THE OHIO
JOURNAL OF SCIENCE

(CONTINUATION OF THE OHIO NATURALIST)

Official Organ of ihe

OHIO ACADEMY OF SCIENCE

and o( the

OHIO STATE UNIVERSITY SCIENTIFIC SOCIETY

VOLUME XVIII, 1917-1918

EDITORIAL BOARD

Editor Frederic H. Krkckek

Associate Editor James S. Hine

Associate Editor J. V. Pahk

EDITORIAL BOARD

T. G. Phillips Agricultural Chemistry V. H. Davis Horticulture

F. W. Ives Agricultural Engineering W. A. Kxight Industrial Arts

F. L. Laxdacre Anatomy K. D. Schwartzel Mathematics

J. H. ScHAFFXER Botany Horace Judd Mechanical Engineering

C. B. Harrof Ceramic Engineering Joxathax Formax Pathology

J. U. WiTHROW Chemistry A. D. Cole Physics

F. H. Exo Civil Engineering R. J. Seymour . Physiology, Physiological

J. B. Park Farm Crops Chemistry and Pharmacolog.\'

X. \V. ScHERER Forestry E. R. Hayhurst ..Public Health and Sanitation

T. M. Hills Geology J. S. Hixe Zoology and Entomology

Ohio Academy of Science

R. A. BuDiNGTox Zoology G. D. Hubbard Geology

M. E. Stickne Y Botany S. J. M. Allen Physics

F. C. Waite Anatomy and Physiology

OHIO STATE UNIVERSITY

COLUMBUS

rif>^'

THE

Ohio Journal of Science

I'lTil.ISHKD BY THE

Ohio State University Scientific Society
Volume XVIII NOVEMBER, 1917 No. 1

TABLE OF CONTENTS

FoRRMAN AND Rekd— Tumors ill Dogs. II 1

WAi/roN — The Axial Rotation of Aquatic Microorgonisins and its Significance (>

Napper — Concretionary Fonns in the Greenfiekl Limestone 7

Rice— Report of the Twenty-seventh Annual ■Meeting of the Ohio Academy

of Science 14

TUMORS IN DOGS.

II. A Fibro-blastoma of the Alveolar Border of the Jaw
Contaming Giant Cells, (a. Giant Cell Epulis).

By Jonathan Forman and Carlos L Reed.
(From the Laboratories of Pathology and Physiology of the Ohio State University)

According to Wolff, ('13) the dog is more frequently
afflicted with sarcoma than any other lower animal. Frohner
observes that sarcomas in the dog occur most often in the
subcutaneous tissues. This author also mentions "Epulis
sarkomatosa" as presenting itself frequently on the upper jaw
of dogs and less frequently on the lower jaw. The specimen
described here is not presented because of the rariety of the type,
but because it apparently throws some light on the nature of the
giant cells frequently seen in epulides.

Epulis is a topographical term signifying any growth upon
the gums. Used in this broad sense, it may cover tumors of
bone, connective tissue, epithelium, inflammatory conditions,
and simple hypertrophies of the gums such as are occasionally
encountered in pregnancy. The term, however, has become
restricted to those tumors which are seated upon the gum
or the upper edge of the alveolar border and which are of
mesenchymal origin. In man three types have been described;

2 The Ohio Journal of Science [Vol. XVIII, No. 1,

first, the fibrous epulis; second, the giant cell epulis; and third,
the endothelial variety described by Whitman ('14) and Ivy
('15). The term epulis, however, seems to us to be a poor one
and we would prefer to see it replaced by the correct name of
whatever type of tumor it happens to be.

The so-called giant cell sarcoma or epulis of the jaw has been
the subject of controversy. Some authorities claiming that
it is a benign growth while others believe it to be malignant.
Several causes have led to this confusion: first, the use of the
term epulis, which does not define the type of growth; second,

Fig. 1. A Photograph of the Tumor.

too much importance has been attached to the presence of
giant cells but without due attention to their nature; third,
the giant cell fibro-sarcoma may have been confused with the
endothelial variety described by Whitman ('14).

The tumor upon which this paper is based occurred upon
the upper jaw of a dog which came into the Laboratory of
Physiology during the school year 1915-16. The dog was an
adult mongrel shepherd whose age was unknown. The accom-
panying figure gives an adequate idea of the size and location

Nov., 1917]

Tumors in Dogs

of this growth. It measures ;^ x 2 x 15 cm. and is attached by
a pedicle 1 cm. in diameter to the gum margin between the
incisor teeth. It is firm, and covered by a mucous membrane
of normal color, except at the extreme tip, where the covering
has taken on a warty appearance. The tumor is not ulcerated
at any point.

Microscopal examination reveals a fan-shaped section cov-
ered except at the point of attachment of the tumor by a normal
appearing epitheUum. The outer half of the tumor is composed
of normal appearing fibro-blasts. Just beneath this in the
connective tissue is a zone of giant cells.

Fig. 2. Giant cells containing reticular fibrils.

The distinctly neoplastic cells are more closely packed in
the pedicle of the tumor. While there has occurred an elabora-
tion of inter-cellular substances in this area, they are not so
rich in amount as in the older portion of the tumor. There is,
however, a sufficient degree of differentiation on the part of the
connective tissue cells composing the tumor in this growing
portion to place it among the benign fibroblastomas.

Giant cells are generally thought to be produced by fusion
of cells or else by division of the nucleus without a corresponding
division of the cell body. Mallory ('11) has called attention
to the presence of both types in sarcomas. Those, which are the
product of an incomplete multiple mitosis of the tumor cells, he
calls "The true tumor giant cell." These are much less
frequently found than the other type and are indicative of
the rapid rate of growth of the tumor. Since a careful search
fails to reveal any evidence of mitosis in any of the nuclei
of the giant cells seen in this specimen, it is probable that
these belong to the fusion type of giant cell.

It is usually stated that the type of giant cells due to fusion
is most frequently found in tumors associated with bone.

4 The Ohio Journal of Science [Vol. XVIII, No. 1,

although at times, as in this specimen, all the neoplastic cells
are fibroblasts and there is no evidence of either bone or cartilage
formation. This tumor, however, arose from or near the
periosteum. From this close association with bone, it has been
suggested that these giant cells were identical with osterclasts.
This type, Mallory says, (loc. cit.) "Is due to endothelial
leukocytes invading tumors, especially those involving bone,
and fusing to form foreign body giant cells. They are not tumor
cells (although the tumors containing them are the ones which
receive the name giant cell sarcomas) and usually signify only
erosion and disintegration of bone." In addition to this
source, the origin of osteoclasts has been assigned to reticular
cells and to osteoblasts.

It would be of interest then to determine if possible whether
the giant cells in this fibroblastoma were really brought about
by a fusion of cells as they appear to have been and also whether
the cells which entered into their formation were tumor cells
or were cells that had infiltrated the tumor from some outside
source.

Sections from this specimen were impregnated with silver
after the method of Maresch-Bielskowsky. By using this
method. Dr. James H. Warren has studied the nature of the
giant cell seen in early tubercles ('17). This technique as
employed b}^ Ferguson ('12) would appear to be well established
as a specific stain for reticular fibrils. These fibrils are
abundant in this tumor. This is not surprising when it is
considered that this is a tumor of rather slow growth arising
in the connective tissue of the jaw. Some of these fibrils appear
to leave the cytoplasm, and are seen free, while others bear
only a faint rim of cytoplasm. The giant cells are especially
interesting when stained with this silver impregnation. In
many of the smaller giant cells, which contain only a few
irregularly placed nuclei, the arrangement of fibrils together
with the outline of the protoplasm suggests that the giant
cells have been formed by the fusion of the reticular cells.
Even in some of the larger giant cells, reticular fibrils can be
seen in the cell body. These facts together with the complete
absence of any evidence of mitosis would make it appear that
these cells have been formed by a fusion of the reticular cells
rather than by the fusion of invading cells or by multiple
mitosis of the tumor cells.

This specimen represents a neoplasm of slow growth in
close association with bone. Since there are normally reticular

Nov., 1917] Tumors in Dogs 5

cells at the point of origin of this tumor and since there is a
close association of reticular fibrils with the giant cells as well as
with the tumor cells, it seems indicated that the giant cells in
this tumor are produced by the fusion of tumor cells and for
this reason present a tendency to differentiate as do the cells
from which the tumor arose.

That this is not the only mode of origin of fusion giant
cells seen in new growths is apparent from the description of
the origin of such cells by Whitman ('14). An examination of
the specimens of giant cell sarcoma in the Museum of Pathology
at the Ohio State University has shown that some of the giant
cells originate from endothelium. This agrees with Mallory.
It is the behavior of the more undifferentiated cells that deter-
mines the clinical course of the growth of the tumor. Giant
cells such as are seen in this specimen become, therefore, of
secondary interest for they do not determine the classification
of the tumor as a fibroblastoma. They, however, indicate
that the tumor is relatively benign. This is in agreement with
Ivy's conception ('15) that the presence of these giant cells
indicates that the tumor is probably not malignant.

SUMMARY.

This tumor arising in the alveolar process of the upper jaw
of a dog is a rather slowly growing fibroblastoma, which con-
tains giant cells.

By the application of the Maresch-Bielskowsky technique
of impregnating with silver it is established that the tumor cells
possess black argentiferous fibrils and that the giant cells
present in this tumor are produced by the fusion of cells which
also have these same fibrils in close association with them,

BIBLIOGRAPHY.

1. Ferguson, 1912. The Application of the Silver Impregnation Method of

Bielskowsky to Reticular and Other Connective Tissues. Am. Jour. Anat.
Vol. XII, p. 277.

2. Fohner. (Cited by Wolff).

3. Ivy, 191.5. Mesothelial Tumors of the Jaws. Jour. Am. Med. Assn. LXIV, 40.

4. Mallory, 1911. Giant Cell Sarcoma. Jour. Med. Res. Vol. XIX, p. 463.

5. Means & Forman, 1915. Precancerous Conditions of Face and Mouth. Dental

Summary, XXXV, p. 945.

6. Means & Forman, 1916. Epulis. Dental Summary. XXXVI, p. 494.

7. Means & Forman, 1917. A Clinico-Pathologic Study of Early Malignant Con-

ditions of Face and Mouth. Jour. Am. Med. Assn' Jan. 20, 1917. p. 180.

8. Warren, James H., 1917. The Giant Cell in Tuberculosis. Jour. Med. Res.

Vol. XXXVI. No. 2. May, 1917.

9. Whitman, 1914. Giant Cell Endothelioma of the Gums. Jour. Med. Res.

Vol. XXIV, p. 465.

10. Wolff, 1913. Lehre von der Krebs Krankheit, Jena. 1913.

THE AXIAL ROTATION OF AQUATIC MICROORGANISMS

AND ITS SIGNIFICANCE.

L. B. Walton.

Some studies concerning the axial rotation of aquatic
microorganisms occupying my attention during the last two
years, have yielded results which it is believed are of decided im-
portance in connection with the principles underlying evolution.

In general it may be stated that the positively phototactic
free swimming forms of the northern hemisphere rotate clock-
wise, assuming the observer in front of the advancing organism,
and the negatively phototactic forms counter-clockwise, while
in the southern hemisphere there are reverse conditions. The
apparent exceptions thus far noted have been found closely
allied to northern forms and may have been introduced sub-
sequent to the origin of the southern forms.

The evidence although far from complete is gradually
accumulating that the characteristic is one which has been
gradually impressed upon the organisms through the rotation
of the earth and the apparent path of the sun from east to west.
Any attempt to account for it on the basis of natural selection
can scarcely gain credence. While the possible effects of the
angular velocity of the earth or of certain electrical conditions
may be considered, it does not seem that the result could in
this way be explained. Experimental attempts to reverse
or to diminish the rapidity of the rotation of the organisms
have thus far failed.

The problem is an interesting one from the geophysical as
well as the biological point of view, and many interesting
principles come under consideration. Among these may be
mentioned the Pendulation Theory, Bipolar Distribution,
The Twining of Plants, The Tropism Theory, etc. Similarly
the direction of the spirals in Spirochaeta, Spirulina, Arthro-
spira, Spirogyra, etc., are of interest.

Modern genetics throws light on the distribution but not on
the origin of the character producing genes or factors. These
units have in general long been in existence. It is to some
such principle as the one suggested that we must look for
evidence sufficiently definite to overcome the prejudice which
has arisen against the old acquired character ideas as to the
fundamental origin of characters.

Nov., 1917] Concretionary Forms 7

There is need of studying, in this connection, the behavior of
microorganisms near the equator as well as in the southern
hemisphere, particularly in an experimental way, before definite
conclusions may be drawn.

Kcnyon College, Gambicr, O., Oct. 15, 1917.

CONCRETIONARY FORMS IN THE GREENFIELD

LIMESTONE.*

Charles W. N.'Xpper, Greenfield, Ohio.

The locality of this study is at Greenfield, in southwestern
Ohio. The workings are the Rucker Quarries. The rock is a
Silurian dolomite, the quarry face above and below water-level
measuring sixty (GO) feet. Characteristic features divide the
exposure into two parts, called the Gray Stone for the lower
twenty(20) feet and the Buff Stone for the upper forty (40)
feet.

In this paper the term "concretionary force" is employed
in the sense and with the idea of the aggregation of rock material
into various forms which are distinctly different from the
surrounding stratification.

THE GRAY STONE — THE ISOLITH.

Herein the concretionary force is manifested in one single
form. This stone is very evenly bedded, the ledges maintaining
horizontal regularity unless there is interference from large,
irregular, unstratified masses over which they bend.

The quarrymen call these masses "horsebacks" or "blisters"
and this has caused the writer to search for possibly a better
name. While producing a disturbance somewhat similar,
the terms laccolith and bathoUth can not be used for they refer
to conditions produced by the intrusion of one kind of rock,
igneous, into another, hence two kinds of rock material are
involved. In our instance, the masses causing disturbance
and the strata disturbed are of one and the same rock material.
Therefore I suggest and have used the term "isolith" for them.

*This paper, accompanied by lantern slide views and an exhibit of forms
described, read before the Geological Section of the Ohio Academy of Science,,
meeting in Columbus, at the Ohio State University, April 7, 1917.

8

The Ohio Journal of Science [Vol. XVIII, No. 1,

These isoliths have 8 x 10 x 12 feet as average dimensions
and appear as large ovals. They are a hindrance to quarrying,
being difficult to break up and causing uneveness in adjacent
ledges. Operations downward usually cease when they are
reached.

The first isoliths studied were of solid texture and not
fossiliferous, and I was then disposed to regard them as a form
separate and distinct from the brecciation well known as occurring

Fig. 1. An Isolith. Curving Strata Clearly Shown.

in the Monroe formation, of which the Greenfield Limestone
is the basal member. But later finding isoliths of brecciated
structure and fossilferous, and unable to observe the resumption
of stratification beneath, I am now rather of the opinion that
both forms, to some degree, are manifestations of the same
force of aggregation. Why then I continue to mention isoliths
among concretionary forms, will appear in the correlation I
shall present further on in this paper.

Isoliths afford us two interesting features — ledges can often
be traced through them and come out on the other side — they
sometime contain a fauna not characteristic of the immediately
surrounding ledges.

Nov., 1917J

Concretionary Forms

9

JUL

rn:

r7~i

r 1° ■" I

un

Glacial Drift Material — 10 to 15 feet.

Concrt'lionary Whorls (rare)

Waffles
Concretionary Pebbles
Masses

'Cup and Cone'
Concretions

Isoliths

Buff Stone
40 ft.

Gray Stone
20 ft.

V

Total

Section

60 ft.

V

Section at Rucker Quarries, showing position of Concretionary Forms.

I

10

The Ohio Journal of Science [Vol. XVIII, No. 1,

THE BUFF STONE.

The largest number of concretionary forms are found in
the Buff Stone — the lower fifteen (15) feet containing "cup and
cone" concretions, and the upper twenty-five (25) feet having
the concretionary "masses, pebbles, waffles, and whorls."

The " Cnp and Cone'' Concretions. — In the lower ledges of
the Buff Stone, especially in two or three strata, these forms
are found plentifully and continue so horizontally throughout
their zone.

k

Fig. 2. "Cup and Cone" Concretion in Position in Buff Stone Ledge.

As a standard form, the "cup and cone concretion" consists
of a cap, a cone, a cavity and its filling. From this we find
every possible variation by the irregular combination of these
parts. Additionally we have the "double" variety, two cones
with one common cavity. The accompanying drawings show
a perfect form and the variations. In length they are one to
ten inches and one to five inches in diameter at the largest part.
Usually stems or cones are round in cross section, although
frequently found oval.

The filling of the cavity presents an interesting study
and may be — the cavity partly or tightly filled with small
lenticular pieces of the same rock material appearing as crushed
and pressed into it — partly or completely filled with "rock tar, "

Nov., 1917J

Concretionary Forms

11

the result of carbonaceous material draining in and solidifying —
lined or filled with quartz crystals, through which sphalerite
may be disseminated — containing well formed crystals of
sphalerite — or filled with a brownish, carbonaceous clay. These
last two are very rare forms. These cavities are objectionable
in building stone.

We now pass to the concretionary forms found in the upper
ledges of the Buff Stone.

Concretionary "Masses'". — Usually these forms appear as a
distinct, continuous ledge five to thirty inches thick and two
to five in number. They rarely disturb adjacent layers, there-
fore "contorted ledges" is a term more suggestive of their
structure.

Fig. 3. "Concretionary Pebbles."
Sketched from a piece 5 x 12 x 3 inches.

Concretionary "Pebbles. " — Frequently within the "contorted
ledges" are found bunches or clusters resembling a conglomerate
of rather large pebbles. These pebbles are usually round, about
one inch in diameter and other sizes and shapes occur. When
broken for cross section, they reveal a ring around ring structure
somewhat similar to that of an onion.

Concretionary "Whorls." — A rare form in the Rucker
Quarries, but frequently found in other exposures of the thin-
bedded upper Greenfield Limestone. They appear as fine lines
drawn around a center then joining into one another. May
prove to be a fossiliferous form.

How could rock material aggregate into these various forms,
especially in strata so evenly bedded? How could this occur
while all forms and strata are of the same rock material? How

12 The Ohio Journal of Science [Vol. XVIII, No. 1,

were the ledges arched and the space filled in? Or are the
forms contemporaneous with their surrounding strata? Why
out of the six forms only two are frequently found in other
exposures of the same limestone in the same locality?

The "cup and cone" concretions with their cavities often
filled with rock pieces crushed in, suggest pressure as a cause.
But then why are they not found in the lower Gray Stone where
pressure would be greatest? Why do they occur most frequently
in a few ledges near the middle of the exposure? Always being
vertical, seldom inclined and never horizontal, has this definite
position any importance? The lithological differences between
the Gray and Buff stones, have they any bearing on the prob-
lems involved? These important questions I desire to submit
for further, careful consideration.

CORRELATION.

I have found specimens which are suggestive that there
may be a relationship between these and other concretionary
forms. If this can be sustained, then we have made quite an
advance towards the explanation of the common force which has
operated, what it is and how it has operated.

At the center of an unbrecciated, unfossiliferous isolith
which had been bisected, I found a large mass of sphalerite,
well crystallized and weighing thirty (30) pounds. Its position
was suggestive that it had been the nucleus of the isolith' s
aggregation, a striking similarity to where concretions are
formed with fish bones, leaves, etc., as nuclei.

Embedded in a fossiliferous, unbrecciated isolith I have
found concretionary "pebbles" with characteristic ring around
ring structure in cross section. This seemingly joins my lowest
and next to highest forms as the result of a common force.

The stem of the "cup and cone" concretion sometimes
has a structure somewhat similar to the "cone in cone" form.
As pressure is now regarded the probable cause of the latter,
any relationship that can be established with the former is
evidence of some corroborative character for the pressure
theory. The crushed-in particles in the cavity are ever strongly
suggestive of pressure.

In the disturbed ledges near the contact with the isolith,
I have found smoothed, grooved portions. This is evidence

Nov., 1917] Concretionary Forms 13

of pressure and movement, either the isolith upward or the
ledge downward. Can such occurrence be of use and value to
the pressure theory?

The arches of the concretionary "waffles" sometimes
become so high, large and shaped, as to be suggestive of a
transition into the "pebble" form.

In the gorge of the Cuyahoga River near Akron, just below
the electric power-house dam, in the bluish sandstone, I found
two ordinary and familiar round concretions about eight (8)
inches each in diameter and not requiring more than twenty (20)
inches of space in length over all. Midway between these, I
found a concretion very similar to the "cup and cone" type.
It has a cap, a stem or cone and an oval end suggestive of a
cavity. If this similarity has merit, then we have joined an
additional form to the series.

These observations are presented in the hope that they will
be found to be clews of some value and assistance in arriving at
a full and complete understanding of the concretionary force
and the forms resulting from its operation.

We now see that the Greenfield Limestone has features of
intense interest which its usual regularity would seem to
preclude.

REPORT

OF THE

TWENTY-SEVENTH ANNUAL MEETING

OF

THE OHIO ACADEMY OF SCIENCE

The Twenty-seventh Annual Meeting of the Ohio Academy
of Science was held at Ohio State University, Columbus, Ohio,
April 6 and 7, 1917. In the absence of the president. Professor
F. O. Grover, of Oberlin College, Professor Stephen R. Williams,
of Miami University, vice-president for Zoology, presided over
the meeting. Fifty-one members registered in attendance.

GENERAL PROGRAM

Friday, April 6

9:00 A. M. — Business Meeting.

10:00 A. M. — Reading of Papers in General Session.

11:45 A. M. — Illustrated Lecture by Professor Robert F. Griggs, of Ohio
State University: "Recent Explorations of Mt. Katmai,
Alaska." In Place of Presidential Address on "Some
Phases of Present Da}" Biological Unrest," by Professor
F. O. Grover, of Oberlin College, whose attendance was
prevented by illness.

12:15 P. M. — Lunch, Ohio Union.
1 :30 p. M. — Reading of Papers in General Session.
3:00 p. M. — Reading of Papers, in Sectional Meetings.
5:45 p. M. — Dinner, Ohio Union.

7:30 p. M. — Public Lecture by Professor James R. Withrow, of Ohio
State University: "The Relation of War to Chemistry
in America."

Saturday, April 7

9:00 A. M. — Adjotirned Business Meeting.
10:00 A. M.— Reading of Papers in Sectional Meetings.

14

Nov., 1917] Report Ohio Academy of Science 15

MINUTES OF BUSINESS MEETINGS

The first business session was called to order by Vice-
President Williams at 9:00 A. M., on Friday, April 6. An
adjourned session was held at 9:00 on the following morning.

The appointment of the following committees for the
meeting was announced by the chair:

Committee on Membership — M. E. Stickney, A. W. Smith,
F. L. Landacre.

Committee on Resolutions — E. L. Moseley, L. B. Walton,
C. G. Shatzer.

Committee on Necrology — G. D. Hubbard, Herbert Osborn,
A. D. Cole.

The following Auditing Committee was elected by the
Academy: L. B. Walton, C. G. Shatzer.

The following Nominating Committee was elected by the
ballot of the Academy just before the close of the first business
session: L. B. Walton, G. D. Hubbard, A. D. Cole.

Report of the Secretary

The following report by the Secretary, read by the Acting
Secretary, was received and ordered filed:

Cambridge, Mass., March 31, 1917.
To the Ohio Academy of Science:

The work of the Secretary for the past year has consisted mainly
of routine work, largely in co-operation with the Executive and Program
Committees.

A brief report of the Twenty-sixth Annual Meeting, 1916, was
published in Science, July 28; the full report appeared in the Ohio
Journal of Science for January. Notices of the present meeting
have been sent to the leading dailies in Columbus, Cleveland, Cin-
cinnati, and Toledo.

The officers of the Academy were notified of their election; and the
relevant portions of the report of the Committee on Resolutions were
transmitted to all persons and organizations receiving recognition in
that report.

The Program Committee is to be congratulated on securing the
services of Prof. Withrow for the Friday evening address. Prof.
Withrow is giving this very timely address at serious personal incon-
venience. His loyalty to the Academy and the spirit of national
patriotism by which he is actuated should receive the fullest recognition
of the Academv.

16 The Ohio Journal of Science [Vol. XVIII, No. 1,

The gratitude of the Academy is also due to Prof. Griggs for his
kindness and telegraphic promptness in filling in the gap in the program
caused by the illness and absence of the President.

Two minor matters concerning the make-up of the program may well
be mentioned here. Some members may be disappointed to find their
papers scheduled for sectional meetings rather than general sessions;
this has been necessitated by the presentation of an unusual number of
titles of a general character. There have been presented, also, an
unusually large number of titles unaccompanied by specifications as
to time needed. This causes no inconsiderable complication in the
arrangement of the program. A uniform allowance of ten minutes has
been made in such cases ; for the future the secretary would recommend
the reduction of this allowance to five minutes.

During the past year the Secretary has prepared a card catalog
of the membership of the Academy, past and present, with record of
dates of admission and withdrawal. Some inaccuracies (due largely
to variation in spelling and to change of name on the part of the ladies)
are still to be corrected; but the following summary is approximately
correct, and may be of interest to the members:

Ex-Members 455

Present Members 265

Total 720

The preparation of this catalog has made possible the record in the
published membership list of the dates of admission to the Academy.
The Secretary will be grateful for the correction of any errors which
may have slipped into this record.

Preliminary steps have been taken this year looking toward the
establishment of sections for Psychology and Chemistry; and it is
hoped that another year may see these sections organized. The
secretary has also very recently received the suggestion of the desirability
of a section for the medical (or near-medical) sciences — Anatomy,
Physiology, and Pathology. The program of this meeting shows the
justification for this suggestion.

Some complications have arisen because of the Secretary's temporary
absence from the state, but it is confidently expected that the very
efficient work of Prof. Krecker as chairman of the Local Committee and
of Prof. Westgate as Acting Secretary for the meeting will prevent
any loss to the Academy.

Respectfully submitted,

Edward L. Rice, Secretary.

Report of the Treasurer for the Year 1916-17
The report of the Treasurer was received as follows, and
referred to the Auditing Committee, whose report is appended.

For the year since our last annual meeting, the receipts, including
balance from last year, have amounted to $590.73, and the expenditures
to $451.63, leaving a cash balance of $139.10.

Nov., H)17j Report Ohio Academy of Science It

RECEIPTS

Balance from last year $298.13

For sale of publications 22.(50

Membership dues 270.00

$590.73

EXPENSES

Miscellaneous expenses % 4S.N8

ISO subscriptions to the Ohio Journal of Science 180.00

Publishinj; Annual Report of the 25th meeting 222.75

Balance, April G, 1917 139.10

$590.73
Respectfully submitted,

Jas. S. Hine, Treasurer.

April 7, 1917.
The Auditing Committee has verified the receipts and expenditures
of the Treasurer of the Academy, Professor J. S. Hine, and finds that
they are correct.

L. B. Walton
C. G. Shatzer.

Report of the Executive Committee

The report of the Executive Committee was received as
follows and ordered filed.

Cambridge, Mass., April 2, 1917.
To the Ohio Academy of Science:

The Executive Committee has held but one meeting during the
year, on January 27. The officers of the Academy were invited to meet
with the Committee. Profs. Hine, Walton and Rice, of the Com-
mittee were present, together with Profs. Foerste, Osbom, Schaffner and
Westgate by invitation.

It was decided to hold the annual meeting on April 6 and 7, in
Columbus.

The Program Committee was instructed to arrange an independent
session of the Academy for Saturday morning, and to arrange for a joint
meeting with the Ohio College Association for Friday evening only
in case of agreement upon a distinctively scientific address. (When
Prof. Withrow accepted the invitation to speak at the Friday evening
meeting, an invitation was extended to the Ohio College Association to
join with the Academy; but other arrangements had already been made.)

Dr. Mendenhall was appointed to fill the vacancy in the Board of
Trustees of the Research Fund caused by the death of Prof. Lazenby;
and Prof. Fenneman was requested to serve as chairman of the Board.

18 The Ohio Journal of Science [Vol. XVIII, No. 1,

Prof. Wcstgate was appointed as Acting Secretary to serve during
the absence of the Secretary.

The following communication from the Ohio State University
Scientific Society was presented by the Secretary:

Columbus, May 29, 1916.
Professor E. L. Rice, Secretary Ohio Academy of Science,
Ohio Wesleyan University, Delaware, Ohio:
Dear Professor Rice: — At the last meeting of the Ohio State University
Scientific Society it was unanimously voted that the Ohio Academy of Science
be requested to name "a representative of each section of science represented in
the Academy (Zoology, Botany, Geology, Physics), to serve on the Editorial
Board of the Ohio Journal of Science," such representatives (four in all) not to be
members of the Ohio State University Scientific Society.

Will you be so kind as to notify such officers of the Academy as should receive
this announcement.

