

Part I. Seasonal History and Habitats

Seasonal Fluctuations in Abundance 1664

Number of Generations Per Year 1666

Overwintering 1669

Dispersal and Favored Habitats 1669

Nesting Habitats 1670

Nectar Habitats 1675

Studies of Marked Individuals 1676

Part II. Relations With Flowers

Kinds of Flowers Visited 1679

Nectar Gathering 1682

Flower Constancy in Nectar Gathering 1683

Pollen Gathering 1684

Rapidity of Pollen Gathering 1686

Constancy in Pollen Gathering 1687

Part III. Miscellaneous Activities

Relation of Adult Activity to Environmental Factors 1689

Flight 1690

Miscellaneous Notes on Behavior 1691

Nocturnal Resting Places 1691

Male Reactions to Females 1691

Longevity of Adults 1692

Part IV. Nesting

Finding Nesting Sites 1694

Nesting Sites 1695

Nest Structure 1697

Number of Cells Constructed 1699

Cell Structure 1700

Duration of Trips 1710

Petal and Leaf Cutting 1714

Nest Construction 1719

Provisioning 1720

Egg Laying 1721

Flight Patterns Around Nests, etc 1722

Changes in Behavior 1724

Part V. Growth and Development

Technique of Study 1727

Egg 1728

Larva 1728

Cocoon 1732

Pupa 1735

Emergence 1735

Summary of Developmental Stages 1735

(1660)

Michener: Biology of a Leafcutter Bee 1661

Part VI. The Cuckoo Bee, Coelioxys page

Seasonal History and Habitats 1737

Finding Megachile Nests 1737

Egg Laying 1738

Larva 1738

Cocoon 1742

Emergence 1742'

Part VII. Natural Enemies of Megachile brevis

Summer 1743

Winter 1743

Part VIII. Possible Importance of This Bee As a Pollinator 1745

1662 The University Science Bulletin

INTRODUCTION

This work was undertaken in order to provide detailed informa-
tion on the bionomics of one of our commonest North American
leaf cutter bees. Such information should contribute to the study of
comparative behavior of bees, which ought to be pressed with the
same vigor as that of comparative morphology, for it will doubtless
improve our understanding of the relationships of various bee
groups. Such information will also serve as a biological basis for
efforts to increase the effectiveness of this bee as a pollinator of alfalfa
and other crops. The work was begun primarily with this latter
idea in mind for Megachile brevis is usually the commonest wild bee
in Kansas alfalfa fields that is effective in tripping the flowers. How-
ever, since the relationships with the environment were found to be
exceedingly complex and to vary from season to season, observations
were made on all aspects of behavior that could be investigated
with the hope that information of practical value in the effort to
increase the abundance of these bees could be obtained.

The work was carried on over a period of three years, 1950
through 1952. Much of it was done in the vicinity of Lawrence,
Kansas, but long series of observations at other points in the state
were made, as indicated in the text.

The observations are not those of any one person; field informa-
tion was gathered by several persons other than myself. Among
these I wish to mention particularly Jimmy R. White, Alvaro Wille,
Wallace E. La Berge, and Dr. Roland L. Fischer. Without their
help the studies here reported would have required many more
years.

Megachile brevis Say is a member of the subgenus Litomegachile.
It is one of a group of very closely allied species. Mitchell (1936)
separated the more distinctive species of Litomegachile but re-
tained three forms as "varieties" of brevis. One of these "varieties,"
nupta Cresson, is presumably nothing but a rare color variant of
brevis proper. If this is true, the name nupta ought to be placed in
the synonymy of brevis.

Another of the "varieties" does not seem to intergrade with brevis
and occupies a range from North Carolina and Mississippi to south-
ern Florida. It is entirely sympatric with brevis and must therefore
be regarded as a closely related but distinct species, M. pseudo-
brevis Mitchell.

Michener: Biology of a Leafcutter Bee 1663

The third "variety," onobrychidis Cockerell, has a wide range from
Nebraska and Texas to the Pacific. Over this entire area it appears
to intergrade completely with bees having characteristics of brevis
proper. It is therefore probable that a western subspecies, M. brevis
onobrychidis Cockerell, should be recognized, although some indi-
viduals of this subspecies are indistinguishable from brevis proper.

The bee upon which the series of observations described below
is based is therefore properly called Megachile brevis brevis Say.
The species ranges from the Atlantic to the Pacific and from southern
Florida and northern Mexico to Quebec and British Columbia. It
is thus one of our most widespread bees.

Although the details of the life history will be explained later, it
seems necessary to give certain background information here. Mega-
chile brevis, like other members of the family Megachilidae, is a non-
social insect having no worker caste. Each female constructs cells
made from pieces of leaves and petals, hence the name leafcutter bee.
These cells are placed in any of a wide variety of situations such as in
hollow weed stalks, in curled leaves, or in holes in the ground. They
may be single or in series, placed end to end. Each is provisioned
with a viscous mass of honey and pollen, sufficient to provide food
for the entire larval development. An egg is then laid on top of the
food mass and the cell is closed with a cap made from more pieces
of petals and leaves. In summer the time required for growth and
development from egg laying to emergence of the adult is about a
month, but larvae reaching maturity after about the middle of
August cease their development, passing the winter as mature
larvae which pupate and emerge as adults in the spring. The males
emerge at about the same time as the females and live about as long
as the latter, a situation unusual among bees.

1664 The University Science Bulletin

PART I. SEASONAL HISTORY AND HABITATS
Seasonal Fluctuations in Abundance

Megachile brevis can be seen on flowers throughout the warm
season of the year, in Lawrence from May 10 to October 1, and
doubtless earlier in very mild springs and later in warm fall seasons.
In southern Florida there are collecting records as early as February
9 and as late as November 22; it is possible that the species is active
throughout the year there.

Because of the mobility of populations of this bee, it is difficult to
obtain satisfactory data on the fluctuations in abundance during
the season. A particularly attractive patch of flowers may result
in a concentration of bees at almost any time, yet for miles around
there may be very few bees. When the attractive patch ceases to
bloom, the bees disperse, with the result that there is an apparent
(but almost certainly not a real) drop in population. Moreover,
the observers were often intent on watching nests or known con-
centrations of bees with the result that serious attempts to estimate
bee abundance over large areas were not made. Furthermore,
such factors as temperature, wind, and sunlight so affected the
activities of the bees that two days' observation rarely gave com-
parable results. In spite of these difficulties considerable informa-
tion on population fluctuations was gathered. For comparative
purposes, this was converted to average number of Megachile brevis
seen per half hour in the field.

In the vicinity of Lawrence this bee is not especially common and
in many places, particularly early in the season, not a single speci-
men would be seen. However, we soon learned the most satisfactory
places to look and the data is largely based on observations made
in these places, although as indicated above, this introduces irregu-
larities due to the ability of the bees to concentrate in favorable
areas.

In any event, near Lawrence, the first M. brevis were seen on
May 15 in 1950 (a not entirely fresh female which must have been
active for several days), May 22 in 1951, and May 26 in 1952. A
very fresh male in the Snow Entomological Museum was collected
near Lawrence on May 10, 1949. During May and the entire
month of June, the bees were scarce. This was particularly so in
the very cool, wet June of 1951. This was the wettest June on
record, with a total rainfall of 11.65 inches, resulting in great floods.
During this month there was no day recorded when the average

Michener: Biology of a Leafcutter Bee

1665

number of M. brevis observed per half hour exceeded four. During
the dry and relatively warm June of 1952, the bees were but little
more abundant, probably because of the small number produced
during 1951, but in the more normal but cool June of 1950, there
were days when ten bees per half hour could be observed.

Both in 1950 and 1952, marked increases in abundance were noted '
soon after July 1. In the warm dry season of 1952, this was par-
ticularly noticeable and by the last week in July areas existed where
one could hear the sound of Megachile flight at all times during the
day and could see as many as 60 individuals ( amount of duplication
unknown) per half hour. In 1951, however, doubtless because of
continued cool wet conditions and presumably poor reproduction
during the extreme June weather, no marked increase in Megachile
abundance was noted during July.

In August, 1950 and 1951, particularly the former, increases in
Megachile populations were noted, so that averages of 40 bees
observed per half hour were sometimes recorded in 1950, 20 in
1951. Although no more adults probably emerge after the first
week in September,1 this abundance was maintained through Sep-

Fig. 1. Diagram showing estimated average seasonal fluctuation in adult
abundance of Megachile brevis, together with probable seasons of adult emer-
gence and of egg laying.

tember each year. In 1952, the records suggest a different picture,
for by August 15, the bees seemed much scarcer than in late July
and early August, and this situation prevailed through September.
The year 1952, by contrast to 1951, was one of the dryest growing

1. A specimen emerged in the laboratory on September 5, 1952, and a very fresh
specimen was seen in the field on September 11, 1950.

1666 The University Science Bulletin

seasons on record and therefore by middle and late summer flowers
were very scarce. There is no doubt that this scarcity resulted in an
apparent reduction in the bee population by dispersing it as well
as in a real reduction due to the scarcity of food supplies. Counts
made in early September, 1952, in the most favorable locations and
in good weather, showed only six to ten Megachile per half hour.

Based on the three years of experience which included one extra-
ordinarily wet and cool summer (1951) and one hot and extra-
ordinarily dry summer ( 1952 ) , it seems certain that in an ordinarily
warm summer with enough precipitation to grow a normal crop of
flowers, Megachile brevis populations in the Lawrence area would
increase during the summer in approximately the manner shown
in figure 1. It seems possible that in the characteristically drier
regions of western Kansas, where the predominant late summer
and fall flowers are yellow composites rarely utilized by this bee,
the fall increase in abundance might not occur, as was the case
in 1952 in Lawrence.

Number of Generations Per Year

The number of generations per year doubtless fluctuates with
the length of the active season. Even in a single locality it is appar-
ently not a regular or constant matter. The only generation which
is relatively easily understood is the first. All bees active in May
and early June seemed rather fresh. Because of the scarcity of this
generation it is impossible to say how long the adult emergence
period is, but the impression gained from the apparent increase in
abundance of bees on flowers during May is that the emergence
may continue until about the end of May. Nest construction and
egg laying doubtless begin within a few days after emergence of
the adults and continue for nearly their entire lives, probably at
least a month. In the climatically more or less normal spring and
early summer of 1952, a perfectly fresh female believed to be of
the second generation was seen on June 17, and an exceedingly
badly worn female, doubtless a remnant of the first generation, on
July 3. In summer the egg to adult period is about a month; in
the cooler weather of spring it is somewhat longer. Thus second
generation individuals would be appearing more or less continuously
from about June 17 until the beginning of August.

Considering the earliest offspring of each generation (thus the
maximum number of generations per year), we conclude that
second generation individuals emerging as early as June 17 could
probably have mature third generation progeny as early as July

Michener: Biology of a Leafcutter Bee

1667

20 or 25. (These would thus be emerging at the same time as
the last second generation individuals.) These third generation
individuals could have adult progeny of the fourth generation by
the end of August or early September. Their progeny would hiber-
nate and emerge in the spring as the first generation of the following
year.

A
B
C

Fig. 2. Diagram showing the way in which generations become mixed,
largely because of the long reproductive period of these bees. A represents
a seasonal cycle of 4 generations, in which the earliest eggs of each generation
give rise to the following generation. B represents a seasonal cycle of 2 genera-
tions in which the last eggs of each generation give rise to the following
generation. C represents the presumably commonest situation of three genera-
tions. Solid lines represent immature stages; dotted lines adults. Generations
of adults are numbered.

?. :

MAY

JUNE

JULY

AUG.

SEPT.

By contrast, considering the last offspring of each generation
(thus the minimum number of generations per year) we conclude
that second generation individuals emerging in late July would lay
some of their last eggs well after the first week in August. Such
eggs would not develop into adults until spring; thus there would
be but two generations during the year. It is quite likely that
some such second generation bees would survive until late August
and thus be in flight at the same time as the earliest fourth generation
individuals mentioned in the preceding paragraph. In short, it
seems that in a year of ordinary temperatures near Lawrence,
Kansas, there may be as few as two generations or as many as
four; most probably go through three generations. After the first,
the generations become completely mixed as shown in figure 2,
because of the rather long periods ( probably about a month ) during
which adults are active and laying eggs. One can therefore observe
most adult activities, such as nest construction and provisioning,
at all times from May until the first of October.

Studies of ages of population samples during 1952 support these
statements. Age was judged by the amount of tattering of the
wing margin; the condition of specimens was recorded as fresh,
fair, or poor by comparison with drawings prepared in advance
and carried in the field. Many of the bees were liberated after
examination. About equal numbers of each sex fell in the various

1668 The University Science Bulletin

categories, indicating that wing wear of males is about equivalent
to that of females. All specimens (15) collected in May and on
June 1 and 2, were fresh. Five specimens collected on June 7,
and five more on June 13, were fair. On June 17, nine specimens
were fair, two poor, and one ( presumably second generation ) fresh.
Increasing numbers of fresh specimens were observed later in June
and on July 3, a small sample was classified as follows: one very
poor (remnant of first generation), three fresh, three fair. As bees
became more abundant it became possible to obtain larger samples
for this purpose and during July and August 290 specimens were
examined to determine the amount of wear of the wings. Of these
111 were classified as fresh, 166 as fair, and 14 as poor. During
these months there was no particular time when one category was
more abundant than at other times; the summer population was a
complete mixture of age groups. The small number of bees in poor
condition was due to the extremely tattered wings required for this
classification; only a few bees achieved sufficient wear (and sufficient
age?) to be so classified.

The data for the exceedingly cool and wet spring and early sum-
mer of 1951 indicated that weather conditions influenced the rate
of development and hence the number of generations. Two
freshly completed nests of three cells each were found, one on May
27, one on May 29. Each was kept and the time of emergence
of adults was July 11 and 12. Unfortunately all six bees which
emerged were the parasite, Coelioxys. However, later in the sum-
mer the developmental period of the Coelioxys is about the same
as that of Megachile; therefore there is good reason to believe
that the cool weather slowed the developmental period of both
forms from about a month to about 45 days. This surmise is sup-
ported by the field observations which show that, while very few
bees could be found, only tattered individuals (presumably first
generation) were seen during the first week in July. Their num-
ber was even smaller than during June, so that it appeared that the
first generation nearly died out before the appearance of the second
generation. Several fresh individuals were seen in the field during
the second week in July. In a year such as this it would seem
that the maximum possible number of generations would be three
and that two might be very common. The small number of genera-
tions, in addition to the reduced reproductive activity of individual
bees due to frequent periods of cloudy weather, rain, and low
temperatures, would account for the low populations observed
during 1951.

Michener: Biology of a Leafcutter Bee 1669

Overwintering

As can be seen from figure 1, a very large proportion of the normal
year's production consists of the overwintering individuals, and
relatively few of these individuals survive to emerge the following
spring. The reasons for the high winter mortality are discussed
elsewhere. Overwintering occurs as fully fed mature larvae or
prepupae. The factors which cause the cessation of development
at this stage are not obvious, for temperatures are still high, some-
times very high, in the third and fourth weeks of August. Yet
larvae which reach maturity at that time remain in that stage.
They do not pupate and soon emerge as adults, as would mature
larvae earlier in the season even at much lower temperatures.

The data concerning the time at which diapausing begins are as
follows: An egg laid August 5, 1952, produced an adult on Septem-
ber 5. An egg laid approximately August 7, 1952, produced an
adult at an unknown date in early September. Larvae from these
eggs must have been spinning cocoons about August 16 and 18,
and proceeded with their development to become adults. Eggs
laid about August 17, and on August 20, 1952, developed into indi-
viduals which went into winter as mature larvae.

Twelve eggs ( in several nests ) which were laid on various dates
from August 7 to 13, 1950, did not produce adults that year but
went into winter as mature larvae. Six other larvae which had
reached maturity and were beginning cocoon spinning from August
16 to 21, 1950, also went into winter as mature larvae.

Nests are still being provisioned as late as October 1; therefore
all of the eggs laid from about August 7, until October 1, go to
make up the overwintering generation, or first generation of the
following year. It is not known exactly when pupation of this gener-
ation occurs but it is in the spring with adults emerging during
the last two thirds of May.

In nests kept in the laboratory at room temperature during the
winter the emergence is spread out over a long period, from October
to March. Under conditions of continued warmth the diapause is
apparently broken very irregularly and at widely different times.

Dispersal and Favored Habitats

As has already been intimated, Megachile brevis, unlike many soli-
tary bees, is a highly mobile species. Since it has a long season of
adult activity with several generations per year, the places which
are particularly favorable at one time of the year are entirely differ-

1670 The University Science Bulletin

ent from those that are favorable at other times. Not only do suc-
cessive generations disperse from their regions of emergence and
concentrate in the places that happen to be attractive at the moment,
but it seems certain that a bee in the midst of its life and of its nest
making activities may leave one area to go to another, perhaps
several miles distant.

Strong indications exist of a constant shifting about of populations.
Concentrations, largely of males, occur in areas of good nectar
supplies even though nesting sites and pollen supplies are not
available.

There are probably virtually no areas in eastern Kansas where
Megachile brevis cannot occasionally be found. There appear to
be no natural habitats which are favorable enough to attract large
numbers of Megachile throughout the season of activity. The fol-
lowing sections on favorable areas where most of our studies were
made are presented in order to indicate the characteristics of the
places that are favorable at different seasons and in order to present
the evidence for dispersal.

There are two types of favorable habitats which cause concen-
trations of the bees. One is called a nesting habitat, for it provides
nesting sites and pollen sources as well as nectar. So far as is known,
all plants utilized for pollen also provide nectar; at least the bees
thrust their proboscides into the flowers as though sucking nectar
at the same time that they collect pollen. The other is called
a nectar habitat. It is a place where the principal flowers are highly
attractive as nectar sources but are not used for pollen. Nesting
sites may not be available in the vicinity. Male bees sometimes
concentrate in such situations, and some female bees are found there.

Nesting Habitats

Although the first Megachile brevis emerge in the vicinity of
Lawrence in the first third of May, there are no known areas of
concentration until the last third of May. At that time (actual
observations dated May 24 to June 5) Amorpha fruticosa comes into
bloom in low moist areas where there is no shade and often the bees
concentrate on it in considerable numbers for this season. Area 1,
where many spring observations were made, is one mile east, one
mile south of Lawrence. It is a swampy spot about two acres in
area full of Amorpha fruticosa. A railroad embankment along one
side of it is covered with small rose bushes which provide leaves
for nest making and with dead stalks of various large weeds, such
as Helianthus and Ambrosia, which provide favorable nesting sites.

Michener: Biology of a Leafcutter Bee 1671

The Amorpha is the only source, so far as known, for pollen and
nectar at this place. In early June, when this species of Amorpha
ceases to bloom freely, the Megachile disappear in spite of the
presence of certain flowers sometimes used by Megachile ( e. g.
Trifolium x ) . At no subsequent time in the year, in spite of periodic
observations throughout the summers of 1951 and 1952, were many
Megachile seen in the region of Area 1, although very occasional
individuals sucking nectar from Teucrium and Verbena were seen on
a hill one fourth of a mile away during August.