Sincerely yours,

R. J. Seymour, Secretary.

Owing to the near approach of the annual meeting, no formal
action was taken by the Committee, but the appointment of rep-
resentatives was left to the Academy as a whole.

In November, 1916, the Executive Committee decided by corre-
spondence to join with other organizations in extending an invitation to
the Central Association of Science and Mathematics Teachers to meet
in Columbus next Thanksgiving. The following letter from the Pres-
ident of that Association should be presented to the Academy :

Columbus, March 21, 1917.
E. L. Rice, Secretary, The Ohio Academy of Science, Delaware, Ohio:

My Dear Mr. Rice:— You will recall that your organization joined with a
number of other organizations of the state in extending to the Central Association
of Science and Mathematics Teachers an invitation to hold the next meeting in
Columbus. The Central Association will meet in Columbus, November 30 and
December 1, 1917. I take this means of suggesting to you that the Association
will deeply appreciate any co-operation you may find it possible to give.

There' will be a general meeting on Friday morning to which all are invited
and at which a Science address will be made in addition to a general educational
address. On Friday afternoon the sectional meetings will be held. On Friday
evening there will be a banquet followed by detailed reports of plans for science
and mathematics work in "The High School of To-morrow." On Saturday
morning the general business meeting is held, followed by excursions to various
places of interest in or near Columbus.

The Association invites you to be present and to take part in any and all of
these meetings. The inspiration of this large body of teachers should be of great
benefit to every similar organization in the state. Let us make this the biggest
Science and Mathematics meeting ever held in Ohio.

Plan now to be present at the November meeting.

Yours very truly,

Marie Gugle, President.

This invitation has been acknowledged l^y the Secretary.

A single application for membership has been approved by the
Executive Committee, and will be presented later for the ratification of
the Academy.

Respectfully submitted,

Edward L. Rice,

Sec y for Committee.

Nov., 1917] Report Ohio Academy of Science 19

Report of the Publication Committee

The report of the Publication Committee was received as
follows and ordered filed.

The committee has published the Report of the Twenty-fifth
Annual Meetinji:, consisting of 202 pages and also the Report of the
Twenty-sixth Annual Meeting, consisting of 26 pages. The first
report was pubHshed at a cost of $222.75, and the twenty-sixth, which
was a i^artial reprint froin the Ohio Journal of Science, at a cost of
$42.00.

The method of publication of the Proceedings of the Annual Meetings
was not fully detennined at the last meeting. The Executive Com-
mittee having been given power to act on its own initiative. After
correspondence with mcml^ers of the Executive Committee it was
thought best to publish the Annual Report as heretofore, the first
twelve pages consisting of list of officers and members, being set up
separately, and the Secretary's Report reprinted from the Journal of
Science. It was not thought necessary to reprint the President's
address, since this -will now be published regularly in the" Ohio Journal
of Science.

The committee recommends that the Proceedings of the Acadeni}-
be handled in the regular way as heretofore, and that the Annual Report
mainly a reprint from the Ohio Journal of Science, consisting of lists
of officers and members, and the Secretary's Report be continued. The
committee strongly urges that the publication of special papers be
continued also and that if possible at least one be published each year.

Respectfully submitted,

John H. Schaffner, Chairman,
L. B. Walton.

Report of the Emerson McMillin Research Fund

The following report of the Trustees of the Research Fund
was received and referred to the Auditing Committee, whose
report is appended.

April 5, 1917.

RECEIPTS

Balance April 21, 1916 $308.41

Check from Emerson McMillin 250.00

$558.41

EXPENDITURES

Forest B. N. Brown $ 46.25

B. H. Wells 40.00

W. L. Du Bois 21.50

Stephen R. Williams 41.69

Wm. M. Barrows 19.60

Balance in Bank 389.37

$558.41

N. M. Fenneman.

20 The Ohio Journal of Science [Vol. XVIII, No. 1,

April 7, 1917.

No report on the McMillin fund by the Auditing Committee is pos-
sible, inasmuch as vouchers, paid checks, and bank book are not
submitted.

L. B. Walton,
C. G. Shatzer.

Report of the Library Committee

For the Library Committee, Mr. Reeder, of the Ohio State
University Library, presented the following report, which was
received and ordered filed.

To the Ohio Academy of Science:

The Ohio State University Library begs to report that it has cared
for all the accessions to the Library of the Academy, which have been
received during the year 1916-17. The number of publications is not
large, aggregating about one hundred (100) items.

During the year the annual reports of the twenty-fifth and twenty-
sixth meetings of the Academy, for 1915 and 1916, were pubHshed.
Copies were mailed to all members whose names appeared in each
report and to all exchanges.

The sale of publications for the year has amounted to $22.60. This
sum has been turned over to the Treasurer.

The librarian desires to raise three questions on which he would like
a vote of the Academy:

1. Is there any policy regarding the number of copies of a Special
Paper which may be presented to the author? Request was m.ade
during the year, by an author, for copies of his publications. Under
existing arrangements the only disposition is by sale, after distribution
has been made to the membership and to the exchanges.

2. Is there any policy regarding the reprinting of Special Papers,
which are nearing exhaustion? Special Paper No. 15, "Trees of Ohio
and Surrounding Territory," by J. H. Schaffner, is almost gone, there
being only seven copies in stock. A request for eighteen copies for use
in a university had to be turned down.

3. Is the Academy ready to take a further step in the management
of its library and turn its collection over to the Ohio State University
for incorporation with the other sets and books on scientific subjects?

In the original agreement between the Academy and the University,
adopted November, 1914, it was stated in article 3, that "for the time
being" the library of the Academy should be kept as a separate col-
lection. This has been done. In article 4, it was stated that "ulti-
mately it is mutually agreed that the library of the Academy shall be
classified with, and distributed in the University Library. "

The question is: Is the Academy ready to take this step at this
time? The Universitv Library is readv.

Nov., 1917] Report Ohio Academy of Science 21

As bearing on this proposition, the following statements are
submitted :

(a) The Ohio Journal of Scicnee is receiving a subsidy from the
University. In return for this annual grant, the University Library
receives 250 copies of each issue for exchange purposes. At the present
time, some 230 societies and periodicals are on the exchange list.
Furthennore, the Biological Club has turned over to the University
Library all the exchanges received by the Ohio Naturalist from 1900 to
1915. This material has been bound, catalogued and classified by the
Library.

{b) Some confusion exists between the two sets of exchanges, those
received by the Academy and those received by the Library through
the Ohio Journal of Science. If the proposed step is taken, the two sets
of exchanges can be unified and duplication eliminated, both in sending
and in receiving.

(c) If the proposed step is taken, the duplicates found in the
Academy's collection can be used to secure other sets not now in the
possession of either the University or the Academy, or they can be sent
to other Ohio libraries as the Academy may decide.

{d) The University Library has been purchasing from its book
budget, many new sets and filling in the gap in existing sets. If the
Academy's collection is turned over, a number of its sets can be com-
pleted through purchase, something the Academy cannot afford to do.
Furthermore, there will be no chance for duplication or waste in money,
when the complete sets are purchased by the University of which the
Academy has a partial or incomplete file.

(e) The University Library is unifying other exchanges received and
distributed by the University, such as the Ohio History Teacher's
Bulletin, the Contributions from the Department of Zoology and
Entomology, the Doctoral Dissertations, and the proposed series of
studies to be issued by the Graduate School. If the Academy agrees
to the proposition put forward, all readjustments can be made while the
work is being done on these other series, and the entire question of
exchanges can be settled.

(/) The result of favorable action on the part of the Academy will
be advantageous to both parties, because the members of the Academy
will get improved library facilities through a better handling of its own
sets and exchanges and the University Library will be in a better
position to unify its collections, to plan its purchase of sets and of
''fill ins" and to develop systematically, its entire system of exchanges.

Respectfully submitted,
April 6, 1917. C. W. Reeder.

Mr. Reeder was requested to make specific recommendations
on the subjects treated in this report. This was done at the
second business session. In accordance with the recom-
mendations of Mr. Reeder, the following actions were voted
by the Academy:

22 The Ohio Journal of Science [Vol. XVIII, No. 1,

1. That fifty copies of eacli special publication of which
copies are now on hand or which may be published in the
future shall be given free to the author. (It is understood that
any copies already given shall count on this number.)

2. That the library of the Ohio Academy of Science be
turned over to the Ohio State University for incorporation in
The Ohio State University General Library.

3. That a special card catalog be kept of the books thus
received by the Ohio State University Library from the Academy.

4. That a suitable printed plate, to be paid for by the
Academy, be put in the books given to the Ohio State Uni-
versity Library.

Report of the Committee 07i Legislation

An oral report was presented by the chairman, Dr. Menden-
hall. The report was received and the Committee continued.

Report of the Committee on the Ohio Journal of Science
No report was presented. The question at issue has prac-
tically solved itself, and the general satisfaction with the
present relations of the Academy and the Ohio Journal of
Science renders further work by the committee unnecessary.

Report of Committee o?i Catalog of Scientific Journals
The following report was presented by Mr. Reeder, of the
Ohio State University Library, for the Committee on Catalog
of Scientific Journals. The report was received and ordered
filed.

The Committee on the Union List of Scientific Periodicals begs
leave to submit the following report :

Twenty-one Ohio libraries are now pledged to assist in the union
catalog. Three additional libraries have agreed to join the movement
during the past year — Cincinnati University, Kenyon College, and
Otterbein College. Of these co-operating libraries, complete or partial
deposits have been made by nine, namely:

Case School of Applied Science.

Kenyon College.

Lloyd Library.

Municipal Universit}^ of Akron.

Obcrlin College.

Ohio Archaeological and Historical Society.

Ohio State University Librar}\

Otterbein College.

Wittenberg College.

Nov., 1917] Report Ohio Academy of Science 23

All cards so far deposited, have been arranged in one alphabet,
after each card has been marked to show the owner library. In order
to ascertain the work done during the year by the co-operating liVjraries
which had not made deposits, a request was sent to each one asking for
a statement of ]jrogress. Following are excerpts from the replies
received :

Adelbert College of Western Reserve — Beginning in October, an assist-
ant has gi\-en about half-time to the preparation of the records. The
periodicals of the Reference Department and the 000 numbers of the
Circulating Department are practically completed, making about 350
titles. Work is now proceeding on the lOO's.

Antioch College — The librarian expects to get the records completed
wathin a reasonable time.

Cincinnati University — This library contains the exchange deposit
of the American Association for the Advancement of Science. Not
much, if anvthing, can be done on the records until the work eases up a
little.

Cleveland Public Library — Cards for about 550 titles are ready.
These are for such titles as happen to be catalogued or recatalogued, so
nothing has been done along any special line.

Hebrew Union College — No progress has been made in the preparation
of the records.

Miami University — The staff work has been interrupted by sickness
and leaves of absence, so no progress whatever has been made.

Ohio State Library — The larger part of the records of periodicals has
been corrected and brought down to date, so the deposit is nearly ready
for submission.

Ohio University — Members of the staff have been away, so nothing
has been done on the records.

Ohio Wesleyan University — Owing to lack of sufficient staff" to give
it attention, not much progress has been made. However, the list of
cards has been prepared for the bound volumes down to J. It is barely
possible that a special worker can be put on these records and complete
them in the near future.

Toledo Public Library — Press of work has prevented progress on the
records.

Young Men's Mercantile Library Association — Few of our magazine
sets have ever been more than arranged on the shelves. Fift^'-one
sets have been shelf-listed and catalogued and cards for these will be
forwarded within the near future. Thirty-four sets have been shelf -
listed only and ^\dll be catalogued as time pemiits.

No word was received from the Cincinnati Public Library and
Hiram College.

The Academy is aware that the College Section of the Ohio Library
Association is co-operating in this work. A full and detailed report was
made at its meeting on October, 1916. It was voted at that time to
continue the compilation on the lines already laid down and rush the
union list to completion as quickly as the time and resources of each
individual library permitted.

24 The Ohio Journal of Science [Vol. XVIII, No. 1,

However, a phase has developed to the Ohio situation which needs
thorough consideration. The university libraries of the middle west
have projected a Union List which is to include Nebraska, Minnesota,
Wisconsin, Iowa, Illinois, Michigan, Ohio and the metropolitan districts
of St. Louis and Chicago. The plans for this list have been laid by the
H. W. Wilson Co., the estimates of printing made and the copy from
nearly all the libraries is ready. Consideration is now being given to
the time limit, so the publication may proceed.

Here are some problems for Ohio:

(1) Shall we ask for the inclusion of the Ohio Union list in the
proposed "North Central List?"

(2) Is the Academy ready to finance the inclusion of its list, on
the basis of what other co-operators will pay?

(3) If the Academy does not finance the proposition, shall we ask
each library to finance its own contribution to such a list, or buy its
quota of volumes?

(4) When the "North Central List" is pubhshed, will there be
need for an exclusive Ohio List?

The committee recommends that, in spite of the complication of the
situation, work be continued on the distinctively Ohio List and that
the relation of the Ohio proposition to that of the universities of the
Middle West be held in abeyance until many of the details of the larger
union list are definitelv settled.

C. W. Reeder.
April 6, 1917.

The following actions were voted by the Academy:

1. That the recommendations contained in the report of
the Committee on Catalog of Scientific Journals be adopted.

2. That the Committee on Catalog of Scientific Journals
be discharged, and that the work formerly done by this com-
mittee be transferred to the Library Committee.

Election of Officers

From the double slate of nominations prepared by the
Nominating Committee, the following officers for 1917-18 were
elected by the ballot of the Academy:

President — Professor F. L. Landacre, Ohio State University,
Columbus.

Vice-President for Zoology — Professor C. G. Shatzer, Wittenberg
College, Springfield.

Vice-President for Botany — Professor Freda Detmers, Ohio State
University, Columbus.

Vice-President for Geology — Professor G. F. Lamb, Mt. Union College,
Alliance.

Nov., 1917] Report Ohio Academy of Science 25

Vice-Presidenl for Physics — Professor Samuel R. Williams, Oberlin
College, Oberlin.

Vice-President for Aiialoniy, Physiology, etc. — Professor R. J.
Seymour, Ohio State U^i^'ersity, Columbus.

Secretary — Professor E. L. Rice, Ohio Weslcyan University,
Delaware.

Treasurer — Professor J. S. Hine, Ohio State University, Columbus.

Elective Members of Executive Committee — Professor C. G. Shatzer,
Wittenberg College, Springfield; Professor Frank Carney, Denison
University, Gran\-illc.

Member of Publication Committee — Professor L. B. Walton, Kenyon
College, Gambier.

Trustee of Research Fund — Professor Herbert Osborn, Ohio State
University, Columbus. (The resignation of Professor M. M. Metcalf
was i)resented, and the Chair was instructed to appoint a trustee to fill
the vacancy until the next Annual Meeting. Chair appointed Professor
M. E. Stickney, Denison University, Granville.)

Member of Library Committee — ^Professor F. 0. Grover, Oberlin
College, Oberlin.

On nomination from the floor, the following were elected
to represent the Academy on the editorial board of the Ohio
Journal of Science:

Zoology — Professor R. A. Budington, Oberlin College, Oberlin.

Botany — Professor M. E. Stickney, Denison University, Granville.

Geology — Professor G. D. Hubbard, Oberlin College, Oberlin.

Physics — Professor S. J. M. Allen, University of Cincinnati,
Cincinnati.

Anatomy, Physiology, etc. — Professor F. C. Waite, Western Reserve
University, Cleveland.

Election of Members

The Membership Committee reported ten names for election

to membership; one additional name, previously approved by

the Executive Committee and marked with * in the following

list, w^as presented for ratification. All were elected, as follows:

Brooks, Clyde, Physiology, Physiological Chemistry, Pharmacology, O. S. U.,
Columbus.

Cameron, Hazel C, Pathology, 1234 Highland St., Columbus.
*Eggleston, H. Ray, Botany, Zoology, Paleontology; Marietta College,
Marietta.

Faulkner, Herbert C, Entomology, Biology; 2 Val. View Boulevard,
Ashtabula.

Huber, H. E., Biology; Ohio Northern Universit}', Ada.

Htigger, Carl Claron, Pathology; 710 N. Park St., Columbus.

Kisliuk, Max, Jr., Zoology, Entomology, Botany; Botany and Zoology Bldg.,
O. S. U., Columbus.

McCaughey, W. J., Mineralogy; Lord Hall, O. S. U.. Columbus.

Orcutt, Ruth; Botany, Zoology; Granville.

Speaks, John C, Ichthyology, Ornithology; Columbus.

Yampolsky, Cecil, Botany, Zoology, Genetics; Toledo Universit3^ Toledo.

26 The Ohio Journal of Science [Vol. XVIII, No. 1,

The Report of the Committee on Necrology

The following report was presented by the Committee on

Necrology :

Not often does it fall to the Academy to record the loss of two leading
members and past officers within one year. Within three days, Sept. 11
and Sept. 14, there occurred the deaths of Professor Charles Smith
Prosser and Professor William Rane Lazenby. In many points their
lives had run in similar ways; both bom as farmer's sons in central New
York, both graduates of Cornell University, both of necessity in part
working their way through, both retained for a number of years by the
University as assistants in their respective fields, both subsequently
called to Ohio State University as heads of their respective depart-
ments, both loyal members of the Sigmi Xi, and both active through
their entire scientific careers in painstaking research and conscientious
teaching.

Professor Lazenby was born in 1850 at Bellona, N. Y. He was
graduated froin Cornell University in 1874, at which time he won the
Ezra Cornell prize in Agriculture. He was Instructor in Horticulture
and Botany, '74-77 and Assistant Professor, '77-81 at Cornell; Professor
of Botany and Horticulture, '81-'92, of Horticulture and Forestry,
'92- '08, and of Forestry at Ohio State University from that date to his
death. He was founder and for five years director of the Ohio Experi-
ment Station, an institution which has been a very important factor
in the agricultural development of his adopted state.

He was an active member of many scientific organizations, among
which are:

American Association for the Advancement of Science (twice
Secretary and once Vice-President of Section I) ;

Society Horticultural Science;

American Pomological Society;

American Forestry Society (Vice-President many years) ;

Society for Promotion of Agricultural Science (Secretary five
years and President two years) ;

Ohio Academy of Science (founder and President, '02) ;

Ohio Forestry Society (President, '04 to date of his death).
His activity in these organizations grew out of a keen interest in
them, their members and their ideals, an interest which expressed
itself in wise counsel, frequent contributions to their programs, and
discussions of the papers of others.

Teaching was his main business during the college year, but his
summers were always spent in travel and investigation; several were
spent in Europe, studying her forests and forestry methods.

His publications include many contributions both to Scientific
Journals and State reports and to the semi-popular and popular press.

In the words of a close friend at Cornell: "While he found his great
interest in life, the mastery and development of his special field in
science, it was the human side of him that had the strongest hold on his
friends and colleagues. He never lost his interest in the struggles

Nov., I*) 17] Report Ohio Academy of Science 27

of students with limited means, and in a quiet way extended aid to many
of them. He never lost an opjjortunity for service to his friends or
others in need; sympathy, helpfulness and loyalty were his characteristic
qualities as a man and friend; and the loss to all of us who knew and
loved him is irreparable." — J. H. Comstock, Science, December 29,
1016, p. 913.

Hundreds of students now scattered to all parts of the country
have sat at his feet, and can testify to the quiet, unassuminj^ scholarship
and sympathetic instruction to which they were there exposed. Many
have attained to eminence in horticulture and forestry through the
working out of the interest and enthu.siasm he engendered.

As a member of the Academy he was practically always at its
meetings with an important contribution to make. His presentation
was always interesting and plain. Continually alert for op]:)ortunities
by which the Academy could serve the Commonwealth and Science, he
became the channel through which we received our annual, financial aid
for research from our loyal patron and life member, Mr. Emerson
McMillin. He served on the trustee committee from 1901 to the time
of his death and wisely discharged his duties.

If any difficult problem arose in the affairs of the Academy he was not
lacking in efficient counsel nor was it hastily or injudiciously given.
Thus we shall miss his thoughtfulness and wisdom.

Professor Prosser was born in ISOO at Columbus, New York. He was
graduated from Cornell Uni\'ersity in 1S83, was a fellow one year and
instructor to 18SS; received his master's degree in 1886 and returned
twenty years later and took his doctor's degree in 1907. One year
before Union College gave him an honorary Doctorate of Science.

He was professor of Natural History at Washburn, Kansas, '92-'94, of
Geology at Union, '94-'99, Associate Professor of Historical Geology,
'99- '01 at Ohio State University and Professor and Head of the Depart-
ment from then to the date of his death. For four years after leaving
Cornell University he was Assistant Paleontologist on the U. S.
Geologic Survey, Assistant Geologist '92-'9o and '00-' 16. He was a
member of the N. Y. Geological Survey, '95-'99; Kansas Geological
Survey, '96, Chief Appalachain Division, Maryland Geological Survey,
'98-'01; Assistant Geologist, Ohio Geological Survey, '00-'16.

He was a member of the leading Geological and Scientific Societies
of America and of some foreign organizations, among which are the
f ollomng :

International Congress of Geologists, London, 1888;

International Geographic Congress, New York, 1904;

Fellow American Association Advancement Science (Vice-
President, Section E, at the time of his death);

Society of Naturalists;

Society of Vertebrate Paleontologists;

Geological Society of America;

Paleontological Society of America;

'National Geographic Society;

Geologische Verein;

Washington, D. C, Kansas and Ohio Academies of Science.

28 The Ohio Journal of Science [Vol. XVIII, No. 1,

While teaching was the chief business of Dr. Prosser during prac-
tically his whole public life, his vacations and all spare time during the
school year were devoted to research. He loved it and was willing to
sacrifice much to attain his goals in his scientific career.

His contributions were always marked by most painstaking and
precise accuracy in the fields they covered and thus left little for anyone
else to do when he had finished.

Among his publications are extensive papers and reports on the
geology and paleontology of New York, Maryland, Kansas, and Ohio.
They are largely stratigraphic and paleontologic, and are particularly
strong in the correlations which they set forth.

The State Geologist of New York, with whom Professor Prosser
had long been associated, gives a beautiful estimate of him in the
following words:

"There never was a more loyal, a more devoted, a more sensitive
spirit. His attitude of mind was puritanic in its simplicity and in its
practices, and, left to himself he could never suspect another of indirect-
ness or duplicity — a quality of which he contained not a grain. When con-
fronted by the broader bearings of his science and the natural sequences
of its greater propositions, he held himself somewhat carefully aloof.
Yet this simplicity of heart which would not let him go far afield,
also made him extraordinarily conscientious in his scientific work.
It would not be fair to him to say that he had a genius for details, but
it would be eminently right to assert that he sought intimately and
faithfully for the exact construction of every observation he made so
far as that had to do with the theme in hand. This mental method led
him to a precision of manner and gave him a certain formality which
was seldom dismissed under the most informal circumstances." —
J. M. Clarke, Science, October 20, 1916, p. 559.

Many men and women have gone out from under his training, in
Kansas, New York, and Ohio, who can trace their geological ideals and
beginnings to him. Some have attained to eminence in geologic and
paleontologic research or in teaching and uphold the high standards to
which he was devoted.

(Signed) Geo. D. Hubbard, Chairman,
Herbert Osborn.

Report of the Committee on Resolutions

The following report was presented by the Committee on
Resolutions, and adopted by the Academy.

J. The Academy wishes again to express its gratitude for the
continued generosity of Mr. Emerson McMillin, and to put on record
its appreciation of the value of his gifts to the Research Fund in the
stimulation and extension of research on the part of the membership of
the Academy.

2. The Academy desires to express sympathy for Professor F. 0.
Grover, whose illness, brought on by overwork, has prevented him from

Nov., H)l<j Report Ohio Academy of Science 29

attending and presiding as president at this meeting. It is to be hoped
that he may offer for pubhcation the address he had expected to give.

3. Professor Robert F. Griggs is entitled to the thanks of the
Academy for his very interesting illustrated lecture on his recent
''Explorations of Mount Katmai, Alaska."

4. The thanks of the Academy are given to Professor James R.
Wi throw for his timely and valuable lecture on "The Relation of War
to Chemistry in America. "

5. The Academy thanks the Local Committee and the Professors
and Ofhcers of the Ohio State Univcrsit}- for providing convenient
rooms for the general and sectional meetings and for their efforts in
ministering to the comfort of all members in attendance.

6. The Secretary of the Academy, Professor E. L. Rice, deserves
much credit for making detailed arrangements for this successful meet
ing, although other duties have necessitated his absence from the State.

E. L. MOSELEY,

L. B. Walton,
C. G. Shatzer,

Commitlee.

Section for Physiology, Anatomy, Bacteriology and Pathology

A petition, signed by sixteen members, was presented,
requesting the Academy to establish a new section for Phy-
siology, Anatomy, Bacteriology, and Pathology — the exact
name of the section to be determined later.

The petition was granted; and a vice-president and rep-
resentative upon the Editorial Board of the Ohio Journal of
Science were elected for the new section.

Constitution and By-Laws

The Acting President was instructed to appoint a committee
of three to codify the Constitution and By-Laws. This committee
was requested to consider the desirability of lengthening the
terms of office of the president and secretary. (This action
to be interpreted as notice of amendment, so that definite
action may be taken at the next meeting.)

The Acting President appointed the following committee:
Secretary E. L. Rice, ex-of!icio; F. C. Blake, Frank Carney.

The Academy directed the Executive Committee to print a
new edition of the Constitution and By-Laws.

30 The Ohio Journal of Science [Vol. XVIII, No. 1,

Time Limit for Papers

It was voted by the Academy that, in case titles of papers
are sent in without specification of the time needed, a time limit
of five minutes shall be set by the Program Committee.

Adjournment

The meeting adjourned without determining the place of the
next meeting.

Scientific Sessions
The complete scientific program of the meeting follows:

Presidential A ddress

Some Aspects of Present Day Biological Unrest, F. O. Grover
(Omitted because of the absence of the President)

Lectures

The Relation of War to Chemistry in America,

James R. Withrow

Recent Explorations of Mount Katmai, Alaska,

Robert F. Griggs

Papers

1. Eskers in the Dayton Area. 20 min. (Lantern) Aug. F. Foerste

2. The Preservation and Utilization of Wild Life in Ohio. 20 min.

Herbert Osborn

3. The Anatomy of an Eight-Legged Pig. 15 min. (Lantern),

Stephen R. Willi.\ms

4. The Origin of the "Reserve Cells" (Neoblasts) in the Regeneration of

Tubifex. 15 min. (Lantern) F. H. Krecker

5. Prospective and Retrospective Methodology in relation to the Problem

of Evolution. 15 min. (Lantern) L. B. Walton

6. Mutations by Interference :— A Study in Heredity. 12 min. . . A. B. Plowman

7. Drainage Developments in Southeastern Ohio. 20 min. . .Geo. D. Hubbard

S. Furs in the Wild and in the Shop. 20 min Katharine Dooris Sharp

9. A Suggested Method for the Distinctive Description of Dipterous

Larvae. 7 min C. L. Metc.\lf

10. Additional Records and New Species of Ohio Homoptera. 10 min.

Herbert Osborn

11. Some New or Interesting Insect Predators. 10 min C. L. Metc.\lf

Nov., 191 7J Report Ohio Academy of Science 31

12. Notes on the Protista: — (a) A New Species of Eulreptia; (h) A New

Classification of the Protista. 10 min L. B. Walton

13. The Remains of tlie Porcupine foimd in an Ohio Prehistoric Village

Site. 8 min Wm. C. Mills

14. Indians of Licking County and Earthworks. 10 min H. A. Alby.n

15. Suggestions for the Making of a Key to the Seeds and Seed-like Fruits

of Ohio Plants. 10 min John H. Schaffner

16. Ray Tracheids of Conifers. 7 min FkiiDA Det.viers

17. The Nature of Sexual Hereditary Expression in tiie Sporophyte. 10 min.

John H. Schaffner

18. The Correlation of the Niagara Formations of Ohio, Indiana and Ken-

tucky. 10 min Aug. F. Foerste

19. Pottsville and Allegheny Limestones in Muskingum County, Ohio.

15 min Wilber Stoit

20. Further Data on the Maxville Limestone. 5 min G. F. Lamb

21. Bearing of the Glacial Period on Theories of Evolution. 10 min.

G. Frederick Wright

22. Some Glacial Features in Central Ohio. 5 min G. F. Lamb

23. Some Segments of the Maumee Shoreline near Columbus Grove, Ohio.

10 min. (Lantern) Frank Carney

24. A Physiographic Map of North America. 15 min W. N. Thayer

25. The Nature of Beatricea undulata. 5 min W. H. Shideler

26. Symposium on Modern Physics: —

A. Hall Eft'ects in Metals and Alloys,

Professor A. W. Smith, Ohio State University

B. Magnetostriction and its Bearing on Magnetic Theories,

Professor Samuel R. Williams, Oberlin College

C. A Review of Recent Work in X-Rays,

Professor S. J. M. Allen, University of Cincinnati

27. Spermatogenesis of Branchipus vernalis. 7 min Rollo C. B.\ker

28. Origin of Pre-Auditory Ganglia in Rana. 7 min Ralph A. Knouff

29. The Origin of Cartilage from Ectoderm in the Urodeles. 10 min.

F. L. Landacre

30. Amphibian Blood Transfusion. 10 min R. A. Budington

31. Observation on the Cytology of the Exudation seen in Lacunar Tonsilitis.

10 min Hazel C. Cameron

32. A Contribution to the Origin of the Macrophages seen in the Exudation in

Pneumonia accompanying Typhoid Fever. 10 min. . .Carl Claron Hugger

33. The Occurrence of Eosinophilic Leucocytes and the Granule Cells of

Paneth in the Vermiform Appendix of Man. 5 min James H. Warren

34. A Study of the Incidence of Pathological Conditions of the Thyroid

Gland among Dogs in the City of Columbus. 10 min. (Lantern),

C. I. Reed

35. Nephroblastic Tumors in Rabbits. 10 min. (Lantern) Ernest Scott

36. Notes on the Formation of Giant Cells in Tumors. 5 min.

Jon.\than Forman

37. Mudballs and Mudflows in the Ordovician Rocks in the Vicinity of

Cincinnati. 10 min. (Lantern) Walter H. Bucher

32 ■ The Ohio Journal of Science [Vol. XVIII, No. 1,

38. Intraformalional Pebbles of the Ordovician and Silurian Formations of

Ohio. 10 min Aug. F. Foerste

39. Conditions of Deposition in Devonian and Penn.sylvania Times. 10 min.

Lewis G. Westg,\te

40. The Weathering of Wisconsin Gravels in the Little Miami Valley near

Cincinnati, Ohio. 10 min Walter H. Bucher

41. Some More of the Saluda Question. 10 min W. H. Shideler

42. Lake Abram. 3 min. (Opaque Projection) Wesley N. Speckman

43. Concretionary Forms in Greenfield Limestone. 10 min C. W. Napper

44. Natural Gas in Ohio. 10 min John Bownocker

45. The Radioactivity of Smoke. 10 min S. J. M. Allen

46. Physical Factors concerned in the Measurement of Blood Pressure.

15 min. (Lantern) Clyde Brooks

47. The Inheritance of Spotting in Dutch Rabbits. 20 min. (Lantern).

H. D. Fish

48. Unconformity between the Mississippian and Pennsylvanian in the

Grand River at Parkman, Ohio. 10 min Geo. H. Colton

Demonstrations

1. A Freak Leptinotarsa decemlineata C. L. Metcalf

2. Mudballs and Pebbles from the Ordovician Rocks of the Vicinity of

Cincinnati, illustrating Paper 37 Walter H. Bucher

3. Pictures and Specimens to illustrate Paper 40, on the Weathering of

Wisconsin Gravels Walter H. Bucher

Lewis G. Westgate,

Acting Secy during Meeting,

Edward L. Rice,

Secretary.