In general, although it is occasionally a nectar source, sweet
clover (Melilotus) is not a pollen source for Megachile brevis. On
two occasions, however, among hundreds of patches of this plant
examined, small patches of freshly blooming Melilotus officinalis
have been found to be rather heavily used ( considering the scarcity
of the first generation) as pollen sources. One of these, Area 2,
is four miles north of Garnett, Kansas, and bees were observed there
on May 22, 28, and 29, 1951, each day abundant enough that about
six could be seen per half hour of observation. This patch was on
high dry land in a weedy roadside area where there were plenty of
old Helianthus stalks for nesting sites. On June 10, only a single
Megachile was seen there; presumably the others had left in favor
of areas of Psoralea floribunda half a mile away or because the sweet
clover became less attractive as it became older. The second area
where bees were seen using Melilotus officinalis as a major pollen
source was along the road through the sand hills four miles south
of Garden City, Kansas, on June 12, 1952.

After the Amorpha fruticosa is no longer in full bloom the principal
plant attractive to Megachile brevis as a pollen source is Psoralea
floribunda, or in certain areas in southeastern Kansas (e. g., Galena
and three miles east of Baxter Springs), Psoralea psoralioides eglan-
dulosa. The former in particular is a widespread and common
prairie plant. It occurs often in small prairie patches remaining
along roadsides or railroads, but is especially abundant in large
prairie acreages. In eastern Kansas the prairies are limited to high
ground, usually hill tops. Thus the bees of the first generation,
when the Amorpha fruticosa is no longer in bloom, apparently must
often move distances of several miles from low wet areas to prairies
covered with Psoralea floribunda.

The most studied prairie of this sort, Area 3, located three miles
southwest of Ottawa, Kansas, is several square miles in extent. A

1. Except for the list in the section on kinds of flowers visited, generic names only are
used in mentioning flowers unless two species of a genus are discussed in this paper.

1672 The University Science Bulletin

few trees grow in the gullies but even there the herbaceous vege-
tation consists largely of prairie species. The Psoralea foribanda
begins to bloom there at about the beginning of June and each year
Megachile have been collected there between June 2 and 6. There
are no earlier prairie flowers attractive to Megachile in this area.
The Psoralea remains in bloom and serves as both a pollen and
nectar source until near the end of June. In the latter part of that
month Amorpha canescens, another prairie plant, is sometimes also
used as a pollen and nectar source. As soon as the Psoralea is no
longer in abundant bloom, however, the bees disappear from the
prairie, even though Amorpha canescens, Petalostemon, and other
seemingly suitable plants are in full bloom there. In Area 3, there
are no suitable weed stalks for nesting, and the bees nest under bits
of sod, under dried cow dung, or in fence posts. They use for nests
leaves and petals of the small prairie rose as well as of various
other prairie plants.

Area 4 is a floristically similar prairie of only a few acres extent
located one and one half miles west and one half mile south of
Lawrence. Any part of it is within a few hundred feet of such
nesting sites as old Ambrosia and Helianthus stems. In this area,
too, Megachile activity, although always slight, was limited to the
season when Psoralea floribunda was in bloom, except for the one
observation described in the following paragraph.

One of the first bits of evidence concerning migration came from
the following series of observations: On August 3 and 7, Area 4
was carefully inspected for Megachile because of the excellent
growth of seemingly suitable flowers such as Petalostemon. Not a
specimen of M. brevis was seen. On these days, however, a few
M. brevis were seen in a pasture one fourth mile away which was
heavily grown up with Vernonia, from which the bees were gathering
pollen as well as nectar and with Symphoricarpos, which was used
only for nectar. On August 15, in the morning, this pasture was
mowed and all the flowers in it cut. By noon that day several M.
brevis of both sexes could be seen in Area 4, sucking nectar from the
Petalostemon and from Solidago flowers. The next day, and subse-
quently, none could be found there. The inference was that the
bees dispersed from the mowed pasture, some of them stopping
for nectar in Area 4, but not staying since the flowers there were
of rather unattractive sorts.

During the latter part of June and the first half of July the Mega-
chile brevis seem very much dispersed with no areas of abundance

Michener: Biology of a Leafcutter Bee 1673

on native vegetation. A very few remain in the prairies where
they collect pollen of Amorpha canescens; others are attracted by the
first flowers of Vernonia. Introduced alfalfa ( Medicago ) is the only
plant which seems to attract rather large numbers of Megachile
brevis at this season. They were studied in numerous areas on
alfalfa, but the only place worthy of special comment is known as
Area 5. This is a two-acre patch of alfalfa surrounded by brushy
and weedy creek bottoms and roadside vegetation. It is located
near Blue Mound, southeast of Lawrence, Kansas. It was in full
bloom when first visited on July 14, 1950, and was nearly out of
bloom by August 4. During this period Megachile brevis was abun-
dant collecting pollen and nectar, particularly so about July 20,
when the hum of several Megachile could be heard at all times and
at one time an estimate of 80 bees seen per half hour of observation
was made. In late July, as the flowering became poorer, the Mega-
chile became much scarcer.

One of the principal sources of pollen in the latter part of July
and throughout most of August is Vernonia. This plant comes up as
a perennial weed in poorly cared-for pastures, particularly in low
but not swampy lands. Patches of it are very common; a few
Megachile brevis can be found in most of them, many in some. The
Vernonia patch most fully studied is Area 6, a pasture of many acres
9 miles south and 3 miles east of Lawrence. It had grown up in
brush and small trees, and then a few years ago it was cleared
except for certain large trees. The resulting cut brush and trees
were placed in large piles scattered over the pastures. Weeds com-
ing up in these piles were out of reach of cattle and matured and
died, leaving many old weed stalks for nesting sites, in addition to
scattered weed stalks elsewhere in the pasture. Sumac (Rhus)
bushes around the pasture served as an important pollen source
in 1950, especially as the Vernonia went out of bloom in mid-August.
In 1951, the Vernonia was just coming into bloom at the end of July
and continued until the end of August but in the warmer summer
of 1952 it was blooming by mid-July and nearly out of bloom by
mid-August.

Area 7 is an abandoned pasture six miles southeast of De Soto,
Kansas. Since the principal pollen source is Vernonia, the area is
attractive to Megachile at the same season as Area 6. The situation
differs from that of Area 6 principally in the great abundance of
dead Ambrosia and Helianthus stalks throughout the area, forming
suitable nesting places, and in the profusion of Symphoricarpos

1674 The University Science Bulletin

which provides nectar from late July into September. Lythrum in a
low area also serves as a nectar source. Thus as long as the Vernonia
is in bloom this is an important nesting habitat; indeed in 1952, the
only season that this area was observed, Megachile was more abun-
dant here than at any other place found in eastern Kansas, possibly
because of the combination of a good pollen source and many
nesting places.

On July 25, 1952, most of the Megachile observed in this area were
working on Vernonia flowers. By August 8, many were visiting
Symphoricarpos for nectar, as was observed on repeated visits there-
after. By August 20, the few stragglers among Vernonia plants were
still being visited for pollen but most of the bees in the area were
visiting Symphorocarpos for nectar. Although, as indicated above,
the abundance of Megachile in this area in 1952 was doubtless due
to Vernonia and the numerous nesting places, the availability of
Symphoricarpos nectar may be the factor that kept the bees in the
area almost until the last Vernonia was out of bloom. By August 10,
the bees were nearly gone from a Vernonia patch about a mile away
which lacked the Symphoricarpos.

Area 8 is a somewhat similar pasture two miles east and eight
miles south of Lawrence. It lacks the numerous nesting sites of
Area 7, and the Vernonia is rather sparse; as at Area 6 an important
pollen source is bushes of Rhus. Because of the abundance of
Symphoricarpos, and in some places of Lythrum, the area is also a
nectar habitat. Observations began in this area on August 4, 1951,
when both Vernonia and Rhus were being used as pollen sources. It
seemed that Rhus was most visited in the mornings. As this was
a high dry area, the Vernonia ceased bloom earlier than in Area 6,
so that by mid-August most of the pollen collecting was from Rhus,
which in turn ceased its blooming in late August. Thereafter the
only Megachile hrevis seen in the area were a few sucking nectar
from various flowers which are not used for pollen.

After the Vernonia of the pasture areas and the Rhus of nearby
fence rows and waste land stop blooming, there is a short season
when there is no really suitable abundant pollen source for Mega-
chile hrevis in eastern Kansas. The bees apparently disperse (see
records of dispersal from Area 8 in the section on studies of marked
individuals below) and can be seen sucking nectar from a very
wide variety of flowers.

In 1950, a pasture (Area 9) three miles northwest of Lawrence
was the site of intense activity of Megachile hrevis. On September

Michener: Biology of a Leafcutter Bee 1675

5, when the place was first visited the bees were collecting pollen
from both Gutierrezia and Trifolium. This activity very likely
started because of the shortage of more suitable flowers. At no
other place were these flowers found to be used except as an occa-
sional nectar source. Moreover, the same flowers growing in the
same place were not seen visited for pollen by Megachile brevis in
1951 or 1952. In 1950, however, pollen gathering continued until
the Gutierrezia ceased flowering about September 25.

The usual pollen source after the first week in September is
various species of purple Aster which bloom along roadsides and
in pastures. In Area 10, four miles northwest of Lawrence, Aster
began to bloom about September 6, 1950, and continued until
October 7, although no Megachile were seen after October 1, doubt-
less because of cool weather. They were collecting pollen on
October 1, however.

Nectar Habitats

It seems likely that Megachile brevis will, on occasion, suck nectar
from almost any flower from which this bee is physically equipped
to obtain nectar. Moreover, as already stated, it will obtain nectar
from probably all the kinds of flowers used as pollen sources. There
are, however, certain kinds of flowers not or rarely visited for pollen
but very attractive to these bees as nectar sources. Patches of such
flowers seem to hold wandering bees, especially males, for varying
lengths of time. It is the resulting concentrations of bees not active
in nest making that characterize the nectar habitats.

Areas 7 and 8 described above became nectar habitats when the
pollen sources in them failed. The bees obtained the nectar prin-
cipally from Symphoricarpos, to a lesser extent from Lythrum, occa-
sionally from Pycnanthemum, Bidens, and various other plants. In
each of these areas there was a considerable period when both nest-
ing bees and those which appeared to be only sucking nectar could
be found.

A small seepage space in Area 6 is a nectar habitat, often full of
male bees and of some females which show no evidence of nesting
in the area. Here the nectar sources are numerous and attractive
before, during, and after the pollen producing plants (Vernonia,
Rhus) of this area are in bloom. The plants producing the nectar
are Lythrum, Lippia, Verbena, Lycopus, Trifolium, Pycnanthemum,
Teucrium, Ludwigia, and Bidens. One or another of this series of
plants is in bloom from early July to early September, and at least
a few Megachile can be found there throughout that season.

1676 The University Science Bulletin

The only area studied which was strictly a nectar habitat was
Area 11. This is a swampy meadow about 50 yards long and 25
yards wide. It is located nine miles south of Lawrence. It is sur-
rounded by a corn field, a bit of prairie containing neither flowers
nor nesting places likely to attract Megachile brevis, and a sowed
grass pasture. Thus there are practically no nesting sites. The
meadow was full of Lythrum in bloom through the entire month of
August, 1951, and until mid- August in the dry summer of 1952.
Megachile brevis was the commonest bee on these flowers.

The presence of an occasional nesting Megachile in this area is
shown by the record of a female cutting leaves there. Such scat-
tered nesting individuals can be found almost anywhere, irrespec-
tive of the areas of concentration discussed above.

Studies of Marked Individuals

The preceding sections show clearly that Megachile brevis is
found in a wide variety of situations and that it concentrates in dif-
ferent places at different seasons. The conclusion is obvious that
considerable dispersal occurs. To shed additional light on the mat-
ter, several marking studies were undertaken. Only two such
studies produced significant data and even in these cases the data
are meager. Both of these studies were made during August, 1951.

During the period August 4 to 20, inclusive, 148 Megachile brevis
were marked in Area 8. Weather permitting, the area was visited
and specimens marked every other day.

Marking was done by means of quick drying paints, spots of
which were applied by means of a fine brush to bees captured in a
net. All individuals marked on any one day were marked with the
same color on the same part of the body. If a bee were recaptured
on a subsequent day, it was marked with the color of that day, so
that a bee might acquire several spots of color if it were recaptured
several times. There was no evidence that the paint injured the
bees. Sometimes it probably wore off, for bees were occasionally
seen from which most of the paint had disappeared. However,
there is no reason to believe that bees which had thus lost their
identity played any large role in the results described below.

No doubt due to the change in this area from a nesting habitat in
early August to a nectar habitat in the later part of the month, a con-
siderable change in the Megachile population was noted. Thus
from August 4 to 12 a total of 103 bees were marked, of which 67
( 13.8 per day of marking ) were females, 36 ( 7.2 per day of mark-
ing) males. From August 16 to 20, a total of 45 bees was marked,

Michener: Biology of a Leafcutter Bee 1677

of which 13 (4.3 per day of marking) were females, 32 (10.7 per
day of marking) males. Thus the number of males marked per day
increased as the area became a nectar habitat, perhaps largely due
to the decrease in abundance of females which allowed the marker
more time for capturing and marking males. At the same time the
number of females decreased very greatly. Each day of marking
usually amounted to about three hours in the field, with variations
due to weather conditions. The ages of the bees varied greatly
throughout the month; there was no evidence of the bees in the
nectar habitat averaging younger or older than those in the nesting
habitat.

Recoveries of marked bees were remarkably few. Even early
in the month when pollen collecting females were often marked,
only two females were seen after marking; one first marked on
August 6, was marked again on August 10 and 16, another first
marked on August 4, was marked again August 8. Since nest
making females spend most of their time either collecting pollen
or pieces of leaves, it seems that had they remained in the vicinity
they would have been seen again in greater numbers. The few
nests located in and around Area 8 at this time consisted of only
one or two cells. The impression which we acquired is that, with
Vernonia, the principal pollen source, rather scarce, the female
nesting bees remained only long enough to finish one or two cells
and then moved on. It should be pointed out that while Rhus was
an important pollen source, we have never seen this plant alone
support a population of these bees; we have always seen it used
merely to supplement Vernonia as the latter nears the end of its
flowering season.

Two additional females were recovered from among individuals
marked after the middle of the month. One was merely seen in
Area 8 two days after marking. By good fortune, however, one
which was marked in Area 8 on August 16 was seen again on
August 30 near Area 1 at a distance of 5.5 miles from the point
where it was marked. This lends strong support to the idea of the
great mobility of these bees. On both occasions this female was
merely sucking nectar, not gathering pollen, the first time from
Symphoricarpos, the second time from Teucrium. As already ex-
plained, late August is a period of poor pollen supplies for Megachile
brevis in this region, for Vernonia and other midsummer pollen
sources are waning, but purple Aster, the main autumnal pollen
source, is not yet in bloom. It is quite likely that many Megachile
spend this time wandering about.

1678

The University Science Bulletin

Among males eight individuals were recovered, twice as many
as among females although there were 12 less males than females
marked in total. If significant, this difference between the sexes
indicates that the males moved about less than the females. This
is quite likely true, for because of Symphoricarpos Area 8 was ap-
parently an attractive nectar habitat. Females, being probably
motivated by the need for pollen with which to provision nests,
would not be expected to remain long in a nectar habitat.

One male (the only one recovered more than once) was marked
on August 6, recovered August 16, 18, 20, and seen on August 22 and
28. All five times after August 6, this bee was found in or around
a single clump of Symphoricarpos bushes, showing that this indi-
vidual, probably unlike most, stayed in or returned to a very re-
stricted locality for at least 22 days.

The other marking study was conducted at Area 11, the small
isolated Ly thrum patch with practically no nesting sites or pollen
sources nearby. On August 17, 1951, in one and one half hours,
55 male and four female bees were marked (all in the same way).
An hour later a 15 minute survey of the area revealed 19 marked
males and 4 unmarked ones. Assuming that the ratio of marked to
unmarked bees in the area was as the ratio of marked to unmarked
ones seen in the 15 minute survey, it was calculated that there
were 12 unmarked male bees in the area, or a total of 67 males.
Their age distribution was wide, as usual at this season. Of the
55 males marked, 5 were fresh, 45 in good condition, and 5 in poor
condition.

Table I shows the results of similar 15 minute surveys conducted
on other dates. The consistent scarcity of females in this nectar
habitat is evident, as is the rather rapid disappearance of marked
individuals and appearance of unmarked ones. This seems to
provide additional evidence of mobility in this species.

Table I. — Number of marked (M) and unmarked (U) Megachile brevis
found in 15 minute surveys of Area 11 on various dates in August, 1951.
Marking was done on August 17.

Number
marked

August
17

August
20

August
22

August
29

August
31

Male

Michener: Biology of a Leafcutter Bee 1679

PART II. RELATIONS WITH FLOWERS
Kinds of Flowers Visited

Megachile brevis is commonly regarded as a highly polylectic
species. For example Mitchell (1936) writes, "There are apparently
few flowers which it will not visit." This is probably quite true
insofar as visiting flowers for nectar is concerned, although definite
preferences are exhibited for certain nectar sources.

In its pollen collecting, Megachile brevis is clearly rather re-
stricted, as shown in Table II. There is some systematic significance
to the limitations of the list; for example whole families like the
Rosaceae are unrepresented. Yet very diverse plants such as cer-
tain of the Leguminosae and certain of the Compositae are much
used. It is interesting to note that the principal pollen sources are
blue or purple flowers, although such flowers may be systematically
in families as diverse as the Leguminosae and the Compositae.
Certain less important pollen sources are white or greenish, and
one rather minor source, Gutierrezia, is yellow. This is interesting
in view of the large number of yellow composites of the region.
Many of these, such as various species of Helianthus, are regularly
visited by many bees, including various species of Megachile, but
they are only rarely utilized by Megachile brevis even for nectar.

These observations show clearly that Megachile brevis, in its
pollen collecting, is far more restricted than such forms as Apis
and most species of Halictus and Lasioglossum. This leads to con-
sideration of the meaning of the terms oligolectic and polylectic.
These terms were used by Robertson and subsequent workers,
oligolectic bees being those which visit few kinds of flowers and
polylectic ones being those which visit many kinds. Mere visiting
for nectar, however, is usually not of great biological significance,
and most records of visits of bees to flowers are of far less value
than if additional information as to whether or not pollen was
gathered were also recorded. In most bees the males and nectar
sucking females visit most readily the kinds of flowers from which
pollen is also gathered by the species. However, they may visit
almost any flowers which provide nectar. It is in their pollen
gathering that many bees are specific.

The many highly oligolectic forms known to gather pollen from
only a single genus ( or even species ) of flower, fall at one end of a
spectrum, with highly polylectic forms (e.g. Apis) at the other
end. Megachile brevis falls in the midst of this spectrum.

1680

The University Science Bulletin

Table II. — Kinds of flowers visited in Eastern Kansas by Megachile brevis.
(x marks under pollen and nectar sources indicate relative importance; from
x — one or two records only, to xxxx — of major importance. ) Presumably all
pollen sources are also nectar sources but this is not indicated in the table.
Localities of information not obtained in vicinity of Lawrence, Kansas, are
indicated under "remarks."

Scientific name

Common
name

Pollen
source

Nectar
source

Season

Color

Remarks

Alismataceae
Sagittaria

Arrowhead

July

white

Euphorbiaceae
Croton

Croton

August

white

Verbenaceae

Verbena micrantha

Verbena

late July

white

Verbena

July and
August

purple

Lippia lanceolata
var. recognita

Fogfruit

July

whitish

Labiatae

Lycopus americanus

Water
Horehound

late July
and August

whitish

Pycnanthemum
flexuosum

Mountain
Mint

XXX

August

whitish

Teucrium canadense

Wood Sage

August

purple

Leguminosae
Astragalus

Loco weed

June

whitish

Amorpha
fruticosa

False Indigo

XXX

May and
early June

whitish

Amorpha
canescens

Lead Plant

mid-June to
mid-July

purple

Petalostemon

Prairie
Clover

July and

August

white
purple

Medicago
sativa

Alfalfa

July and
August

purple

Melilotus
alba

White Sweet
Clover

July and
August

white

Melilotus
officinalis

Yellow
Sweet
Clover

May and
June

yellow

Locally
used as an
important
pollen
source

Psoralea

floribunda

Wild
Alfalfa

XXX

June

purple

Psoralea

psoralioides
eglandulosa

June

purple

South-
eastern
Kansas

Lespedeza
violacea

Bush
Clover

August

purple

Strophostyles
helvola

Wild Bean

late Aug.,
early Sept.

red

Tephrosia
virginiana

Goat's Rue

June

red and
yellow

Michener: Biology of a Leafcutter Bee

1681

Table II — Concluded

Scientific name

Common
name

Pollen
source

Nectar
source

Season

Color

Remarks

Trifolium
repens

White
Clover

XXX

July to
Sept.

white

In one

locality,

one season,

important

pollen

source

Lythraceae

Lythrum alatum

Winged
Loosestrife

xxxx

July and
August

purple

Onagraceae

Ludwigia alternafolia

False
Loosestrife

August

yellow

Anacardiaceae
Rhus copallina

Sumac

August

whitish

Found used
only near
failing
Vernonia
patches

Umbelliferae
Zizia aurea

Meadow
Parsnip

late May
early June

yellow

Rubiaceae

Diodia teres

Buttonweed

August

pinkish

Caprifoliaceae

Symphoricarpos
orbiculatus

Buckbrush

XXXX

August

whitish

Compositae

Coreopsis grandiflora

Coreopsis

June

yellow

South-
eastern
Kansas

Bidens

polylepis

Beggarticks

late Aug.,
early Sept.

yellow

Helianthus annuus

Sunflower

August

yellow

Silphium perfoliatum

Cupplant

August

yellow

Rudbeckia
serotina

Black-eyed
Susan

July

yellow

Ratibila
pinnata

Yellow
Coneflower

July

yellow

Helenium autumnale

Sneezeweed

August

yellow

Gaillardia
pulchella

Gaillardia

June

yellow-
orange

Garden City
Kansas

Gutierrezia

dracunculoides

Broomweed

Sept.

yellow

In one

locality,

one seaaon,

important

pollen

source

Atter

Aster

xxxx

Sept.

purple

Solidago

Goldenrod

late Aug.

yellow

Eupatorium
perfoliatum

Boneset

August

whitish

Vernonia
interior

Ironweed

xxxx

late July
and August

purple

1682 The University Science Bulletin

It should be clear that the terms oligolectic and polylectic are
usually significant only in relation to pollen collecting, and that
even in that reference they are only comparative terms.

Nectar Gathering

Bees of both sexes regularly insert their proboscides into flowers,
evidently for nectar. This activity occurs throughout the lives of
the adults. Individuals which had only been out of their nest
for a few minutes were liberated in the midst of a patch of Lythrum.
One of each sex alighted on Lythrum flowers after flying less than
two feet from the point of liberation and inserted its proboscis in the
ordinary and seemingly expert fashion. At the other extreme, very
tattered and old individuals of both sexes are often seen sucking
nectar.

Males alight on flowers almost solely in order to obtain nectar.
Often in the midst of actions interpreted as searching for females
they were seen to alight and suck from flowers.

Females probably suck nectar from every flower used as a pollen
source. Rarely they stop only for nectar in the midst of pollen
collecting. We have a record ( August 9, 1950 ) of two females seen
in Area 6 with a little Vernonia pollen on their scopas, but sucking
nectar from Ludwigia flowers. Perhaps in the course of gathering
pollen they wandered away from the Vernonia into the nectar habitat
and sucked some nectar before going on to more Vernonia. More
frequently, bees with full pollen loads are seen to stop at certain
flowers and suck nectar only. This is most noticeable when nectar
is gathered from kinds of flowers not used as pollen sources. Thus
near Lawrence on August 19, 1950, two females with full loads of
pollen from unknown but distant sources were seen to return to
the vicinity of their nests. Before they went into their nests each
stopped to suck from several flowers of Diodia. Such sucking may
occur before the bees leave the vicinity of the pollen source, as
shown by the following example : Near Galena, Kansas, on June 12,
1951, females were gathering pollen from Psoralea psoralioides
eglandidosa. One with an apparently full pollen load was seen to
leave the Psoralea and go to a Coreopsis head blooming among the
Psoralea. There it sucked from several disk flowers, flew on to three
more Coreopsis heads, to a Psoralea spike, to a Coreopsis head, to a
Psoralea, to a Coreopsis, and then away. The impression was that
having obtained a full pollen load, the bee sucked nectar from any
convenient source, perhaps to fill its crop. That this is a common
action is indicated by the frequency with which bees having full

Michener: Biology of a Leafcutter Bee 1683

pollen loads were seen sucking nectar but not gathering pollen.
Thus bees with full loads of Wins pollen were several times seen
sucking from flowers of Teucrium or Vernonia.

Female bees, while looking for a nesting site, will usually stop
and suck from a flower of almost any kind if it is close to their
line of flight, and if a patch of flowers is encountered, they may
visit several of them before going on.

It is interesting to note that nectar gathering female bees visiting
alfalfa trip most of the flowers from which they suck. On various
occasions females were observed to trip from all to about 70 per
cent of the flowers visited. In spite of the fact that they do less
work than pollen collecting individuals, they usually visit fewer
flowers per minute than those which are collecting pollen. Thus
one female (August 5, 1950) was observed to suck from 10 flowers
per minute (average of three minutes). She tripped 21 of the 30
flowers visited. Another sucked from an average of 12 flowers per
minute (in over three minutes observed) and of 34 flowers which
she was watched to visit, she tripped 24. Pollen collecting females
regularly trip every flower.

Males, on the other hand, often trip few or none of the alfalfa
flowers visited. It would be interesting to know if each learns inde-
pendently how to suck the nectar without tripping the flowers, as
appears to be true of honeybee workers.

Flower Constancy in Nectar Gathering

There is no evidence of the slightest tendency for bees of either
sex to restrict themselves to particular kinds of flowers as nectar
sources, even for short periods of time, if several equally favorable
sources are available. The following records are selected from
among many to exemplify this statement: On August 13, 1951, a
female which appeared to be searching for a nesting place was
watched as it chanced to approach the nectar habitat in Area 6.
As it encountered the low growing flowers in this area, it started
going from flower to flower sucking nectar and in three minutes
was seen to suck from Lythrum, Vernonia, Croton, Trifolium, and
Sagittaria. It then wandered away continuing its original activity.
Another female was seen the next day in the same place under
otherwise similar circumstances to visit Trifolium, Croton, Ludwigia,
and Lythrum.

In the average area, however, there is one flower distinctly more
attractive as a nectar source than the others. It may or may not
be a pollen source, as well.

1684 The University Science Bulletin

Pollen Gathering

The exact mechanics of pollen gathering by female Megachile
brevis is difficult to determine because of the rapidity of the action
and the minute quantities of pollen obtained from any one flower.
The following observations were made near Cherryvale, Kansas, on
June 13, 1951, where numerous bees were visiting flowers of
Psoralen floribunda. Corroborative observations have been made
repeatedly on this flower.

The female bee, on landing on the flower spike, puts her proboscis
into an individual flower. This opens the flower, exposing the
stamens and pistil. The front tarsi are rubbed on the anthers occa-
sionally during the process. The bee moves rapidly over the spike,
working several of the flowers in this manner, then goes on to
other spikes. Occasionally, after the bee has visited several flowers,
the front tarsi contact the middle legs. The details are not apparent
because of the rapidity of the movement but presumably pollen
gathered by the front legs is transferred to the middle legs. At other
times the middle tarsi can be seen to brush the venter of the thorax,
where pollen must sometimes be brushed off of the anthers, or the
middle legs may even be bent up over the thorax to comb the meso-
scutum. Presumably the pollen gathered from the body and front
legs is transferred by the middle legs to the hind legs, but this was
not observed; perhaps it occurs in flight.1 After visiting 10 or 12
flowers the bee alights on a new spike of flowers and rubs the scopa
(pollen collecting hairs of the abdomen) with one rear basitarsis,
or often with one after the other. At the same time the abdomen is
bent upward several times, almost spasmodically, then back to its
normal position. This action opens up the spaces between the
groups of scopal hairs on each sternum, and pollen seems to be in-
troduced into the spaces from the rear. Because of the way in which
pollen is placed on the scopa by the hind basitarsi, a bee which is
starting to gather a load of pollen may be seen to have the pollen
of the scopa arranged in the V-shaped pattern shown in figure 3,
each arm of the V representing the zone rubbed by the rear basi-
tarsus of that side of the body.

On alfalfa the pollen collecting is very similar to that on Psoralea.
When the bee inserts her proboscis, the alfalfa flower is tripped, that
is, the column of stamens and pistil rapidly move up from their

1. Leg movements which probably accomplished transfer of pollen from leg to leg were
noted in flight between flowers of Vernonia. They were also noted among sluggish indi-
viduals gathering pollen from alfalfa. Sluggish individuals are better for such observations
than highly active ones.

Michener: Biology of a Leafcutter Bee

1685

■Si

mk?

Fig. 3. Under surfaces of abdomens of females of Megachile brevis, showing
the pollen collecting scopa. a, without pollen; b, with a little pollen; c, with a
full pollen load.

original position in the keel of the flower to a position against the
banner. In this process the bee's proboscis is often trapped between
the column and the banner and the bee can be seen to pull loose,
then reinsert the proboscis beside or behind the column. In this
process also pollen is doubtless brushed off the stamens onto the
under side of the thorax and head of the bee. Moreover, as the bee
presses its head down into the flower, its face comes in contact with
the stamens of the tripped flower. Thus pollen is probably scat-
tered over the bee, especially over its head, more thoroughly by
alfalfa than by Psoralea. Perhaps for this reason, bees were seen to
brush their heads more when collecting from alfalfa than when col-
lecting from Psoralea. After visiting several flowers a bee will
cling to a flower with middle and rear legs only while brushing the
front part of the body, especially the face, with the front legs. Other
activity was as has been described for Psoralea pollen collecting.

On one occasion it was noted that when a female Megachile came
to an alfalfa flower which had already been tripped, she imme-
diately went on to other flowers.

The behavior of the bee is, of course, quite different on different
flowers, particularly on those whose flowers occur in flat heads
(Compositae) or in large masses (Rhus). On Gutierrezia a pollen
collecting female tends to work with her head near the center of
the flower, rotating her body around the flower head. On Rhus the
female is noteworthy for being constantly on the move with a sort of
wriggling motion over the large surface made up by the many small
flowers. She appears to feed the pollen back from the front legs
to the middle legs, thence to the hind legs and the scopa, rubbing

1686 The University Science Bulletin

the scopa with the hind legs and at the same time raising the ab-
domen, just as has been described for pollen collecting on Psoralen
except that the bee is able to do all this without stopping her motion
over the flowers. Sometimes while flying from one flower mass to
another or while hovering the bee rubs the two hind legs together.

Rapidity of Pollen Collecting

Details on the duration of pollen collecting trips will be pre-
sented in the section on nesting activities. The present section
concerns rates of flower visiting, that is, flowers visited per minute.

The data were gathered by watching individual pollen collecting
bees over periods as long as possible and recording the number of
flowers visited each half minute of observation. The figures were
then converted to flowers visited per minute.

Most of the data were obtained by observing bees collecting
pollen from alfalfa. There is some evidence that the rate of col-
lecting is higher at higher temperatures, but due to the variability
found at any one temperature, the evidence is not conclusive. The
greatest amount of data collected in any one temperature range
(82°-85° F.) was gathered during July 1950. A total of 121 half-
minute counts gave a minimum rate of 9.4 flowers visited per
minute, a maximum of 24, and an average of 15.7. Of the flowers
visited, over 95 per cent were tripped and therefore presumably
pollinated and utilized as pollen sources. The others were merely
probed by the proboscis but for some reason not tripped. These
results agree reasonably well with more extensive data on this
subject gathered by Franklin (1951) in Kansas and Linsley and
MacSwain (1947) in California.

Data from Psoralea floribunda, a plant often called wild alfalfa
because its flowers are similar to those of alfalfa, taken in the same
temperature range indicate a higher rate of visiting flowers, pos-
sibly because tripping, required for alfalfa, is unnecessary for
Psoralea. A total of 87 half-minute counts gave a minimum rate
of 12 flowers visited per minute, a maximum of 30, and an average
of 22.0.

Information on other flowers is slight, but we have records of 16
to 18 Gutierrezia disk flowers probed per minute, 18 to 36 Trifolium
flowers visited per minute, and 10 to 11 Aster disk flowers probed
per minute.

The variation in rates of visiting single kinds of flowers indicated
in the preceding paragraphs is not due entirely to irregularities in
flower abundance in the fields studied. There is evidence that at

Michener: Biology of a Leafcutter Bee

1687

the same time, place, and temperature different individuals work
at different rates. Some appear sluggish, and repeatedly rest on
leaves in the midst of their pollen collecting activities.

To verify this impression, data on individual bees in a single
alfalfa field were gathered. The results are shown in table III.
Similar data, obtained in a single patch of Psoralea floribunda, are
presented in table IV.

Table III. — Rates of flower visiting by individual pollen gathering bees in a

single alfalfa field.

Ree

Flowers visited per minute

Minutes of

number

Minimum

Maximum

Average

observation

19.3

15.7

16.3

11.3

Table IV. — Rat

es of flower visiting by individual pollen gathering bees in a
single patch of Psoralea.

Ree

Flowers visited per minute

Minutes of

number

Minimum

Maximum

Average

observation

25.7

18.3

3.5

20.7

Constancy in Pollen Collecting

Because of the small number of acceptable pollen sources usually
available in any one area, individual pollen constancy is more diffi-
cult to judge than nectar constancy. However, one ordinarily
observes pollen collecting females visiting flower after flower of the
same species, and not deviating to visit other flowers, even for
nectar, until a full load of pollen is gathered.

Exceptions to this general observation do occur when a bee
stops using one flower and starts visiting another, but one never

1688 The University Science Bulletin

sees erratic visiting of various kinds of flowers as with nectar gath-
ering. Evidence for changes in pollen sources are as follows:

Different kinds of pollen are sometimes found in a single cell.
Changes may occur in the midst of a pollen collecting trip as shown
by the following two observations: In September, 1950, in Area 10
Gutierrezia and Trifolium were intermixed, both serving as pollen
sources. Most pollen collecting females seemed constant to one
flower or the other so far as we could observe, but on September 10,
a bee which had been collecting from Gutierrezia flowers was seen
to start visiting Trifolium. From then on it gathered pollen from
Trifolium constantly, as long as it could be watched (three or four
minutes). On August 11, 1950, in Area 6 a female was watched
collecting Rhus pollen. The posterior part of its scopa was covered
with Vernonia pollen, showing that it had changed its flower during
a pollen collecting trip.

Another observation made in Area 6 concerned a bee which was
nesting in a hollow dead weed stalk. For several days it had been
observed bringing in the pale pollen of Vernonia, and had con-
structed three cells with that pollen. This plant, however, was be-
coming gradually older, with fewer fresh flowers, although there
were still a great many. In the midst of provisioning the fourth cell,
the bee stopped using Vernonia pollen and began using Rhus.

Additional information on flower constancy (or lack of it) was
obtained in recording the activities of certain individual bees near
their nests. The color of the pollen brought in was usually uniform
from trip to trip. However, the bee recorded as "d", figure 8, used
yellow Rhus pollen throughout the period of observation except that
after her second pollen collecting trip for the second cell recorded
in figure 8, she came in with white (Vernonia?) pollen. Nest "i"
(figure 9) was provisioned with Vernonia pollen on August 11. On
August 12, Rhus pollen was used, except that after the eighth and
twelfth pollen collecting trips on that day the pollen seemed white
and was probably from Vernonia. On August 14, the same bee
used bright yellow Rhus pollen except for the sixth and seventh
trips, which resulted in collections of white (Vernonia?) pollen.

Michener: Biology of a Leafcutter Bee 1689

PART III. MISCELLANEOUS ACTIVITIES
Relation of Adult Activity to Environmental Factors

Very little significant information on this subject has been gath-
ered. Much better bee subjects for such investigations are those
with more permanent nests, where activity can be watched over long
periods. Activity appears to be dependent upon certain relation-
ships of temperature, light intensity, wind, and possibly other factors.
On a cool day, no wind and strong sunlight are necessary to produce
activity. On a cloudy day, no wind and high temperature are neces-
sary for activity. On a windy day, sunlight and high temperature
are necessary. With regard to these various factors, different bees
probably have quite different thresholds of activity, for even on a
rather poor day one may find a few active bees.

Quantitative data concerning these matters is almost nonexistent
for Megachile brevis. It has been seen active at temperatures ( taken
in the shade of the observer's body at the level of the flowers being
visited, usually one to two feet above the ground ) of 76° to 100° F.
Flight below 80° F. was rarely observed in summer, but in Septem-
ber, when generally lower temperatures prevail, it was more com-
mon. On August 21, 1950 (a clear day) a female was seen to start
her work when the temperature near the nest reached 80° F.; on
another such occasion the temperature reached 83° F. before the
bee flew.

On three different cloudy mornings it was noted that activity did
not begin until the temperature reached 85° F. Twice this was not
until 11:00 a. m. or later. There is a little evidence that once the
bees start to work they continue at temperatures lower than those
necessary to activate them.

The hours of activity during the day vary not only with weather
but of course with the season. In early August in clear weather
they have been seen active as early as 8:30 a. m. and as late as 5:20
p. m. On July 27, a bee returned to its nest at 6:07 p. m. It had
left 17 minutes earlier. In general, morning activity does not involve
many individuals until 9:00 a. m. or later even on an ideal day.

This species has been observed flying in winds strong enough to
roll a bee over and over on the ground if it happened to attempt to
land in a gust. This sort of behavior was rare near Lawrence, in
eastern Kansas, but in western Kansas where high winds are almost

76—8663

1690 The University Science Bulletin

continuous in certain seasons, it is common. Perhaps this is be-
cause the bees are forced into activity in spite of the wind.

Flight

Megachile brevis is a bee that flies very rapidly with a distinct
high pitched buzz audible on calm days for as much as ten or twelve
feet. It is usually more easily found by means of this sound than by
sight, although the sound is easily confused with that of certain
bombyliid, nemistrinid, and tachinid flies as well as with that pro-
duced by certain of the other small species of Megachile. No differ-
ence was noted between the sound produced by males and females.

In full flight, as when females fly to and from their nests, the bees
fly straight and at such a speed that the human eye can scarcely
catch sight of a bee as it goes past. It is to be remembered, how-
ever, that the bee is dull colored and rather small, so that it is not
easily seen. One bee was timed as making the 150 foot distance
from her nest to the place where she was cutting leaves in 15
seconds. This is a rate of under 7 miles per hour. Considering that
the bee quite obviously flies rather slowly as she starts her flight,
gathering speed over a distance of 6 or more feet, and slows down
over a similar distance at the end, the actual maximum speed is
probably over 7 miles per hour but under 10. The bee is a notice-
ably faster flier than is Apis, which, however, has been reported to
fly from 5.6 to 13 miles per hour by various authors.