Date of Publication, November 30, 1917.

THE

Ohio Journal of Science

PUBLISHED BY THE

Ohio State University Scientific Society

Volume XVIII DECEMBER, 1917 ' No. 2

TABLE OF CONTENTS

Leonard — The Astereae of Ohio 33

Adkins — Some Ohio Heteroptera Records 58

Claassex — Second Alphabetical List of the Lichens Collected in Several
Counties of Northern Ohio 62

THE ASTEREiE OF OHIO.

Emery C. Leonard.

Mostly erect geophilous perennial or occasionally annual
herbs with stems branching at the top or in a few cases scapose.
Leaves alternate, sessile, petioled, or clasping. Flowers bispor-
angiate, carpellate, or staminate, on a common receptacle,
forming heads subtended by an involucre of few to many
bracts imbricated in one or more series. Pappus of capillary
bristles, teeth, or scales, or sometimes wanting. Ray flowers
present in most cases. Achenes glabrous or pubescent.

Synopsis of the Astereae.

I. With 3-ellow ray flowers or with rays absent (white in Solidago bicolor).

1. Pappus of scales, not capillary bristles Grindelia.

2. Pappus of capillary bristles with or without an outer series of shorter ones.

a. Pappus double, an inner series of capillary bristles and an outer series
of short ones; heads large. Chrysopsis.

h. Pappus a single series of capillar}^ bristles; heads small.

(1) Receptacle alveolate; rays mostly fewer than the disk flowers.

Solidago.

(2) Receptacle fimbrillate; rays more numerous than the disk flowers;

heads in a corymbose paniculate inflorescence. Euthamia.

II. Ray flowers present, not yellow but some shade of pink, blue, purple, white

or greenish.
1. Pappus a crown of aw'ns or none, never of typical capillary bristles.

a. Pappus none or a mere crown of minute bristles; receptacle conic. Bellis.

b. Pappus a crown of few awns; receptacle flat or convex. Boltonia.

33

34 • The Ohio Journal of Science [Vol. XVIII, No. 2,

2. Pappus of numerous capillary bristles.

a. Pappus a single series of capillary bristles.

(1) Bracts of the involucre two to many series.

(a) Involucre narrow, its bracts firm; rays few, white.

Sericocarpus.

(b) Involucre turbinate or hemi-spherical, its bracts mostly thin;

rays usually more numerous, white to purple. Aster.

(2) Bracts of the involucre mostly in one or two series; heads mostly

long pedicled.

(a) Rays longer than the disk flowers. Erigeron.

(b) Rays not longer than the disk flowers; heads very small,

panicled. Leptilon.

b. Pappus distinctly double, the outer series shorter than the inner.

(1) Leaves lanceolate-ovate to obovate; rays white. Doellingeria.

(2) Leaves narrowly linear; rays violet. louactis.

Key to the Genera.

1. Ray flowers yellow. 2.

1. Ray flowers not yellow. 5.

2. Leaves punctate, linear or linear-lanceolate; heads small, corymbose, some-

what glutinous. Eiithamia.

2. Leaves not punctate in our species. 3.

3. Pappus in 2 series, the inner of long, the outer of short somewhat chaffy

bristles; stem leaves entire. Chrysopsis.

3. Pappus of nearly equal bristles; leaves mostly serrate or dentate. 4.

4. Pappus of 2-8 coarse deciduous bristles; heads large, sometimes glutinous.

Grindeha.

4. Pappus of numerous capillary bristles; heads comparatively small, not

glutinous. Solidago.

5. With single flowers on leafless scapes; pappus of the disk flowers none. Bellis.

5. Flowers on leafy stems, pappus present on the disk flowers. 6.

6. Pappus of the disk flowers 2-5 scale-like bristles. BoUonia.

6. Pappus of the disk flowers of numerous capillary bristles. 7.

7. Ray flowers 4-5. 8.

7. Ray flowers numerous or rarely none. 9.

8. Flowers in terminal cymose panicles. Sericocarpus

8. Flowers in crowded clusters forming a spike or thyrsus. Solidago (bicolor).

9. Pappus in a single series. 10.

9. Pappus double, the outer series of short bristles. IL

10. Bracts of the involucre imbricate in several series. Aster.

10. Bracts of the involucre in but 1 or 2 series. Erigeron.

11. Ray flowers inconspicuous, white; annuals. Leptilon.

11. Ray flowers conspicuous. 12.

12. Rav flowers 25-200, narrow, bracts of the involucre very narrow. Erigeron.

12. Ray flowers 8-15. 13.

13. Rays white; leaves lanceolate to oblong, pinnately veined. Doellingeria.
13. Ra^'s violet; leaves narrow, one nerved; peduncles covered with bracts.

lonactis.

Grindelia Willd. Gum-plant.

Coarse perennial or biennial herbs with smooth stems,
branched above and sometimes woody below. Leaves alter-
nate, sessile or clasping, mostly spinulous dentate. Heads
either large and solitary at the ends of the branches; involucre
hemispherical, its bracts with green points and imbricated in
several series; ra}^ and disk flowers yellow. Pappus of 2-8
deciduous bristles. Achems short and glabrous.

Dec, 1917] The AsterecB of Ohio 'So

1. Grindelia squarrosa (Pursh) Dun. Broadleaf Gum-
plant. An erect jilant, (> in. -2 ft. high. Leaves oblong to
oblong-spatulate, obtuse, more or less clasping at the base,
sharply spinulose dentate, 3^-13^ in. long and \i-Y2 iri- wide.
Heads ^-13<£ in. broad, glutinous; involucral bracts linear-
lanceolate; rays sometimes absent. In dry soil. June to
September. Hamilton County. From the west.

Chrysopsis (Nutt.) Ell. Golden-aster.

Branching perennial or sometimes biennial wooly or hairy
fall flowering herbs with alternate, sessile, entire or slightly
serrate leaves. Heads rather large and generally solitary at
the ends of the branches; involucre hemispherical, of linear
bracts imbricated in several rows, the outermost being the
shortest; both ray and disk flowers yellow; ray flowers carpel-
late; disk flowers bisporangiate or only staminate. Pappus
double, the outer series of short chaffy scales, the inner series
of long capillary bristles. Achenes flattened, oblong-linear or
obovate, pubescent.

1. Leaves parallel-veined, lower stem leaves, elongate-linear, entire; involucre
campanulate. C. gramitiifolia.

1. Leaves pinnately-veined, lower stem leaves oblong-lanceolate, slightly serrate;
involucre hemispherical. C. mariana.

1. Chrysopsis graminifolia (Mx.) Ell. Grassleaf Golden-
aster. Slender pubescent plants corymbosely branched above.
Leaves grasslike, shining, 3-5 nerved, entire, the basal leaves
4-12 in. long, \-\ in. wide, stem leaves smaller, the uppermost
reduced to erect awl-shaped bracts. Heads several or numerous,
about \ in. broad; bracts of the involucre glabrate. In dry
soil. August to October. No specimens.

2. Chrysopsis mariana (L.) Nutt. Maryland Golden-
aster. Stout herbs 1-2^ ft. high, pubescent with silky hairs
when young, becoming smooth with age, corymbosely branched
above. Lower stem leaves oblanceolate to spatulate, narrowed
into a petiole, 2-4 in. long and mostly 1 in. wide; upper leaves
oblong to lanceolate, sessile, 1-2 in. long. Heads numerous,
f-1 in. broad, on glabrous peduncles; bracts of the involucre
acute, glandular, viscid-pubescent. Dry soil. August to Sep-
tember. Hocking, Jackson.

36 The Ohio Journal of Science [Vol. XVIII, No. 2,

Solidago L. Goldenrod.

Perennial herbs with simple or little branched stems, often
somewhat woody at the base. Leaves alternate, toothed or
entire. Some species with nodding tips. Heads several in
a terminal or axillary panicle, thyrsus, or cymose, corymbose or
capitate cluster; involucre oblong or narrowly campanulate, of
appressed bracts imbricated in several series, the outer suc-
cessively shorter; both ray and disk flowers yellow or rarely
white; disk flowers mostly bisporangiate, ray flowers sometimes
carpellate. Pappus of simple even capillary bristles.

Key to the Species.

1. Heads in axillary clusters, in a terminal spike-like thyrsus, in a typical
thyrsus either with normal or secund branches; not flat- topped. 2.

1. Heads in a terminal corymbiform or thyrsoid cyme forming a flat-topped

inflorescence. 20.

2. Heads in axillary clusters along the stem or the upper forming a spike-like or

somewhat branched thyrsus. 3.

2. Heads in a terminal thyrsus or panical either simple or much branched, often

secund. 8.

3. Involucre bracts with prominently spreading herbaceous tips. 5. sqiiarrosa.

3. Involucre bracts without spreading tips, wholly appressed. 4.

4. Rays white; lower leaves mostly obtuse, more or less pubescent. S. bicolor.

4. Rays yellow. 5.

5. Stem densely pubescent; ovulary glabrous. S.hispida.

5. Stem glabrous or sparingly pubescent. 6.

6. Stem leaves broadly ovate, acuminate, sharply and coarsely serrate, some-

times glabrous; ovulary pubescent. 5. flexocialus.

6. Stem leaves lanceolate, or ovate-lanceolate. 7.

7. Stem leaves acuminate, lanceolate; ovulary pubescent. 5. caesia.

7. Stem leaves merely acute, broadly lanceolate or oblong-lanceolate; ovulary

glabrous or nearly so. S. erecta.

8. Heads in a slender more or less wand-like thyrsoid panicle, not secund on the

branches or only slightly so; leaves not triple veined. 9.

8. Heads in a large usually dense panicle, secund on its slender or recurved

branches. 11.

9. Inflorescence an oblong thyrsus; growing in bogs and on wet shores.

5. uliginosa.
9. Inflorescence a pyramidal or thyrsiform panicle with numerous erect racemes;

upland species. 10.
10. Lower leaves ovate to broadly oval, serrate. S. speciosa.

10. Lower leaves lanceolate to oblong-lanceolate, entire. S. rigidiiiscula.

11. Leaves triple nerved with a pair of lateral veins much stronger than the

others. 12.

11. Leaves not triple nerved. 14.

12. Leaves rather narrowly lanceolate, the two lateral veins very prominent. 13.

12. Leaves oblanceolate or spatulate-oblong, the 2 lateral veins obscure; stem

very pubescent at the base. ' 5. nemoralis.

13. Stem minutely pubescent above; involucre less than 3^ in. high. 5. canadensis.

13. Stem glabrous throughout except in the inflorescence, often glaucotis; invol-

ucre more than '« "''• liigh. 5. serotina.

14. Stem densely' pulK>scent; leaves more or less so. 15.

14. Stem glabrous or merely pubescent above. 16.

15. Leaves rugose-veiny beneath, sharply serrate, oval to oblong-lanceolate.

5. rugosa.

15. Veins not prominent; leaves sometimes more or less triple nerved, somewhat

serrate, oblanceolate to spatulate-oblong. 5. tiemoralis.

Dec, 1917] The AsterecE of Ohio 37

16. Leaves verv rough on the upper surface, serrulate; stem strongly angled.

5. patula.
11). Leaves minutely roughened on the upper surface; stems terete or nearly so. 17.

17. Main branches of the inflorescence few, usually leafy-bracted at the base,

widely divergent, very slender. S. ulmifolia.

17. Main branches of inflorescence numerous, rather crowded, spreading, recurved

or ascending. 18.

18. Leaves firm, tapering gradually to the base, the uppermost chiefly entire. 18.

18. Leaves thin, the lower and middle stem leaves rather abruptly narrowed to a

petiole; basal leaves mostly serrate. i'. arguta.

19. Leaves ovate-lanceloate or' oblong-lanceolate; panicle usually longer than

broad; rays 2-8. S. neglecta.

19. Leaves lanceolate or oval-lanceolate; panicle usually as broad as high; rays

8-12. S. juncea.

20. Leaves ovate-oblong, mostly rough on both sides; flowers comparatively

large. -5- rigida.

20. Leaves lanceolate or oblong-lanceolate, glabrous or nearly so; flowers com-

paratively small. 21.

21. Leaves oblong-lanceolate, serrulate. 5. ohioensis.
21. Leaves lanceolate or narrowly lanceolate, entire. S. riddellii.

1. Solidago squarrosa Muhl. Stout Goldenrod. Stem
stout, simple, or rarely branched, glabrous, mostly pubescent
above, 2-5 ft. high. Leaves glabrous or sHghtly pubescent, the
lower and basal ones ovate, oval, or broadly spatulate, acute or
obtuse, 1-4 in. wide; the upper leaves oblong, sessile, acute,
entire or slightly toothed. Heads numerous, in a narrow
terminal often leafy thyrsus, sometimes 12 in. long; rays 10-16,
showy; involucral bracts green with acute or obtuse, usually
strongh^ recurved tips. Achenes glabrous. In rocky soil.
August to September. Ashtabula, Lake, Cuyahoga.

2. Solidago caesia L. Wreath Goldenrod. A plant with a
round, slender, glabrous, often glacous, blue or purplish stem,
branched or simple, 1-3 ft. high. Leaves lanceolate to oblong-
lanceolate, sessile, acuminate at the apex, narrowed at the base,
glabrous, sharply serrate, 3-5 in. long, \-\\ in. wide. Heads
in axillary clusters or racemes, often forming a stout terminal
thyrsus; involucral bracts obtuse and appressed. Achenes
pubescent. Woods and thickets. August to October. General.

3. Solidago flexicaulis L. Zig-zag Goldenrod. Stem
glabrous, angled, zig-zag, usually simple. Leaves thin, ovate,
acuminate at the apex, narrowed at the base into margined
petioles, pubescent or glabrous beneath, sharply serrate, 2-7
in. long, 1-4 in. wide, the upper-most often lanceolate and
entire. Heads in stout axillary clusters or rarely in a narrow
terminal thyrsus; bracts of the involucre obtuse to acute>
appressed. Achenes hirsute-pubescent. In rich woods. July
to September. Eastern Ohio, as far west as Cuyahoga, Fair-
field, Jackson and Lawrence Counties; also in Ottawa County.

38 The Ohio Journal of Science [Vol. XVIII, No. 2,

4. Solidago bicolor L. White Goldenrod. Stem rather
stout, hirsute-pubescent, sometimes glabrous, simple, or
branched, ^-4 ft. high. The basal and the lower leaves obovate
to broadly oblong, mostly obtuse, narrowed into long margined
petioles, dentate to dentate-crenulate, sometimes pubescent,
2-6 in. long, 1-2 in. wide; the upper leaves smaller, oblong to
lanceolate, often acute, mostly sessile and entire. Heads
crowded into a narrow terminal thyrsus, 2-7 in. long or clustered
in the upper axils; involucral bracts whitish, obtuse; rays
white. Achenes glabrous. In dry soil. July to September.
Columbiana, Geauga, Cuyahoga, Summit, Wayne, Erie, Fair-
field, Vinton, Jackson, Lawrence.

5. Solidago hispida Muhl. Hairy Goldenrod. Stem densely
pubescent or hirsute, simple or branched, 1^-3 ft. high. The
lower and basal leaves oval, obtuse or acute, petioled, pubescent
on both sides, mostty dentate, 2-5 in. long, 1-2 in. wide; the
upper leaves smaller, oblong, sessile, acute, dentate or entire.
Heads crowded in a dense terminal thyrsus, also often in
racemose clusters in the upper axils; involucral bracts yellowish,
obtuse. Achenes with a few appressed hairs or glabrous.
In dry soil. August and September. Ottawa, Lake.

6. Solidago erecta Pursh. Slender Goldenrod. A slender
rarely branched plant, 2-3 ft. high, glabrous or puberalent above.
Leaves firm, glabrous on both sides; ciliolate on the margins;
the lower and basal leaves oblong to oval, obtuse, crenate-
dentate, the upper lanceolate to oblong lanceolate, acute and
usually entire. Heads in a narrow terminal thyrsus, sometimes
with a few clusters in the upper axils ; rays light yellow ; bracts of
the involucre obtuse. In dr}- soil. August-September. Fair-
field, Hocking, Meigs.

7. Solidago uliginosa Nutt. Bog Goldenrod. A rather
stout plant with simple glabrous stem, 2-4 ft. high with the
branches of the inflorescence somewhat pubescent. Leaves
oblong-lanceolate to lanceolate, firm, glabrous, mostly ciliolate or
scabrous on the margins, few-veined, acute to acuminate, the
lower and basal ones 4-9 in. long, |-1^ in. wide, somewhat
serrate and narrowed into petioles; the upper, smaller, sessile
and entire. Heads in a terminal, oblong, dense thyrsus with
appressed branches; bracts of the involucre linear-oblong,
obtuse. Achenes glabrous. August-September. In swamps
and bogs. Lucas, Portage, Stark, Wayne, Licking, Franklin.

Dec, 1917] The Asterece of Ohio 39

8. Solidago speciosa Nutt. Showy Goldenrod. Plants
with stout stems, glabrous below and often rough above, 3-7
ft. high. Leaves firm and glabrous, the lower and basal ones
ovate to broadly oval, 4-10 in. long, 1-4 in. wide, acute to
obtuse at the apex and long petioled, dentate, pinnately-
veined; the upper smaller, oblong to oval, acute at both ends,
crenate-dentate or entire, sessile or short petioled, rough
margined. Heads in a large terminal thyrsus of ascending,
often leafy branches; bracts of the involucre oblong and very
obtuse. Achenes glabrous or slightly pubescent. In rich
woods. August-September. Lucas, Franklin, Fairfield and
Lawrence.

9. Solidago rigidiuscula (T. & G.) Port. Slender Showy
Goldenrod. vStem rather slender, glabrous below and rough
pubescent above, simple, 2-4 ft. high. Leaves lanceolate to
ovate-lanceolate, entire or the basal ones often crenate, strongly
ciliolate on the margins, 1-5 in. long, \-l in. wide, sessile or
the lower often narrowed to petioles. Heads generally in a
narrow dense, simple or branched thyrsus. In dry soil. August-
October. Erie, Wyandot, Wood, Lucas, Fulton.

10. Solidago rugosa Mill. Wrinkle-leaf Goldenrod. Stout
scabrous hirsute plants, 1-7| ft. high, simple oi" often branched
at the top. Leaves scabrous or pubescent, oval, oblong-
lanceolate or ovate-lanceolate, acute, acuminate or sometimes
obtuse at the apex, narrowed or obtuse at the base, rugosely
veined on the lower surface, serrate, 1-4 in. long, \-l\ in.
wide, sessile or the lower tapering into petioles. Heads secund
on spreading or recurving often leafy branches of usually
a ver}' large panicle; bracts of the involucre linear, obtuse.
Achenes pubescent. In dry woods. July-November. Rather
general.

11. Solidago patula Muhl. Roughleaf Goldenrod. A
stout rigid plant with usually a simple stem, glabrous below
and very rough above, pinnate-veined; the lower and basal
ones very large, 3-16 in. long, 1^-5 in. wide, oval to eliptical,
narrowed to margined petioles; the upper similar, oval to oblong,
sessile, acute, slightly serrate or entire. Heads secund on
widely spreading and recurving branches of a loose panicle.
Rays small; bracts of the involucre linear-oblong, obtuse.
Achenes pubescent. In swamps. August to September. Rather
general.

40 The Ohio Journal of Science [Vol. XVIII, No. 2,

12. Solidago ulmifolia Muhl. Elmleaf Goldenrod. A very
slender glabrous or puberulent plant, 2-4 ft. high, simple or
with arched puberulent branches. Leaves thin, oblong to
eliptical-lanceolate, acute or acuminate at the apex and base,
sharply serrate, pinnately veined, slightly pubescent, the
lower and basal ones wider, 3-5 in. long, 1-1^ in. wide, narrowed
to margined petioles, the upper smaller and sessile. Heads
secund on the usually few elongated and often leafy branches
of the panicle; ray flowers small, deep yellow; bracts of the
involucre oblong-lanceolate, obtusish. Achenes pubescent. In
woods and copses. July-September. Rather general.

13. Solidago neglecta T. & G. Swamp Goldenrod. Stem
glabrous or slightly rough above, simple, rather stout, 2-4 ft.
high. Leaves firm, the lower and basal ones lanceolate to
oblong-lanceolate, large, sometimes 12 in. long, acute, closely
serrate or serrulate, tapering to margined petioles, rough on the
margins; the upper smaller, lanceolate, acute, sessile, serrate
or nearly entire. Heads more or less secund on the short
branches of the thyrsoid panicle; rays 3-8, small; bracts of
the involucre thin, linear, obtuse. Achenes glabrous or nearly
so. In swamps and bogs. August-September. Wood, Madison,
Fairfield.

14. Solidago juncea Ait. Plume Goldenrod. Stem glabrous
or nearly so throughout, rigid, rather stout, single or branched
at the top, 1^-4 ft. high. Leaves glabrous or rough, lanceolate
to oval-lanceolate, acute to acuminate, serrate or entire, the
upper ones smaller and sessile. Heads secund on the recurved,
branches of the usually spreading panicle; rays 7-12, small;
bracts of the involucre oblong to ovate-oblong, obtuse or
acute. Achenes glabrous or sparingly pubescent. In dry or
rocky soil. June-November. Rather general.

15. Solidago arguta Ait. Cut-leaf Goldenrod. Stem simple,
rather stout, glabrous or sparingly pubescent above, 2-4 ft.
high. Leaves thin, pinnately veined, the lower and basal ones
broadly ovate-oval, short-acuminate, 3-16 in. long, 1-5 in.
wide, narrowed to margined petioles or subcordate, sharply
and coarsely serrate; the upper leaves smaller, sessile, ovate to
oblong, acute or acuminate, more or less serrate. Heads
secund on the lateral racemose branches of the terminal often
leafy panicle; rays 5-7, large; bracts of the involucre oblong,
obtuse. Achenes glabrous or nearly so. In rich woods. July
to October. Erie County.

Dec, 1917] The Asterece of Ohio 41

1(). Solidago canadensis L. Canada Goldenrod. Slender
glabrous or pubescent plants, 1-5 ft. high. Leaves 3-nerved,
linear-lanceolate, 2-5 in. long, J-^ in. wide, entire or serrate
with somewhat appressed teeth, acuminate at the apex, narrowed
at the base, sessile, or the lower petioled, glabrous or pubescent.
Heads secund on the spreading branches of the often large
panicle; rays 4-G, short; bracts of the involucre thin, linear,
acutish. Achenes small and glabrous. Hillsides, thickets and
banks of streams. August to October. General and abundant.

17. Solidago serotina Ait. Late Goldenrod. A stout
glabrous often glaucous plant, 3-8 ft. high. Leaves lanceolate
to oblong-lanceolate, thin, the lowest petioled, glabrous on
both sides or pubescent beneath, more or less rough-margined,
3-6 in. long, ^-^ in. wide, acuminate at the apex, narrowed at
the base. Heads crowded on the spreading or recurved branches
of the usually long leafy, often puberulent panicles; rays 7-15,
rather large; bracts of the involucre oblong, thin and obtuse.
Achenes finely puberulent. In moist soil. August to Sep-
tember. General.

18. Solidago nemoralis Ait. Gray Goldenrod. A slender
erect, or sometimes depressed and prostrate ash-gray plant,
^-2 ft. high, finely and densely pubescent. Leaves thin,
roughened, the basal and lower ones ovate-lanceolate or spatu-
late, petioled, obscurely triple-nerved, obtuse to acute, crenate-
dentate, 3-6 in. long, ^-1 in. wide; the upper becoming gradually
smaller, oblanceolate or linear-oblong acute or entire; heads
secund on the spreading or recurving branches of the terminal
usually one-sided panicle; rays 5-9; bracts of the involucre
linear-oblong. Achenes pubescent. In dry soil. July-Novem-
ber. General.

19. Solidago rigida L. Stiff Goldenrod. A stout simple or
branched plant with densely and finely, pubescent, hoary stem,
often becoming glabrous below with age. Leaves thick, fiat,
rigid, ovate to oblong, obtuse, pinnately veined, roughened on
both sides or usually only beneath; the lower and basal long-
petioled, sometimes 1 ft. long and 3 in. wide, entire or serrulate;
the upper sessile, clasping and rounded or sometimes narrowed
at the base, 1-2 in. long. Heads many-flowered, in a dense
terminal compound corymb, the clusters sometimes slightly
secund; rays 6-10, large; involucre broadly campanulate with
oblong obtuse bracts, the outer ones pubescent. Achenes

42 The Ohio Journal of Science. [Vol. XVIII, No. 2,

glabrous, 10-15-nerved. In dry, gravelly, sandy, or rocky
soil. August to October. Erie, Ottawa, Lucas, Defiance,
Auglaize, Madison, Franklin, Lawrence.

20. Solidago ohioensis Ridd. Ohio Goldenrod. A very
smooth, simple-stemmed plant, 2-3 ft. high. Leaves firm,
pinnately veined, flat; the lower and basal ones elongate-
lanceolate, or oblong-lanceolate, obtuse, long-petioled, serrulate
the end or towards entire, often 1 ft. long ; the upper smaller ; those
of inflorescence often bract-like. Heads 15-25 flowered, numer-
ous, in a terminal compound corymb; rays 6-9, small; bracts
of the narrow campanulate involucre oblong, obtuse and glabrous.
Achenes glabrous, 5-nerved. August to September. Stark, Erie,
Wyandot, Franklin, Champaign, Clark, Montgomery.

2L Solidago riddellii, Frank. Riddell's Goldenrod. A
stout plant, glabrous or slightly pubescent above, 1-3 ft. high.
Leaves numerous, thick, glabrous, entire, acute at each end; the
lower and basal ones long-petioled, elongate, lanceolate, some-
what triple-nerved, conduplicate, often 1 ft. long, ^-| in. wide;
the upper smaller, sessile, usually clasping at the base, con-
duplicate and somewhat falcate. Heads 20-30 flowered, numer-
ous, in a dense corymb; rays 7-9, narrow; bracts of the oblong-
campanulate involucre broadly oblong to obtuse. Achenes
5-nerved, glabrous. August to September. Lucas, Fulton,
Wyandot, Franklin, Ivladison, Clark.

Euthamia Nutt. Fragrant Goldenrod.
Perennial herbs with corymbosely branched stems. Leaves
linear to linear-lanceolate, entire, sessile, punctate, 1-5-nerved.
Heads very numerous and small. Clustered in a large, corym-
bose, convex-topped inflorescence; bracts of the involucre
obtuse, sometimes glutinous, appressed, imbricated in several
series; ray-flowers carpellate and small; disk-flowers bisporangi-
ate. Pappus of simple even capillary bristles. Achenes villous-
pubescent.

1. Leaves distinctly 3-5-nerved; heads 20-27 or more flowered. E. graminifolia.

1. Leaves 1-nerved, with a pair of indistinct lateral ones; heads rarely over

20-flowered. -E. tenuifolia.

1. Euthamia graminifolia (L.) Nutt. Bushy Fragrant
Goldenrod. A paniculately much branched or rarely simple
plant, glabrous or roughish-pubescent, 2-4 ft. high. Leaves
numerous, linear-lanceolate, acute to acuminate at both ends,

Dec, 1917] The Asterece of Ohio 43

1-5 in. long, about -3 in. wide, 3-5-nerved, rough pubescent
on the margins and nerves of the underside. Heads sessile
in glomerules, arranged in a flat-topped corymb; ray flowers
12-20; disk flowers 8-12; bracts of the ovoid-campanulate or
subcylindrical involucre yellowish, oblong to oblong-lanceolate,
slightly viscid. In moist fields and roadsides. July to Sep-
tember. General.

2. Euthamia tenuifolia (Pursh) Greene. Slender Fragrant
Goldenrod. Slender, glabrous, somewhat resinous plants,
branched above, 2-4 ft. high. Leaves narrowly linear, entire,
acuminate, sessile, narrow at the base, 1-nerved or with a
pair of faint lateral nerves, punctate with minute resinous
ducts, 1-lG - ^ in. wide, with often smaller ones crowded in the
axils. Heads in glomerules forming a dense nearly flat-topped
corymb; rays 6-12; disk flowers 4-6; bracts of the oblong-
campanulate involucre oblong. In dry, sandy soil. August to
October. Erie, Lucas, Cuyahoga, Lake.

Bellis (Tourn) L. Daisy.

Tufted herbs with scapose or branched stems. Heads rather
large and solitary at the ends of the scapes or branches, involucre
hemispherical or broadly campanulate, of 1-2 series of nearly
equal, imbricated, herbaceous bracts; ray flowers pink or white,
carpellate; disk flowers yellow, bisporangiate. Pappus none
or a ring of minute bristles.

1. Bellis perennis L. European Daisy. Small plants
with one or more naked pubescent scapes, 1-7 in. high. Leaves
basal, obovate, obtuse, slightly dentate, 1-2 in. long, narrowed
to margined petioles, pubescent and hirsute. Heads ^-1 in.
broad; ray-flowers numerous, linear, white, pink, or purple;
bracts of the involucre oblong, obtuse and usually purple. In
waste places and occasionally spontaneous on lawns. April to
November. Lake, Cuyahoga. From Europe.

Boltonia L'Her. Boltonia.

Perennial herbs with striate or angled glabrous stems and
alternate sessile, entire leaves. Heads rather large and numer-
ous, paniculate or solitary on the ends of the branches. Invo-
lucre hemispherical or broadly campanulate, of scarious-
margined bracts, imbricated in several series, the outer being
somewhat shorter than the inner series; ray flowers carpellate;

44 The Ohio Journal of Science [Vol. XVIII, No. 2,

disk flowers bisporangiate. Pappus a series of short scales with
2-4 long bristles.

1. Boltonia asteroides (L.) L'Her. Boltonia. A rather
stout plant, 2-8 ft. high. Leaves lanceolate to oblong-lanceo-
late, sessile, 2-5 in. long, \-\ in. wide; the upper linear-lanceolate
and smaller. Heads with white, pink, or purple rays; bracts
of the hemispherical involucre lanceolate to oblong-lanceolate,
acute or acuminate. Pappus of small bristly scales, often with
2-4 slender bristles nearly as long as the achene. Achenes
obovate to oval. In moist soil. July to September. Erie,
Ottawa, Lucas, Auglaize, Paulding, Defiance.

Sericocarpus Nees. Whitetop Aster.

Perennial herbs with erect branching stems. Leaves alter-
nate and sessile. Heads rather small, in terminal cymose
panicles; involucre oblong-ovoid or campanulate, of coriaceous
bracts with herbaceous or squarrose tips, imbricated in several
series, the outer being the shorter, ray flowers white, carpellate;
disk flowers mostly bisporangiate, often purplish. Pappus
of many capillary bristles, the outer often shorter.

1. Leaves entire, linear, 3-nerved; stem striate, glabrous. 5. linif alius.

1. Leaves dentate, ovate to oblong, piimately veined; stem not striate, pubescent

to glabrate. 5. asteroides.

\. Sericocarpus linifolius (L.) B. S. P. Narrowleaf White-
top Aster. Slender plants with glabrous striate stems 1-2^ ft.
high. Leaves linear to linear-spatulate, spreading, scabrous
margined, faintly 3-nerved, thick, entire, obtuse at the apex,
narrowed at the base, 1-2 in. long, \ in. or less wide, sessile
or the lower on short margined petioles. Heads clustered in
2's to 6's at the ends of the cymose branches; rays about \ in.
long; involucre about \ in. high, its bracts oblong, obtuse, the
outer with spreading green tips, the inner scarious, and often
ciliolate at the apex. Pappus white. In dr}', usually sandy
soil. June-September. No specimens.

2. Sericocarpus asteroides (L.) B. S. P. Toothed Whitetop
Aster. Stem pubescent or glabrate, slightly angled, 1-2 ft. high.
Leaves pubescent or glabrous, ciliate, pinnately veined, faintly
3-nerved; the basal ones obovate or spatulate, dentate or entire,
2-4 in. long, l-\\ in. wide, narrowed to margined petioles; the
upper smaller, oblong-lanceolate, dentate or entire, acute or
obtuse. Heads densely clustered in a rather loosely corymbose

Dec, 1917] The Asterece of Ohio 45

inflorescence. Involucre campanulate, its bracts oblong, ciliate,
pubescent, the outer with green reflexed tips. Pappus brown
or white. In dry woods. July-September. Cuyahoga, Sum-
mit, Wayne, Holmes, Fairfield, Hocking, Jackson, Gallia,
Lawrence.

Aster L. Aster.
Fall blooming herbs with branching stems, mostly perennial.
Leaves alternate. Heads of various sizes forming a corymbose
or paniculate inflorescence; involucre hemispherical, cam-
panulate, or turbinate, of various kinds of bracts, imbricated
in several rows, the outer usually shorter; ray flowers white,
pink, purple, blue, violet or rarely yellow; disk flowers bispor-
angiate, usually yellow, becoming brown or purple. Pappus
of numerous, slender bristles, scabrous or denticulate and mostly
in one series. Achenes mostly flattened and nerved.

Key to the Species.

1. Lower stem and basal leaves cordate and definitely petioled. 2.

1. None of the leaves cordate and definitely petioled. 12.

2. Stem leaves cordate-clasping. A . undulatiis.

2. Stem leaves petioled or sessile, not clasping. 3.

3. Ra^'s white. 4.

3. Rays blue or violet. 5.

4. Leaves thick, rough; inflorescence glandular. A. macrophyllus.

4. Leaves thin, smooth, inflorescence not glandular. A . divaricattis.

5. Plant not glandular. 6.

5. Branches of the inflorescence glandular. A. macrophyllus.

6. Leaves entire or nearly so, thick and firm. 7.

6. Leaves mostly sharply serrate, thin. 8.

7. Leaves glabrous above or nearly so. A. shortii.

7. Leaves rough-pubescent on both sides; the upper bract-like. A. azureus.

8. Involucre H-J^ in. high, bracts appressed, with conspicuously colored tips. 9.

8. Involucre Y^-V-z in. high, bracts loose, without conspicuously colored tips. 10.

9. Cordate leaves mostly on smooth petioles, rough. A. cordifolius.
9. Cordate leaves mostly on winged petioles, smooth. A. lowrieaims.

10. Heads loosely paniculate or subcorymbose. A. lindleyanus.

10. Heads densely racemose-paniculate. 11.

11. Stem glabrous or nearly so. A. sagittifolius.

11. Stem densely short pubescent. A. dnimmondii.

12. Stem leaves or some of them with more or less clasping cordate bases, none

of them both cordate and petioled. 13.

12. Stem leaves without cordate and clasping bases, sessile, or petioled. 22.

13. Stem rough or hirsute-pubescent. 14.

13. Stem glabrous or only sparingly pubescent above. 18.

14. Leaves or some of them serrate. A. puniceus.

14. Leaves entire. 15.

15. Heads J^-1 in. broad; leaves large, slightly clasping. A. oblongifolius.

15. Heads 1-2 in. broad; leaves strongly cordate and clasping. 16.

16. Branchlets of the inflorescence glandular viscid; involucre hemispherical.

A . nova-anglics.

16. Branchlets of the inflorescence not glandular viscid; involucre turbinate. 17.

17. Leaves thick and firm, very rough, oblong to oval. A. patens.
17. Leaves thin, roughish, oblong-lanceolate. 18.

46 The Ohio Journal of Science [Vol. XVIII, No. 2,

18. Stem leaves sharply serrate. 19.

18. Stem leaves entire or nearly so. 21.

19. Leaves tapering to the base. A. puniceus.

19. Leaves not tapering to the base. 20.

20. Leaves abruptly contracted into margined petioles, often enlarged at the base.

A. prenanthoides.

20. Leaves strongly cordate clasping, bracts green tipped. A.Icevis.

21. Bracts of the involucre with rhomboid green tips. A. lavis.

21. Bracts of the involucre narrow, not folicaeous, leaves narrowly linear.

A . junceus.

22. Involucral bracts without green tips, leaves narrow, entire. A. paniculattis.

22. Involucral bracts with green tips. 23.

23. Heads in more or less 1-sided racemes. 24.

23. Heads not in 1-sided racemes, mostly paniculate. 26.

24. Stem leaves oblong to lanceolate, serrate or mostly so. 25.

24. Stem leaves linear-lanceolate to linear, nearly entire; stem glabrate.

A. vimineus.

25. Stem pubescent or glabrate. A. lateriflorus.

25. Stem villous; leaves narrowly lanceolate, thin. A. hirsuticaulis.

26. Involucral bracts appressed, acute. 27.

26. Involucral bracts spatulate, mostly ciliate, somewhat spreading, at least

the outer obtuse; heads numerous.

A. miiltifloriis.

27. Heads solitary at the end of very small leafy branches. A. dumosus.

27. Heads paniculate. 28.

28. Stem leaves lanceolate, serrate or entire. 29.

28. Stem leaves linear-lanceolate to lanceolate, mostly entire. 31.

29. Heads ^A-% in. broad. 30.

29. Heads 3^-M in. broad, stem leaves narrowly lanceolate. A. tradescanti.

30. Leaves firm, rough or roughish; rays often purplish; involucral bracts acute.

A. salicif alius.

30. Leaves thin, smooth; rays chiefly white; involucral bracts acuminate.

A. paniculatus.

31. Heads scattered, l^-^i in. broad; the upper leaves linear. A. faxoni.

31. Heads numerous, 34~K in. broad; the upper leaves subulate. 32.

32. Stem smooth; leaves linear-lanceolate. A. ericoides.
32. Stem densely villous; leaves oblong-lanceolate to lanceolate.

A . ericoides var. platyphyllus.

1. Aster divaricatus, L. White Wood Aster. Tufted
plants with slender, assurgent, somewhat zig-zag stems, 1^-2^
ft. high, becoming glabrate with maturity. Leaves thin,
ovate-lanceolate, serrate with sharp spreading teeth, acuminate
or acute at the apex and cordate at the base, on slender petioles;
inflorescence leaves smaller, ovate and acute. Heads f-1 in.
broad, in a flatfish forked corymb of long slender divergent
branches; rays 6-9, linear, white; disk turning brown; bracts
of the involucre mostly obtuse, ciliate, with inconspicuous green
tips, the outer short and oblong, the inner linear. In open
woodlands and thickets, in rather dry soil. September-October.
Meigs, Franklin, Fairfield, Lorain, Erie.

2. Aster macrophyllus L. Largeleaf Aster. Rough plants
from long, thick rootstocks; stems reddish, smooth, often
grandular-viscid in the inflorescence. 2-3 ft. high. Basal

Dec . , 1017] The A sterecB of Oh io 47

leaves broad, cordate, with a large irregular sinus, thick,
rough and harsh above; teeth broad curved and pointed, on
long petioles; the upper leaves smaller and narrower, oblong,
on short winged petioles; the uppermost sessile and acute.
Heads about \ in. high on rigid thickened peduncles forming
a broadly corymbose irregular inflorescence; rays often 16,
lavender or violet, rarely pale; bracts of the involucre con-
spicuously green-tipped, the outer short-ovate, the inner
elongated and linear, sometimes roseate tinged on the margins.
In moderately dry soil, in shaded places. August. Rather
general; no specimens from the southwestern counties.

3. Aster shortii Hook. Short's Aster. Slender plants
paniculately branched above, with smooth or roughish stems,
2-4 ft. high. Leaves thick, smooth above, minutely pubescent
beneath; the lower and basal ones ovate to ovate-lanceolate,
acute to acuminate at the apex, cordate at the base, dentate
or entire, 2-6 in. long, 1-2 in. wide, on slender petioles; the upper
leaves lanceolate, entire, rounded at the base, sessile or on short
petioles; inflorescence leaves small and bract-like. Heads
numerous, rays 10-15, linear, violet blue, | in. or less long;
bracts of the broadly companulate involucre linear, acute,
pubescent, with green appressed tips, imbricated in several
series. Pappus tawny. On banks and along edges of woods.
September-October. From Franklin and Montgomery Counties
southward; also in Lake and Ottawa Counties.

4. Aster azureus Lindl. Azure Aster. Stem slender, stiff,
rough, branched above, 1-4 ft. high. Leaves thick, entire,
rough on both sides; the basal ones ovate to ovate-lanceolate,
acute to obtuse at the apex, somewhat cordate at the base, on
slender naked petioles; upper leaves tapering to short petioles
or sessile, lanceolate or linear; inflorescence leaves reduced to
small appressed bracts. Heads numerous; rays 10-20, bright
blue; bracts of the turbinate involucre glabrous, linear-oblong,
acute, imbricated in several series, with green appressed tips.
Pappus tawny. Along borders of woods. August-October.
Franklin, Wood, Fulton, Erie.

o. Aster cordifolius L. Common Blue Wood Aster. Bushy
much branched plants with glabrous or rarely pubescent stems,
1-5 ft. high. Leaves thin, rough, often with scattered hair
above and on veins beneath; lower and basal ones ovate-
cordate, on slender scarcely winged petioles 2-5 in. long; the

48 The Ohio Journal of Science [Vol. XVIII, No. 2,

upper smaller, ovate-lanceolate, sessile or on short petioles.
Heads numerous, small; rays 10-20, bracts of the turbinate
or cylindrical involucre linear, obtuse, green-tipped, appressed.
Woods and thickets. September-December. Rather general.

6. Aster lowrieanus Port. Lowrie's Aster. Glabrous
branched plants 1-4 ft. high. Leaves thickish, firm, often
greasy to the touch; the basal ones on slender petioles, ovate to
ovate-lanceolate, mostly cordate, acute or obtusish, serrate or
increased, 2-6 in. long; stem leaves ovate to oblong, often
cordate, on winged petioles; the uppermost lanceolate. Heads
rather few, loosely panicled; ray flowers light blue; bracts of
the turbinate involucre obtuse, appressed. In woods. Sep-
tember to October. Lake, Cuyahoga, Auglaize, Fairfield,
Hamilton.

7. Aster lindleyanus T. & G. Lindley's Aster. Glabrous
or sometimes pubescent plants, usually stout, 1-6 ft. high and
branched above. Leaves rather thick, glabrous or slightly
pubescent on the lower veins; basal ones acute or acuminate to
the apex, cordate at the base, sharply serrate, ovate, on slender
naked petioles; upper leaves ovate to ovate-lanceolate, serrate
or entire, sessile or with margined petioles; inflorescence leaves
smaller, linear-lanceolate. Heads often few; rays 10-20, blue
or violet; bracts of the broadly turbinate or hemispherical
involucre lanceolate, acute, rather loosely imbricated, glabrous
or nearly so, with green tips. Pappus white. In open places.
August to October. Wayne, Franklin.

8. Aster drummondii Lindl. Drummond's Aster. A stout
finely and densely canescent plant, branched above and 2-5 ft.
high. Leaves thin, ovate, acuminate, rough above and cane-
scent below; the lower and basal ones cordate, on slender
naked petioles, sharply toothed, 2-4 in. long; the upper cordate
or rounded at the base, usually on margined petioles; those
of the inflorescence smaller, sessile, entire or nearly so. Heads
on racemose branches; rays 8-15, blue; bracts of the turbinate
involucre linear, slightly pubescent, acute to acuminate, with
green appressed tips. Pappus whitish. In dry soil, borders of
woods and on prairies. September to October. Madison
County.

9. Aster sagittifolius Willd. Arrowleaf Aster. Stout or
slender herbs, glabrous or slightly pubescent above, 2-5 ft. high,
with a paniculate inflorescence of ascending branches. Leaves

Dec, 1917] The AsterecB of Ohio 49

thin, slightly glabrous or roughened above, glabrous or pubescent
below, the lower and basal ones cordate or sagittate, ovate,
lanceolate, sharply serrate, acuminate, 3-6 in. long, on slender
naked or narrowly margined petioles; the upper lanceolate,
sessile or on short margined petioles, serrate or entire;
inflorescence leaves very small. Heads crowded and racemose;
rays 10-15, light blue or purplish; bracts more or less glabrous
with slightly spreading green tips. Pappus whitish. In dry
soil. August to October. Rather general.

10. Aster undulatus L. Wavy-leaf Aster. Plants with
stiff, rough, pubescent stems, divergent above, 1-3^ ft. high.
Leaves rough on both sides, pubescent beneath, dentate,
undulate or entire, acute to acuminate, the lower and basal
ones ovate, cordate, 2-0 in. long, on margined petioles; the
middle ones ovate-lanceolate or oblong, with margined petioles
dilated and clasping at the base; the upper sessile and clasping;
those of inflorescence small and bract-like. Heads numerous
and racemose, often secund on spreading branches; rays 8-15,
pale blue to violet; bracts of the broadly turbinate involucre
linear, oblong, pubescent, acute or acutish with appressed green
tips. Pappus whitish. In dry soil. September to October.
Wayne County.

11. Aster patens Ait. Late Purple Aster. Slender, rough,
divergently branched plants, 1-3 ft. high. Leaves ovate-
oblong to oblong-lanceolate, rough and somewhat rigid, strongly
cordate or auriculate-clasping at the broad base, entire, acute,
or the lowest obtuse, 1-3 in. long, those of the branches smaller
and bract-like, the margins rough-ciliate. Heads 1 in. or more
broad, solitary at the ends of the branches; rays 20-30, purplish
or deep violet ; bracts of the broadly turbinate involucre linear-
oblong, finely pubescent and somewhat glandular, imbricated
in several series, with spreading, green, acute tips. Pappus
tawny. Achenes pubescent. In dry open places. August to
October. Portage County.

12. Aster phlogifolius Muhl. Thinleaf Purple Aster. Plants
resembling Aster patens Ait, but usuall}^ taller. Leaves larger,
lanceolate to oblong-lanceolate, thin or membranous, acuminate
at the apex, auriculate-clasping at the base, roughish above,
pubescent beneath, usually narrowed below the middle, often
6 in. long. Heads usually numerous, 1-2 in. broad, panicled
or racemose on the branches of the inflorescence; rays numerous.

50 The Ohio Journal of Science [Vol. XVIII, No. 2,

purplish-blue; bracts of the involucre lanceolate, glabrate,
somewhat loose, with herbaceous tips. In woods and thickets.
August to September. Wayne and Portage.

13. Aster novae-angliae L. New England Aster. Stout
leafy plants with hispid, pubescent stems, corymbosely branched
above. Leaves lanceolate, entire, rather thin, acute, pubescent,
2-5 in. long, ^-1 in. wide, clasping the stem by an auriculate or
broadly cordate base. Heads numerous, 1-2 in. broad, clustered
at the end of the branches; rays 40-50, linear, violet purple,
rarely pinkish, red or white; bracts of the hemispherical invo-
lucre linear-subulate, rather unequal, spreading, pubescent;
more or less glandular viscid. Pappus reddish- white. Achenes
pubescent. In fields and swamps. August to October. General.

14. Aster oblongifolius Nutt. Aromatic Aster. Stem much
branched, hirsute-pubescent, 1-2^ ft. high. Leaves crowded,
oblong to oblong-lanceolate, sessile by a broad partly clasping
base, usually rigid, entire, acute or macronate at the apex,
rough on both sides, rough-margined, stem leaves 1-2 in. long
and \ in. or less wide; those of the branches smaller. Heads
1 in. broad; corymbose on the divaricate or ascending branches;
rays 20-30, violet purple, rarely rose-color; bracts of the hem-
ispherical involucre much imbricated, glandular, aromatic,
linear to linear-oblong, with green, acute, spreading tips.
Pappus light brown. Achenes hoary with gray pubescence.
On prairies and bluffs. August to October. No specimens.

15. Aster puniceus L. Purple-stem Aster. Stout reddish
plants with hispid, rigid, hairy or glabrous stems, corymbosely
branched above, 3-8 ft. high. Leaves lanceolate to oblong-
lanceolate, acuminate, sessile, sharply serrate or entire, usually
rough above, pubescent on the midrib or glabrous rough above,
pubescent on the midrib or glabrous beneath, 3-6 in. long,
§-1^ in. wide. Heads usually many, 1-1§ in. broad; rays 20-40,
violet, purple or pale, rarely white, showy; bracts of the hemi-
spherical involucre linear-oblong, alternate, imbricated in
about 2 series, glabrous or ciliate, green, loose, spreading, nearly
equal, sometimes broadened. Pappus nearly white. Achenes
pubescent. In swamps. July to November. Rather general.

16. Aster prenanthoides Muhl. Crooked-stem Aster.
Flexous much-branched plants with the stem glabrous or
pubescent in lines above, 1-2 ft. high. Leaves thin, oblong to
ovate-lanceolate or lanceolate, sharply and coarsely serrate,

Dec, 1917] The Asterece of Ohio 51

scabrous above, glabrous or nearly so beneath, .3-8 in. long,
f-1^ in. wide, acuminate at the apex, abruptly narrowed below
to a broad margined entire petiole with an auriculate clasping
base. Heads usually numerous, 1 in. or more broad; rays 20-30;
violet; bracts of the hemispherical involucre acute, green,
spreading, imbricated in 3-4 series, the outer shorter. Pappus
tawny. Achenes pubescent. August to October. In moist
soil. Rather general.

17. Aster laevis L. Smooth Aster. Stem usually stout,
glabrous and often glaucous, 2-4 ft. high, branched or simple.
Leaves entire or serrate, glabrous, slightly roughened on the
margins, the basal and lower gradually narrowed to winged
]Detioles, the upper sessile and usually clasping, lanceolate,
oblong-lanceolate or ovate, acute or obtusish at the apex,
1-4 in. long, \-2 in. wide; those of the branches small and bract-
like. Heads usually numerous, about 1 in. broad, in a close
panicle; rays 15-30, blue or violet; bracts of the campanulate
involucre rigid, ovate, appressed, green-tipped, imbricated in
several series. Pappus tawny. Achenes glabrous or nearly
so. Usually in dry soil. September to October. Rather
general.

IS. Aster junceus Ait. Rush Aster. Very slender simple
or little branched herbs w4th stems glabrous or pubescent
above, 1-3 ft. high. Leaves firm, glabrous, roughish mar-
gined, entire, or with a few distinct teeth, acute to acuminate
at the apex, sessile by a broad clasping or slightly cordate base,
3-6 in. long, \-\ in. wide. Pleads about 1 in. broad, rather
distant, paniculate; rays violet to white; involucre hemispherical,
its bracts glabrous, linear, very acute, imbricated in 3-4 series,
the outer shorter. Pappus pale. In swamps and bogs. July-
September. Licking. Wayne.

19. Aster lateriflorus (L.) Britt. Starved Aster. Stem
pubescent or nearly glabrous, slender, divergently branched,
1-5 ft. high. Basal leaves ovate, slender petioled; stem leaves
broadly lanceolate, or oblong lanceolate, acuminate, serrate,
2-5 in. long, ^-1 in. wide; leaves of the branches smaller,
oblong to linear-oblong. Heads \~\ in. broad, usually numer-
ous, crowded short pedicled or sessile, arranged on the branches
in onesided racemes; rays numerous, short, white or pale; disk
flowers purple; bracts of the turbinate involucre obtuse to
acutish, imbricated in about 4 series, with short, green, appressed

52 The Ohio Journal of Science [Vol. XVIII, No. 2,

or with slightly spreading tips. Pappus white. Achenes
sHghtly pubescent. In dry or moist soil. August to October.
Rather general.

20. Aster hirsuticaulis Lindl. Rough-stem Aster. Stem
slender, erect, 1^-3 ft. high, pubescent nearly to the base.
Leaves thin, glabrous above, usually pubescent on the midvein
beneath, serrate, with a few appressed teeth, or entire, Hnear-
lanceolate to lanceolate, often G in. long, \-\ in. wide, the
sessile or basal ones spatulate and petioled. Heads more or
less unilateral on the branches, densely or loosely clustered or
sometimes few in the axils of the leaves; rays white; bracts
of the involucre linear-lanceolate, acute or acuminate, imbri-
cated in 3-4 series. In woods and thickets. August to
October. Warren, Auglaize.

21. Aster vimineus Lam. Small White Aster. Slender
bushy plants, divergently branched above, 2-5 ft. high. Stem
leaves linear-lanceolate, entire or with a few fine sharp teeth
in the middle, 3-5 in. long, \-\ in. wide, acuminate at the apex,
narrowed to a sessile base; leaves of the branchlets much smaller.
Heads very numerous, small, usually densely racemose-secund
and short pedicled; rays numerous, white to purphsh; bracts
of the broadly turbinate involucre linear, acute, green-tipped,
imbricated in 3 or 4 series. Pappus white. In moist soil.
August to September. Wayne County.

22. Aster multiflorus Ait. Dense-flowered Aster. A much
branched, bushy plant, rough pubescent or scabrous, 1-7 ft.
high. Leaves rigid, sessile or slightly clasping, rough, ciliate;
those of the stem \ to 1^ in. long; those of the branches very
small and crowded. Heads densely crowded, nearly sessile and
often sHghtly secund on the branches; rays 10-20, white;
involucral bracts coriaceous, ciliate or pubescent, imbricated in
3 or 4 series, with green obtuse or mucronate spreading tips.
Pappus brownish. Achenes puberulent. In dry open places.
August-November. Lucas, Erie, Gallia.

23. Aster dumosus L. Bushy Aster. Glabrous or nearly
so throughout, rather stiff and viscid, usually paniculately
branched, 1-3 ft. high. Leaves firm; the basal ones spatulate,
dentate; those of the stem, linear to linear-lanceolate, entire,
acute or obtuse, 1-3 in. long, \-\ in. wide, roughish margined,
often reflexed, those of the branches very numerous, small and
bract-Hke. Heads terminating the minutely leafy branches

Dec, 1917] The Asterece of Ohio 53

and branchlets, usually numerous; rays 15-20, blue to pale
violet or white; bracts of the broadly campanulate involucre
linear subulate, obtuse or acutish, appressed, green-tipped
imbricated in about 4 series. Pappus white. Achenes pub-
escent. Sandy soil. August-October. Eric County.

24. Aster salicifolius Lam. Willow Aster. A rather slender
paniculately much branched, leafy plant, 2-5 ft. high, glabrous,
roughish or pubescent above. Leaves firm, lanceolate to linear-
lanceolate, roughish or rough margined, acute at the apex,
sessile or slightly clasping at the base, entire or sparingly
dentate, glabrous or nearly so. 2-4 in. long, \-\ in. wide, the
lower sometimes petioled ; those of the branches smaller. Heads
numerous; rays numerous, violet, violet purple or sometimes
white; bracts of the involucre linear-oblong, appressed, imbri-
cated in 4-5 series, with acute or obtusish tips. Pappus white.
Achenes pubescent. In moist soil. August-October. Wayne
County.