Among flowers or elsewhere when the bees fly only short distances
there is no difficulty in following their flight. Bees of either sex
visiting flowers can be distinguished by sound from other bees be-
cause of the frequent brief interruptions of the flight while they rest
on flowers. The characteristic buzz is therefore interrupted 9 to 36
times per minute, depending on the particular bee, kind of flower,
etc.

Females searching for nesting sites can usually also be recog-
nized by sound as they fly a sinuous or zigzag course close to the
ground, investigating sticks and other possible nesting places. The
flight is continuous except for irregular and often rather long inter-
ruptions while a bee crawls into a hole. In full flight to and from
the nest, female bees fly two and one-half to three feet above the
ground if there are no obstructions, but we have seen them fly over
trees twenty feet high. They seem to do this in preference to flying
through any considerable amount of shade.

Michener: Biology of a Leafcutter Bee 1691

Miscellaneous Notes on Behavior

The following notes on various activities seem worth recording
because they may be clues to consistent behavior patterns which
may eventually be recognized:

One male was observed near Lebanon, Missouri, August 4, 1951,
flying about plants of Lespedeza violacea. Frequently it lit on a
leaf and rested for several minutes holding its forelegs up and
forward.

Both sexes are often seen brushing various parts of the body
with their legs. This often occurs when the bees are working
flowers and probably serves to get scattered pollen off of parts of
the body. We have frequently seen the eyes brushed.

In marking a male bee with quick drying enamel, paint was in-
advertently got on the wings of the right side. It hardened, sticking
them together. The bee could not fly. After some struggling it
hooked the inner hind tibial spur between the two wings and pushed
outward, straightening the leg. After repeating this several times
the bee managed to peel the paint off of the wings and free them
so that it could fly.

Nocturnal Resting Places

As will be clearly shown later, females constructing nests rest in
their nests at night. No positive field evidence is available on the
night resting places of males or of females not constructing nests.
We do know from numerous observations that they do not return
to the nests in which they have developed and from which they have
emerged to spend the night or for any other purpose.

In cages males and females crawl into curled leaves or similar
protection to pass the night. It seems probable that they do like-
wise in the field.

Male Reactions to Females

Curiously enough mating has never been observed in this bee.
Probably it occurs very soon after emergence.

Males are often seen flying from flower to flower, not alighting
but hovering for a moment over each flower, then going on to the
next. The males may be very persistent in this activity or they
may stop at intervals to suck nectar from certain of the flowers.
We have considered this activity to be the search for females.

Males are occasionally seen to fly rapidly at females from a dis-
tance of a few inches and strike them. Females, so far as have
been observed, repulse the males under such circumstances. Sev-

1692 The University Science Bulletin

eral times we have observed females which were collecting pollen
disturbed by males pouncing upon them.

Slightly more elaborate approaches to females have also been
observed. A male sometimes hovers for several seconds two to
five inches above and behind a female on a flower, then somewhat
slowly, or at other times rapidly, descends upon the female. In all
cases observed, the female merely flew away. If she only flies to
another flower an inch or so away, the male may repeat his ap-
proach.

Longevity of Adults

The oldest marked bee ( a male ) of which we have a record was
last seen 22 days after the time of marking. However, there is
considerable evidence that the bees actually live more nearly a
month. The last eggs which will develop into fall adults are laid
in early August; adults from these emerge in early September;
such adults (both sexes) are active until the beginning of October
(see figure 1) when they are probably killed by cold. There is
some evidence that at least in cool weather longer adult life may
occur. In the extraordinarily cool spring and summer of 1951, the
second generation did not appear until the second week of July.
Therefore, the disappearance of old and tattered first generation
individuals could be observed without the complicating factor of
second generation bees. Probably all first generation individuals
had emerged by the last week of May, yet some still survived in
the first week in July. All were females, suggesting that females
survive longer than males.

Aging is shown in this species of bee in a number of ways. The
wings become very much tattered in both sexes as the bees become
older. This was the principal character used to judge age in the
observations on age composition of populations reported under
"Number of Generations Per Year." Figure 4 shows the amount
of wear that occurs in wings.

The pubesence, quite ochraceous on the dorsum of the thorax in
young males and slightly so in females, loses its reddish color, the
pale hairs becoming grayish white with age. This fading must
occur very quickly in females, for they are not often seen in the
field with ochraceous pubescence. Males, however, seem to retain
this color for at least a week after emergence. Very old individuals
often seem blacker than less worn ones because some of the pale
hairs are broken off. This is especially true of the dorsal abdominal
hairs.

The mandibles of females (but not males) show much wear.
Figure 5 illustrates this.

Fig. 4. Fore wings illustrating wear of outer margin, a, fresh; b, an ex-
ample of the class called fair; c, an example of the class called poor; d, one of
the most worn individuals studied.

Fig. 5. Mandibles of females illustrating wear, a, fresh; b, well worn.

1694 The University Science Bulletin

PART IV. NESTING
Finding Nesting Sites

As mentioned in the section on "Flight," females searching for
nesting sites are rather easily recognized. This behavior must
occupy a considerable part of their time for they are often seen
apparently searching. They fly in a wandering flight, often cover-
ing an area rather thoroughly, but at other times moving on in one
general direction even though zigzagging considerably. In bare
areas they usually fly one or two inches above the ground, in grassy
places just over the grass tops, sometimes going down among the
blades of grass. In weedy or brushy areas they commonly go down
into the vegetation and are sometimes out of sight for minutes at a
time as they fly or walk about. They rarely go into thoroughly
shaded places, however. They alight and investigate all sorts of
small cavities. Although they sometimes nest among green leaves,
there seems to be a tendency for them to investigate darker objects,
such as brown curled leaves, and especially sticks lying on the
ground. The following quotations from our notes exemplify this
behavior: "August 13, 1951 [Area 8]; 10:30 a.m. Saw one female
which appeared to be looking for a nesting site. She was flying low
over the grass, stopping here and there on leaves or [to suck] from
flowers. Once she stopped for several minutes and crawled up and
down and around a stick lying on the ground. Eventually she flew
off." "August 14, 1951 [Area 6]; 10:00 a. m. One female (without
pollen) watched for 15 minutes while she presumably searched for
a nesting place. She flew about in the vegetation, especially where
it was dense, occasionally stopping to walk about on some stems.
When she found an old dead stick, she spent a long time crawling
around it flying off a few inches and then returning to it. It had no
hole in it and she finally left. As she flew about in her wandering,
somewhat zigzag flight, she occasionally came upon a flower and
would stop and suck from it."

The following observations provide the only concrete evidence
available that the type of behavior described above is actually
searching for a nesting place. "August 29, 1951 [Area 8]; 2:00 p. m.
[A female was flying over the grass in the usual manner of one
presumed to be searching for a nesting site. She happened to ap-
proach me.] I held out a rubber tube (inside diameter 9 mm.)
just above the ground. She went into the tube, then flew a foot or
so away, then re-entered, went in and out five times in five minutes,

Michener: Biology of a Leafcutter Bee 1695

three times in the next five minutes, flying variable distances away,
sometimes out of sight, between her visits. Ten minutes after she
first entered the tube, she started bringing pieces of yellow petals
for the nest. She brought many of them in rapid succession, but
stopped and was not seen again after 4:00 p. m. [possibly because of
the heat in the tube in the direct sunlight.]"

Another occasion when the "searching flight" was seen to lead to
nest establishment was on August 11, 1950 (Area 6); 11:00 a. m. A
female Megachile brevis was seen flying over the small rocks piled
along the roadside. Once she left these rocks and went into the
weedy pasture, searching along sticks, then returned to the rocks.
At intervals she would crawl into spaces among the rocks. Finally
she stayed in such a space for two minutes then left and flew all
about the area (orientation flight?) well above the ground in a
manner quite different from the searching flight. After this flight,
she returned, then left again, flying about. This was repeated three
times, one of the flights taking the bee 50 feet away. (Perhaps
because of the uniform appearance of the roadside, these orientation
flights were more extensive than usual. ) The bee then left and
after ten minutes ( about 20 minutes from the time she first entered
the nesting place ) returned with a piece of a petal. She proceeded
to construct a cell, as will be described later.

Rather different observations concerning nest establishment were
made in an alfalfa field near Hutchinson, Kansas, July 26, 1950.
Usual nesting sites, such as old weed stalks, were scarce in the area.
A female was seen crawling in and out among the dense foliage of a
deformed, fasciate plant of Erigeron canadensis. This bee left but
soon returned to crawl around more among the leaves. After 50
minutes of this coming and going, the bee seemed to select a place,
for she came back to the same spot among the leaves of the weed
three times in about as many minutes, then started to bring leaves
for the cell.

It is perhaps significant that among the three cases of nest estab-
lishment that we observed, the longest period required to start the
work of cell construction was in the most abnormal nesting situation
(among leaves of Erigeron).

Nesting Sites

Mitchell (1936) and Hicks (1926) both suggest that the wide
variety of nesting sites reported for M. brevis by various workers
(Reed, 1871; Packard, 1892; Rau, 1916, 1922; Hicks, 1926) may in-
dicate that more than one species was identified as brevis. In view

1696 The University Science Bulletin

of the similarity of various species of Megachile this may have
been the case, but the differences in the nesting sites do not neces-
sarily indicate this. Even greater variability in nesting sites has
been observed in Kansas populations of Megachile brevis than was
previously recorded in the literature for this species. At the season
of the year when the bees are largely in the prairie patches using
Psoralea floribunda as a food source, their nesting sites will often
necessarily be different from the old weed stalks favored when the
populations are in other places. Perhaps the mobility of this bee,
correlated with the fact of its several annual generations, forces it
to retain catholic tastes in nesting sites. That we are not dealing
in Kansas with two or more related species utilizing different sites
is shown by the actions of females searching for nesting places.1
They will sometimes search along sticks and weed stalks, then
search bare ground, looking into holes, then crawl into curled leaves,
as though all these sites were potential nesting places to a single bee.

The great variability in nest sites observed in Megachile brevis is
not surprising in view of the literature concerning other species of
this genus. For example, Megachile centuncularis Linnaeus of
Europe has been recorded as nesting in the soil, in rotting wood
(Baysson, 1902), beneath rocks (Bellevoye, 1884), in hollow stalks
and bamboo (Hardouin, 1945), and in various other situations.
Smith as early as 1855 recorded a great diversity of nesting sites
for this species.

As Table V shows, the nesting sites most commonly chosen, in our
experience, are hollow dead weed stalks, old cornstalks, and the
like, lying on the ground. I believe that most of the nests in our
area are in such places but probably not as preponderant a percent-
age as the table at first suggests. Nests are most easily found by
splitting dead stalks; it is only by observing bees that one finds nests
in the soil, under prairie grass, and in like situations. The second
column (nests found by observing bees) probably gives the best
comparative data on the various sorts of nesting sites.

In summary, it is evident that almost any sort of small cavity may
be used for nesting by this bee. The bees apparently never ex-
cavate holes. When they nest in stalks, it is almost inevitably in
pithy stalks hollowed out by various borers and subsequently broken
to expose the hollow. There is some evidence that the bee may
clean loose debris out of such a hollow or even enlarge the hollow.
The diameter of the cavity holding the cells is sometimes slightly

1. Because of the superficial similarity of various Megachile species, specimens have
been collected frequently in all areas of study and identification checked by means of the
characters detailed by Mitchell (1936).

Michener: Biology of a Leafcutter Bee

1697

larger than the same hollow where bees have not entered it, and bits
of freshly removed pith may be found beneath the entrance.

We have never found a nest in deep shade of woods or even under
a large tree or in heavy shade produced by a luxuriant growth of
tall weeds. Nests are most commonly found in old broken pieces of
stalk lying about where the grass and weeds are short.

The pieces of stalk may be well hidden by short grass and weeds
and shaded by them, or may be entirely exposed.

Table V. — Nesting sites of Megachile brevis, and numbers of cells in nests.

Site

Nests

found by

observing

bees

Total
nests
found

Number of cells per nest

Mir

Maximum

Average

Dead horseweed stalks, prostrate.
erect. . . .

Dead pokeweed stalks, prostrate.

Dead sunflower stalks, prostrate,
erect

Dead cornstalks, prostrate

Dead thistle stalks, prostrate,
erect

Rase of dead ironweed stalk, prostrate

Dead curled boneset leaf

Termite hole in garage door

Among dense leaves of living fasciate
muletail weed

Among small rocks on ground.

Under dry cowchips

Under mat of prairie grass

In holes in ground

22
3

14
4

30
2

7
1
3
2
4

2
2

1
3

3.7
2

3.4
3

4.2
6

1.7

1.1
1
2
1
1 .5

The plants listed in table V are as follows: Horseweed (or giant
ragweed), Ambrosia trifida; pokeweed, Phytolacca decandra; sun-
flower, Helianthus annuus, H. tuberosus; corn, Zea mays; thistle,
Cirsiam sp.; ironweed, Vernonia interior; boneset, Eapatorium
perfoliatum; muletail (or horseweed, fleabane), Erigeron cana-
densis.

Nest Structure

Nests typically consist of several cells placed end to end and
snugly fitted into a tubular hollow. Sometimes only a single cell is
located in a hollow. When there are several cells, of course, the
first constructed is the one farthest from the entrance into the cavity
while the last constructed is the one closest to the entrance. The

1698 The University Science Bulletin

position of the cells in a long hollow ( such as a hollow stalk ) varies
greatly. Sometimes, even when the hollow was many centimeters
long, cells were found only near the entrance, blocking the hollow
and leaving most of it inaccessible and useless. At other times bees
go deep into hollow stalks to construct cells. The last cell con-
structed may be flush with the open end of the hollow or may be
deep in the hollow. Among 46 nests (in stalks) whose structure
was recorded in detail, one had the last cell constructed 35 cm. from
the entrance into the stalk, three had the last cell flush with en-
trance into the stalk, and the others were intermediate in this regard,
the average distance from the entrance to the last cell constructed
being 3.5 cm.

The cells are ordinarily tightly fitted together so that if a stalk
containing a nest is carefully split, all the cells can be removed
as a single unit. This is because the base of each cell fits inside of
the apex of the preceding one. The elongate pieces of leaves and
petals which form the cup of any cell extend well beyond the cap
of that cell, leaving a hollow into which fit the elongate pieces which
form the cup of the next cell. Often the cells of a series are so
closely associated that it is necessary to break the series in order
to determine how many cells are present.

Occasionally cell series constructed by two bees, or by one bee at
different times, are found in the same stalk. This situation can be
recognized by the old leaf and petal material of one series, com-
pared to fresh material of the other. Sometimes in such cases there
is an unoccupied space of one to several centimeters between the
two series of cells.

As shown in table V, the nests are commonest in horizontal hol-
lows ( e. g. in prostrate stalks ), so that the cells are usually horizontal.
They are not infrequently vertical, however, with the caps upward,
as in hollows of standing stalks exposed by breaking the stalk. One
nest was found in which the bee entered a hole in the side of a
standing stalk and turned upward, there constructing a series of
vertical cells with the caps downward.

Most nests (except those few in which the last cell is flush with
the entrance to the hollow) are closed at the entrance with a plug
made of pieces of petals or leaves. These pieces are more or less
round, like those used to cap cells, and are loosely or firmly fitted
into the opening. The space between the last cell and the plug
is usually empty although in one nest there was an additional plug
of eight pieces of leaves just beyond the last cell and separated

Michener: Biology of a Leafcutter Bee 1699

from it by a millimeter or two. The distance from the last cell to
the usual entrance plug varies from 2 mm. to several centimeters.
Entrance plugs (in 15 nests in which they were studied) ranged
from a thick plug of 20 pieces of leaves and petals to a thin one
consisting of only four pieces of rose petals and another thin one of
four leaflets of Lespedeza violacea. The average number of pieces-
in the 15 plugs studies was 9.3. Most of them consisted of leaves
or petals of the sort used as "fillers" in cell construction but five
contained pieces of rose leaves in addition to leaves of Lespedeza or
alfalfa, making much firmer plugs.

About one third of the cell series studied were provided with no
entrance plug. Sometimes this was probably due to the death
of the bee or to our collecting the nest for study before her work
was completed, as indicated by an incomplete last cell. In other
cases, however, the last cell was complete and it appears that some-
times Megachile brevis does not construct an entrance plug. In few
cases was an entrance plug found where only a single cell had been
constructed.

Number of Cells Constructed

The number of cells in any one nest depends in part on the size
of the space available. A small cavity may provide room for only
a few cells. The right hand column of table V illustrates this point;
the average number of cells per nest is higher in stems, where the
long cavities provide plenty of room, than in the miscellaneous small
cavities listed in the lower part of the table. The small number
of cells in nests in ironweed stalks is due to the very short basal
region large enough to hold Megachile cells.

The number of cells constructed by any single female is unknown.
A series as long as 11 cells has been found, but there is no reason
to believe that this is a maximum. Few nesting places provide
space for very many cells; as shown in table V many single cells
were constructed, even in hollow stalks where there would have
been space for several cells. Since the bees move freely from place
to place, a bee may construct a few cells in one place and others
several miles away. Packard (1868) reports a single Megachile
(identified, probably incorrectly, as centuncularis Linnaeus) that
constructed 30 cells. Perhaps females of M . brevis construct simi-
larly large numbers, scattering them in various places.

Study of figures 8 and 9 shows that these bees can construct and
provision 1 to 1.5 cells per day. If a bee lives for 30 days, it might
presumably construct 30 cells, even allowing for some bad weather.

1700

The University Science Bulletin

Cell Structure

The present section contains a summary of the data obtained by
dissecting 100 cells of Megachile brevis from 48 nests. It is well
known that cup portions of cells of many leaf cutter bees are made
of elongated pieces of leaves which bend inward at one end ( called
the base of the cell) to close it. After provisioning the cell and
laying an egg, the bee closes the other end by means of round pieces
of leaves, which are called the cap.

Megachile brevis follows this general pattern, using elongate
pieces of leaves and petals for the cup and circular pieces for the
cap. The pieces for the cup are broadly overlapped, as shown in
figures 6 and 7. The pieces which form the cap are all of about the
same size and shape, slightly larger than the diameter of the hollow
in the cup, so that when pressed into place their edges turn up
slightly and the fit is snug (figure 6).

base of next

base of eel I

Fig. 6. Longitudinal section of cell (diagrammatic, to show make up of
walls). Solid lines represent leaves; dotted lines, petals; broken line, mass of
provisions.

Fig. 7. Cross-section of cell. Explanation as for figure 6.

In gathering the data on cells, the series were broken into the
component cells. If round pieces adhered to the base of a cell, they
were transferred to the preceding cell and considered part of the
cap of that cell. This procedure is ordinarily justifiable, for the
cap of one cell and the base of the next are in close contact (see
figure 6). However, round pieces are sometimes found at the base
of the first cell in a series. In such cases they must have been the
first leaves cut and had nothing to do with any cap.

Great diversity is shown among various individuals of Megachile
brevis as to the manner of cell construction. Part of this diversity
is due to the size of the hollow in which the bee nests. The inside
dimensions of the cells are quite constant, 4.2 to 5.2 mm. in diameter,
8.2-10.0 mm. in length. The dimensions of the hollows used are by
no means as constant, ranging from just over 5 mm. in diameter to
10 mm. in diameter. A bee nesting in a small hollow needs to use
only a few leaves and petals to construct a cell cup having the

Michener: Biology of a Leafcutter Bee 1701

proper inside dimensions while a bee nesting in a large cavity must
carry into it many leaves and petals to form the thick walls neces-
sary to obtain the proper inside dimensions. For example, one bee
nesting in a small hollow in a weed stalk used only eight pieces to
construct a cup (nest 20, table VI) while another bee nesting in
a space among leaves of a fasciate Erigeron canadensis used 41
pieces. ( nest 23, table VI ) . This is the bee whose record is shown
as nest f, figure 9. From this record it can be seen that she really
brought to the cell about 58 pieces but some were dropped through
the leaves to the ground and lost, a thing which never happens in
ordinary nesting places. The outside length of a cell is also variable,
depending in large part on how far the walls of the cup extend
beyond the cap. Most cells are from 11.5 to 14 mm. long but rarely
they reach 17 mm. in length.

Additional diversity in construction is due to individual variations.
Two bees working at the same place constructing nests only 10 feet
apart in cavities of similar size may use quite different materials.
Moreover, the cells in any one series are usually fundamentally
similar but often markedly different from those in other series. As
will be explained later, the bees tend to gather their construction
materials largely from small "cutting places." No doubt the some-
what chance selection of these places determines in large degree
the characteristics of the cells.

Almost all individuals of Megachile brevis make use of two or
more different materials in constructing the cups of their cells. If
the cavity is appreciably larger than the inside diameter of the
cell, filler petals or filler leaves are the first materials to be brought
in. All petals used are thin and satisfactory as fillers. The leaves
most commonly used are leaflets of Lespedeza violacea or Medicago
(alfalfa). These are thin, easily cut leaves often placed rather
loosely in the cavity. If petals are used they often are cut first so
that they form the outside of the cup, with filler leaves inside of
them, although they may be intermixed to a certain extent. Major
irregularities in filler material sometimes occur. Thus the second
cell in nest 3, table VI had 14 Lespedeza leaflets on the outside, fol-
lowed by five pieces of Cassia petals on one side of the cell, followed
by nine more Lespedeza leaflets. As shown by the first eleven nests
in table VI, leaves are more commonly used than are petals as fillers.
Yet there are nests in which petals are used extensively for fillers
(nests 12, 13, and 14, table VI.) and others in which petals are
used exclusively or nearly so ( nests 15, 17, 22, 23, table VI ) . Occa-

1702 The University Science Bulletin

sionally nests are found in which the filler material is omitted en-
tirely; see, for example, nests 18, 19, 20, and 21, table VI.

Inside of the filler materials are almost always placed a few firm
leaves, usually cut from rose or sometimes from Symplioricarpos.
These leaves are ordinarily the ones which provide much of the
strength of the cell wall. In many species of Megachile, these are
the only sort of leaf used in cell construction but in M. brevis they
are usually very few in number. There are nests in which firm
leaves constitute a large majority of the pieces used (e.g., first
three cells of nest 13, and nests 18 to 22, table VI). On the other
hand there are nests (25 to 30, table VI) in which firm leaves are
altogether omitted. In nest 21, almost the whole of both cells was
constructed of Symphorocarpos leaves. These leaves are a little
thinner and more flexible than most rose leaves and presumably in
this instance served both as firm leaves and filler leaves. This would
explain the large number used. In nest 22, however, the 22 firm
leaves recorded in table VI were all rose.

Lining petals are the last placed in the cell before pollen collect-
ing begins. They are rather consistently present. Their number
is often difficult to count in old nests, however (hence the many
"-f-" signs in table VI), because they become soft and mushy, or
later very delicate and brittle, and because at least those of the cap
are often eaten by the bee larva. Lining petals are ordinarily from
the same kind of flower as filler petals although they are occasionally
mixed. Rarely, as in certain cells in nests 9, 10, 13, and 21, lining
petals are absent and the pollen is placed in direct contact with
firm leaves. Sometimes there is only a single lining petal in the cup,
so that no complete lining exists (see nests 5, 9, and 21, table VI).
Nest 30 is remarkable in that it contained no leaves, therefore no
distinction between lining and filler petals existed. Rather than
arbitrarily making a division, they are all recorded in table VI as
lining petals. In three cells thin leaves, like those used for fillers,
formed the lining or were among the lining petals. In table VI
these leaves are listed in the column for lining petals but marked
with an asterisk.

Like the materials used for the elongate pieces of the cup, those
used for the round pieces of the cap are varied. Lining petals are
quite consistently present, although occasionally absent (see nest
13 ) . Filler leaves are also rather consistently present but sometimes
absent. Firm leaves and filler petals are frequently absent.

In the cups, the materials are arranged in a fairly consistent man-

Michener: Biology of a Leafcutter Bee

1703

Table VI. — Materials used in the construction of cells.

Each horizontal row indicates a single cell, the figures representing the number of
pieces of each material present. The first column at the left gives nest numbers, in order
to show the way the cells were associated to form nests. The averages in the last row are
based upon exact numbers and ignore "-J-" and "?" signs. A "-(-" indicates that the
material was present but that the number of pieces was unknown. A "?" indicates that it
could not be determined whether a material was present or not. In the "lining petals"
column, a "*" marks figures representing thin leaves, such as those used for fillers. A
blank space indicates absence of a type of material while a " — " indicates lack of information.

Cup

Cap

Nest
No.

Lining
petals

Firm
leaves

Filler
leaves

Filler
petals

Total

Lining
petals

Firm
leaves

Filler
leaves

Filler
petals

Total

5 + 2*

—

•>

—

1704

The University Science Bulletin

Table VI. — Continued

Cup

Cap

Nest
No.

Lining
petals

Firm
leaves

Filler
leaves

Filler
petals

Total

Lining
petals

Firm
leaves

Filler
leaves

Filler
petals

Total

—

Michener: Biology of a Leafcutter Bee

1705

Table VI. — Continued

Cup

Cap

Nest
No.

Lining
petals

Firm
leaves

Filler
leaves

Filler
petals

Total

Lining
petals

Firm
leaves

Filler
leaves

Filler
petals

Total

—

5 + 1*

—

1706

The University Science Bulletin

Table VI. — Concluded

Cup

Cap

Nest
No.

Lining
petals

Firm
leaves

Filler
leaves

Filler
petals

Total

Lining
petals

Firm
leaves

Filler
leaves

Filler

petals

Total

—

Average

2.81

5.5

9.9

3.1

22.1

2 3

1.1

2.5

0 4

6.3

1. Nest 30 omitted from this calculation, since many of the 33 petals might best be
called fillers.

ner. Thus the vertical columns of table VI are arranged from left to
right in the order in which the materials are ordinarily found from
the inside to the outside of the cell; however, considerable mixture
often occurs so that a firm leaf may be inside a lining petal or filler
leaves and petals may be somewhat intermixed. For the caps inter-
mixture is much more the rule, the only generalization possible
being that the lining petals are consistently on the inside.

The most complicated cells studied were the last three in nest
13, figure VI. All three of these cells contained four different kinds
of materials, and their arrangement was far from standard. An
example (the fourth cell) will be sufficient. From the inside out
it contained the following: two lining petals (Cassia), seven firm
leaves (Rose), six filler leaves (Lespedeza), two firm leaves (Rose),
one firm leaf (Sympliorocarpos) , two filler leaves (Lespedeza), and
15 filler petals (Cassia).

The variability and complexity of the cells of Megachile brevis are
not entirely unexpected in view of what is known of other species.
Complex cell walls, consisting of layers of different materials, are
well known in other species. For example several species put a
layer of mud between outer and inner layers of leaves, others put
a layer of masticated leaf material between outer and inner layers

Michener: Biology of a Leafcutter Bee 1707

of leaves. Great variation in construction materials from individual
to individual is also recorded for other species and Markowsky
(1933) even records a nest of M. centuncularis Linnaeus made with-
out the side walls of the cells but with only partitions between the
cells made of pieces of leaves. That the construction of the cells
of any one nest tends to be similar in M. centuncularis, as in M.
brevis, is illustrated by Markowsky's anomalous nest mentioned
above and by Grandi's ( 1934 ) figures of the numbers of leaf
pieces used in five cells of a single nest.

Nests for the numerical analysis shown in table VI were selected
at random from those available. They were collected at various
seasons of the year. The arrangement of the nests from one to 30
is intended to place nests having similar characteristics together;
the commonest types are at the beginning of the table, the unusual
ones near the end. All of these nests were in stalks except for num-
ber 11 which was in a hollow under a mat of prairie grass, number
22 which was in a space among pebbles, and numbers 23 and 25
which were in spaces among the dense leaves of fasciate plants of
Erigeron canadensis.

The sizes of the leaves used in different parts of cell cup con-
struction vary. The alfalfa leaflets used in nest 25, table VI varied
from 7 mm. to 14 mm. in length. In other cell cups (not included
in table VI) Lespedeza leaflets from 7 to 11 mm. in length and Tri-
folium leaflets 5 to 11 mm. in length were used as filler material.
The smallest pieces were near the bottom of the cell.

Leaf pieces cut most of the way around ( as rose leaves ) also vary
in size. The long firm pieces of rose leaves varied in one cell from
4.5 mm. wide and 10 mm. long to 5 mm. wide and 14.5 mm. wide.
In another the extremes in sizes were 5 x 11 and 6 x 13 mm. In an-
other cell in which Sumphorocarpos leaves were used, the smallest
long pieces were 5 x 10 mm., the largest 9 x 15 mm. In this cell the
Sumphorocarpos served both as firm leaves and filler and the large
pieces were consistently on the outside of the cell, sizes becoming
progressively smaller toward the inside.

From the above information the impression is inevitable that as a
bee cuts leaves or petals, it selects the texture and cuts the size
appropriate to the needs of the moment in fashioning its cell.
The same conclusion has been independently reached by other
authors (e. g., Popovici-Baznosanu, 1907) working on other species
of Megachile. This suggests that there are delayed reactions in-
volved, the bee reacting while cutting the leaf to stimuli received
while in its nest.

1708

The University Science Bulletin

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Michener: Biology of a Leafcutter Bee

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1710 The University Science Bulletin

It is well known (see, for example, Malyshev, 1935) that senile
bees sometimes behave in unusual ways. No evidence of this was
obtained in the present study because no one bee could be followed
through "its entire adult life. However, some peculiar structures
made by Megachile brevis were found. The most remarkable was
in a nest of two cells, the lower cell of which was normally pro-
visioned and contained a growing larva. The upper cell, that is
the second one to be constructed, consisted of a normal cell cup
which was filled, not with the usual provisions, but with 40 round
pieces of leaves of Symphorocarpos.

Duration of Trips

Much of the information on the timing of the construction and
provision activities of Megachile brevis is summarized in figures
8 and 9. In these figures is presented the information on the com-
ings and goings of nine different bees ( a to i ) watched continuously
for more or less extended periods.

Bee "a" started to construct a cell between some small roadside
rocks at Area 6 on August 16, 1950, at 11:00 a.m. The record
(figure 8) consists solely of the 39 trips in two hours and 25 min-
utes required for the bee to construct the cup ( bottom and side walls
of the cell) from pieces of petals and leaves. For some reason which
is not known, the bee never provisioned the cell.

The bee lettered "b" was first seen at 10:55 a. m. on August 21,
1952, as it flew out of an old cornstalk lying on the ground near
Area 6. It was observed for a little less than an hour and a half,
after which it never returned. It was neither constructing nor
closing a cell, but was constructing a plug at the entrance to a series
of cells that it had previously built and provisioned. This is, there-
fore a record of this final activity which often follows cell construc-
tion.

The bee lettered "c" was first seen at 1 : 19 p. m. on June 19, 1951,
as it left a cell situated in a small hollow just below the surface of the
ground near Lawrence, Kansas. The bee was last seen thirty-eight
minutes later when it left the nest, so far as we could tell, for the
last time. The record is presented, in spite of its brevity, since it is
the only information available on the closing of an underground
cell. The space was completely occupied by the one cell and no
more cells could have been constructed here.

The bee lettered "d" was first observed at 2:12 p. m. on June 30,
1951, as it left a nest in a curled leaf of Eupatorium perfoliatum in

Michener: Biology of a Leafcutter Bee 1711

Area 6. The bee brought in about 14 leaf pieces in the next hour,
then started collecting pollen. It brought in its ninth pollen load
and ceased work for the night at 5:39 p. m. The next morning it
resumed activities at 8:43. Observations ceased six hours and
twenty minutes later as the bee was provisioning a second cell. The
unusually long period at the nest (5 minutes) just before the last
departure observed was due to the fact that the curled leaf had
wilted some in the hot sun and the bee had trouble getting into her
nest.

Nest "e" (figure 9) was found in a fasciate Erigeron canadensis
plant in an alfalfa field at Hutchinson, Kansas. The bee was seen
to enter the nest at 11:45 a. m. on July 26, 1950. The bee was col-
lecting alfalfa pollen at the time, and in due course laid an egg and
closed the cell with petals and leaves. Observations were con-
tinued for a total of 3 hours and 10 minutes.

Nest "f" was established in the same fasciate Erigeron as nest "e".
The record shows every visit in the construction, provisioning and
sealing of a cell. The first leaf was brought at 4:16 p. m. and the
cell was partly constructed when the last trip of the evening termi-
nated at 6:07 p. m. Activity began next morning at 9:35 a. m. and
observations continued until the bee was caught at 6:02 p. m. after
having apparently completed closing the cell.

Nest "g," also in a fasciate Erigeron near Hutchinson, Kansas, was
located at 12:23 p. m. on August 2, 1950, as the bee entered the
nest. It was observed through a series of pollen collecting trips
to the beginning of closing of the cell two hours and ten minutes
later, at which time the bee was taken to verify its identity.

Nest "h" was found on August 9, 1950, at 9:38 a.m. as a bee
entered a hole in a dead prostrate Ambrosia stem in Area 6. The
bee was carrying pollen when first seen, but soon after brought
in petals and leaves to close the cell and to construct the cup of the
next cell, and by shortly after noon was carrying pollen for the
next cell. At 2:39 p. m. it started on the first trip to get petals to
close this cell, the second cell to be closed this day. Petals and
leaves were brought in until the last trip of the afternoon which
ended at 5:30 p.m. Next morning activity started at 8:53 a.m.
with bringing of more petals and leaves. At 9:36 a. m. the first pollen
collecting trip started, the last one for this cell terminating at 12:19
p. m. Petals and leaves for closing the cell were then brought, and
more of both were used to close the hole in the stalk. The bee was
last seen about the nest at 2:12 p. m.; two hours of watching after
that time indicated that the bee had left permanently.

1712 The University Science Bulletin

The bee lettered "i" was found on August 11, 1950, at 12:35 p. m.
as it flew from its nest in a piece of Ambrosia stalk lying in the grass
in Area 6. The first petal to cap the cell was brought in 35 minutes
later, at 1:10 p. m. and by 3:30 p. m. the cup of the next cell in the
series was complete. Pollen was carried into the cell until 5:03
p. m., the last trip of the day. Next morning it was cloudy and the
bee did not leave the nest until 11:15 a. m. and provisioning of the
cell started the previous afternoon was not complete until 3:29 p. m.
The bee then brought in a few petals and leaves, after which it
seemed to cease work; it had not returned to the nest for the evening
when the observer left at 5:53 p. m. The next day, August 13,
threatened rain all day. The bee was seen carrying leaves, although
she had only a few suitable periods during the whole day. She
must have completed the next cell cup in the series, however, for
after she left the nest the next day for the first time, at 10:45, she
returned with pollen. Provisioning was complete and the first petal
to form the cap of the cell was brought in at 1:10 p. m. on August 14.

A study of figures 8 and 9 shows some interesting things about
the timing of the trips to and from the nest. The trips at the time
pieces of petals and leaves are being brought to the nest are very
irregular in length, varying from 20 seconds to two hours and 26
minutes. Sometimes considerable series of very short trips are
made, for example there is a series of ten trips in ten minutes shown
in the record of nest c. Among the data shown in figures 8 and 9,
plus other records, are 335 timed trips which ended by a piece of
leaf or petal being brought to the nest. Of these, 93 were trips of
less than a minute in duration, 203 were trips of one to five minutes
in duration, 20 were trips of six to ten minutes, 8 were of seven to
fifteen minutes, three were trips of 15 to 20 minutes, and seven
were trips over 20 minutes in duration.

There is no evidence as to what the bees were doing on the very
long trips; possibly they might merely suck nectar. It is apparent
from figures 8 and 9 as well as from examination of nests, that leaves
usually are gathered in series preceded or followed by series of
petals; it is unusual that a petal is brought in after a leaf and fol-
lowed by another leaf, or vice versa. There is no evidence that
the bees take longer to obtain leaves than petals or that the first of
a series takes longer to obtain than others. As is fully confirmed by
the study of nest structure presented earlier, figures 8 and 9 show
that the last additions to the cup (innermost layers) are ordinarily
petals as are the first or innermost layers of the cap.

Michener: Biology of a Leafcutter Bee 1713

The length of time spent in the nest between such trips is short,
ranging from as little as 10 seconds, in cases where it was later
found the bee had been merely using leaves or petals to fill excess
space in the nesting cavity, to 10 minutes. In the latter cases the
bee apparently merely rested in the nest. In most cases where the
bee was continually active two minutes was the maximum period
spent in the cell between leaf or petal gathering trips.

The foraging trips, by contrast, are of relatively uniform duration
for any one nest. Thus for nests e, f, and g the trips are rather short,
four and one-half to ten minutes, except for one of 23.5 minutes.
This suggests that pollen can be collected rather quickly from
alfalfa. By contrast, pollen collecting from Rhus and Vernonia
seemed to require a longer time per trip, as can be seen, for example,
from nest "d," in which the shortest trip was 10.3 minutes, the
longest 29 minutes.1 For this nest the pollen source (Rhus) was
not near the nest ( it was at least one-fourth mile away and this may
account for the long duration of the trips. The length of time spent
in the nest between pollen collecting trips was one to two minutes
in nearly every case.

After provisioning and before bringing in further petals and
leaves, a pause in outside activity consistently occurs. This is the
egg laying time. The bee is in the nest from 4.5 to 9 minutes
( average of 13 observations, 6.6 minutes ) at this time.

Study of figures 8 and 9 suggests that unless the bee is gathering
pollen, the last trips in the late afternoon tend to be rather long and
to result in nothing visible being brought back to the nest.

It is also evident that when a nest is completed, it is not always,
if ever, summarily abandoned; the bee often returns once or twice,
carrying nothing, and buzzes about the nest or alights there before
going on. In one such instance (nest f) it was noted that the bee
was acting as though looking for a new nesting site. This would
suggest that after one nesting cavity is filled or abandoned, the bee
goes on immediately searching for another.

Besides irregularities of action already noted, there are occasional
trips for which there seems to be no explanation whatever. An
example is seen near the beginning of the record of nest "g". After
bringing in a load of pollen, the bee left, was gone for some 45 sec-
onds, returned with nothing visible, then left on another pollen
collecting trip. On another occasion a bee in the midst of con-

1. Temperature does not seem to be well correlattd with duration of pollen collecting
flight.

1714 The University Science Bulletin

structing the cup of a cell left the nest and rested on a leaf ten feet
away for a full minute, then returned to the nest, and entered it,
then left and soon brought back another leaf. The number of such
irregular trips is very small, as is shown by the small number of
"o" marks on figures 8 and 9.