25. Aster paniculatus Lam. Panicled Aster. A pan-
iculately much branched plant with a glabrous or sometimes
pubescent stem, 2-8 ft. high. Leaves lanceolate to linear-
lanceolate, acuminate at the apex narrowed to a sessile or
slightly clasping base, glabrous, roughish-margined; those of
the stem slightly serrate or entire, 3-6 in. long, \-\ in. wide;
the upper branch leaves gradually smaller. Heads numerous;
rays numerous, white or faintly tinged w4th violet; bracts of
the involucre linear-lanceolate, acuminate, appressed, green-
tipped and imbricated in 4-5 series. Achenes minutely pubes-
cent. In moist soil. August to October. General.

26. Aster tradescanti L. Tradescant's Aster. Stems slender,
paniculately branched, 2-5 ft. high; the branches often pubescent
in lines. Stem leaves linear-lanceolate to lanceolate, acuminate
at the apex, narrowed to a sessile base, 3-6 in. long, \-\ in.
wide, glabrous or nearly so on both sides, sharply serrate at
the middle, or entire. Heads numerous and racemose on the
branches; rays numerous, white or nearly so; bracts of the
hemispherical involucre linear, acute, appressed, green-tipped,
imbricated in 4-5 series. Pappus w^hite. Achenes minutely
pubescent. In fields and swamps. August to October. Rather
general.

27. Aster faxoni Porter. Faxon's Aster. A rather stout
glabrous plant, paniculately or corymbosely branched, ^-5 ft.

54 The Ohio Journal of Science [Vol. XVIII, No. 2,

high. Basal leaves oblong to spatulate, obtuse, dentate or
entire; stem leaves lanceolate to linear-lanceolate, acute or
acuminate at the apex, narrowed to a sessile base or a margined
petiole, mostly entire, 2-5 in. long, |-f in. wide, those of the
branches gradually smaller. Heads rather few; rays numerous,
bright white or rarely purplish; bracts of the hemispherical
involucre linear-lanceolate, acute, green-tipped, imbricated
in about 3 series, the outer shorter. Pappus white. Achenes
minutely pubescent. On moist cliffs. August-September. Vinton
County.

28. Aster ericoides L. White Heath Aster. Usually bushy
paniculately branched plants with glabrous or hirsute stems,
branchlets often secund. Leaves firm; the basal ones spatulate,
obtuse, dentate, narrowed to margined petioles, glabrous or
ciliate; stem leaves linear to linear lanceolate, acute entire,
1-3 in. long, \-\ in. wide; those of the branches bractlike and
numerous. Heads very numerous; rays 15-25, white or tinged
with rose, bracts of the campanulate or hemispherical involucre
coriaceous, lanceolate, abruptly acute, green-tipped, imbricated
in about 3 series. Pappus white. Achenes finely pubescent.
In dry soil. September to December. General.

var. Aster ericoides platyphyllus, T. & G. A broad-leaved
densely villous relative of Aster ericoides found in the western
half of the state, east to Erie, Franklin and Meigs Counties.

29. Aster ptarmicoides (Nees.) T. & G. Upland White
Aster. Slender tufted plants with rigid corymbosely branched
stems, usually rough above, 1-2 ft. high. Leaves linear-lanceo-
late, 1-3 nerved, entire or with a few distant teeth, firm, shining,
rough margined, acute, narrowed to a sessile base, or the lower
petioled; the lower leaves 3-6 in. long, \-\ in. wide; the upper
smaller and bract-like, f-1 in. broad; rays 10-20, white, bracts
of the involucre linear oblong, obtuse, the outer acutish,
appressed, nearly glabrous, green, imbricated in about 4 series.
Pappus white. Achenes glabrous. In dry or rocky soil. July
to September. (Ottawa County — Moseley Herbarium).

Erigeron L. Fleabane.
Branching or scapose herbs with alternate or basal leaves.
Heads solitary or in a naked corymbose pedunculate inflores-
cence, some species with nodding tips. Involucre hemi-
spherical, cylindrical or campanulate, of narrow equal bracts
imbricated in several series; ray flowers white or pink, bispor-

Dec, 11)17] The Asterecc oj Ohio 55

angiate; disk flowers yellow and bisporangiate. Pappus of
slender scabrous or denticulate bristles in one series or some-
times a shorter additional outer series.

Heads 1-13-2 in. broad, few; stem simple, pubescent. E. pulcliellns.

Heads J^-1 in. broad, many; stem branched. 2.

Ravs 100-150, mostly purple or violet. E. philadelphiciis.

Rays consideral)ly less than 100, white, sometimes purple tinged. 3.
Stem leaves linear lanceolate, the upper entire, the lower seomwhat dentate.

E. ramosus.
3. Stem leaves lanceolate, all but the extreme upper ones sharply serrate. E. annus

1. Erigeron pulchellus Mx. Showy Fleabane. Slender
simple villous plants, perennial by stolons and offsets, 10-24 in.
high. Basal leaves tufted, spatulate or obovate, somewhat
wedgeshape at the base, narrowed into slender margined
petioles, obtuse at the apex, 1-3 in. long, \-2 in. wide, dentate,
stem leaves partly clasping, oblong, lanceolate or ovate, acute,
entire or slightly serrate. Heads 1-6, on slender peduncles;
rays numerous, white or purple, \-^ in. long; bracts of the
depressed hemispherical involucre linear, acuminate, villous.
Pappus simple. Achenes nearly glabrous. On hills and banks.
April to June. General.

2. Erigeron philadelphicus L. Philadelphia Fleabane.
Slender, soft pubescent or sometimes nearly glabrous plants,
with nodding heads, perennial by stolons and offsets, mostly
branched above, 1-3 ft. high. Basal and lower leaves spatulate
or obovate, dentate, obtuse at the apex, narrowed to short
petioles, 1-3 in. long; the upper stem leaves clasping, obtuse
or acute, often clasping at the base, entire or sometimes dentate.
Heads several or numerous, ^-1 in. broad, on the slender
peduncles of a corymbose, paniculate inflorescence, the peduncles
thickened at the summit; rays 100-150, light rose purple to
pinkish; bracts of the depressed hemispherical involucre linear
and mostly scarious margined. Pappus simple. Achenes
pubescent. In fields and woods. April to August. General.

3. Erigeron annus (L.) Pers. White-top Fleabane. Annual
erect corymbosely branched plants sparingly pubescent with
spreading hairs, 1-4 ft. high. Leaves thin; the lower and basal
ones ovate to ovate-lanceolate, obtuse, petioled, usually coarsely
dentate, 2-G in. long, 1-3 in. wide; the upper sessile or short-
petioled, oblong to linear-lanceolate, acute or acuminate,
sharply dentate at the middle, those of the branches narrow
and often entire. Heads rather numerous, |-f in. broad.

56 The Oh'io Journal of Science [Vol. XVIII, No. 2,

short peduncled; rays 40-70, linear, white or commonly tinged
with purple, bracts of the hemispherical involucre hispid.
Pappus double, the inner a series of slender fragile deciduous
bristles, often wanting in the disk flowers, the outer a persistent
series of short partly united scales. In fields. May to Novem-
ber. General.

4. Erigeron ramosus (Walt.) B. S. P. Daisy Fleabane.
A plant resembling Erigeron annus but often lower and the
pubescence more appressed. Basal and lower leaves spatulate,
usually serrate; stem leaves linear-oblong to linear-lanceolate,
mostly entire. Rays white or purplish, sometimes minute
or wanting; bracts of the involucre glabrous or nearly so. In
fields. May-November. General.

Leptilon Raf. Horseweed.

Annual or biennial herbs. Heads small, in a racemose, or
paniculate inflorescence. Involucre campanulate, of narrow
bracts imbricated in 1-2 series. Ray flowers carpellate, white,
shorter than the disk flowers. Disk flowers bisporangiate.
Pappus of numerous bristles, in 1 series.

1. Leptilon canadense (L.) Britt. Common Horseweed.
Erect wand-like plants with leafy hispid-pubescent or glabrate
stems, 3-10 ft. high, the larger plants much branched. Leaves
often pubescent or ciliate, the basal and lower ones spatulate,
petioled, incised, dentate or entire, 1-4 in. long, obtuse to acute;
stem leaves linear, mostly entire. Heads usually numerous,
about \ in. broad; rays numerous, white, shorter than the
pappus; involucre campanulate, its bracts linear, acute, glabrate,
the outer shorter. In fields and waste places. June-November.
General.

Doellingeria Nees. Flat-top Aster.

Perennial, branching, leafy herbs. The upper leaves large,
mostly entire, acute or acuminate; lower leaves much reduced;
basal leaves none. Heads numerous, in a corymbose inflores-
cence; involucre campanulate to hemispherical, of thin
appressed, often scarious margined bracts imbricated in several
series; ray flowers carpellate, white and not very numerous;
disk flowers bisporangiate, white to greenish. Pappus double,
the outer of short bristles, the inner of long capillary bristles.

1. Stem leaves lanceolate to ovate; heads mostly numerous. D. umheUata

1. Stem leaves obovate, heads commonly few. D. infinna.

Dec, 191 7j The Aster ece of Ohio 57

1. Doellingeria umbellata (Mill.) Nees. Tall Flat-top
Aster. Corymbosely branched plants with striate stems,
glabrous or pubescent above, 1-8 ft. high. Leaves lanceolate to
oblong-lanceolate, ascending, glabrous above, pubescent on the
veins beneath, membramous, acuminate at the apex, short
pctioled at the base, stem leaves 5-G in. long, ^-1 in. wide; the
upper leaves sessile, hispid-margined. Heads numerous, in
terminal compound corymbs; rays 10-15, white; bracts of the
broadly campanulate or hemispherical involucre lanceolate,
usually pubescent, ovate, often imbricated in 3-4 series, the
outer shorter. Pappus usually white. Achenes slightly pubes-
cent. In moist soil. July-October. Rather general.

2. Doellingeria infirma (Mx.) Greene. Infirm Flat-top
Aster. Slender sparingly branched plants with round glabrous
or roughish stems, l|-3 ft. high. Leaves entire, hispid-mar-
gined, glabrous above, sparingly hispid on the veins beneath,
the lower obovate, small, obtuse, usually sessile, the upper
larger, oblong-lanceolate acute, 2-5 in. long, 1-1^ in. wide.
Heads few, about 1 in. broad, in a spreading corymbose cluster;
rays 8-15, white; bracts of the broadly campanulate involucre
oblong-lanceolate, obtuse, sparingly pubescent, imbricated in
about 4 series. Pappus tawny. Achenes nerved and glabrous.
In dry, usually rocky soil. August to September. Portage
County.

lonactis Greene. Stiff leaf Aster.

Low branching perennial herbs. Stem leaves numerous,
narrow, entire, the basal ones none. Heads rather large and
showy, terminating the stem and branches; involucre of
appressed coriaceous bracts imbricated in several series; ray-
flowers carpellate, violet; disk flowers violet. Pappus double,
the inner of long capillary bristles, the outer shorter.

1. lonactis linariifolius (L.) Greene. Stiffleaf Aster. Very
leafy corymbosely branched plants with stiff tufted puberulent
or scabrous stems, ^-2 ft. high. Leaves linear or spatulate,
spreading, one nerved, mucronate at the apex, f-l| in. long,
about \ in. wide, sessile, those of the branches much smaller and
somewhat appressed. Heads several, about 1 in. broad,
terminating the branchlets; rays 10-15, violet, rarely white,
with entire, dentate, or laciniate tips; bracts of the broadly

58 The Ohio Journal of Science [Vol. XVIII, No, 2,

turbinate involucre linear-lanceolate, keeled, green on the
back, appressed, imbricated in 4-5 series, the inner obtuse, the
outer acute. Pappus bristles in two series, tawny. Achenes
silky. In dry or rocky soil. July to October. Adams,
Hocking.

SOME OHIO HETEROPTERA RECORDS.

W. S. Adkins,
Department of Anatomy, University of Illinois, Chicago.

I am under obligations to Mr. W. J. Gerhart for the use of
his collection and for other assistance; Mr. H. H. Knight
has kindly determined some of the Miridse; and my thanks are
due to Miss Mildred M. Constiner for assistance in collecting.
The year is 1917 when not otherwise stated.

Family Scutellerid/E.

Thyreocoris nitiduloides (Wolff). Springfield, August 11.
Thyreocoris pulicarius (Germar). Springfield, August 11; Yel-
low Springs, August 14.

Family Pentatomid.e.

Peribalus limbolarius Stal. Lagonda, August 16; Springfield,
August IS; Trenton, September 26; Cincinnati, May 3.

Trichopepla semivittata (Say). Cincinnati, May 3; Springfield,
August 20.

Mormidea lugens (Fabr.). Springfield, August 11, 13, 22
(grasses).

Euschistus euschistoides (Voll.) Springfield, August 15 (nymph)
17, 23 (on catnipj, June 18, 1916; Trenton, September
15, 16, 27 (nymphs).

Euschistus tristigmus (Say). Springfield, August 14-15 (nymphs
abundant).

Euschistus variolarius (P. B.) Springfield, August 11-26; La-
gonda, August 16; Springfield, June 18, 1916; Cincinnati,
May 3 (mullein). Trenton, September 26.

Coenus delius (Say). Yellow Springs, August 12 (spearmint).

Hymenarchys nervosa (vSay). Springfield, June 18, 1916.

Neottiglossa sulcifrons (Stal). Springfield, August 11.

Dec, 191; ] Some Ohio Ileteroptera Records 59

Cosmopepla bimaculata (Thomas). Springfield, June 18, 1916,
July 1"), August 11-25; nymphs abundant on mullein and
various labiata?, August lo.

Thyanta custator (Fabr.) Trenton, May 25 (on mullein).

Acrosternum hilaris (Say). Springfield, August 15 (nymphs, on
blackberry); Clifton, August 22, 1915; Trenton Sep-
tember 25.

Banasa dimidiata fSay). Clifton, August 18 (on wild grape).

Podisus maculiventris (Say). Trenton, September 15.

Family Coreid.e.

Acanthocephala terminalis (Dall.) Springfield, August 13,
(grasses); August 15 (nymphs, blackberry).

Anasa tristis (De Geer). Trenton, October 1.

Megalotomus 5-spinosus (Say). Springfield, August 15.

Alydus pilosulus H. S. Springfield, August 13; Trenton,
vSei)tember 1().

Harmostes reflexulus (Say). Springfield, August 11-25.

Corizus lateralis (Say). Lagonda, August 20; Springfield,
August 11-24; Yellow Springs, August 12 (on smartweed
and Malva rotundifolia) ; Trenton, September 15, 23.

Family Neidid.'E.
Jalysus spinosus (Say). Springfield, August 15.

Family h\GJE\DM.

Oncopeltus fasciatus (Dall.) Springfield, June 18, 1916

(Asclepias syraica).
Lygaeus kalmii Stal. Springfield, August 16, 25; June 18, 1916

(milkweeds).
Ortholomus longiceps (Stal). Springfield, August 16.
Geocoris uliginosus (Say). Trenton, September 23.
Phlegyas annulicornis Stal. Springfield, August 11-24.
Oedancala dorsalis (Say). Springfield, August 13, 15; Yellow

Springs, August 12-14.
Myodochus serripes Oliv. Springfield, August 16.

Family PHYMATiDiE.

Phymata erosa (Linn.) vSpringfield, August 13-20; Yellow
Springs, August 14; Trenton, September 12.

Phymata erosa var. wolfii Stal. Springfield, August 13-22;
Yellow Springs, August 14; Trenton, September 12, 26.

60 The Ohio Journal of Science [Vol. XVIII, No. 2,

Family Reduviid.^.

AchoUa multispinosa (De Geer). Springfield, August 11, 13,
20, 22; Yellow Springs, ' August 14; Clifton, August 22,
1915; Trenton, September 15; nymphs and adults on elms
and other shrubbery, Snyder Park, Springfield, August
11-25.

Sinea diadema (Fabr.) Springfield, August 11-25 (nymphs);
Trenton, September 15 (nymph, alfalfa), 16 (nymph,
ragweed) .

Family Anthocorid^.
Triphleps insidiosus (Say). Springfield, August 15-16.

Family Mirid.^.
Miris dolobratus (Linn.) vSpringfield, June 18, 1910; July 17;

Yellow Springs, June 6, 1916, August 12 (grasses).
Stenodema trispinosum Rent. Springfield, August 11; Yellow

Springs, August 12;*Lagonda, August 16 (grasses).
Neurocolpus nubilis (Say). Springfield, August 11-25 (poke-
berry, tall weeds).
Adelphocoris rapidus (Say). Springfield, June 18, 1916, July 18,
August 11-25; Yellow Springs, August 12, 14; Lagonda,
August 16; Trenton, September 6, 12 (goldenrod), 15, 16
(ragweed, alfalfa), 23, 26.
Stenotus binotatus (Fabr.) Springfield, June 18, 1916; Aug. 15.
Garganus fusiformis (Say). Yellow Springs, August 12; Spring-
field, August 11-25.
Paracalocoris scrupeus (Say) var. Springfield, June 18, 1916.
Poeciloscytus basalis Reut. Springfield, August 11-24, June
18, 1916; Yellow Springs, August 12; Lagonda, August 16;
Columbus, June 1, 1916.
Horcias dislocatus (Say). Springfield, June 6, 18, 1916.
var. coccineus (Emm.) Springfield, June 18, 1916.
var. marginalis (Reut.) Springfield, June 18, 1916.
var. goniphorus (Say). Springfield, June 18, 1916, July 8.

Both the red and the orange forms are found,
var. affinis (Reut.) Springfield, June 18, 1916.
var. nigrita Reut. Springfield, June 6, 18, 1916.

These varieties were found on tall grass in an open field, and
on ragweeds. There are two other forms which do not
belong to any described variety.

Dec, 1917] Some Ohio Ileteroptera Records 61

Poecilocapsus lineatus (Fabr.) Springfield, June 18, 1916,

July 8, 14. Columbus, June 1, 1916. Both the green and

the yellow forms were found.
Capsus ater (Linn.) Springfield, June 18, 1916.
Lygus pabulinus (Linn.) Springfield, August 15.
Lygus belfragei Rent. Springfield, August 23.
Lygus pratensis (Linn.) Springfield, June 18, 1916, August

11-25; Yellow Springs, August 12, 14; Lagonda, August 16;

Columbus, June 1, 1916; Cincinnati, May 3; Trenton,

September 6 (red clover, ragweed, thistle), 12 (goldenrod),

15, 16 (ragweed, alfalfa), 23, 26.
Lygus pratensis var. oblineatus (Say). Yellow Springs, August

12.
Lygus plagiatus Uhler. Springfield, August 11, 16; Trenton,

September 26.
Lygus campestris (Linn.) Yellow Springs, August 12; Spring-
field, August 16, 23.
Lopidea confluenta (Say). Springfield, August 11, 15, 20

(locust).
Orthotylus flavisparsus (Sahlbg.) Springfield, August 24;

Trenton, September 23.
Ilnacora stalii Reut. Springfield, August 11-25 (Artemisia);

Trenton, September 6 (catnip), 12, 15 (alfalfa), 16 (ragweed).
Ilnacora malina (Uhler). Springfield, June 18, 1916.
Reuteroscopus ornatus (Reut.) Springfield, August 11-25;

Yellow Springs, August 12; Lagonda, August 16; Trenton,
September 6 (sweet clover), 15, 16 (ragweed).
Plagiognathus politus (Uhler). Springfield, August 24, 29;

Trenton, August 6, 16, 23.

SECOND ALPHABETICAL LIST OF THE LICHENS COL-
LECTED IN SEVERAL COUNTIES OF
NORTHERN OHIO.

Edo Claassen.

Since June 1912, when the first list was pubhshed in the
Ohio Naturahst, 23 Hchen species, not yet Ksted, have been
collected. The name of the counties, where the collections
were made, are again abbreviated as before: C stands for
Cuyahoga, E for Erie, G for Geauga, L for Lake, O for Ottawa,
S for Summit and St for Stark.

Several of these lichens, such as Arthopyrenia macrospora,
Arthopyrenia quinqueseptata, Bilimbia trachona, Lecanora
tartarea, Lecidea myriocarpoides and Lecidea pycnocarpa, seem
to be quite rare. It is hoped and expected, that in coming years
some more species can be added, mainly by excursions to
localities not as easily accessible as those which usually have been
visited, for instance the islands of Lake Erie.

List of Lichens:

Acarospora cervina fuscata (Schrad.) Fink. On sandstone: C.
Amphiloma lanuginosum, (Hoffm.) Nyl. On base of tree: C,

on rock: C, L.
Arthopyrenia macrospora Fink. On bark (maple) : C.
Arthopyrenia punctiformis fallax (Nyl.) Fink. On bark

(poplar) : C.
Arthopyrenia quinqueseptata (Nyl.) Fink. On bark (ash,

maple) : O.
Bacidia endoleuca (Nyl.) Kukx. On bark (sycamore) : C.
Bacidia inundata (Fr.) Koerb. On sandstone: C.
Biatorella moriformis (Ach.) Th. Fr. On dry and rotten wood:

C, St.
Bilimbia trachona Flot. On limestone: O, on sandstone: C.
Buellia myriocarpa (Lam. and D. C.) Mudd. On rotten wood: C.
Buellia spuria (Schaer.) Arn. On sandstone: C.
Cladonia delicata (Ach.) Floerke. On rotten wood: C, E, G, S.
Cladonia macilenta Hoffm. On rotten wood: C.
Lecanora dispersa (Pers.) Floerke. On old bark: E.
Lecanora subfusca campestri.s Schaer. On sandstone: C.
Lecanora tartarea (L.) Ach. On rock: G, L.

62

I

Dec, 1917] Second List of Lichens 63

Lecidea contigua convextella (Wainio) Fink. On granite
boulder: C.

Lecidea lactea (Flot.) Schaer. On sandstone: C.

Lecidea myriocarpoides Nyl. On old wood: L.

Lecidea pycnocarpa Koerb. On rock (conglomerate) : G, L.

Placodium pyraceum (Ach.) Fink. On old wood and sand-
stone: C.

Rhizocarpon alboatrum (Hoffm.) Th. Fr. On bark (maple) : O.

Theloschistes polycarpus (Hoffm.) Tuck. On bark: C.

OHIO ACADEMY OF SCIENCE

Annual Meeting

At a meeting of the Executive Committee, held in Columbus
on December 8th, it was decided to hold the next Annual
Meeting of the Academy in Columbus, May 30 and 31, 1918.
The usual detailed notice will be issued later.

Saturday, June 1, will be available for field trips for such of
the sections as may desire to hold them, the late date being
unusually adapted to this purpose.

A more prominent place on the program will be given to the
exhibition of specimens, microscopic preparations, drawings,
models, scientific apparatus, etc., than at previous meetings. It
is expected that definite hours will be assigned for such dem-
onstrations, when the exhibitors will be present to give any
desired explanations.

Edward L. Rice, Secretary.

Date of Publication, December 31, 1917.

THE

Ohio Journal of Science

PUBLISHED BY THE

Ohio State University Scientific Society

Volume XVIII JANUARY, 1918 No. 8

TABLE OF CONTENTS

With ROW — The Relation of War to Chemistry iii America G5

ObkrholseR — New Light on the Status of Kmpidonax Trailii (Audubon)... 85

ScHAFFNKR — Additions to the Catalog of Ohio Vascular Plants for lit 17 ',)9

THE RELATION OF WAR TO CHEMISTRY IN AMERICA.*

By James R. Withrow.

War is an evil beyond the power of language to express.
To kill one's neighbor or one's enemy is so repellant a thought
that one cries out in horror at the idea and instinctively wants
to refuse to have any part in action or government which
involves such baseness irrespective of the provocation. It is
only with the greatest difficulty that we persuade ourselves to
act together in any such capacity except in spontaneous defense.
Were it not for the religious emphasis upon our duty to support
the civil magistrate in the execution of righteous law, and
therefore to resist aggression against such law, we would find
little ground to stand upon in our present crisis, except it be
the desire to bring annihilation upon the philosophy which
gave rise to this world war.

It has become more and more apparent that we have been
dealing with a power in the case of Germany that is as unscru-
pulous as her acts are unmanly and cruel, and that the com-
plaints of her opponents against her since 1870 have probably
not been overdrawn. The pall of horror and indignation which
fell upon us during the invasion of Belgium and France was

*The annual address before the Ohio Academy of Science, Columbus Meeting,
evening of April 6, 1917.

65

66 The Ohio Journal of Science [Vol. XVIII, No. 3,

relieved when the Marne gave hope that there was still power
enough in the world to frustrate the dream of the bully. This
pall has been slowly growing upon us again, however, in spite
of the persistent efforts of the German propaganda amongst us
to conceal and belie the reports of the damnable conduct of
their armies and government at home and in the hapless
countries for a time at their mercy. Because of these things we
see men everywhere bowed down and depressed as it becomes
clearly demonstrated that science, mental endowments and
education are no specifics against a wicked heart. These
things we really knew before but refused to believe. They are
demonstrated to us now by appalling examples so that the
whole thinking community has become so mentally and spir-
itually depressed that one has great difficulty in going about
one's normal work, health is damaged and continued research
is a matter of great difficulty. A nation of unusual opportuni-
ties, great mental endowment and development in science seems
to have become the willing or at least easily manipulated pawn
in the hands of the unscrupulous statesman. We have not for-
gotten that it was a chemist Ostwald, in the early days of the
war when he was acting as spokesman for Germany to men of
science throughout the world, who was quoted, when Germany
was in the flush of her initial victories over Belgium, as saying,
the world had outgrown the idea of freedom for little or weak
peoples.

War, therefore, is a universal mental depressant and as such
alone, must damage progress in science. It saps national energy
and material resources. It destroys the life of the younger
generation of scientists and in large part, the student material
from which the scientists of the future are recruited. It inter-
feres with systematic research in many lines by mentally
depressing the workers, placing insuperable difficulties in their
path and at times by destroying priceless work, records and
literature. Certainly war is not desirable to science, even if
we could restrain our detestation of it and all its works.

Bitterly as we may condemn war, we would be wrong, how-
ever, to claim that science stagnated or declined in war time.
Since war requires brains, science is of course utilized, and since
the demand is inexorable, science must produce, and when
science and engineering are producing, they grow. We have
come to learn that modern war is a scientific business under-

Jan., lyicSj The Relation of War to Chemistry 67

taking. It involves the use of all vital human endeavors, and
therefore to varying extent, of all applied science. On the one
hand, it involves the utilization of medical science to maintain
physical efficiency and speedily repair damage to the fighting
machine. On the other hand, it involves the utilization of
agricultural, physical and chemical science in feeding and
clothing the whole military and naval establishment, and manu-
facturing the equipment, armament, and "concentrated energy"
or explosives consumed by the fighting force. It is stated that
it requires three men in the shops to maintain one man in the
army and seven men for one in the navy. It is evident, there-
fore that it is the applied portions of science that are most used
and hence that grow most under war's influence. It is common
experience, however, that the stretching into new domains and
the striving for new goals by applied science enriches the feeding
ground of unapplied science and uncovers fertile fields for the
patient and quiet research w^hich follows and which often
becomes the very backbone of science itself. These results are
scarcely visible and will not mature in any event, for years
after the war, so that we can see at present little good effect
upon unapplied science and we feel quite certain that the reverse
influences have the upper hand.

Although it would not be wise at present even if we had the
time to go into detail in discussing this subject, nor would it
profit you particularly, yet it may be useful to emphasize
certain points of view which come sharply to our attention when
we attempt to survey the field.

war's damage to unapplied chemistry.

We could scarcely expect to estimate the retarding effect of
the war on chemistry, because we cannot pry into it deeply
and broadly enough to prove our impressions, for research is
partly in the minds of scientific w^orkers. However, certain
signs of influences actually exist which tend to weaken and
retard progress. The American Chemical Society of some 9,000
members, the largest chemical society in the world, publishes
twice a month the journal "Chemical Abstracts. Its editorial
offices are in the Chemistry Building, a few steps from the one
in which we are now assembled. "Chemical Abstracts" has
for some years covered the field of chemistry by abstracts more
thoroughly than any foreign journal of the kind. It reviews

68 The Ohio Journal of Science [Vol. XVIII, No. 3,

some 600 journals from all parts of the world and there is spent
upon it by the American Chemical Society an annual budget of
over $40,000. It is easily seen that this must be the most pow-
erful and most important agency for research in chemistry, and
perhaps also in any science, that exists in the world. Some
quantitative idea of the evil effect of the war upon chemistry in
general may be gotten from its effects upon this colossal agency
for assisting applied as well as unapplied research in chemistry.
Inquiry to the editor of "Chemical Abstracts" developed
that the effect of the war on current chemical literature as
reflected in "Chemical Abstracts" may be shown approximately
by the following statement:

"Total No. of abstracts published (patents included):

In 1913—25,971 In 1915—18,449

In 1914—24,388 In 1916-15,784"

These figures are fairly representative of actual publication
of original papers of more or less direct interest to chemists
as Chemical Abstracts has continued closely to approach
completeness in spite of war conditions. The quality of the
papers being published is somewhat below the normal standard.
Not a great many foreign chemical journals have entirely
ceased publication since the war started, but all show more or
less marked decrease in the number of pages turned out. Most
of the French and German journals are published much less
frequently than in normal times, two or more numbers being
grouped under one cover. Apparently no important English,
Italian or Russian chemical journal has ceased publication
since the war started. The following list includes the journals
concerning which we have uncertain information, but which
indicates that they have probably ceased publication due to the
war :

English (one journal) :
Chemical World.