Petal and Leaf Cutting

Leafcutter bees are widely known to use rose leaves freely in
constructing their cells. The broad range of leaves and petals used
by Megachile brcvis is something of a surprise, although other
species are known to use both petals and leaves.1 Table VII lists
the various kinds of leaves and petals which we have seen cut by
this species. Certain European authors ( e. g. Hardouin, 1945 )
have overemphasized specificity in the kinds of leaves cut by species
of Megachile such as M. centuncularis Linnaeus, a species similar
to M. brevis in many ways. This emphasis is surprising in view of
the lists of different kinds of leaves cut by various species provided
in Friese's (1923) review and in much earlier papers referring to
M. centuncularis, for example by Buysson ( 1902 ) and Ferton
(1896).

As was shown in more detail in the section on nest structure, the
various kinds of leaves do not fill the same needs. The petals used
on the outside of the cell and the thin leaves such as those of the
Leguminosae serve merely to fill up extra space. The firmer leaves,
such as those of rose, give the cell its shape.

If one examines a region where Megachile brevis is nesting,
scattered leaves or petals from which a piece has been removed by
a bee will be seen. However, most of the cutting is found to be in
small areas, for example a particular branch of a rose bush or a
patch of Oenothera flowers a couple of feet square. There are no
evident differences between such places which are favored for
cutting and similar bushes or patches a few feet away.

The following notes made on June 14, 1951, in Area 3 illustrate
this and other points: '"Occasional rose petals cut by Megachile
could be found scattered over the entire prairie. Most of the
cutting, however, is concentrated in small areas two to four feet in
diameter. In these places both petals and leaves are cut. The
number of cuts in each little area is such that all may have been
made by a single bee. One similar area where strawberry leaves

1. The author has examined a nest of Megachile montivaga Cresson made in a green
weed stem near Eagle Rock, Los Angeles County, California. The cells in this nest were
made of the pink petals of Phlox and leaves of poison oak, Rhus diversitoba, which had be-
come red with fall coloring.

Michener: Biology of a Leafcutter Bee

1715

Table VII. — Kinds of leaves and petals utilized in nest construction.

(x indicates a single observation or a single nest, xx several observations or nests, xxx indi-
cates repeated observation in many nests. )

Scientific name

Common name

Leaves

Petals

Remarks

Polygonaceae

Polygonum opelusana

Smartweed

Polemoniaceae

Phlox (cultivated)

Phlox

Solonaceae
Petunia

Cultivated Petunia

Rosaceae

Fragaria virginiana

Fragaria

Potent ilia simplex

Rosa suffulta

Rosa

Wild Strawberry
Strawberry
Cinquefoil
Prairie Rose

Cultivated Rose

X
X
X
XXX

XXX

Important source of leaf

pieces.
Important source of leaf

pieces.

Cassiaceae

Cassia chamaecrista

Partridge Pea

XXX

Important source of petals

Fabaceae

Medicago sativa

Lespedeza violacea

Lespedeza virginica
Trifolium repens

Alfalfa
Bush Clover

Bush Clover
White Clover

XX
XXX

X
X

Whole or nearly whole
leaflets used.

Important source of whole
or nearly whole leaflets.

Nearly whole leaflets.

Whole or nearly whole
leaflets.

Lythraeeae

Lythrum alatum

Winged Loosestrife

Crassulaceae

Penthorum sedoides

Ditch Stonecrop

Oenotheraceae
Ludu'igia

alternafolium

Oenothera speciosa

False Loosestrife

White Evening
Primrose

Leaves bitten but piece
not carried away.

Aceraceae

Acer sp.

Cultivated Maple

Leaves of young seedling.

Caprifoliaceae

Symphoricarpos
orbiculatus

Buckbrush

Compositae
Zinnia sp.

Silphium perfoliatum

Cultivated Zinnia
Cupplant

X
X

1716 The University Science Bulletin

(Fragaria virginiana) were being cut was found, and another
where leaves of Potentilla simplex were being cut. Strawberries
and roses were scattered over the whole prairie." Nevertheless
bees observed cutting returned unhesitatingly to their little patches.

It seems likely that an advantage of this tendency to return to a
single place for leaves and petals is that it makes possible rapid
gathering of these materials without the need to search on each
trip.

We have seen alfalfa leaflets cut only three feet from the nest,
and have seen pieces of rose leaves and leaves of Lespedeza
virginica cut 150 to 200 feet from the nest. In the latter cases rose
and Lespedeza leaves could have been found within ten feet of the
nest. The time spent in flights of such lengths is negligible com-
pared to that that might be spent in searching.

Figures 8 and 9 show that petal and leaf collecting trips away
from the nest vary greatly in length but that the periods of time at
the nest are comparatively constant. Observations of individual
bees made at cutting places show that the length of time spent in
cutting is rather constant, but that there is great irregularity in the
length of time away from the cutting place. This indicates that
the bees may spend considerable time on some trips before getting
to the cutting place, for after cutting a leaf they always fly directly
toward the nest. This time is perhaps spent in sucking nectar or in
searching for new leaf or petal sources.

The bees cut leaves and petals with remarkable speed. The cut-
ting is done by means of the mandibles (figure 5). We noted the
cutting of a round piece of rose leaf such as is used for capping cells
in 3 or 4 seconds, and timed the cutting of the longer pieces used
for cup walls at 13 to 30 seconds.

Sometimes a bee cuts the first leaf it alights on but very often it
flies on to leaf after leaf, as though for some reason dissatisfied. We
watched one bee alight on 38 leaflets of Lespedeza violacea before
finally cutting one off. As shown in figure 10, the bees very com-
monly start to cut into a leaf, and may have a piece half cut off,
before abandoning it and going on to another. This hesitancy is
far more noticeable when bees are not working in a repeatedly
visited cutting place, but occurs even in such cutting places.

The cutting process begins with the bee resting on the edge of the
leaf ( or petal ) with the legs of one side clinging to the upper surface
of the leaf, those of the other side clinging to the under surface.
The bee cuts into the edge of the leaf with its jaws, cutting in a
smoothly curved line irrespective of veins in the leaf ( e. g. in small

Michener: Biology of a Leafcutter Bee

1717

rose leaflets it cuts through the midvein, figure 10). As it cuts, the
bee turns its body so that a piece of standard shape (long for the
cup, round for the cap) is cut out. The middle part of the bee's
body moves very little, merely rotating as the head and jaws de-
scribe an arc while the cutting goes on. As the cut is made the bee
shifts its footing so that it is clinging to the piece being cut. At
least in the case of an elongate piece, the piece is curled as it is cut
by bending the edges away from the body of the bee. This is done
as the bee shifts onto the under surface of the piece being removed,

14b

Fig. 10. Rose leaflets cut by Megachile hrevis. a, leaflet with two incom-
plete cuts and one long piece removed; b, small leaflet with one long piece
removed, showing that the bee cut across the midvein; c, leaflet with round
piece removed.

Fig. 11. Leaflets of Lespedeza virginica showing the ways in which long
pieces are obtained from leaflets of various sizes, a and b are common, c rare
since leaflets are rarely so large.

Fig. 12. Leaflets of Lespedeza violacea showing ways in which they may be
cut whole or nearly so for use as filler leaves.

Fig. 13. A female Megachile brevis showing the way in which she holds a
piece of a leaf.

Fig. 14. Diagrams showing the way in which a bee places long leaf pieces
to form the cell cup. The circles represent the hollow (or already partly con-
structed cup) seen from the open end. The leaf piece is represented by the
broken lines. The bee, headed into the cell, is represented in black. The bee
carries the leaf into the cell in the manner shown in a, moves sideways, as
shown in b, until she gets past the edge of the leaf, as in c.

1718 The University Science Bulletin

so that as the six tarsi hold the edges of the piece away from the
bee's body, the center is bowed against the sternum. In every in-
stance observed carefully (about 30 cuttings of leaves, three of
petals ) the piece was held in this fashion with the original under sur-
face (of leaf) convex against the venter of the bee. As will be
shown later, this results in the under surface of the leaf being outer-
most in the complete cell. Cutting continues around the arc char-
acteristic of the type of piece being removed, and at the instant
that the piece is severed the bee takes flight. The details of the
cutting process must be rather constant among a large group of
species of Megachih; see for example Ferton's (1893) notes on
leaf cutting in M. centuncularis Linnaeus. Sizes and shapes of
pieces removed will be discussed in more detail under nest structure.
When the bees are cutting small leaves, leaflets, or petals which,
because of size, require a minimum of cutting, the cutting pro-
cedure is surprisingly little modified. Thus when a bee alights on
a leaflet of Lespedeza violacea, she starts to curl it in the usual way
as she works around the edge of the leaf with her jaws, not cutting
it. When she reaches a point on the edge of the leaflet where the
normal cutting arc enters the leaflet, she cuts into the leaflet and
cuts it off ( figure 12 ) . This action suggests that cutting from larger
leaves is the primitive behavior pattern, modified in Megachih
brevis until more small leaflets and petals are used than pieces of
larger ones. The pieces of larger ones are, of course, more ac-
curately shaped and it is perhaps significant that the firmer leaves
responsible in most cases for the shape and firmness of the cell are
always cut from moderate-sized leaves (e.g., rose) and are not
merely severed leaflets.

As shown in figure 11, the leaflets of Lespedeza virginica are of an
intermediate size so that, while a piece may be cut out as with rose,
more often the tip of a leaflet is cut off and dropped, then the jaws
of the bee follow the leaf edge to a point near the base where they
cut into the leaf and across it.

In one instance a bee was seen to cut off and carry away a small
triangular piece of rose leaf left between two normal sized pieces
removed previously. It is possibly significant that this was done
on the last of at least ten trips of the bee to a particular cutting area.
Once the bee has cut off the leaf piece, it usually flies from one
to ten feet and alights on a leaf or on the ground, remaining there as
though resting for 45 to 80 seconds ( average of 10 observations, 61
seconds). While resting the bee holds the leaf as previously de-
scribed, curled under her body, the claws holding its margins

Michener: Biology of a Leafcutter Bee 1719

( figure 13 ) . Just before flying off toward the nest, the bee, in every
instance observed, rubbed her abdomen against her closed wings
several times. This brief period of inactivity following cutting is
often omitted but only when bees are cutting either soft and easily
cut petals or leaflets which required little more than severing.
Eleven cuts of Lespedeza virginica leaflets by a single bee 1 were
observed. Of these only two leaflets were large enough to require
cutting around three sides; after these two the bee rested. The
others were smaller leaflets requiring less cutting (see figure 11)
and the bee did not rest but flew directly to the nest, 150 feet away.
Data on how far from the nest bees will go to obtain leaves and
petals are very scanty. We have seen them cut within a few feet
of the nest and we have seen pieces of petals brought from 300 feet
away. In view of the diversity of leaves and petals that can be used,
they are probably usually obtained rather near the nest.

Nest Construction

Ordinarily the cells of Megachile brevis are placed in hollows
where the actual manner of building the cell cannot be observed.
We have been fortunate in finding certain nests whose cells were
incompletely enclosed, enabling us to observe certain things about
the construction process. Three of these nests were in spaces
among the leaves of fasciate plants of Erigeron canadensis but one
was in a curled leaf of Eupatorium perfoliatum. In all cases the
construction technique was the same. In making the cup of the
cell, elongate pieces of leaves or petals are used. The bee brings
in the piece of leaf or petal curled, with the edges held away from
the body, as already described (figures 13 and 14). She carries
the leaf into the cavity in this position, entering head first, then
she releases it and works laterally around the cavity until she gets
past one edge of the leaf, as shown in the diagrams (figure 14).
Additional leaves and petals are added in the same manner so that
the cell wall is constructed from the outside inwards. After the
bee has placed a leaf or a petal in a partially constructed cell, she
works in and out and around, the abdomen moving in small, quick
(respiratory?) movements. The bee may move in and out only
twice, or up to five times. While doing so, she is mouthing the
leaves, particularly their edges, as can be seen occasionally when
she backs almost out of the cell to mouth the outer portions of the
leaves. At such times it can be seen that the apex of the proboscis

1. With an observer at the nest and another at the cutting place, there was no doubt
about the bee being the same one each time she was observed cutting even though she was
unmarked.

1720 The University Science Bulletin

is applied to the leaves although the proboscis is scarcely unfolded.
Probably this activity adds the invisible quantities of adhesive
apparently used in cell construction. The leaves stick together
only very feebly, yet they are not entirely loose when a nest is
exposed. Some species of the genus evidently stick the leaves to-
gether much more firmly (see footnote 1 in Ferton, 1898) while
others may use no adhesive material at all. When this activity is
finished, the bee backs out of the cell and flies away.

After the cell is complete, provisioned, and an egg laid, the bee
caps the cell, using round rather than elongate pieces of petals and
leaves. The cap is placed well down within the mouth of the cup.
When a bee is putting in this plug, she hangs on the edge of the
cup with her rear legs, the body in the cell and braced against the
inner walls with the other legs, then she pushes the round piece of
petal or leaf into the opening with her head and mandibles. (This
closing behavior was carefully noted only once, and may vary more
than indicated here. )

Provisioning

The partially exposed cells described in the preceding section
provided an opportunity for observations of provisioning activities.
When a bee returns to her cell with the scopa (the long hairs on
the under side of the abdomen) full of pollen, she quickly enters
the cell head first and remains in this position for 20 to 45 seconds
(average of 14 observations, 28 seconds). While in this position
the apex of her abdomen, which is visible inside the cell, can be
seen to tremble slightly. During this time the bee is no doubt
placing nectar in the cell. After this the bee quickly backs out of
the cell, turns around, and backs in. When she backs out the
pollen can still be seen in the scopa. Occasionally as she backs in,
the beginning of probable combing motions by the rear legs to
remove pollen from the scopa can be seen. She remains in the
cell in this position from 40 to 65 seconds (average of 12 observa-
tions, 50 seconds), her head visible within the opening of the cell.
Just before leaving the cell, the bee moves from side to side a little
and often brushes the head and eyes with the front legs; then she
crawls out and quickly flies away. As she leaves it can be seen
that the scopa is clean.

Unless the cell is vertical, the bee consistently enters the cell with
her ventral surface down during the provisioning process, not
rotating on her long axis within the cell as when the cup is being
constructed.

Michener: Biology of a Leafcutter Bee 1721

Observations of the accumulating provisions in the cell, made
while the bee was away, gathering more, show them to be firm
(not sticky as in opened cells) and with the surface flat, at right
angles to the long axis of the cell and smooth as though tamped
down.

The distance from which Megachile brevis will gather pollen is
unknown but we have observations of pollen gathering from a few
feet to one quarter mile from the nest, and circumstances often
suggest that they may go much farther than this.

Egg Laying

The behavior connected with egg laying was observed five times
in cells which were sufficiently exposed that some details of the bee's
activities could be seen. In each case the procedure was as de-
scribed below. Minor variations in timing occur, of course, as can
be judged by the variations in the length of the egg laying period
shown in figures 8 and 9.

On returning from her last pollen gathering trip, the bee enters
the cell head first presumably to regurgitate nectar, then backs out
and backs into the cell and brushes the pollen from the scopa, just
as after any other pollen collecting trip. She then comes out of the
cell head first, quickly turns around at the entrance and re-enters,
head first. She now works for about 50 seconds, moving about in
the cell and often turning the body on a longitudinal axis so that
she may be clinging to the roof of the cell (if it is horizontal) part
of the time. During this period the apex of the abdomen (the only
part visible) is sometimes bent downward in an unusual way. The
bee then backs out of the cell, turns around, and backs in, remaining
there about 45 seconds. It is during this period that the egg is laid
on the surface of the mass of provisions. She then comes out head
first, turns around, goes in head first again, once more working
around the cell and curling the apex of the abdomen downward.
This continues for about two minutes (in one case five minutes),
after which the bee backs out and flies away, soon to reappear with
a petal and start capping the cell.

In an ordinary nest in a weed stalk none of this activity is visible,
for the hole in the stalk is large enough to allow the bee to turn
around outside the cell but inside the stalk. It is interesting that
in the megachilid genus Hoplitis (subgenus Alcidamea) precisely
similar enterings and re-enterings occur at egg laying except that

77—8663

1722 The University Science Bulletin

the diameter of the hole in which the cells are constructed is so small
that the bee must come all the way out of the nest in order to turn
around.

Flight Patterns Around Nests and Leaf-gathering Places

The most noticeable feature of the arrival at and departure from
a nest is its rapidity. When an observer first takes his place near a
nest, the returning bee may be somewhat disturbed and may fly
about the vicinity or about the observer for a short time before
entering the nest, but when she becomes used to the observer and
her activities are undisturbed, she ordinarily flies directly toward
the nest, often with a little zigzag motion in the last few feet of the
approach (figure 15). If the nest is among grass or weeds the bee
may or may not have an invariable route through the obstructing
vegetation; the flight through it is always slow, however, compared
to flight in the open. One nest located under grass on an open
prairie, where landmarks are presumably poor for the bees as they
are for us, was always approached from one side, the bee swinging
around and then zigzagging broadly in the last eight or ten feet of
the approach (figure 15). The same bee regularly approached a
leaf cutting place on the prairie directly, with no zigzagging what-
ever, perhaps because no great precision was required, the leaf
cutting area being two or three feet in diameter. The most exten-
sive searching flight which we observed in the approach to a nest
occurred in the case of a bee which nested among small rocks along
a roadside. Similar small rocks had been dumped for 100 yards
along the road and the rocky surface looked much the same every-
where. The bee was rarely able to approach its nest directly and
often, particularly if it had been away for over five minutes, flew
up and down over the rocks, often going as much as 20 feet in the
wrong direction before finally narrowing its field of search and then
zigzagging toward the nest. Such behavior is very striking compared

cr^ —

I5a 15b 16

Fig. 15. Patterns of arrival at nests, a, the common pattern, in which the
bee scarcely slows down to zigzig before entering the nest; b, a pattern in which
a bee regularly circled and zigzigged rather broadly on the way to the nest.

Fig. 16. Flight pattern of departure from a cutting place.

Michener: Biology of a Leafcutter Bee 1723

to the vast majority of nests studied, to which direct approaches
were made by the bees, probably because brushy and weedy situa-
tions usually provide adequate landmarks. Similar difficulty in
locating a nesting site was recorded by Bau (1916) for a female
nesting in a railroad tie. The bee searched the wrong tie. Presum-
ably all the ties along the track were much alike.

In leaving the nest in favorable weather the bee comes out, im-
mediately takes wing and is gone, usually in an essentially straight
course. Occasionally in cool weather, or at the first departure in
the morning, the bee crawls out of the nest and rests for a few sec-
onds to a minute or more before flying. It was noted that in leaving
her nest on the open prairie, the bee mentioned above regularly
spent about two seconds zigzagging or circling over the nest site.
The same bee, leaving a leaf cutting place on the prairie usually
zigzagged only slightly (figure 16).

Orientation flights around a newly discovered nesting site were
observed three times, and about a newly discovered cutting place
once. These flights did not follow any consistent pattern such as
flights of increasing length from the site or flights in increasingly
large circles around it but involved merely irregular flights over the
site, and in the case of the nesting places, several returns from many
yards away.

From the usual nest, the pollen source is more or less definitely
in a particular direction, while sources of petals and leaves are in
other directions. The bee, under such circumstances, often (al-
though by no means always) uses particular routes as she leaves
the nest and returns to it. The routes of departure may or may
not be identical to the routes for returning. Our notes contain
numerous references to the exact way which a bee, during any one
phase (e.g., pollen gathering), follows a certain course, identified
for us at least by bushes, weeds, and other fixed objects. When
several observers could be got in the field, it was sometimes possible
for them to take stations along the line of flight from the cutting
place to the nest and see that the line of flight was followed quite
exactly time after time for the whole distance ( 150 feet ) .