German (eight journals) :

Bohmische Bierb. Technikum.

Deut. med. Wochschr. Zentr. Exp. Med.

Leipziger Farber-Ztg. Z. exp. Med.

Silikat-Z. Z. Hyg.

Jan., 1918] The Relation of War to Chemistry

69

Fren'ch (thirty-one journals) :
Ann. Mines.
Arch. biol.

Arch, intern, pharaiacodyn.
Arch, intern physiol.
Arch. med. exp.
Betterave.
Brass, malt.
Bull. sci. phamiacol.
Bull. see. franc, min.
Bull. see. geol. France.
Bull. soc. ind. Amiens.
Bull. soc. ind. min.
Bull. soc. ind. Mulhouse.
LeCuir.
L'Engrais.
J. d'agr. trop.

AusTRi.VN (one journal) :
Oester. Z. Berg. Huttenw.

Belgi.an (seven journals) :

Bull. acad. roy. med belg.
Bull. sci. acad. roy. belg.
Bull. soc. chim. belg.
J. phann. soc. d'anvers.

J. des. fabr. sucre.

j. physique.

J. sci. math. phys. nat.

Radium.

Mon. ceram. verr.

Mon. teint.

Nord Brass.

Papier.

Petit brasseur.

Rev. chim. appl.

Rev. chim. ind.

Rev. electrochim.

Rev. viticulture.

Rev. gen. mat. color.

Sucrerie Ind. colon.

Rev. intern, pharni.
Sucrerie Belg.
Chimiste.

The cost of publication of Chemical Abstracts has been
increased by about 10% as a result of the war. This is chiefly
due to the increased cost of paper. The same percentage
increase will enter into the cost of the Decennial Index to
Chemical Abstracts, which is about to be issued by the American
Chemical Society at a cost of over $30,000.

Need we go further for evidence of the ill effects of war upon
science? Certainly it takes little insight to see that this
stoppage or at least side-tracking of the wheels of chemical
research will be felt in this science for years to come.

war's relation to applied chemistry.

In considering the applied side of chemistry let us remember
that war is essentially engineering. Its object is to overcome
natural and artificial obstacles. It must therefore get results
which are deliberately selected at the will of those directing the
war. It insists, therefore, that everyone and everything must
produce. Its main agents are engineering and applied chemistry,
the engineering, because it struggles with the problems of space

70 The Ohio Journal of Science [Vol. XVIII, No. 3,

and time and material for tools and weapons, and applied
chemistry, because it is a necessary handmaiden to efficient
engineering, and in addition furnishes the source and vehicle
for convenient and effective handling of energy in the most
concentrated forms. The chemical energy of the modern
high explosive is the strong right arm of the fighting force and
without it, armies are but chaff. With British control of the
seas, German armies with all their numbers, thorough equipment
and splendid miiltary power, would have been impotent in a
few weeks or months without the chemical ability to get nitric
acid from atmospheric nitrogen instead of Chilean nitrate,
for without nitric acid high explosives and even smokeless
powder are impossible.

The time at our disposal is too brief to touch on all the
divisions of applied chemistry. Much progress, for instance,
has been made in the domain of the special branch called
engineering chemistry, which involves among other things, the
chemical investigation of materials for alloys, shrapnel, aero-
planes, submarines and other war supplies. It would be
unwise now that we have become involved in the war, to deal
publicly with some of the improvements in this field, for they
are vital as well as interesting. Some of us have followed
the policy during the last three years of not even discussing
with our colleagues or students such innovations of military
importance in this and the allied countries as have come to
our attention, which might by any means percolate into
Germany. The branch of applied chemistry known as metal-
lurgy, in which this country is perhaps the most highly
developed in the world, also renders distinct service in war
time because it is vital to engineering and in the production of
arms and ammunition.

We, however, will emphasize more particularly the twin
fields of industrial chemistry and chemical engineering, because
in the nature of things this field is less popularly known even
among chemists. Industrial Chemistry is that branch of
chemistry which uses all the rest of chemistry and much
engineering, for the furtherance of production of chemical
substances, or, the use of chemical means or methods for
manufacturing any material of commerce. Chemical engineer-
ing is that branch of engineering and industrial chemistry which
applies engineering principles and methods to chemical man-

Jan., 1918] The Relation of War to Chemistry 71

ufacturing or production. Because their aim is production,
these two fields have been largely dominated by war con-
ditions for the past three years. On them the war has had two
mutually antagonistic effects, the one retarding or injuring
and the other developing and benefitting.

war's damage to chemical industry.

The main factors vital to success in any chemical industry
are:

1. Thorough knowledge of an assured market.

2. Possession of at least one well studied and workable
chemical process and chemical ability to handle it economically
under varying raw material and finished product markets.

3. Possession of engineering ability to carry out and
maintain in operation the chemistry involved in the process.

4. Sufficient margin of profit to attract capital and business
confidence in chemical and engineering ability in meeting the
problems of the field.

Anything or anyone who weakens or strikes at any of these
four factors is an enemy of chemical industry and does damage
to it.

When war was declared in Europe stagnation set in at once
in the chemical industries and indications of disaster were the
rule in many of them. Petroleum refining, turpentine, rosin
and wood products among others were, hard hit because we
are strong exporters and such industries as mixed fertilizer
manufacture also, because we import heavily of potash. This
stagnation could not last long, since the chemical industries
underlie the whole fabric of modern industrial development
and civilization, and production is necessary to life. Eventually,
therefore, the chemical industries were forced to resume oper-
ations but great uncertainty as to markets rendered operations
difficult and held back many changes in processes and equip-
ment, rendered necessary by changes in source or k'nd of raw
materials. The nature of these industries is often such that
the failure of supply of one chemical raw material even if used
in but limited amounts may prove fatal by rendering the
product unsatisfactory to the market if indeed it is not entirely
valueless. A good illustration of the vital importance of
accurate knowledge of the market in these chemical industries
is furnished by the dye situation, where we had recently the

72 The Ohio Journal of Science [Vol. XVIII, No. 3,

anomalous condition of bitter complaint of shortage by con-
sumers simultaneously with utter inability of some producers
to market their product, and still other producers with large
contracts for product and inability to produce due to poor
deliveries or failure of equipment. These difficulties do much
harm, since they tend to discourage capital and it must not be
forgotten that industrial chemical development is im.possible
without capital. German chemical manufacturers understood
this clearly when they organized American branches of their
color works, eliminating American employees to conceal the
market and its peculiarities, and placing all their business in
the hands of "American citizens" of German name. Then
when the U. S. Bureau of Foreign and Domestic Commerce
attempted last September to publish the amounts of each
dye consumed in this country, they vigorously protested that
their rights as American citizens were being infringed by
encouraging competition. The uncovering of this octopus to
public gaze should be set down to the war's credit. It has
long been a familiar animal to many industrial chemists.

Another evil effect of war, a common one now greatly
intensified, is the discouragement of capital by failure of hasty
and ill-advised manufacturing projects. Successful specu-
lators and others have been influenced by the potential earning
capacity of industrial chemistry and have jumped into projects
with little study and no experience. Often such capital has
not known enough to employ chemical engineers, but has put
growing works into the hands of electrical and mechanical
engineers whose general engineering sense has not always saved
them from physical disasters that chemical experience would
have avoided. Such engineers and capital and sad to say,
m_any chemists, who either, lacking entirely in manufacturing
experience or having had manufacturing experience, though
they acquired no sense of responsibility to protect capital
against hazard from decisions without basis in experience,
have been the easy victims of the machinery and equipment
company who needs but to see a plant, or a picture in a book,
and they will design you one while you wait. There not being
the proper engineering check, such plants fail at times with
regretable loss of life as well as capital and confidence in things
chemical, or if they succeed (because the process is simple and
well known) the plant can be counted upon to cost from 50
to 100 or more per cent, higher than it should.

Jan., 1918] The Relation of War to Chemistry 73

The equipment companies and their engineers are not
necessarily dishonest. They sell equipment, and who but
they are responsible if they do not sell you enough equipment
when you consult them for advice in designing your plant?
They, therefore, sell you enough. Experienced engineers
will often cut the estimates of such equipment manufacturers
in half.

Another illustration of how this situation works out in
practice might be given in the case of benzol refining. This
is an important matter in modern high explosive manufacture.
Some little time ago the best text ever written in English on
Industrial Chemistry contained a chapter by a chemical engineer
who had ample opportunity of observing the best American
practice (which happens to be second to none in the world).
His chapter on this subject, therefore, is a classic, but in
illustrating the text he did not reproduce details of stills, for
instance, with engineering exactness but allowed the artist
who made the drawings considerable leeway to his imagination.
In fact, he left out entirely a vital feature in the construction
of such stills. Were it not for the loss of efficiency and the
expense involved, you would be greatly amused if you could go
with me to a number of the refineries built in this country in the
last two years under war pressure by machinery companies, for
good engineers who were not themselves experienced in this
industry, but who needed the industry as one of the links in
their larger operations. In every case the stills were built
exactly patterned after the picture in this text and in no case
were they efficient or as nearly efficient as was possible, if a
little thought regarding the use to which they would be put
had been given them. They were built to sell, not to operate.

The same capital, newly invested in chemistry, is also the
easy victim of another evil which is necessarily costly to indus-
trial chemistry and is a heavy blow to the whole science. This
evil is the ignorant or unscrupulous chemist. The great dif-
ference between industrial chemical research and other chemical
research is that the former must produce results on the prob-
lem in hand while the latter may ramble if necessary into less
dif^cult fields. When inexperienced capital is seeking chemical
assistance the first individual it meets who claims to be a
chemist is assumed to be competent to handle any problem
without inquiry into his past experience. This same capital

/4 The Ohio Journal oj Science [Vol. XVIII, No. 3,

would scarcely employ a bridge engineer to design a dynamo,
yet plant after plant for chemical manufacture has been con-
structed in the last two years in this country with no more
intelligence than this. As a result literally millions have been
squandered and lost in these unsuccessful plants. But unfor-
tunately, enough such plants are successful, that their authors
escape the penalty of their dishonesty, and therefore, the evil
persists and continues. Plants have been constructed for the
manufacture of high explosives by engineers who knew nothing
of the business, resulting in great loss of property and even life
from their final destruction, or in abandonment where they
proved unprofitable. I have heard of plants erected for the
concentration of sulfuric acid in which a battery of stills for this
purpose costing in the neighborhood of a quarter of a million
dollars was placed in operation without even a single experiment
preliminary to erection, on the type of material to be used, and
not even a trial run on one of the stills before all were placed in
operation. The first day they operated was the last day, for
they all went into solution in the acid.

Men who were or claimed to be chemists have read how
simply some reactions described in the general chemistries
work, and designed a plant upon their nerve or — as they
thought — common sense, and found to their consternation
that under the conditions they made for themselves the reaction
did not proceed at all, or they were so inexperienced in large
scale operations that they could not recognize what they had
when the work was under way. Others have so far lost their
heads by publicity" or financial possibilities, even though good
chemists, that they have assumed that what could be done
with raw material from one source could be equally w^ell done
with it when from another source, provided they merely proved
its actual presence in the new product. Ignoring the whole
history of chemical as well as industrial chemical development
that the chemical environment profoundly affects chemical
reaction, no adequate confirmatory studies were made before
capital to the extent of hundreds of thousands of dollars has
been induced to invest in such guesses, with disastrous results
to capital and grave loss of confidence in chemical research.
These things are in large part due to or at least the losses could
only be so heavy under war pressure. Processes which gave
every promise of success have been hurried into failure or near

Jan., 191SJ The Rchilion of War to Chemistry 75

failure by undue publicity giving premature capitalistic con-
fidence in them and it is with profound regret that we see the
passing for the time at least of such things as toluol from
petroleum, which more attention to study may still make
useful in war emergency at least.

These are outlines of some of the evil influences due to or
accentuated by war. They are in part of such a technical
or professional nature that they should not be imposed
upon your attention unless it were to protect you against
misunderstanding the just criticism of the results of these evils
and to emphasize that we do not consider war an unmitigated
blessing if we should appear enthusiastic about the progress
that has been made in war time. Then too, we should always
ponder more over our lapses. The successes can take care of
themselves.

PROGRESS IN APPLIED CHEMISTRY IN WAR TIME.

There is indeed another side than the evil we have been
discussing. There has been much real progress. The evils
mentioned are largely growing pains. Engineering and its
services to mankind have been long appreciated to some extent
at least. Chemistry is less easily understood. The everywhere
present applications of chemistry pass unnoticed for the most
part in everyday life. Probably the greatest contribution to
science, therefore, of the present war, is the awakening of the
average mind to the power and value to mankind of that group
of phenomena which we study as chemistry. This is probably
because we most easily grasp and appreciate applications
rather than generalizations, and the use of chemistry in war has
been a revelation to the general public.

In other ways also this war has affected a development in
chemistry and its applications which has outstripped any
influence since the modern foundations of the science were laid
over a century ago. It will be many years before the influence
will mature and become apparent or measurable. Neverthe-
less, we do not crave progress or development at such a price
as war.

We must recognize, however, that severe disturbances are
very effective in dislocating fetishes, for instance. So, one of the
phases of this struggle which is noteworthy is the public awak-
ening to consciousness of the power of chemistry and the

76 The Ohio Journal of Science [Vol. XVIII, No. 3,

universal distribution of the ability to use it promptly and
effectively, as against the old idea, that this power and this
ability is possessed by a chosen few. An illustration or two
will perhaps show that this latter idea is still too prevalent.

I have met manufacturers since the war whose operations
were brought to a full stop by lack of some raw material or
other who complacently accepted their fate on the ground that
they could not get a German chemist. They had no bias in
favor of Germany at all. They just thought it was a matter of
common information that chemists were domestic animals
imported from the Black Forest. Would you believe that some
of these manufacturers were engineers graduated from some of
our large colleges of engineering and not men without education?

In such a time as this we see that our keeping quiet about
the progress and development of American chemistry in years
gone by, was criminal, for much harm results from lack of
information as well as from misinformation. There are always
patriotic Germans and others who praise their country's
achievements to us and as I pointed out last year in your Ohio
Journal of Science, we are glad to see this and our University
teachers of chemistry have been lavish in their praise, partic-
ularly of German chemistry. They, however, are not rendering
very good service to the community when, as they should, they
give such praise if they fail to make a real effort to find out also
what is going on in their own country. We university professors
feel abused if it is inferred that we are not well informed, yet
we innocently assume as the only modern development in
chemistry the latest tale of achievement from a German dye
advertisement and these lads know how to use the educated
public and university chemistry professors as well, in furthering
their advertising propaganda. Much good work has been done
by the "Journal of Industrial and Engineering Chemistry" in
publishing a series of articles by authorities, on what the
American Chemist has done for the individual industries.
Time only will eradicate the evil. Only a short time ago I was
at a banquet of a society of engineers in an eastern city. The
professor of chemistry from a nearby university was an invited
speaker. He was a revered and respected man among American
chemists and a man of affairs, too, but he lived in the dark ages
of chemical achievement. He spent half of his time telling how
wonderful chemistry was and how great the achievements of

Jan., 1918] The Relation of War to Chemislry 11

foreign chemistry in particular, and not one word of American
chemistry. Yet in his own city in the last three years has
sprung up a chemical industry that is marvelous, and which he
did not know existed. In his own line, organic chemistry, was
a plant for making certain organic materials used in war by a
series of steps that has no counterpart in chemical literature for
the magnitude and conception of its chemical engineering
operations. It is not only the largest scale upon which all of
its many operations have ever been conducted but its chem-
istry is a series of highly interesting adaptations and develop-
ments. When peace comes again, if that plant still prospers it
will be a useful aid in the solution of one of our most important
engineering problems of this generation. x\mericans are not
wizards that they do in two years what it took German chemists
decades to work into. Such things are only done where the
ability exists and the power born of experience in solving similar
chemical problems, is possessed. It is not right to our students,
you who teach, to praise the competitors of our compatriots and
never stir yourselves to be informed on what our own country-
men are doing, even if, the foreign achievements are served up
to us, ready to teach as paid advertisement of German dye-
makers. The German general staff has learned, if others have
not, that German chemical achievement which is great indeed,
is no sign that equal ability does not exist elsewhere. The Allies
and America improvised a munitions industry in two years to
match their machine of forty years preparation. Such an
achievement is only the natural result of our present industrial
chemical development in America and the Allied countries.
There is nothing in the rate of American industrial chemical
development of which any American need be ashamed.

The progress in industrial chemistry and chemical engineer-
ing in the last three years itself, in this country, has been
wonderful. Let me protest, however, that this is no ground for
the philosophy which I understand obtains in some quarters,
that war is a desirable, natural, logical or sort of evolutionary
benefit. All this progress is in spite of war. War could force us
to do nothing we did not possess capacity for before. Because
war changes the normal relations between supply and demand,
cost and selling price, gives us opportunities to do only what we
could do anyway, if the same demand arose from any other
cause.

78 The Ohio Journal of Science [Vol. XVIII, No. 3,

Industrial chemical tendencies during the war have been
governed by unusual demands for chemicals from abroad in
addition to war drains, healthy home requirements, new
demands from industries formerly supplied from abroad or
forced to use new raw material by scarcit}^ or high prices,
together with speculation, raising prices to unusual levels.
This resulted in expansion of existing plants, rapid installation
of new ones, hasty perfecting of new processes already slowly
maturing and the seizing of opportunities to profit by high
prices through erection of small plants for the production of
special chemical materials and through the development of
processes hitherto existing as possibilities, only, in the minds of
chemists. This has greatly extended also the supplying of
chemical construction materials and machinery and has
increased the opportunities for the rapid development of inven-
tions in this line. The progress made here alone has been as
great as has been accomplished in many individual decades in
the past. The importance of this is apparent when we consider
that if the chemical engineer had at his disposal as efficient
apparatus and materials of construction in his plant, as exist in
the chemical laboratories of the present day, or as the mechan-
ical and electrical engineers have in their work, progress in the
arts w^ould be at least a hundred years ahead of its present
development.

The tendency to manufacture at the market is another good
development which has been greatly accentuated by the war.
For some time there has been a growing tendency for manu-
facturers who are large consumers of chemicals to produce
these chemicals themselves. Assisted by gradual price elevation,
this tendency has been greatly encouraged by the invention in
the last two decades of processes and machines of merit which
could find no sale as such, in well established chemical manu-
facturing plants, because they frequently offered insufficient
advantages to warrant discarding those already operating, or
were merely alternative in their character. A good example of
how this tendency to manufacture at the market, works out
normally where the impelling force is merely gradualty advanc-
ing prices, competition preventing excessive elevation, is to be
seen among others, in the case of bleach for paper manufacturing.
Consumers of alkali and bleach, such as progressive paper
manufacturers, operating on a large scale, and others have

Jan., 1918] The Relation of War to Chemistry i\)

experimented for years with inventions for the electrolytic
production of these materials from common salt. Our present
high development in this branch of chemical industry is in no
small degree due to these individual efforts, many of which
during the past twenty years have been eminently successful.
High prices and poor deliveries in the last two years have
forced matters to a head in this direction. Where formerly we
had a few large chemical plants manufacturing caustic soda and
chlorine for bleach by electro-chemical means, we now have
distributed throughout the country a great number of concerns
who have added to their equipment a plant for the production
of these products. The operation of these units under widely
diverse conditions will greatly enrich our chemical engineering
experience. A number of cell types are obtainable which operate
economical^. Some of these are well advertised in the current
literature, but some though equally successful, such as the
Allen-Moore, Gibbs, and Nelson Cells, are not so well known.
The cell portion of such a plant is only a fraction, however, of
the equipment required and it is important that the rest of the
plant should be properly designed. The simpler and more
durable, therefore, the design of apparatus, the more satis-
factory the entire equipment will be. There has been placed
in operation in some eight plants recently a total of nearly 2,000
cells of one type alone, with a daily capacity of 200,000 pounds
of chlorine gas. Some plants constructed this year cost as much
as a half million dollars. These will be valuable for defense, for
we use much chlorine in making guncotton or nitro cellulose
for mines and smokeless powder.

This use of alternative inventions is valuable in encouraging
new invention and much industrial chemical investigation, and
alleviates to some extent the ill effects of unwarranted increase
in selling prices.

Progress in Chemical Engineering may be illustrated perhaps
best by the progress in acid making equipment. High pressure
manufacturing of chemicals and difficulty of obtaining supplies
has brought rapid improvements and development of chemical
engineering materials by compelling large scale experimentation
of new products and substitutes. To resist corrosion by acid
and other chemicals, pottery or so-called chemical stoneware,
glass and natural stone apparatus have been used heretofore.

80 The Ohio Journal of Science [Vol. XVIII, No. 3,

This necessitated small sized apparatus, and meant in the case
of stoneware a manufacturing time of about two months for
the clay working, drying and cooling after firing. Attempts
have been made for many years to replace this material by metal.
Platinum, silver and gold are used in special cases, but while
these metals can be made into any size apparatus, cost is
prohibitive for most uses. Two classes of alloys have now been
developed: rare metal alloys, such as tungsten, chromium,
or nickel irons, and more recently the cheaper and more resistant
silicon-iron alloys. Extensive trials in the last two years have
shown the usefulness of these alloys though they do not possess
quite the resistance of stoneware to corrosion. They are known
under varying trade names, such as durion, made here in Ohio,
tantiron, and ironac. They are very resistant to all strengths
of sulfuric and nitric acids and are used with great satisfaction
in their manufacture and permit plants to run for months
without shut down. The success of the modern tower system
displacing platinum for concentrating sulfuric acid has been
largely due to the use of pipes and fittings of this alloy.

Early in 1915 the demand for nitric acid for war purposes
increased to enormous proportions resulting in extensions to
old nitric acid plants and the erection of new ones larger than
the world had ever seen. Deliveries on stoneware jumped
to six months and even longer and had the production of
nitric acid been dependent upon stoneware alone, as a few years
ago, it would have been greatly curtailed and the story of the
great war would have been different. As these alloys can be
secured on short notice, the same as cast iron, chemical manu-
facturers do not hesitate if a still should run wild and froth
sodium sulfate into the condenser to direct workmen to break
the connections at once with a hammer and allow the expelled
material to flow on the floor, thus preventing the wrecking of
the condensing apparatus. New castings can replace the broken
one at once. Such extravagant handling of the material
would not be possible under the usual slow deliveries with
stoneware. This freedom from risk of damage to condensers
and the making of condensers themselves of this material,
enables stills to carry a heavier charge and be operated at
greater speed. Where the old equipment charged 2000 lbs.
once or twice in twenty-four hours, these war-time stills operate
on 0000 lbs. of nitre, plus 4000 lbs. of sulphuric acid, charging

Jan., 1918] The Relation of IVar to Chemistry 81

three times per twenty-four hours. The alloy is somewhat
brittle, but very much less so than chemical stoneware. It is
easy to see these silicon-iron alloys are a boon to the acid
industries and thousands of tons of castings are in use and new
chemical processes are possible and now in operation, too,
which could not exist before, because of lack of suitable material
of which to construct apparatus. Some of these new processes
are having a decided value in the Allies' campaigns. No
single development in many decades has had as much influence
as this one has, and will have, for it is only in its infancy.

I need not weary you with other illustrations of progress
though much has been accomplished in many lines and radically
new chemical processes developed. The most wonderful and
greatest chemical works I have ever seen have been erected in this
country since the war began and the best of them were coal-tar
dye and synthetic organic chemical works. Reasonable progress
has been made in American laboratory glass and porcelain.
After the war we are going to be independent of importation in
gross coal-tar products and practically if not entirely, in
ammonia for fertilizers. We are also weeding out the unneces-
sary use of potash where it replaces soda due to our own careless
teaching of chemistry in speaking of and using potassium
compounds where sodium serves as well. German potash
exporters and others, such as for Saxony manganese, after
the war will have an expensive campaign to win us back to
these former unwarranted uses of their product.

The relation of chemistry to national defense has been rendered
clear by the war, a service of no mean magnitude.

Explosives and asphyxiating gas manufacture are dependent
upon labyrinthian chemical engineering operations. It is
obviously necessary for adequate preparedness that this country
should be self-contained and not dependent upon importation
for such supplies as nitric acid, toluol and sulfuric acid for
defense. We have the sulfur and pyrites for sulfuric acid.
The toluol and other coal-tar products we have ample for our
usual needs, but in time of war toluol becomes the basis of
"T. N. T." or trinitrotoluol, one of the most effective high
power military explosives. The erection of new coke oven
plants has but partially met the demand for toluol in the last
two years. In defending ourselves this would be too slow,
for such installations are difficult to get under successful opera-

82 The Ohio Journal of Science [Vol. XVIII, No. 3,

tion in less than a year. A large and well established dye
industry, therefore, is vital for defense, for it would produce a
bigger demand for coal tar products and toluol production in
peace times and its operations are quickly convertible into
ones for producing high explosives. It is to be hoped therefore
that the German alliance with our textile manufacturers may
be broken up during this war so that Congress will be less
helpless in fostering this dye industry as a matter of defense
than it has been in the past. The expense of storing within
the country nitrate of soda imported from Chile, adequate
for the nitric acid of munitions production in case of war,
w^ould tie up millions. The Government will establish a
plant to make nitric acid from the atmosphere. The Norwegian
process (electric arc) is stated to require five times as much
power, a vital factor, as is required in the making of nitric
acid from cyanamide. Germany has installed for making
cyanamide, during the war additional equipment costing
$100,000,000, utilizing over 600,000 horse power and producing
about 200,000 tons per year of nitric acid, requiring the most
feverish activity for a year and a half on the part of her chemical
engineers. We have som^e American suggestions which if
successful will take less power than the German method. Any
method for nitric acid producing ammonia also, is desirable as
an aid to agriculture. Prices asked for power are much higher
than abroad and as the cost of engineering is only about 10% of
the total charges in electric power installation, it becomes
evident that efficient national defense and economic agriculture
depend on more economic banking methods. So in every
instance we are confronted with the problems of peace when
working out national defense. It should be remembered that
our usual source of nitrogen derivatives, the ammonia of by-
product coke, brings with it the indispensable toluol, and no
electrical method does this. Before the Government nitrogen
plant is built, therefore, it should be a matter of serious inquiry
whether the Government's $20,000,000 might not bring the
same result and give a liberal supply of toluol besides, if invested
in by-product coke expansion, for much of our coke is still made
without saving by-products.

It is an open secret that the acceptance of war orders in
this country strained to the breaking point our best organized
chemical industries. The mere request by the allied countries

Jan., 1918] The Relation of War to Chemistry 83

two years ago for our soda, benzol, toluol and our explosives
for only a small portion of their demands, produced a state of
affairs in our industries that was an appalling warning against
the time when we would need such things ourselves, for defense,
and in immensely greater volume.

It is natural in view of the nature of these defense problems
that the engineers and chemists of the country have been
serious in the preparedness movement. Thirty thousand
engineers and chemists of the United States volunteered without
pay to the National Consulting Board for both the Navy and
Army to work on the organization of the industries of the
country for national defense. The result was m.uch more
efficient than any similar organization in the world, for no
government could afford to pay for the expert services involved.
This Consulting Board and its successor, the National Council
for Defense, have assisted the country to become self-contained
for defense, arranged for speedy conversion of industrial plants
into munitions plants and arranged during peace to prevent
useless waste of experienced engineers. Experienced chemical
engineers for instance, like naval officers, cannot be trained in a
day or a year, though the analytical chemical control can be
taught in a few days to any chemist. The mistakes made b}^
Britain in passing through the blockade materials helpful in
explosive manufacture demands that our military authority
and foreign office have at its call as wide a variety of chemical
experience and advice as possible, and every chemist as well as
engineer in this country is now being card indexed.

If we as scientists ask ourselves individually what we can do
to assist in the general defense of our firesides and our ideals,
the answer is do our daily work in whatever field it may be, as.
though it were the most important single thing in the world and
particularly do our utmost to assist productio?i and those directly
engaged in it, whether manufacturing or agricultural . Then
when the government calls upon us for special service we will
be ready to attack the problems which only the military arm
can formulate for us.