On the other hand, we have watched a nest ( d, figure 8 ) , in which
the bee rarely took the same course in leaving on its pollen collecting
trips. The bee flew in a generally westerly direction for pollen but
left in directions as different as north northwest and southwest and
sometimes returned from directions as different as 45 degrees from
the direction taken on the outward journey. The pollen source

1724 The University Science Bulletin

(or sources) was at least one quarter of a mile away. The same
bee, when collecting petals (mostly of Lythrum, one of Cassia?)
and leaves, flew eastward, following approximately the same course
each time. (There is no general rule that bees fly in the same
direction for petals as for leaves; they may fly in opposite direc-
tions. )

The pollen collecting flight patterns of three bees nesting in alfalfa
fields in the region of Hutchinson, Kansas, are interesting. One
of the bees flew consistently northward from its nest, apparently
gathering pollen from a particular part of the field, which, however,
was not different so far as we could see from other parts of the
field. The other two bees (one of which nested within 15 feet of
the above bee and was observed on the same day) flew from their
nests in many directions, apparently at random, for alfalfa pollen.
These bees were able to return directly and quickly, without search-
ing, from any direction and sometimes returned from a direction
differing from the line of departure by as much as 90 degrees.
All three bees flew in particular directions to gardens around nearby
houses for petals and some of their leaves, but they obtained other
leaves from the alfalfa plants around the nests.

Changes in Behavior

Because of the mobility of Megachile brevis populations and the
tendency of the females to construct cells in different places, we
have no records of activities throughout the lives of individual
bees; therefore we have no knowledge of changes which may occur
in behavior during the life of a bee. However, in the brief period
required for construction of single cells some observations on
changes of activities are possible.

It has been noted that when the female bee concludes one phase
of its activity and goes into the next, the change in behavior is
abrupt and absolute. Thus when the bee finishes constructing the
cell cup it promptly changes to pollen collecting behavior, and after
the cell is provisioned and the egg laid, it promptly returns to leaf
cutting behavior. A good example is provided by the data on
nest "i" presented in figure 9. After each of the two egg layings
indicated in this record, the bee, which had been flying in a generally
westward direction for pollen, came out of the nest and without the
slightest delay flew eastward for petals. In each case she was back
to the nest with a petal within one minute. Doubtless because of
previous experience in the area the bee was able to go quickly

Michener: Biology of a Leafcutter Bee 1725

to a petal source. Examination of figures 8 and 9 shows that first
trips after changes in phases of activity average no longer than later
trips. This indicates that for these bees no extensive searching for
materials (petals, leaves and pollen) was necessary. The bees must
have "known" of the sources of the materials at the time they left
the nest. This is suggestive of a delayed reaction to previous con-
ditioning. There is some evidence to support the idea that much
behavior of aculeate Hymenoptera results from delayed reactions
which were first definitely recognized among invertebrate animals
by Baerends' ( 1941 ) working on Ammophila.

There is some evidence, meager but nonetheless suggestive, that
within any one phase of its activities, changes in sources of materials
are made somewhat gradually. The details of a change from col-
lecting Vernonia pollen to that of Rhus were observed on August
11, 1950, in Area 6. The bee had been gathering pale Vernonia
pollen for several days and had provisioned three and one half cells
with it. In the midst of provisioning the fourth cell, as she left on a
pollen gathering trip, she was seen to fly toward some nearby Ver-
nonia and when nearly there turn and fly on a different course
toward some Rhus bushes 100 yards away. She returned with yellow
Rhus pollen. On her next trip, she flew again toward the Vernonia
but quickly veered toward the Rhus. Thereafter she flew directly
toward the Rhus from the nest, completing the fourth cell and pro-
visioning two more with its pollen.

Another instance of gradual change in behavior concerns leaf
cutting operations of a bee observed in Area 3 on June 16, 1951. The
bee had been gathering leaves of Lespedeza virginica at a very
rapid rate, obtaining them from a small cutting place on the prairie.
An observer was at the nest, another at the cutting place, 150 feet
distant. The bee made 11 trips for leaves in rapid succession, being
away from the cutting place for periods of time varying from one
and one-half to three minutes. On the twelfth and thirteenth trips
the bee was away from the cutting place for 7.5 and 6.5 minutes.
Most of this time was spent in the nest but the two observers
noted that her flight time from nest to cutting place nearly doubled
on these two trips although when she arrived at the cutting place,
she came from the direction of the nest. On the fourteenth and
fifteenth trips, the bee left the nest in the direction of the cutting
place, but did not alight there. The observer there (CDM) was
virtually certain that on each trip she passed the cutting place and
then headed in a southerly direction; at any rate an insect that

1726

The University Science Bulletin

looked and sounded like a female Megachile brevis arrived at the
expected time ( indicated by a shout from the observer at the nest )
from the direction of the nest (northwest) and then headed south-
ward. On each of these trips the bee returned to the nest with
a piece of rose petal. There was no indication that she returned
to the nest via the leaf cutting place. On the fifteenth trip the bee
again left the nest, headed toward the leaf cutting place but was
not detected there; on the sixteenth trip she left the nest headed
in a more southerly direction, presumably toward the source of
petals. From each of these trips she returned with a piece
of rose petal.

Michener: Biology of a Leafcutter Bee 1727

PART V.— GROWTH AND DEVELOPMENT

Technique of Study

After the egg is laid in a cell and the cell is capped, the bee goes
on to the construction and provisioning of other cells, giving no
further attention to the egg or resultant larva. The developmental
stages are ordinarily hidden from view. Two principal methods
were used to study them. Cells, removed from the hollows in
which they were constructed, were slit longitudinally on one side
with a sharp blade. They could then be opened as desired for
inspection of the contents. This method has serious disadvantages
for the cells must be kept in a humid atmosphere to prevent desic-
cation of the pollen mass. Fungal growth therefore often occurs
on the leaves, and later on the pollen. Moreover, the cap of the
cell usually falls out. Since the larger larvae often press against
the cap with their middle or posterior portions in order to force
their heads down into the pollen, lack of the cap seems to cause
slow feeding and sometimes larvae seem to have difficulty finding
the food.

A better method consists of moving pollen mass and larva into a
glass tube having an inside diameter of about 5 mm. The tube
should be plugged at each end with absorbent cotton, the cotton
plugs being about 10 mm. apart at their inner ends. Mold rarely
grows in such containers. If they become too dry water may be
added through the plugs.

Another good method consists of filling a Stender dish with
paraffin, then making depressions 5 mm. in diameter and 10 mm.
deep in the paraffin. Young larvae live very well in such containers,
humidity being provided as needed. Visibility is very good as the
lid of the Stender dish can be removed and the contents examined
under a binocular. Older larvae do best in tubes, however.

Obviously all these methods involve considerable artificiality;
this may account for some of the rather great variability noted in
the duration of various stages. The records given below, however,
omit cases where larvae were obviously away from their food for
long periods or where other such avoidable abnormalities occurred.

All records of duration of stages were obtained at room tempera-
ture during August. Most larvae were observed but once each
day, so that the margin of error in timing activities of short duration
is great.

1728 The University Science Bulletin

Egg

The egg is about three millimeters long or slightly less and 0.6
or 0.7 mm. thick (figure 18), soft and smooth on the outside, glis-
tening white in color. It is laid on the top of the pollen mass, stand-
ing up from the surface of the pollen in a slanting position (figure
17). If the pollen mass becomes somewhat liquid, as often occurs,
or if the nest is jarred in carrying it to the laboratory, the egg will
slump down until it lies at full length on the pollen. This does not
appear to influence development and hatching, and it may well be
that the egg is often laid in this position.

It is exceedingly hard to tell when the egg hatches because the
chorion or "shell" is so thin and soft and because it disappears com-
pletely after being shed. However, segmentation, muscular move-
ments and gas filled tracheae can be seen before hatching. As soon
as pollen is visible in the digestive tract, hatching must obviously
have occurred. Three observations from laying time to approxi-
mate hatching time indicate that the egg stage lasts from 3 to 3.5
days.

Larva

The larva (figures 19 to 21) is a legless whitish grub. Dorsally
it is feebly brownish, and the apices of the mandibles are dark
brown. Details of certain anatomical features of the mature larva
are shown in a previous paper ( Michener, 1953 ) .

During the entire feeding period of the larva, from shortly after
it emerges from the egg until feeding ceases on maturity, small
spots which are whiter than the rest of the grayish white body can
be seen through the integument. They have been seen as little as
twelve hours after hatching and may appear sooner, although they
are absent at the time of hatching. They are not part of the cuticle,
as they can be seen moving beneath the cuticle with the movement
of the tissues inside the body. After the larva finishes feeding these
spots disappear and the general color of the larva becomes more
white, less grayish.

The number of larval stadia has not been definitely determined.
The exuviae are very delicate and are probably usually eaten soon
after ecdysis.

In the first day of its life the larva is straight and lies flat on top
of the pollen mass. It has thick projecting folds along each side
of the body which seem to help the small larva, 3 to 4 mm. in
length, to float on the often quite liquid mass of food. At this stage
the larva is unable to move about to any extent. Usually on the

Michener: Biology of a Leafcutter Bee

1729

Fig. 17. Diagrams of cell cups, a, with egg standing up from edge of mass
of provisions; b, with egg as usually found, lying on provisions.

Fig. 18. Egg of Megachile brevis.

Fig. 19-21. Larvae of Megachile brevis, first stage, half-grown, and mature.

second day, after what is probably the first molt, the larva becomes
more curved as it eats into the pollen at one side of the cell, and
the lateral folds are far less prominent. At the beginning of this
stage the larva when straightened out is about 5 mm. long. As it
grows the larva becomes more strongly and permanently curved
( so that it cannot be straightened for measurements of length ) and

1730 The University Science Bulletin

eats the pollen down toward the base of the cell. Thus it has its
head toward the base of the cell while it is eating.

When about half grown, the larva usually produces its first feces.
We have a few records, however, of individual larvae which pro-
duced no fecal material until after the pollen supply was exhausted.

Feces are remarkably variable, not only in the time when the
first ones appear but in their color, for they vary from white through
testaceous to brown and black. Perhaps the color depends on the
kind of pollen being eaten by the larva. They are cylindrical, often
1.0 or 1.5 mm. long, rather firm, the first ones produced usually
being smaller than later ones. They are produced in considerable
numbers, a dozen or twenty often appearing during the first twenty-
four hours of defecation, a total of 40 or 50 or more being reached
at the end of the period.

Since the larva lies in its cell feeding with the head toward the
base of the cell, most of the feces are deposited near the cap. They
are often crushed against the walls of the cell by the movements
of the larva. Sometimes, perhaps regularly, scattered threads of
whitish silk spun about on the cell walls by larvae serve to hold
the feces against the walls and to prevent contamination of the
food by fecal material.

Commonly about one day after the first feces are voided, the
pollen mass is exhausted. In some cases this requires longer, even
up to four days. The average of observations of this period on 15
larvae is 1.9 days, but it is probable that this average is too high
for it is during this latter part of larval feeding that abnormal
lengthening in the larval period due to laboratory conditions ( e. g.
pollen too moist, too dry, moldy, etc. ) is probably greatest. Not
infrequently in the laboratory, and in nature as well, larvae leave
some of the pollen mass at the base of the cell uneaten. This un-
consumed pollen consists of a pad one half to one millimeter thick
in the base of the cell. It is not clear whether it is left because it
has become unsuitable as food or because the larva has finished its
development and needs no more food, but the latter seems probable.

After the larva has finished feeding on the pollen mass, it spends
at least a day before it begins to spin its cocoon. Occasionally this
period extends to three days, although the average of 14 records
of this period is only 1.3 days. During this period the larva moves
about considerably turning so that its head is away from the base
of the cell and rasping the inner walls of the cell with its mandibles
and eating some of the petals which usually form the linings of
cells and which by this time have become soft and mushy. Evi-

■

Michener: Biology of a Leafcutter Bee 1731

dently the larva may reverse its position in the cell more than once
at this time for cells have been opened containing larvae which
had finished all their provisions, turned so that their heads were
toward the cap of the cell, part of which had been eaten, then
turned back so that the head was toward the base of the cell. The
larva eats especially from the cap of the cell, so that petals and
even the inner leaves of the cap become mere rings, for the larva
can get at the centers but not at the margins of the disc-shaped
pieces used in the cap. So far as known this is the first report of
bee larvae eating petals and leaves. One larva was observed eating
petals from the wall of its cell a full day before it had finished
eating pollen. Larvae reared in artificial containers without petals
and leaves seem to develop quite normally, however, in spite of the
lack of this food. Under artificial conditions, with larvae removed
from their cells, the entire period from hatching until the beginning
of cocoon spinning ranges from 5.1 to 13.3 days in summer. As
already suggested, the maximum lengths of time almost certainly
are abnormal.

Perhaps throughout the entire last half of the growth period the
larva is capable of spinning, for slits in the sides of cells are often
found closed by a few strands of whitish silk during this period.

Cocoon spinning requires from less than a day to three days, the
average of 15 individuals whose cocoon spinning was timed being
1.4 days.

The silk is spun from the slitlike salivary opening and is applied
by side-to-side movements of the head and forward parts of the body
of the larva.

The spinning of the cocoon starts with the larva in a position with
head away from the base of the cell. As will be explained later, the
cocoon consists of various layers. Presumably the larva reverses its
position in the cocoon in the process of laying down each layer. As
a result spinning larvae may be found in almost any position within
the cocoon. After the cocoon is complete, larvae consistently take
up a position with the head away from the base of the cell. In de-
scribing cocoons, therefore, the end in the base of the cell is called
posterior, the end toward the cap is called anterior. This orienta-
tion is maintained in the pupal stage so that the emerging adult has
its head directed toward the entrance of the nest.

After the cocoon is complete, there is a period of a day during
which the larva inside the cocoon is able to seal with new silk any
slit made in the cocoon for observational purposes. In one instance
out of ten observed a larva was able to do this on a second day.

1732 The University Science Bulletin

After this the larva appears to be unable to produce silk; at least
it does not mend slits in its cocoon. It remains able to move for
several days, however. The total length of the larval period after
completion of the cocoon and before pupation is three to eight days
( average of seven, 5.2 days ) in summer; individuals of the fall gen-
eration pass the winter in this stage.

Cocoon

The cocoons spun by the larvae are cylindrical with somewhat
rounded ends. A randomly selected group of 15 varied from
4.3 x 8.5 mm. to 5.2 x 10.0 mm. The average width in this group
was 4.75 mm., the average length, 9.06 mm. The cocoon of a par-
tially starved larva was only 3.75 x 8 mm., but since cocoon diame-
ter ordinarily depends on cell diameter, there probably was no rela-
tion between the starvation of this larva and the small diameter of
its cocoon.

The outermost fibers of the cocoon are slender whitish or pale
brown threads which form no continuous layer but attach any feces,
bits of pollen, or other materials to the walls of the cell and which
cause the cocoon to adhere to the inner wall of the cell except some-
times at its anterior end, where there may or may not be a small
space between the anterior end of the cocoon and the cap of the
cell. The outside diameter of the cocoon, therefore, is normally the
same as the inside diameter of the cell.

Inside of these sparse fibers is the outer cocoon. In the Coelioxys
described below it forms a complete layer, but in Megachile brevis
the outer cocoon consists of a cap of coarse red threads covering
the anterior end of the cocoon. Occasionally it is reduced to only
a thread or two or is absent; more often it extends backward over
the anterior end of the cocoon for a millimeter or two, and in one
cocoon (out of 60 examined) the red fibers of the outer cocoon
reached the middle of the cocoon. Thus the outer cocoon is always
lacking from the posterior part of the cocoon.

The red fibers of the outer cocoon are coarser than those used
in any other part of the cocoon. They vary greatly from cocoon
to cocoon in thickness. In one the coarsest of the red fibers were .08
mm. in diameter, the finest .01 mm. in diameter with most of the
fibers about .03 mm. in diameter. In another cocoon the coarsest
were .04 mm., the finest .01 mm., with most of the fibers .02 mm.
or less in diameter. Sometimes irregular thick places on the fibers
occur so that in one cocoon whose thickest fibers were .045 mm. in
diameter irregular bulges reached a thickness of .065 mm. The

Michener: Biology of a Leafcutter Bee 1733

fibers of the outer cocoon are very stiff, lie criss-crossing one another
at irregular angles, the various thicknesses indiscriminately mixed.
The outer cocoon is hard, firm, and thick in contrast to the rather
delicate and thin inner cocoon.

The inner cocoon is firmly in contact with the outer cocoon, and
the two can be pulled apart only with difficulty. Posterior to the
outer cocoon, the inner cocoon is continuous with the sparse pale
threads which are attached to the walls of the cell. These threads
and those of the inner cocoon seem identical; they are fine, .01 to
.001 mm. in diameter, pale brown, criss-cross one another irregu-
larly, with the various sizes intermixed. The inner cocoon is spun
by the larva until it is opaque, but it still appears pale brown.
A few hours later it changes to a dark brown. This presumably
happens when a dark brown liquid (of unknown origin) is spread
by the larva on the inner surface of the inner cocoon. This liquid
impregnates the inner cocoon and quickly hardens, for in every
cocoon opened this portion of the inner cocoon consisted of fibers
imbedded in a hard but flexible, amorphous, translucent, dark brown
material which obviously must have been applied to the fibers as a
liquid. This is the outer layer of the inner cocoon.

The inner layer, which is difficult to separate from the outer, is
highly variable. It is always thinner than the outer layer. It may
consist merely of pale brown fibers like those of the outer layer
applied to the inner surface of the outer layer and giving it a silvery
appearance seen from the inside. These fibers may be impregnated
and joined by the amorphous brown material characteristic of the
outer layer, or this material may be present only in the anterior part
of the inner layer. In over 50 percent of the cocoons the inner layer
does not reach the posterior end of the cocoon, but fades out short
of it. In six cocoons of the 60 studied, the inner layer of the inner
cocoon was absent.

Rarely there are one or more small additional "layers" of inner
cocoon, sometimes mere flakes, outside the outer layer or inside of it,
at the anterior end of the cocoon. These layers are impregnated
with the dark brown amorphous material. Regardless of the pres-
ence of such additional layers, there is virtually always a place
(sometimes conspicuous and sometimes minute) in the center of
the anterior end of the cocoon where the amorphous material does
not completely close the spaces among the fibers of any of the layers
of the inner cocoon. This allows for ventilation of the otherwise
airtight cocoon. This place corresponds to the conspicuous nipple
found on the anterior end of many megachilid cocoons.

1734

The University Science Bulletin

It seems probable that the inner cocoon of Megachile and Coe-
lioxys corresponds to the entire cocoon of Osmia and Hoplitis, that
the reduction of the nipple in Megachile and Coelioxys is associated
with the addition of the outer cocoon, and that the addition of the
tough, hard outer cocoon, especially anteriorly, is advantageous in
protecting against invaders.

Complexity of the cocoon in Megachile may be widespread, for
Micheli (1937) describes the cocoon of Megachile nigriventris
Schenck as consisting of no less than five layers.

Fig. 22. Pupa of Megachile brevis.

Fig. 23. Adult female of Megachile brevis.

Michener: Biology of a Leafcutter Bee 1735

Pupa

The pupa lies in the cocoon with its head away from the base of
the cell, that is toward the entrance of the nest. At first the pupa
is entirely whitish in color but after one or two days the eyes be-
come pink. They become gradually darker during the following
several days. Then other parts of the body begin to darken, the
antennae, tarsi, and tibial spurs being among the last. Finally the
whole pupa becomes black. About a day before emergence of the
adult, the pupa becomes soft and wrinkled in appearance.

The total pupal period ranges from 10 to 12 days (average of
six, 11.2 days).

Emergence

When the adult first emerges it is wet, and it may require half a
day or more to dry off and expand its wings. This occurs within
the cocoon. After this the apparently perfect adult remains in the
cocoon from two to five days ( average of 5 observations, 3.5 days ) ,
then chews its way out of the cocoon.

The adult then soon chews its way out of the cell. In so doing
it commonly chews the cap and much of the cocoon into small bits.
If cells are in a series, it is usual for the bees to emerge at about
the same time, when the uppermost bee emerges. They leave be-
hind them a tube of leaves and petals, the side walls of the cells,
the lower or inner end of the tube being filled with the remains of
cocoons and ends of cells, all broken into small pieces.

Because of the short series often available and especially because
of the high rate of parasitism in long series, no significant data
were gathered on whether females are usually reared in the older
cells and males in the younger cells of a series.

Freshly emerged bees liberated in the field visited flowers im-
mediately for nectar. One female stopped to suck from a Lythrum
flower only one foot from the point where she escaped from the
nest.

Summary of Developmental Stages

Making use of the data presented above, table VIII has been
constructed to summarize the duration of the stages in the life
history of this bee. Total figures from the time of egg laying until
emergence of the adult from the cocoon are 23.9 days for the mini-
mum column, 32.3 days for the average column and 45.8 days for
the maximum column. Probably no bee achieves the minimum or
maximum, as there is no reason to believe, for example, that a bee

1736

The University Science Bulletin

with one stage minimum in duration would have the others mini-
mum also. The average period from egg laying to adult emergence
from the cell in the case of eight undisturbed cells was 32.2 days
(minimum 30, maximum 35). This indicates that the average
figures independently obtained in the laboratory for the lengths
of larval stages are not greatly different from those existing in
nature.

Table VIII. — Estimates of duration (in days) of the stages in the life history
of Megachile brevis under summer conditions.

Minimum

Average

Maximum

egg

3.2

3.5

larva

before starting cocoon

after starting cocoon

5.1
3.9

7.7
6.7

13.3
12

pupa

11.2

adult

before emerging

after emerging

3.5

30 ±

Michener: Biology of a Leafcutter Bee 1737

PART VI.— THE CUCKOO BEE, COELIOXYS
Seasonal History and Habitats

Perhaps the most important natural enemy of Megachile brevis
is a bee of an allied genus, Coelioxys octodentata Say. This species
has previously been recorded as a parasite of M. brevis by Hicks
(1926). Coelioxys is a social parasite; it does not make its own
nests but our species lays its eggs in the cells of Megachile, where
its young larvae kill the young Megachile larvae and then eat the
provisions gathered by the Megachile.

Much less information is available on the seasonal history of this
bee than on that of Megachile brevis. Its season of flight is prob-
ably about the same, at least individuals have been collected near
Lawrence, Kansas, from May 25 to September 25. Like the Mega-
chile, the Coelioxys is scarce early in the season, much more abun-
dant later. The number of generations per year is probably the
same as in the Megachile, for available information on the rate of
development of immature stages indicates that the stages are of
about the same length as in the Megachile. The wings of the
adults become tattered with wear as in Megachile. Overwintering,
like that of Megachile, is in the mature larval stage in cocoons.
Larvae reaching this stage after about the first week in August
remain in this condition through fall and winter, as in the Mega-
chile.

The habitats of the Coelioxys are the same as those of the Mega-
chile. The Coelioxys seems equally mobile, appearing wherever
favorable flowers are to be found, in both the nesting habitats and
the nectar habitats of the Megachile. Coelioxys are often seen
sucking nectar from flowers, although of course they collect no
pollen. Their choice of flowers is about the same as that of nectar
sucking Megachile brevis, although perhaps they visit yellow
Compositae more frequently.

Finding Megachile Nests

Female Coelioxys are sometimes seen flying over the ground or
through weedy places, stopping to fly along every dead weed stalk,
especially those lying on the ground. We have never seen a
Coelioxys discover a Megachile nest but presume that this is the
searching behavior.

1738 The University Science Bulletin

Egg Laying

Several Megachile nests which had been discovered by Coelioxys
females have been observed. The Coelioxys returned to these nests
every few hours and could fly directly to the vicinity of the nest
apparently as easily as the Megachile. The Coelioxys usually alights
on a twig or grass blade and remains there perfectly quiet for min-
utes at a time. Once one was observed to remain in such a posi-
tion for 72 minutes, after which it flew away without actually going
to the nest. Sometimes after the Megachile leaves on a pollen col-
lecting trip the Coelioxys leaves its resting place and hovers at the
nest entrance for a few moments before flying away. The action
has not been observed by us but it seems certain that if the proper
amount of pollen has been accumulated, the Coelioxys enters at
such a time and lays its egg in a cell which is being provisioned.
On one occasion observations were such that it was certain that this
egg laying by the Coelioxys must have occurred during a one minute
period following the departure of the Megachile for her tenth load
of pollen for that cell.

In view of the ability of the Coelioxys to return again and again
to a Megachile nest, having once found it, it is not surprising that
whole series of Megachile cells or large parts of series are para-
sitized, while other series escape entirely.

The egg of the Coelioxys is deposited at the base of the pollen
mass with one end of the egg inserted into the leaf or petal pieces
composing the base of the cell. No doubt the slender apex of the
abdomen of the female Coelioxys is adapted for forcing its way
through the pollen for egg laying. This seems to be the common
manner of ovapositing in Coelioxys (see Ferton, 1896; Graenicher,
1927; and Iwata, 1939) but as Ferton shows, is by no means the
only one.

Larva

The first stage larva is highly specialized with a large, sclerotized
head and huge, sharply pointed jaws (figure 24). Its body is curved,
not straight like the first stage Megachile, and its size is larger than
that of Megachile. On hatching, the larva works its way slowly up-
ward through the soft pollen mass, constantly opening and closing
its mandibles. On reaching the surface of the pollen it continues
this activity and sooner or later kills the Megachile with its jaws,
usually within a day after the Megachile has hatched and while it

Michener: Biology of a Leafcutter Bee

1739

is still quite immobile. After killing the host larva, the Coelioxys
continues to move about through the mass of provisions opening and
closing its mandibles for as much as 24 hours.

Among first stage Coelioxys larvae there is considerable variation
in the size of the mandibles. Possibly more than one species is in-
volved although this is not evident from adults nor is there a clear
division of the first stage larvae studied into two or more separate
types.

There is considerable evidence that there are five larval stadia in
Coelioxys. The development of the larva of both Coelioxys and
Megachile will be discussed in a later paper.

Fig. 24-26. Larvae of Coelioxys octodentata, first stage, half grown, and

mature.

1740

The University Science Bulletin

The presumed second stage larva has short but acute mandibles.
One cell was opened in which the Coelioxys was in this stage but
the Megachile had not been killed. This is an unusual situation,
however.

Subsequent stages look much like those of Megachile (see Mich-
ener, 1953) and feed on the pollen from the surface toward the
base, not burrowing into the pollen any more than Megachile. The
easiest way to distinguish older larvae of Coelioxys from those of
Megachile is the presence of several setae on the outer surface of
each mandible (maximum of two such setae in Megachile) and of

2<? /■"

Fig. 27. Nest of four cells of Megachile brevis in an Ambrosia stalk. Open-
ing at top plugged with a few pieces of leaves (marked by arrow). There is
also a plug of leaves immediately above uppermost cell.

Fig. 28. Similar nest of six cells with plug of leaves above uppermost cell,
which is so close to end of hollow that no open space exists between last cell
constructed and plug.

Fig. 29. Abnormal fasciate plant of Erigeron canadensis. A Megachile cell
is hidden among the dense leaves near the top of this plant.

Michener: Biology of a Leafcutter Bee

1741

a genal projection just behind the mandibular base which is absent
in Megachile. The duration of the larval stages averages less than
that of Megachile, but the number of individuals is so small that
this may not be significant. Defecation often starts well before the
food is gone, as in the Megachile, and the positions taken by the
larva after the presumed second stadium as well as the orientation
of the pupa is as in Megachile.

-X -

Fig. 30. Nests of Megachile brevis in various weed stalks. The scale at the
left is in millimeters, a, entrance (arrow) at center, two series of three cells
each, one above, the other below, entrance. No entrance plug, b, Series of
five cells, no entrance plug, c, A single cell, with plug at entrance of hollow
(shown by arrow), d, Series of four cells, no entrance plug, e, Series of 3
cells, no entrance plug. This series shows how hollows of varying diameters
can be utilized by packing extra leaves or petals into large hollows. This nest
is small in diameter at bottom, large above.

1742

The University Science Bulletin

Cocoon

The cocoon differs markedly from that of Megachile brevis in that
the outer cocoon of coarse red fibers is complete, covering the entire
inner cocoon. Otherwise the cocoon is as in the Megachile, varying
widely in the coarseness of fibers and in the extent, nature, and
even presence of the inner layer of the inner cocoon.

Emergence

Emergence of adults occurs in the same manner as that described
for Megachile brevis.

Fig. 31. Cell of Megachile brevis (marked by X) among leaves of the plant
shown in figure 29.

Michener: Biology of a Leafcutter Bee 1743

PABT VII.— NATURAL ENEMIES OF MEGACHILE BREVIS

Summer

Table IX presents data on the natural enemies of Megachile
brevis as determined from 112 cells collected in July and August.
These cells were left in the field at least until the larvae were half-
grown, that is, long enough so that parasites would probably have
made their attacks. Some destruction by ants and unknown preda-
tors was probably avoided by bringing the cells into the laboratory
before emergence of the adults. Bees were considered to have
survived if by the pupal stage no parasites were evident or if they
went into winter as mature larvae with no evidence of parasites.

It is significant that not a single cell parasitized by Coelioxys was
found in any situation other than dead stalks, and that the Coelioxys
did not appear to search except in dead stalks, where they are often
able to parasitize one cell after another in a series.

Table IX. — Natural enemies of Megachile brevis

(Based on 100 cells collected in July and August)

Percentage of
larvae destroyed

Coelioxys octodentata Say 29

Leucospis affinis Say (Leucospidae) 3

Aprostocctus sp. (Eulophidae) 1 2

Merisus sp. ( Pteromalidae ) * 1

Unknown predators ■ • 5

Unknown causes (including apparent failure of bee to lay egg) 4

Winter

We do not have sufficient data on the causes of winter mortality
in Megachile brevis. The factors listed in table IX are probably
all operative in winter as well as summer. Coelioxys, the principal
natural enemy in summer, is just as important to the overwintering
generation as to any other. There are, however, additional hazards
to the overwintering brood.

Acrobat ants (Crematogaster) destroy many cells of the over-
wintering generation. The dealated queens in the fall burrow into
weed stalks and often open and pass through Megachile cells and
cocoons. Even if this treatment does not destroy the larvae, they
dry out and die when exposed in this way. Colonies of the same
ant are very common in dead weed stalks in the vicinity of Law-

1. Identified by Dr. B. D. Burks of the Division of Insect Detection and Identification,
United States Bureau of Entomology and Plant Quarantine. Both of these species were also
reared from cells parasitized by Coelioxys octodentata. It is possible although improbable
that Coelioxys was the host in all cases.

1744 The University Science Bulletin

rence and often destroy Megachile cells in the fall. Sometimes the
leaf fragments are carried out so that no evidence of the nest
remains.

Another major cause of winter mortality is trampling of nests by
cattle and other animals, a fate much less likely during the brief
period of a summer generation than during the winter months
when protecting green vegetation is gone. Burning of prairie or
pasture areas in spring or fall must also take a large toll of over-
wintering larvae.

Finally it must be admitted that some larvae die in winter for
unknown reasons possibly resulting from weather conditions.

Michener: Biology of a Leafcutter Bee 1745

PART VIIL— POSSIBLE IMPORTANCE OF THIS BEE

AS A POLLINATOR

As explained in the introduction, the study here reported was
designed, among other things, to serve as a biological basis for at-
tempts to increase the effectiveness of a leafcutter bee, Megachile
brevis, as a pollinator of alfalfa and other crops. By no means all
of our findings are encouraging from this viewpoint and serious
practical efforts along these lines have not yet been made. However,
a summary of the principal facts of the life history that relate to the
possible practical importance of this bee follows:

Individuals of Megachile brevis are effective alfalfa pollinators.
Nearly every flower visited by a female while collecting pollen is
tripped, therefore pollinated. Each female averages over 15 flowers
per minute, and the average pollen collecting trip on alfalfa is about
nine minutes long. Thus in each trip about 135 flowers are polli-
nated. If an average of 12 trips are required to provision a cell,
1,620 flowers would be pollinated per cell provisioned. Little is
known of the number of cells a female bee provisions, but if she
provisions 20, she might be responsible for pollination of 32,400
flowers. She may provision more cells; 30 is suggested as a possi-
bility earlier in this paper in connection with other data. By com-
parison, the honeybee rarely trips alfalfa flowers if other pollen
sources are available.

From a practical standpoint Megachile brevis is usually an ineffec-
tive alfalfa pollinator in Kansas and elsewhere because there are
not enough of the bees in the fields. Probably the principal reason
that the bees remain scarce is that there is not a continuous food
supply. This bee passes through several generations per year.
Thus it can build up its numbers during the summer months. How-
ever, a continuing food supply must be maintained for the various
generations by a succession of suitable food plants if this build-up
is to occur. A failure in the food supply causes the bees to disperse
widely, as they are not bound to any permanent nesting site as are
most solitary bees.

One might establish in the vicinity of an alfalfa field a succession
of blooming plants providing pollen useful to the Megachile, such as
false indigo ( spring ) , wild alfalfa ( early summer ) , iron weed ( late
summer), and wild purple asters (fall), with care being given to
have at least some alfalfa in flower at all times when no one of the
other plants is in bloom. Such a combination might attract and

1746 The University Science Bulletin

maintain a large population of this bee. The wild plants could be
mowed at the time the bees are needed for alfalfa pollination. Un-
fortunately the plants in the above mentioned series have different
soil preferences, but some farms have the various necessary condi-
tions. These plants could grow as weeds in pastures adjacent to
the alfalfa fields or in uncultivated borders or strips.

Where such a program is impossible, one may at least attract
some Megachile in advance of the alfalfa seed crop by stands of
uncut alfalfa which would bloom prior to the seed crop or by
stands of wild alfalfa. Either could be cut as the seed crop comes
into bloom.

Such highly attractive nectar sources as winged loosestrife and
buckbrush may also be useful in holding a Megachile population in
the desired area.

Nesting sites can be provided by the old stalks of large weeds
or even by old cornstalks, broken in one or two places, and left
lying on the ground where they will not be completely shaded by
trees, bushes, or growing weeds. Such nesting places should be
protected from trampling by stock and from burning.

Leaves and petals for nest construction can usually be obtained
in almost any situation where the other requirements for survival
are present. Some rosebushes will probably be helpful, since rose
leaves are much used.

Unfortunately most of the usual farming activities are opposed
to the survival and increase of this bee. Sowed pastures free of
weeds, elimination of weedy fence rows and of patches of prairie,
burning of pastures, trampling of ground by stock, and regular
cutting for hay of entire fields so that no alfalfa flowers remain all
contribute toward decreasing the abundance of the bee. On the
other hand, weedy and brushy pasture areas, fence rows and road-
side strips where flowers and weeds can grow unmolested and where
the old stalks will neither be trampled nor burned in winter, and
roadside alfalfa patches which are not or only irregularly cut, all
favor the development of larger populations of this bee.

Michener: Biology of a Leafcutter Bee 1747

LITEBATURE CITED

Baerends, G. P.

1941. Fortpflanzungsverhalten und Orientierung der Grabwespe Ammo-
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Bellevoye, Ad.

1884. Observations sur le Chalicodoma muraria, le Megachilus centun-
cularis, et l'Osmia bicornis aux environs de Metz, Bull. Soc. Hist.
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BUYSSON, R. DU

1902. Nidifieation de quelques Megachiles, Ann. Soc. Ent. France, vol.
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Ferton, Ch.

1893. Sur les moeurs de quelques Hymenopteres de la Provence du genre
Osmia Panzer, Actes Soc. Linneenne de Bordeaux, vol. 45, pp. 231-
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1896. Nouvelles observations sur l'instinct des Hymenopteres gastrilegides
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1897. Nouvelles observations sur l'instinct des Hymenopteres gastrilegides
de France et de Corse, Actes Soc. Linneenne de Bordeaux, vol. 52,
37-50.

Franklin, Woodrow W.

1951. Insects affecting alfalfa seed production in Kansas, Kansas Agricul-
tural Expt. Station Technical Bull., no. 70, pp. 1-64.

Friese, H.

1923. Die europaischen Bienen (Apidae), vii + 456 pp., pis. 1-33, Walter
de Gruyter & Co., Berlin and Leipzig.

Graenicher, S.

1927. On the biology of the parasitic bees of the genus Coelioxys
(Hymen., Megachilidae), Ent. News, vol. 38, pp. 231-235, 273-
276.

Grandi, Guroo

1934. Contributi alia conoscenza degli imenotteri melliferi e predatori,
Boll. Lab. Ent. R. Istituto Superiore Agrario Bologna, vol. 7, pp.
1-144, pis. 1-8.

Hardouin, Robert

1945. Une variation de l'instinct chez la Megachile centunculaire, Bull.
Soc. Entom. France, vol. 50, pp. 10-12.

Hicks, Charles H.

1926. Nesting habits and parasites of certain bees of Boulder County,
Colorado, Univ. Colorado Studies, vol. 15, pp. 217-252.

IWATA, KUNIO

1939. Biology of Coelioxys elongata Lepeletier, Mushi, vol. 12, pp. 34-40.

1748 The University Science Bulletin

Linsley, E. G., and J. W. MacSwain

1947. Factors influencing the effectiveness of insect pollinators of alfalfa
in California, Jour. Econ. Ent., vol. 40, pp. 349-357.

Malyshev, S. I.

1935. The nesting habits of solitary bees, a comparative study, Eos, vol.
11, pp. 201-309, pis. III-XV.

Markowsky, H.

1933. Einige selten Bienen aus der Umgebung Berlins und ein bemerkens-
wertes Nest von Megachile centuncularis L. (Hym. Apid. ), Mitt.
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Micheli, Lucio

1937. Note biologiche e morfologiche sugli imenotteri (Serie VIII), Atti
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Michener, Charles D.

1953. Comparative morphological and systematic studies of bee larvae
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Mitchell, Theodore B.

1936. A revision of the genus Megachile in the nearctic region, part II,
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1868. The home of the bees, Amer. Nat., vol. 1, pp. 364-378, 596-606.

1892. Notes on the nesting habits of certain bees, Psyche, vol. 6, pp.
340-341.

Popovici-Baznosanu, A.

1907. Megachile bombycina Rad. au point de vue biologique, Bull. Soc.
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Rau, Phil

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Reed, E. B.

1871. Notes on Megachile brevis, Say, Canadian Entom., vol. 3, pp. 210-
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1855. Catalogue of British Hymenoptera in the collection of the British
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24-8663

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Date Due

JUL 1973

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