We have touched on sufficient high points to indicate the
character of the influence of the war upon chemistry in America.
Still other points should be discussed, were there time. Hardly
any branch of the science but contributes an important service;
to the national defense as well as our normal benefit.

84 The Ohio Journal of Science [Vol. XVIII, No. 3,

After all is not chemistry and science itself a petty matter
in the presence of this world calamity and the personal suffering
ever upon our minds? Have we not often wondered what we
had done to be spared to this minute from such things? It
may be proper to say we do not quarrel with the German people
as such, but with the ideals and acts of their leaders and govern-
ment. Do not let that point of view toward our neighbor,
however, be used by us to excuse our individual responsibility
for this government and its every act. We are responsible, and
we alone. We have seen conclusive proof in the last three years
that science and education are merely aids and not specifics
against international immorality and that the devotees of
science are as easily misled as others when the leaders too are
scientific. Though this war has long become evident as a war
for privilege and the exploitation of the weak by the strong,
and the doctrine that the state can do no wrong rather than
that the state must do no wrong, let us not deceive ourselves
that our abolition of aristocratic government is a specific for
this malady, for it is not. This is our constant battle still,
even under our form of government.

It has become so evident in this war that the intelligent
and scientific criminal is a terrible menace, and dislodging him
at times such a weary and fatal task, that we must find some
way of preventing our leaders and groups or classes, whether
governmental or industrial, from becoming this kind of danger.

Have we not reached the time when we are willing to turn
to the One who ordained civil government for our good, acknowl-
edge that He ordained it and not we ourselves, and make our
leaders or rulers "Whom God and this people shall choose" —
"men fearing God and hating covetousness?"

The Ohio State University, Columbus.

NEW LIGHT ON THE STATUS OF EMPIDONAX TRAILLH

(AUDUBONj.

By Harry C. Obkkholskr.

Few birds are so puzzling as the flycatchers of the American
genus Empidonax. None of these are more difficult to under-
stand than Empidonax traillii and its closely related forms,
the status of which has long been in dispute. The careful
investigations and reasonable conclusions of Mr. William
Brewster,* have long been accepted as the final word by
ornithologists generall}-; and it is with some degree of reluctance
that the wTiter now reopens the case. Subsequent identification
of breeding specimens, however, from the north central part of
the United States, from Ohio to x\rkansas, Missouri, and Iowa,
has indicated discrepancies in the range of Empidonax traillii
traillii as currently delineated. With the idea, therefore, of
confirming Mr. Brewster's conclusions and settling the identifi-
cation of specimens of this species in the Biological Survey
collection, the writer resolved to investigate the matter de novo;
and to this end has examined a large amount of material; in all,
411 specimens. The rather surprising and, to some extent,
unfortunate results of this study were announced at the meeting
of the American Ornithologists' Union at Philadelphia in 1911,
and therefore they long ago should have been placed on perma-
nent record.

For the preparation of this paper the writer has made use
chiefly of the collection of the United States National Museum,
including that of the Biological Survey. In addition, he is
indebted for the loan of specimens and for other assistance to
Mr. P. A. Taverner, Mr. F. C. Lincoln, Mr. L. J. Kersey, and
Mr. F. Kermode.

Empidonax traillii traillii (Audubon).

Muscicapa traillii Audubon, Birds Amer., folio ed, I, 1828,
pi. 45 (woods along the prairie lands of the Arkansas River,
Arkansas) .

Empidonax traillii alnorum Brewster, Auk, XII, No. 2,
April, 1895, p. IGl (Upton, Maine).

*Auk. XII, No. 2, April, 1895, pp. 159-163.

85

86 The Ohio Journal of Science [Vol. XVIII, No. 3,

Chars, subsp. — Bill rather small; upper surface decidedly
greenish; wing-bars whitish, tinged with greenish or yellowish.

Measurements. — Male:t Wing, 04. 5-78 (average, 71.7) mm.;
tail, 54-62.5 (58.2); exposed culmen, 11-13 (12); tarsus, 15.5-
17.5 (16.7); middle toe without claw, 9-10.5 (9.5).

Female:J Wing, 64.5-75.5 (average, 68.3) mm.; tail, 51.5-
61.5 (55.2); exposed culmen, 10-12.5 (11.6); tarsus, 14.5-17.5
(16); middle toe without claw, 8.5-10 (9.4).

Type locality. — Borders of the woods along the prairie lands
of the Arkansas River, Arkansas.

Geographic distribution. — North America, except the extreme
northern part, south to Central America and northwestern
South America; no authentic record for the West Indies.
Breeds in the Hudsonian, Canadian, Transition, Upper Austral,
and occasionally the Lower Austral zones in North America,
north to Newfoundland, central Quebec, northern Ontario,
northern Manitoba, northwestern Mackenzie, and central
Alaska; west to western and southeastern Alaska and western
British Columbia; south to central southern British Columbia,
northwestern Montana, southeastern Montana, southern South
Dakota, northeastern Colorado, central Arkansas, southern
Kentucky, southern West Virginia, western Maryland, Penn-
sylvania, and northern New Jersey; east to eastern Massa-
chusetts, eastern New Hampshire, eastern Maine, Nova
Scotia, and Newfoundland. Winters south to Ecuador; and
north to northern Colombia, Costa Rica, Nicaragua, and
Yucatan, Mexico. Migrates west to central Colorado, western
Texas, and Oaxaca.

Remarks. — The best characters to distinguish the eastern
form of this species from that of the western United States, from
Colorado to California, are the more greenish upper surface, the
more greenish or yellowish wing-bars, and smaller bill. Every
specimen does not exhibit all these characters to the best
advantage, but enough are usually present to render possible
satisfactory identification. Furthermore, the eastern bird is
somewhat darker above, though this is not more than an average
character.

jThirty-seven specimens, from New England, Illinois, Minnesota, North
Dakota, and Montana.

JTwenty-eight specimens, from New England, Pennsylvania, New Jersey,
District of Columbia, Indiana, Illinois, North Dakota, Yukon, and Alaska.

Jan.. 191S] Status of Em pidouax Tniillii 87

The geographic variation in this subspecies is not great.
Breeding birds from Maine and New Brunswick seem to be
absolutely indistinguishable from those taken at the same time
of year in Maryland and in Mackenzie, Canada. Furthermore,
birds that breed at Fort Keogh, Montana, appear to be prefectly
typical. Birds from North Dakota, South Dakota, Nebraska,
northeastern Colorado, and extreme western Montana are
intermediate, in that there is among them more individual
variation, rendering some individuals difficult to distinguish
from the bird of the far western United States. They are,
however, as a whole, decidedly nearer the eastern race. Like-
wise, breeding birds from Ohio, Indiana, Illinois, Arkansas,
Missouri, and Iowa, as well as from Michigan and Wisconsin,
must be referred to the New England race. This, therefore,
obliterates the zonal distinction between the breeding ranges
of the two forms of this species, which Mr. Brewster* has
delineated.

One of the things brought out by the present study is the
great and very complicated individual color variation that
exists in both geographic races, which greatly increases the
difficulty of identifying individual specimens. The wing-bars
in the eastern bird are usually whitish, more or less tinged with
yellow or greenish, but occasionally these are decidedly brown-
ish, much resembling those of the western form of the species.
The lower parts are sometimes as yellowish below as in Empi-
douax I'irescens, but are usually more grayish or whitish.
Furthermore, the upper parts exhibit at least six more or less
well marked color phases, which, by individual variation, merge
imperceptibly into each other, so that there exists an infinite
variety of coloration. These phases, if such they may be called,
fall naturally into two series, separated in a general way by the
pattern of the upper surface. In one the color is almost
uniform from forehead to upper tail-coverts, and the type of
Muscicapa traillii Audubon is a bird of this kind; in the other
style the pileum and cervix are, in color, conspicuously different
from the back, being either more brownish or grayish and
forming a rather well defined cap. Birds of each one of these
styles exhibit three color phases: (1) dark greenish, which is
apparently the normal coloration, in which the upper parts are

*Auk, XIT, Xo. 2, April, 189."), p. l.J9.

88 The Ohio Journal of Science [Vol. XVIII, No. 3,

decidedly greenish with no yellowish tinge; (2) yellowish green,
which is very much less commonl}^ observed, a condition in
which the greenish of the upper surface is decidedly tinged with
yellowish, particularly on the interscapulum ; and (3) a grayish
phase, in which the olive upper surface is decidedly grayish
with no yellowish and but little greenish tinge. So different
are some specimens of these six phases that they look like
different species. One specimen, an adult from Fort Simpson,
Mackenzie, taken in July, 1861, is so brown that in color it
actually is not with certainty distinguishable from California
birds of this species; in fact, were it from a breeding locality
anywhere within the range of the Great Basin bird, one would
unhesitatingly refer it to that form. It is an example of the
difficulty which attends identification of some of the individuals
of this flycatcher, and shows that single individuals occur
which are practically impossible to assign subspecifically without
assistance from some other source than the color of the plumage.

Seasonal variation further complicates the case. By the
w^ear of the plumage some adults become, by July, very much
paler and more brownish above, as well as somew^hat so on the
lower parts; in which condition examples of the eastern and
northern race (Empidonax traillii traillii) are easily mistaken
for the form of the western United States, hereinafter described
as Empidonax traillii breusteri* A bird taken on June 25,
1894, at Canton, Illinois, is in this condition, and is practically
indistinguishable from worn specimens of the western race.
This specimen was doubtless partly responsible for Mr.
Brewster's reference of the Illinois birds to the western sub-
species. Since, however, breeding birds from all the surrounding
region now prove to belong to the eastern race, Empidonax
traillii traillii, this example is certainly but a w^orn, abnormally
brown individual of the same form.

Birds in juvenal plumage are also sometimes difficult to
decide subspecifically, because those of the eastern race are,
on the upper parts, much more brownish than the adults, and
thus often readily, at least superficially, to be mistaken for
western representatives.

Mr. Brewster's transferrence of the name Empidonax
traillii traillii to the western subspecies was made on the

"Postea, p. 12.

Jan., 1918] Status uf Empido mix Traillii 89

identification of the type specimen, now, as then, preserved in
the United States National Museum. Mr. Brewster mentions*
three specimens in the United States National Museum marked
as types of Muscicapa traillii Audubon; viz., Nos. UGO, 1865, and
2039. The first of these, No. 900, is now not to be found in
the collection, but it was entered as Tyranmila minima [Empi-
donax minimus] in Baird's own handwriting in the original
catalogue of his collection, which formed the beginning of the
National Museum bird register. Without much doubt, there-
fore, "No. 900" in Mr. Brewster's article is a typographical
error for No. 2960, which is still in the National Museum col-
lection, and is one of the two birds still marked with a red type
label. Of these two supposed types of Muscicapa traillii
Audubon, one is No. 1865, U. S. Nat. Mus., likewise the same
number of the original Baird collection ; and it is also one of the
three specimens mentioned by Mr. Brewster. This bird,
together with another, numbered 1866, were entered by Pro-
fessor Baird in his catalogue in January, 1845. On the label
of No. 1865 there is the following inscription in Baird's own
handwriting, the " (N) " indicating that the specimen was
received from Audubon :

Tyranmila trailli Aud.
186.5 rx)

This specimen, considered alone, is without any doubt
referable to the eastern race, and it agrees absolutely with a
specimen. No. 165385, U. S. Nat. Mus., from Mountain Lake
Park, Maryland, collected, July 3, 1899, by E. A. Preble; the
only observable difference, and this slight, being its rather more
yellowish posterior lower parts, very slightly duller wing-bars,
and partly darker, somew^hat more greenish upper surface.
It is in somewhat Avorn breeding plumage, but it is about as
near a match for this Maryland bird as is any bird in our series.
It measures as follows: wing, 73.5 mm.; tail, 58; exposed
culmen, 11.2; tarsus, 18; middle toe wdthout claw, 9. The
original locality of Audubon's Muscicapa traillii, as given by
him,t is the "Skirts of the woods along the prairie lands of the
Arkansas river." Mr. A. H. Howell has recently collected

*Auk, XII, No. 2, April, 1S9.5, p. 160.
tOrnith. Biog., I, 1831, p. 236.

90 The Ohio Journal of Science [Vol. XVIII, No. 3,

breeding birds at Chester and Stuttgart, in Arkansas, which
are for all practical purposes topotypes, and these belong
unquestionably to the eastern form. From the above facts it
is reasonably certain that Nos. 1865 and 1866 of the United
States National Museum catalogue are the original specimens
collected by Audubon near the Arkansas River, and that they
came into Baird's possession and were duly entered by him in his
catalogue. Since in size and color the one now marked No.
1865 in the United States National Museum collection is
found to agree substantially with Audubon's original descrip-
tion, it may reasonably and properly be considered the type.

The second specimen marked as one of the types in the
United States National Museum, and considered as such
by Mr. Brewster, is an adult and bears on its original label
in Baird's own handwriting the following inscription:

Tyrannula trailli? Aud
2960 n

This individual was evidently later received from Audubon
by Professor Baird, as it was not entered in the catalogue until
May, 1846, and then among a lot of other birds from the
Columbia River received from Mr. J. K. Townsend. It will be
noticed that the specimen was doubtfiilly identified by Professor
Baird as traillii; and it was, without any reasonable doubt,
one of the Columbia River examples obtained by Audubon from
J. K. Townsend after the species was described.! That the
Columbia River was its origin is further borne out by the fact
that it is not subspecificially the same as the specimen here
considered the type (No. 1865, U. S. Nat. Mus.), but is identical,
as it should be, with birds from western Washington. It is
apparently a female, though not sexed, and measures as follows:
wing, 66 mm.; tail, 53; exposed culmen, 10.3; tarsus, 16; middle
toe without claw, 9.5.

The third specimen mentioned by Mr. Brewster, No. 2039,
U. S. Nat. Mus., is, as he states, in juvenal plumage, and was
received by Professor Baird also from Audubon. It is, how-
ever, clearly an immature specimen of the Columbia River
bird.

jCf. Birds Amcr., I, octavo ed., 1840, p. 23.5.

Jan., 1918] Status of Em pidonax Traill ii 91

From the above facts it is very evident that the name
Em pidonax traillii traiUii belongs to the eastern race, and that
we must thus revert to the use of this name as it was before Mr.
Brewster transferred it to the western bird; and by this change
Empidonax traillii alnorum Brewster* now becomes a synonym.

Altogether 202 specimens of Empidonax traillii traillii,
from the following localities, have been examined in the present
connection :

Alaska.— Circle (July 12, 1903); Tatondu River, Yukon
River (August 6, 1899); Charlie Creek, Yukon River (June 21,
1903); Nulato.

Alberta. — Smith Landing (June 15, 1901); Fort McMurray
(August 11, 1903) ; 140 miles west of Edmonton (May 30, 1896).

British Columbia. — Vernon (June 5, 1895) ; Penticton (June
25, 1913); Hazelton (July 21, 1913); Okanagan Landing (June
18, 1915, August 18, 1915); Brackendale, Howe Sound (June
13 and 21, 191G, August 30, 1916).

Mackenzie. — Fort Resolution (June 30, 1865; June 22 and
24, 1903; June 19, 1903; July 10, 1903); Fort Simpson (July,
1861); Fort Rae (July 29, 1901); Nahannie River Mountains
(July 13, 1903); Fort Good Hope (June 22 and 23, 1904);
Hay River, Great Slave Lake (June 5, 6, 13, and 17, 1908);
Fort Norman (June 13, 1904; June 30, 1903; July 4 and 24,
1908); Fort McPherson (July 9, 1904).

New Brunswick. — Grand Manan Island (July 4, 1907).

Ontario.— Mount Forest (June 13 and 29, 1892).

Saskatchewan. — Indian Head (June 24, 1895).

Yukon.— Fort Selkirk (July 26, 1899).

Arkansas. — Chester (June 4, 1910); Stuttgart (May 13,
1910).

Colorado.— A valo (June 7, 1909,; Wray (May 21, 1909);
Lake Station, Lincoln County, (May 27, 1905) ; Clear Creek,
near Denver (June 4, 1911).

District of Columbia.— Washington (May 21, 1886; May 14,
1887; May 15, 1889; Sept. 9 and 17, 1890; August 31, 1890;
May 18, 1892; Sept. 13, 1894; August 20, 1896).

Idaho. — Near Fernan Lake (Sept. 1, 1913).

Illinois.— Warsaw (May 3, 1884); Urbana (May 11, 1908);
Areola (May 14, 1908); Richland County (May 21, 1883);
Sugar Creek Prairie, Richland County (June 5, 1885; June 6,

*Auk, XII, Xo. 2, April, 1S9.5, p. 161 (Upton, Maine).

92 The Ohio Journal of Science [Vol. XVIII, No. 3,

1890); Canton (June 25, 1894); Lee (Aug. S, 1907); Olney
(Sept. 10, 1908).

Indiana.— Wheatland (May 21, 1885).

Iowa. — Clay County (July 3, 4, 13, and 17, 1907; August
7, 9, 13, and 19, 1907); Palo Alto County (July 13, 1907).

Maine. — Sebec Lake (June 1, 1907); Upton (June 11, 12,
and 13, 1872); North Haven Island (June 16 and 19, 19(J7).

Maryland. — Laurel (May 16, 1888) ; Mountain Lake Park
(July 3 and 4, 1899).

Michigan.— Isle Royale (July 14, 1905; August 26 and 30,
1905); Porcupine Mountains (July 22, 1905).

Minnesota. — Tower (June 14, 1895).

Missouri. — Grandin (June 3, 1907).

Montana.— Fort Keogh (May 28 and 31, 1889; June 2, 3,
4, and 8, 1889) ; Little Missouri River (June 6, 1916) ; Blackfoot
Agency (June 13, 1895) ; Summit (June 19, 1895) ; Java (June
28, 1895).

New Hampshire. — Coos County (June 21 and 22, 1905).

New York.— Peterboro (May 21, 1886; May 21 and 30,
1887; June 2 and 3, 1887; July 19, 1886) ; Highland Falls (May
13, 1880; May 27, 1876); Kenwood (June 19, 1900).

North Dakota.— Lisbon (May 28, 1912); Pembina (June 3
and 9, 1873) ; Blackmer (June 5, 1915) ; Lidgerwood (June 18,
1915); Oakes (June 4, 1912; June 29, 1915); Kenmare (July
15, 1913); Dawson (July 21, 25, and 27, 1915; August 2, 1915);
Lostwood Lake (August 16, 1915).

Ohio.— Wooster (June 20, 1892).

Pennsylvania.— Carlisle (May 20, 1845; May 31, 1841);
Erie (May 21, 1894); Philadelphia (May 26, 1845).

South Dakota.— Smithville (May 31, 1894); Miner County
(June 30, 1892) ; Evarts (August 6, 1903) ; Forestburg (August
10, 1906).

Texas. — Brewster County (May 8, 1913); Runge (August
24, 1905); Brownsville (August 27 and 28, 1912); Columbus
(Sept. 4, 1905) ; Victoria (Sept. 12, 1906).

Virginia.— Four Mile Run (Sept. 2, 1894; May 10, 1894).

Wisconsin.— Madison (July 24, 1893).

Costa Rica.— Santa Maria de Dota (Feb. 29, 1866; May 11,
1908).

Nicaragua.— San Carlos (Feb. 25, 1892).

Oaxaca.— Huilotepec (May 12, 1895).

Jan., 191S] Status of Em pidouax Traillii 93

Empidonax traillii brewsteri, subsp. nov.*

Em pidouax traillii pusillus AucT. (nee Swainson).

Empidouax traillii traillii Brewster, Auk, XII, No. 2,
April, 1S95, p. 159 (nee Audubon).

Chars, subsp. — Similar to Empidonax traillii traillii, but bill
larger, wing-bars darker, duller, much more brownish (less
yellowish or greenish) ; upper parts decidedly more brownish or
less greenish.

Description.— Type, adult male. No. 158:^^60, U. S. Nat.
Mus., Biological Survey collection, Cloverdale, Nye County,
Nevada, May 31, 1898; Harry C. Oberholser; original number,
57. Pileum brownish olive, the centers of the feathers olive
brown; the remainder of the upper parts between brownish
olive and light brownish olive, but verging slightly toward
olive, and paling on the shorter upper tail-coverts to light
brownish olive; longest upper tail-coverts brownish olive; tail
fuscous, margined externally on the basal portion with brownish
olive; wings fuscous, the narrow external margins of tertials
and secondaries tilleul buff, the margins of the lesser and
median coverts brownish olive, the tips of the greater coverts
between avellaneous and light drab; lores yellowish white, tips
of the feathers olive brown ; the sides of head and of neck rather
light citrine drab, shading to buffy; sides of breast the same,
but somewhat darker; chin and throat dull brownish white;
jugulum and upper breast between grayish olive and deep
olive buff; lower breast and abdomen naphthalene yellow;
lower tail-coverts colonial buff; sides of body naphthalene yellow
with an olivaceous tinge; thighs Saccardo's olive; edge of wing
below, chamois; and lining of wing naphthalene yellow.

Measurements. — Type: wing, 69 mm.; tail, 60; exposed
culmen, 12.5; tarsus, 16.5; middle toe without claw, 9.7.

Male:t Wing, 69-73 (average 70.6) mm.; tail, 56-60.5
(58.1); exposed culmen, 12.5-13.5 (13.0); tarsus, 15.5-17.5
(16.8); middle toe without claw, 9-10.5 (9.6).

Female :| Wing, 63-68.5 (65.7) mm.; tail, 52-60 (56.0);
exposed culmen, 11.5-13 (12.2), tarsus, 15.5-17 (16.3) ; middle
toe without claw, 9-10 (9.4).

*Namcd for Mr. William Brewster.

tThirteen specimens, from Oregon, California, Arizona, New Mexico, and
Colorado.

JTen specimens, from Oregon, California, Arizona, and Utah.

94 The Ohio Journal of Science [Vol. XVIII, No. 3,

Type locality. — Cloverdale, Nye County, Nevada.

Geographic distribution. — Western United States and extreme
southwestern British Columbia, south to Mexico, Central
America, and northwestern South America. Breeds chiefly
in the Lower Austral, Upper Austral, and Transition zones of
North America and Mexico; north to central Wyoming, central
Idaho, northern Washington, and southwestern British Colum-
bia; west to southwestern British Columbia, western Washington,
western Oregon,^ and western California; south to the extreme
northern edge of Lower California, northern Sonora, Durango,
southern New Mexico, and central Texas; and east to north-
eastern Texas, eastern Oklahoma, southeastern and central
Colorado. Winters from northern Colombia to Panama,
Costa Rica, and Nicaragua. Migrates east to eastern Nebraska,
Tamaulipas, and Oaxaca; west to Sinaloa and Guerrero.

Remarks. — There is surprisingly little geographic variation
within the range of this race, but birds from northern Wash-
ington and extreme southwestern British Columbia are some-
what intermediate, though not sufficiently so to cause doubt
about their proper reference to this form. Birds from eastern
Colorado and Texas likewise verge somewhat toward Empidonax
trail Hi trailUi.

There is, as in Empidonax traillii traiUii, a large amount of
individual variation in the present subspecies, including counter-
parts of the same six more or less well-defined color phases.
The two patterns of coloration on the upper parts are even
better indicated. In one of them the upper surface is nearly
or quite uniform; in the other the pileum and cervix are m^uch
more grayish or brownish, and distinctly, even trenchantly,
different from the back. The type of Empidonax traillii
brewsteri is of the nearly uniform style. In specimens of each
of these two patterns there are three definable phases of colora-
tion. The upper parts may in color be either (1) normally
dark and brownish; (2) yellowish brown on all but the pileum; or
(3) distinctly gray. The yellowish brown phase is not so
greenish as the corresponding plumage in Empidonax traillii
traillii, and the gray phase is very much more purely grayish,
less greenish. Furthermore, the lower parts are sometimes very
strongly tinged with yellowish, particularly on the posterior
portion; in other specimens this yellowish tinge is scarcely
noticeable, except on the crissum.

Jan., 191SJ Status of Empidonax TraiUii 95

Seasonal variation does not seem to be quite so strongly-
marked as in Empidonax traillii traillii, though summer birds
are more brownish (less olivaceous) or more grayish, and paler
than in autumn, spring or winter.

Birds in juvenal plumage, compared with the same stage of
Empidonax traillii traillii, are usually lighter on the breast and
more yellowish on the posterior lower parts, but some speci-
mens are very difficult to distinguish.

The distinction between Empidonax traillii traillii and
Empidonax traillii brewsteri has, of course, long been known.
The name Empidonax pusillus (Swainson) was, until Mr.
Brewster's investigations,* used for the latter, during which
period the name Empidonax traillii or Empidonax pusillus
traillii (Audubon) was employed for the eastern bird. The
name Empidonax pusillus was taken from Platyrhynchus
pusillus of Swainson,! which was described in the following
language :

"Olive brown, beneath yellowish white; wings with two pale
bands; tail moderate, even; bill small; head crested.

Maritime parts of Mexico.

Total length, 5^; bill, Ym; wings, 2>^; tail, 23^."

Swainson subsequently identified with this bird a specimen
killed at Carlton House, Saskatchewan, and described and
figured by him as Tyrannula pusillaX, which is, as already stated
by Mr. Brewster, very probably not this species at all, but
Empidonax mifiimus. Even if the Tyrannula pusilla of Swain-
son be correctly identified as Platyrhynchus pits ill us from Mex-
ico, this does not aid in the solution of the identity of the latter,
for the availability of the name must in this case be determined
from the original description or from an examination of the
type. As the above quotation shows, the original description
of Platyrhynchus pusillus Swainson, while pointing toward the
bird we have here described as Empidonax traillii brewsteri,
is not certainly identifiable as such. Professor Alfred Newton
made careful search among the Sw^ainson types still remaining
in the collection of the mmseum at Cambridge, England, but the
type of Platyrhynchus pusillus is not there; nor apparently is it
ever likely to turn up elsewhere. Therefore, as has already

*Auk, XII, Xo. 2, April, 1895, pp. 159-163.

tPhilosophical Magazine, new series, I, No. V, May, 1827, p. 360.

tFauna Bor.-Amcr., II, 1831 (1832), p. 144, pi. XLVI, upper figure.

96 The Ohio Journal of Science [Vol. XVIII, No. 3,

been so well and fully stated by Mr. Brewster,* we are apparently
justified in rejecting this name as unidentifiable. This leaves
the western bird, the Empidonax pusillus of authors and the
Empidonax traiUii traillii of Brewster, without a name, and we
take pleasure in bestowing upon it herewith the name Empidonax
trail! ii brewsteri, in recognition of Mr. Brewster's careful work
on these very difficult flycatchers.

Two hundred and nine specimens of this subspecies have been
examined, from the localities that follow:

British Columbia.— Victoria (June 4, 1892; June 13, 1894);
Comox (July 11, 1895); Gibson Landing, Howe Sound (July
28, 1897).

Arizona. — Nogales (May 23, 1893) ; Santa Cruz River,
Mexican Boundary Line (June 2 and 8, 1893) ; Alpine (August
1, 1915); Fort Whipple (August 10 and 15, 1864); Clifton
(August 19, 1914); Lees Ferry (August 25, 1909); San Ber-
nardino Ranch, Mexican Boundary Line (August 17, 1893;
August 21 and 31, 1892; Sept. 5, 1892); San Francisco River
(August 21, 1914); Pinal County (Sept. 3, 1883).

California.— Stockton (May 11, 1878; July, 1877); Mountain
Spring, Mexican Boundary Line, San Diego County (May 13,
1894); Fort Mojave (May 21, 1861); Jacumba, San Diego
County (May 23, 1894); Saticoy (May 24, 1873); Brawley

(June 4, 1907) ; Kern County (June 8, ) ; Owen Lake (June

9 and 12, 1891); Lone Pine (June 11, 1891); Hornbrook (June
15 and 16, 1897); Tulare Lake (June 20, 1907); Santa Barbara
(June 25, 1875) ; San Bernardino (June 26, 1910) ; Kern River
Lake (July 21, 1904, three nestlings); Donner Lake (July 25,
1900); Grafton (July 28, 1906); mountains near Camp Bidwell
(July 31, 1878) ; Grizzly (August 11, 1900); Marshalls (August
18, 1905); San Clemente Island (August 23, 1894); Fresno
Flat (August 30, 1904); Fort Tejon (August, 1875; Sept. 11,
1857) ; Bear Valley (1880).

Colorado.— AVray (May 22, 1911); Fort Garland (May 28,
1873); Arvada, Jefferson County (June 12, 1904); Loveland
(June 26, 1891; July 26, 1895) ; Grand Junction (June 28, 1893) ;
Colorado Springs (July 1 and 3, 1882) ; Breckenridge (July 2
and 8, 1877) ; Pueblo (July 23, 1874) ; Clear Creek, near Denver
(August 6, 1911); Olivet, Jefferson County (August 8, 1911).

*Auk, XII, No. 2, April, 1895, pp. 100-161.

Jan., 1918] Status of Empidonax Traillii 97

Idaho. — American Falls (June o, 1911); Weiser (June 18,
1913); Birch Creek (July 22, 1890; August 4 and 15, 1890);
Shellev ( Tulv 28, 1911); Van Wyck (August 10, 1913).

Nebraska.— Columbus (Sept. 1, 1892).

Nevada. — Pahranagat Valley (May 23, 1891); Glcnbrook
(May 30, 1889); Cloverdale (May 30 and 31, 1898); Mountain
City (June 12, 1898); Winnemucca Lake (June 19, 1889);
Ruby Valley (August 29, 18GS) ; East Humboldt Mountains
(Sept. 8, 1868).

New Mexico. — Las Vegas (May 23, 1900) ; Santa Rosa
(May 20, 1903); Rinconada (June 1, 1904); Magdalena Moun-
tains (August 30, 1909).

Oklahoma. — Dougherty (August IG, 1892) ; Hartshorn (Aug.
25, 1892).

Oregon. — Homestead (May 31, 1916; June 1, 1916); Salem
(June 4, 1895); head of Whiskey Creek (June 14, 1897); head
of Drew's Creek (June 17, 1897) ; Forest Grove (June 20 and 22,
1903) ; Portland (June 23 and 24, 1897) ; Fort Klamath (June
26 and 28, 1882; July 23 and 28, 1883); Jordan Valley (June
28, 1915); Looking-glass (June 29, 1916); Beulah (June 29,
1916) ; Beaverton (July 7, 1890) ; Reston (July 8, 1916) ; Juntura
(July 8, 1916); Astoria (July 15, 1897); Rockville (July 16,
1915); Riverside (July 20, 1916); Shirk (July 22, 1896); Drew
(July 28, 1916); Diamond (July 29 and 31, 1916); Telocaset
(August 6, 1915); La Pine (August 11, 1914); Anchor (August
12, 1916); McEwen (August 20, 1915).

Texas.— San Angelo (April 8, 1885); Marathon (May 14,
16, 17, and 19, 1901) ; Brewster County (May 19, 1913) ; Tascosa
(May 21, 1894; June 6, 1899); Laredo (May 23, 1866); mouth
of Tornillo Creek (May 24, 1901); Fort Hancock (June 18,
1893); Boston (July 4, 1902); upper Devil's River (July 21,
1902); Texline (August 1, 1903); Pecos River, 55 miles north-
west of Comstock (August 2 and 4, 1902) ; Pecos City (August
28, 1902); near Santa Rosa, Cameron County (Sept. 20, 1891).

Utah.— Mouth of Bear River (May 26 and 28, 1915);
June 7, 1916; August 20, 1915); Antelope Island, Great Salt
Lake (June 4, 1869); Ogden (June 15, 1872); Provo River
(July 25 and 26, 1872) ; Parley's Park, -Wasatch Mountains
(July 28, 1869); Provo (July 26, 1872; August 2 and 3, 1872);
Clear Creek (August 9, 1908).

98 The Ohio Journal of Science [Vol. XVIII, No. 3,

Washington. — Fort Bellingham (May 27, 1905) ; Neah Bay
(June 10, 1897); Prescott (June 18, 1908); Mount Vernon
(June 24, 1897) ; Tenino (June 29, 1897; July 1, 1897) ; Granville
(July 9, 1897); Steilacoom (August 1, 1856); Ellenburg (August
13, 1897); Tacoma (August 20, 1897).

Wyoming.— Stanley (August 23, 1911).

Costa Rica. — San Jose? (1873).

Durango. — Rio Nazas (June, 1853).

Guerrero. — Dos Arroyos (May 4, 1903).

Lower California. — Mount Mayor (May 24, 1915) ; Nacho-
guero Valley (June 1. 1894).

Michoacan.— La Salada (March 18, 1903).

Morelos. — Cuernavaca (April 9, 1908).

Nicaragua.— Sucuya (Feb. 7, 1883); Greytown (Feb. 8,
1892).

Oaxaca. — Tehuantepec (May 20, 1895); Huilotepec (May
11, 1895).

Panama. — Canal Zone (Jan. 14, 1912); Gatun, Canal Zone
(March G, 1911; May 8 and 15, 1911); Tabernilla (April 18,
1911); Porto Bello (May 26, 1911); Cana (May 23, 1912).

Sinaloa. — Mazatlan.

Sonora. — Nogales (June, 1855).

Tamaulipas. — Jaumave (June 2, 1898).

ADDITIONS TO THE CATALOG OF OHIO VASCULAR

PLANTS FOR 1917.

John H. ScHAFFx\iiR.

6G. Tsuga canadensis (L.) Carr. Hemlock. Bainbridge, Ross
Co. E. N. Transeau.

71. Pinus echinata Mill. Shortleaf Pine. Bartles, Lawrence

Co. A. E. Taylor; Ironton, Lawrence, Co. Lillian E.
Humphrey; Marietta, Washington Co. Forest W. Dean.

72. Thuja occidentalis L. Arborvitas. Bainbridge, Ross Co.

E. N. Transeau.

74. Juniperus sibirica Burgs. Low Juniper. Two specimens
growing on "Fort Hill," Berea, Cuyahoga Co. E. L.
Fullmer.

76. Taxus canadensis Marsh. American Yew. Under arbor-
vitae and hemlock trees. Bainbridge, Ross Co. E. N.
Transeau.

250.1. Carex swanii (Fern.) Mack. Swan's Sedge. In woods.
Sunbury, Delaware Co. Frank T. McFarland.

321.1. Uniola latifolia Mx. Broadleaf Spike-grass. Bain-
bridge, Ross Co. E. N. Transeau.

365. Change name of Triticum vulgare L. to Triticum
aestivum L.

452. Syntherisma ischaemum (Schreb.) Nash. Small Crab-
grass. GallipoHs, Galha Co. John H. Schaffner.

874. Viola lanceolata L. Lanceleaf Violet. Ironton, Lawrence
Co. Lillian E. Humphrey.

1214. Ampelopsis cordata Mx. Heartleaf Ampelopsis. Galli-
poHs, Gallia Co. John H. Schaffner.

1323. Change name from Opuntia humifusa Raf. Western
Prickly-pear, to Opuntia opuntia (L.) Coult. Common
Prickly-pear.

1400. Andromeda polifolia L. Wild Rosemary. Akron swamps,
Summit Co. E. L. Fullmer.

1433.1. Convolvulus repens L. Trailing Bindweed. On steep
bank. Hillsboro, Highland Co. Katie M. Roads.

1471. Frasera carolinensis Walt. American Columbo. Barnes-
ville, Belmont Co. Emma E. Laughlin.

99

100 The Ohio Journal of Science [Vol. XVIII, No. 3,

1571. Cymbalaria cymbalaria (L.) Wettst. Kenil worth Ivy.

Berea, Cuyahoga Co. E. L. Fullmer.

2046. Lactuca sagittifolia Ell. Arrowleaf Lettuce. Parma

Twp., Cuyahoga Co. E. L. Fullmer.

Additions made to the Cedar Point flora during the summers
of 1914-1916, by E. L. Fullmer.

Sagittaria brevirosta Mack and Bush. July 6, 1914.
Salsola pestifer A. Nelson. July 27, 1915.
Cerastium velutinum Raf. June 21, 1916.
Agrostemma githago L. June 22, 1916.
Camelina microcarpa Andrz. June 22, 1916.
Thlaspi arvense L. June 25, 1914.
Cucumis melo L. June 8, 1914.
Hypochaeris radicata L. July 23, 1916.
Tragopogon pratensis L. July 27, 1915.
Sonchus arvensis L. July 27, 1916.

Date of Publication, January 31, 1918.

THE

Ohio Journal of Science

PUBMSHED BY THE

Ohio Statk Univkksity Scientific Society

Volume XVIII FEBRUARY, 1918 No. 4

TABLE OF CONTENTS

SCHAFFNiCR — The Expression of Sexual Dimorphism in Heterosporous

Sporophytes 101

Walton— Eutetramorus Glohosus, a New Genus and Species of Alg^e

Belonging to the Protococcoidea (Family Crelhstiidte i 126

Pott — A Study of Cattle "Temperament" and its Measurement 129

THE EXPRESSION OF SEXUAL DIMORPHISM IN
HETEROSPOROUS SPOROPHYTES.*

John H. Schaffner.

The writer has been engaged for a number of years in
studying the evolutionary development of the higher plants.
In the study of the flower, the problems of sexuality and of
sporophyte dimorphism have presented themselves in an
unusual manner, w4th many hints as to the nature of their
hereditar}^ expression.

The most remarkable dimorphism in plants is, of course, the
difference of form and function expressed between gametophyte
and sporophyte. One can hardly conceive of a more funda-
mental difference than exists, for instance, between the two
generations of the ordinary ferns. In the higher plants this
dimorphism is as extensive as sexuality itself. Sexual di-
morphism seems more prominent to us only because we con-
stantly associate our ideas with the sexual dimorphism of the
higher animals.

Aside from the dimorphisms mentioned above there are also
very striking vegetative dimorphisms which have no relation
to sexuality or the alternation of generations. Such vegetative
dimorphisms are usually expressed in different parts of the same
individual. In plants like Megalodonta beckii (Torr.) Greene
and Neobeckia aquatica (Eat.) Britt., the dimorphism is

♦Contribution from the Botanical Laboratory, Ohio State University. No. 99.

102 The Ohio Journal of Science [Vol. XVIII, No. 4,

related to the environment, the submerged leaves being dis-
sected and the aerial leaves of the ordinary broad type, char-
acteristic of closely related genera and species. In Lepidium
perfoliatum L. there is just as abrupt and striking a change
at the middle of the stem although there is no change in the
surrounding medium. The lower leaves are finely dissected,
while the upper are entire, oval in shape, and deeply clasping.

These dimorphisms are changes of states or expressions of
hereditary units in the common tissue of a growing shoot.
It is plain that they have to do with activity and latency rather
than with the presence and absence or the shifting of hereditary
units. This fact is to be kept in mind further on in the dis-
cussion of analogous sexual phenomena. Similar dimorphisms,
although usually not so abrupt, are, of course, very general
in great numbers of rosette plants. There are also cases where
there is an abrupt, new development of shape, structure, and
often color in the leaves just below the inflorescence. This type
of vegetative dimorphism is especially common in the Euphor-
biace^.

Among the fungi and many other organisms another type
of dimorphism is common, which often involves not only the
reproductive cells, but also the surrounding hyphae. In the
Ascomycetae there is generally a decided difference between
the conidial and ascus stages. These differences appear in
different branches of an individual mycelium. The difference
in sexual and nonsexual reproductive parts of such fungi
corresponds to the vegetative dimorphisms mentioned above in
that the phenomena are not associated with shiftings of the
chromosomes. In some cases they are produced at rather
definite stages of the life cycle, in others the change of expression
depends on the environment.

Now what is meant by sexuality? In its simplest form it
is a ph^^siological difference expressed only in the developing
sexual cells. As we go up the scale of plant complexity, we note
that more and more of the tissues are involved until finally
in the extreme forms the entire organism seems to be affected.
From the evidence to be presented it must become apparent
that in at least the vast majority of cases, and probably in all,
the sexual condition is simply a state of the living substance
which may continue for a greater or less length of time before
a neutral state or the opposite sex condition is set up.

Feb., 1918] Sexual Diniorp/iisni 103

Physiological states may be due to chemical or other causes.
Among the conditions in which organisms may be growing or
living, the following may be mentioned : An active or quiescent
state; vigorous or exhausted; sterile or fertile; immunity to
disease at a certain age or susceptibility; specific diseased con-
dition, like cretinism in children, or normal; gametophyte
or sporophyte (without change of chromosome number) ;
female or male; carpellate or staminate; juvenile state or mature
state (like certain Acacias and Junipers) ; water form or air
form; root state or stem state (usually with change of environ-
ment); conidial stage or sexual; protonema or scaly moss
plant; difference of unusual morphological expressions without
any hereditary difference, as for example, two or more very
different types of insect galls on the leaves of celtis occidentalis
L. In the case of the transition from gametophyte to sporo-
phyte and vice versa, there is usually a change in the chromo-
some number which might be assumed to be the cause of the
remarkable change in morphological expression, but there are
many examples in both mosses and ferns, and also in the
flowering plants, where the change takes place in the vegetative
tissues without a shifting of chromosomes. The study of
apogamy and apospory should shed much light on the nature
of sexual and nonsexual states.

Before presenting the evidence in hand on the development
of dimorphism in the heterosporous sporophytes a few general
facts may be given in relation to sex as expressed in the game-
tophyte. The term sex used in its strict sense applies both
to diploid and haploid individuals; among animals to diploid
and among plants more commonly to haploid individuals,
although there are numerous algae in which the life cycle is
essentially similar to that of the animals.

As already stated, there is a gradual development from a
condition where no difference is apparent between the con-
jugating cells except the mutual attraction at a certain period
to highly specialized cells, and further through hermaphordite
organisms with strikingly different sexual organs on different
parts of the same body up to completely unisexual individuals.
In the past it has been assumed that sexualit}^ was evolved
because of some fundamental utility to the organism, but the
doctrine of specific utility as an explanation of origins can have
little place in the theory of any investigator who has carefully

104 The Ohio Journal of Science [Vol. XVIII, No. 4,

studied the facts of plant evolution. The utilitarian view as
developed to a supreme position in the Darwinian theory
is largely discordant with present day views, but still seems to
have a tenacious hold on the philosophy of sex. Things
may be useful but the cause of their origin and persistence is
another question. One can find endless cases where exactly
opposite developments occur side by side in closely related
species and varieties subject to a common environment. For
example, note the barbed involucral bristles of Chaetochloa.
In Chaetochloa verticillata (L.), Scrib. the barbs are retrorse
while in the closely related Chaetochloa viridis (L.) Scrib.
they point outward. In neither case have they any relation to
seed distribution since the bristles remain on the dead inflores-
cence when the spikelets fall. In the genus Bidens, the papus
awns are usually barbed, but here also some species have
retrorse barbs and some outwardly projecting barbs. Since
the awns are persistent on the fruit the retrorse barbs become
useful but the others, if they have any effect must rather hinder
seed distribution. In some species of Bidens the awns have
no barbs whatever. Any number of similar cases could be
cited.

Sex appears in some way to be associated with physiological
and chemical states of the living protoplasm. It is perhaps
most reasonable to assume, at present, that a certain organi-
zation or complexity of the cell is necessary before sexual states
originate. But it is not true on the other hand that these
states are necessarily set up at any stage of the life history
even in organisms that have the essential complexity.

Among plant gametophytes the greatest sexual difference
is shown in the Spermatophyta, between the male and the
female. In the Archegoniata, very striking examples are
certain species of Polytrichum where the mature male plant
has a very different appearance in form and color from the
female. The difference is much greater than is exhibited
by many mammals or even birds. Now, just as in the game-
tophyte generation we find no vegetative dimorphism in the
lower forms but find this becoming more and more pronounced
as we ascend the scale, so also in the evolution of the sporophyte
it is only the extreme forms in the evolutionary series that
show but the one state throughout the entire individual. It is
evident also that the gametophyte is far advanced in evolution

Feb., 1918] Sexual Dimorphism 105

before the appearance of heterosporous sporophytes and that
it shows complete sexual segregation from the Heterosporous
Pteridophytes on up, while as stated, comparatively few
sporophytes have attained complete individual dimorphism.

The gametophytes of Homosporous Pteridophytes exhibit
both hermaphrodite and unisexual species, and in the case of
certain unisexual ferns and horsetails, various experimenters
have shown that the sex can be controlled or reversed after it
is determined. The reversal can be brought about in either
direction, but generally a much larger per cent, of females
can be induced to produce male reproductive organs than
males can be changed to females. This is probably due to
the more specialized nature of the male soma, especially in
relation to nutrition and vegetative growth. It is well to call
attention here again to the fact that when the higher hetero-
sporous plants developed definitely unisexual, gametophyte
individuals, the segregation of the sexes was not accomplished
in or by the reduction division, as one might have expected, but
entirely independently of that process. All the four spores of a
reduction division have the same sexual tendencies and all give
rise either to male or to female gametophytes. Here the
development of males and females is associated with the size
of the spores. It is also interesting to note that in animals
the spermatoc^^tes through reduction give rise to four spermatids
which develop into spermatozoa without any reference as to
whether they contain an allosome or not. If the allosome is
what determines sex, how is it that the two spermatids without
allosomes develop maleness in exactly the same extreme way
as the two which contain the allosome? Certainly the peculiar
morphology of the spermatozoon must be considered maleness,
if there is such a thing. The same reasoning would apply to
the egg cell. In the case of the honey bee, all the oocytes
develop eggs, which we certainly must admit is a female expres-
sion; yet these same eggs without fertilization change their
sexual expression and develop into male individuals. In
animals we cannot test out the hereditary qualities of the
spermatozoa independently as we can the microspores of plants.
If we could we would probably obtain results analogous to those
coming from the androspores of certain species of Oedogonium,
which give rise to dwarf males. In the higher plants than
reduction of chromosomes has nothing to do with the maleness

106 The Ohio Journal of Science [Vol. XVIII, No. 4,

and femaleness of the gametophyte. The determination of the
sex takes place in the vegetative tissues of the sporophyte.
Whatever it is that determines that the given tissue shall
develop as a megasporophyll or a microsporophyll also deter-
mines absolutely the sex of the following generation of game-
tophytes. We are dealing with the establishment of a state
in the cells of a vegetative tissue. A moment's reflection
will show how extremely inappropriate is the application of the
terms, homozygous and heterozygous in relation to sex indivi-
duals. Sexuality is just as pronounced in haploid gameto-
phytes as elsewhere. The males, females and hermaphrodites
of the higher plants can not be "zygous" at all in the normal
life cycle. They are the results of segregation rather than of
conjugation. The vast majority of sporophytes are bispor-
angiate and are of course homozygous or heterozygous in their
chromosome condition, but here the sex is determined in the
vegetative tissue before spores are produced.

In some species of Bryophytes it has been claimed that two
of the cells of the reduction tetrad normally give rise to males
and the other two to females. In such cases the sex must be
determined in the spore mother cells (daughter cells of the
sporocyte). But a sexual state might be influenced by unequal
distribution of the cytoplasm. In great numbers of Bryophytes,
however, the gametophyte is hermaphrodite and the establish-
ment of the sexual state necessarily takes place in the vegetative
cells of the gametophyte. Recently Allen* has reported a
difference in size in one pair of chromosomes in the spore mother
cells of Sphaerocarpos. Even if such a difference can be
associated with a specific sex difference, it does not necessarily
follow that the sex condition is determined by an irreversible
sex factor in the given chromosomes. However, in this case
where the sex individuals are haploid, such a hypothesis would
be much more convincing than in the case of the diploid animals.

In dealing with sexual phenomena in plants, the problem is
always complicated by the presence of an antithetic alternation
of generations. Because of a confusion in terminology coming
from a past period w^hen the nature of the plant life cycle was
unknown, it is often difficult to correlate the meaning of expres-
sions used by different writers, especially of those who disregard

*Allen, Chas. E. A Chromosome Difference Correlated with Sex Difference
in Sphaerocarpos. Science 46 : 466-467. 1917.

Feb., 1918] Sexual Dimorphism 107

modern morphological discoveries. We cannot hope to analyze
hereditary phenomena unless we have a correct understanding
of the morphology and physiology of the parts under considera-
tion. In recent years a consistent terminology has been
developed by morphologists and there is no excuse for not
using it. We have a sporophyte terminology and a game-
tophyte terminology. However, when applied to the new
science of genetics out language is still inadequate, and one is
immediately confronted with the difficulty of expressing
hereditary phenomena in relation to sex when the sporophyte
of heterosporous plants is involved. The homosporous sporo-
phyte having no sexual dimorphism does not raise the issue,
but when passing from homosporous to heterosporous spor-
ophytes the problem presents itself with the common solution
that organs formerly described and defined as without sex
now obtain a radically different treatment, to the confusion
of both the learned and the unlearned,. It seems to the writer
that the way out of the difficulty, at present, is to employ the
sporophyte terminology when structures are mentioned, and
to limit the sexual terminology, as far as possible, to the states
or conditions of special gametophyte phenomena expressed
in the sporophyte. Thus a spore bearing fern leaf is a non-
sexual structure and a sporophyll of Marsilea is still nonsexual
although it produces spores of two sizes. But the difference
between the spores is a sexual difference. A carpel is still a
megasporophyll and a nonsexual reproductive organ of the
same fundamental nature as the megasporophyll of a hetero-
sporous pteridophyte, but it is a sporophyll in which the sexual
phenomena peculiar to the gametophyte are finding expression.
The sexual state is thrown back, so to speak, into a small part
of the sporophyte. It is this spreading of the sexual state,
with an ever increasing area of the tissue involved, that con-
stitutes one of the most interesting aspects of sporophyte
evolution in the higher plants.

So far as the writer knows, Marsilea is the lowest living
genus of heterosporous plants in respect to sexual dimorphism
of the sporophyte. In external aspect the sporophyte shows
no dimorphism, the sporocarps all being alike (Fig. 1). But
the sporangia are slightly different, although the same in general
shape. The most striking difference is in the stalks. The stalk
of the microsporangium is comparatively long and slender.

108 The Ohio Journal of Science [Vol. XVIII, No. 4,

while that of the megasporangium is short and thick. (Figs.
2 and 3). The dimorphism appears very pronounced in the
spores. There is a remarkable difference between the micro-
spores and megaspores in shape and size. There is here also the
difference in spore development, in that the cells from the micro-
spore tetrad all develop while only one of the cells of the mega-
spore tetrad develops. There is only one spore in the megaspor-
angium in marked contrast to the large number in the micro-
sporangium. Here then is a beginning of sexual dimorphism
in the sporophyte. Now how is this difference in morphologic
expression in the sporophyte tissue -coming about? It is not
by any shifting of chromosomes evidently; the tissues have a
common vegetative origin side by side. The two kinds of
sporangia do not even occupy different regions of the modified
leaflet which bears them. It is the setting up of a differential
condition of the same general nature as that which determines
a dimorphic expression in the hard tissue of the sporocarp
and the membranous tissue of the leaf blade on the same
petiole.

Shattuckf found both in the normal plants and in all the cul-
tures that a close examination revealed a homosporous tendency
in that microspores were sometimes formed in the megasporan-
gia, especially in those most distant from the nutritive supply.
He also found that in cases of abortion in the microsporangia
only one spore survives which is about sixteen times as large as
the normal microspore. Shattuck discovered a method by
which the nature of spore development in Marsilea can be con-
trolled to a certain degree.

In the second stage of dimorphic advancement, as is well
illustrated by Selaginella kraussiana (Kunze), the strobili
are bisporangiate and each type of sporangium is confined to
a separate leaf. There are definite microsporophylls and
megasporophylls. In this species of Selaginella there is only
one megasporophyll, at the base of the cone, to about twenty
microsporophylls. The tissues which develop the two types
of sporophylls are definitely located. Dimorphism of spores
and sporangia is present as in Marsilea, but there are usually
four megaspores. When we examine the blades of the sporo-
phylls we also find a slight difference. They are both green and

tShattuck, C. H. The Origin of Heterospory in Marsilea. Bot. Gaz. 49 :
19-40. 1910.

Feb., 1918] Sexual Dimorphism 109

of the same texture, but somewhat different in shape and size.
(Figs. 4 and 5). Here the dimorphic state has spread far
beyond the spores, and sporangia, until it affects the differ-
entiation over a considerable area. The microsporangiate
and megasporangiate states are set up in the incepts of the
sporophylls. In the one case we have male determination,
in the other female. Again, how is this determination brought
about? By the activity and latency, whether only partial or
complete, of hereditary factors. The sexual differentiation
takes place before and outside of the phenomena of the reduction
division and fertilization. It has absolutely nothing to do with
segregation or association of chromosomes or allosomes.

In the gymnosperms generally the dimorphism is expressed
not only in a difference in shape, size and structure of the
sporophylls, but usually also in a difference in color; and since
the cones are usually monosporangiate in the living species, the
dimorphism extends to the floral axis or even beyond. But
as all of these advancing evolutionary stages are also repre-
sented in the Anthophyta, no special consideration will be
given here to the sexual dimorphisms of the gymnosperms.

When we come to the lowest Anthophyta, as in various
genera of the Ranales, there is a considerable advance over the
condition in Selaginella and related plants. Figures 6 and 7
represent a stamen and a carpel of Aquilegia canadensis L.
The dimorphism appears in the shape, size and color of the
sporophylls and in addition the megasporophyll shows that
remarkable secondary sexual character, the stigma, which
becomes necessary here because of the closed condition of the
carpellate blade. The ovulary is also covered with prominent
hairs while the stamen is smooth, a sex-limited character. In
this case the sexual dimorphism expressed in parts of the
sporophyte is about as great in character and degree as is usual
for secondary sexual differentiations either of plants or animals.
But the difference is confined to the sporophylls. The spor-
ophyte as an individual shows but one form and nature and the
dimorphism is developed in closely associated organs arising
from a common tissue. The condition as represented by the
flowers of Aquilegia canadensis L. is the normal state for the
flowering plants. Indeed, if defined in general terms, the
Anthophyta are plants with bisporangiate flowers with here
and there groups or individual species which have advanced in

110

The Ohio Journal of Science [Vol. XVIII, No. 4,

Feb., 1918] Sexual Dimorphism 111

specialization to a greater or less degree toward the monecious or
diecious state. In tracing from the bisporangiate to the
monosporangiate flowers one finds a most remarkable display
of vestigial structures, which are the result of suppression of the
opposite hereditary factors by the distinctive sexual condition
set up in the tissues from which they should develop. This
inhibitory influence is of every degree of intensity in various
species, ranging from cases where the organ is almost normal
to its entire disappearance. In the more primitive, or rather
less specialized species, reversions are very common.

The strictly diecious state is comparatively rare. There are
no original monosporangiate flowers among the Anthophyta.
The few examples, in which all vestige of a bisporangiate con-
dition has disappeared, in nearly every case show a direct
relationship to species or groups with the opposite structures
present either as vestiges or in a normal condition. A few
examples will be given to show the general trend of development
to a typical diecious condition. It is well, however, to caution
against the notion that the monecious condition is a step in the
evolution of the diecious. It may be in some cases but mostly
the development of diecious plants comes about directly
through a succession of more extreme vestiges.

Sagittaria latifolia Willd., a low species of the Helobiae and
closely related to the bisporangiate genus, Echinodorus, is an
example of a plant which has taken but a slight step toward the
monecious condition. In typical cases the inflorescence bears

Explanation of Figures 1-l.j.

Sporocarp of Marsilea quadrifolia L.

IVIicrosporangium from the same.

Megasporangium from the same.

Microsporophyll of Selaginella kraussiana (Kunze).

Megasporophyll from the same strobilus.

Stamen of Aquilegia canadensis L.

Carpel from the same flower, showing hairs on the ovulary, a sex-limited

character.
Staminate flower bud of Cocos nucifera L.
Carpellate flower bud from the same inflorescence.
Diagram of carpellate spikelet of Zizania aquatica L.
Diagram of staminate spikelet from the same inflorescence.
Diagram of a bisporangiate spikelet from the middle zone of the same

inflorescence as Figs. 10 and 11.
Staminate spikelet of Zizania aquatica L., showing lack of awn.
Bisporangiate spikelet from the same inflorescence, showing awn of

intermediate length.
Fig. 15. Carpellate spikelet from the same inflorescence as Figs. 13 and 14,

showing long awn, a sex-limited character.

Fig

1

Fig

2

Fig

3

Fig

4

Fig

o

Fig

6

Fig

i

Fig

8

Fig

9

Fig

10

Fig

11

Fig

12

Fig

13

Fig

14

112 The Ohio Journal of Science [Vol. XVIII, No. 4,

the carpellate flowers below and the staminate above on a
common scape, although the entire inflorescence may be
staminate or carpellate. There is not much difference in the
corresponding perianth segments, but there is a difference in
the receptacle. The staminate flower contains a half dozen
or so of vestigial carpels whose imperfection would hardly be
suspected unless compared with a normal carpel from a car-
pellate flower. The carpellate flower contains much more
reduced vestigial stamens. In general, one can not predict
which set of organs will be the more reduced in any given
species though it is generally true that the carpellate flower is
more apt to retain stamen structures than the staminate flower
is to retain vestiges of the gynecium. This agrees with our
notion that maleness in general is a more extreme condition
than femaleness, but there are many exceptions like Sagittaria.
As to the area involved in the dimorphic state, Sagittaria is
rather extreme in that the inflorescence is divided into definite
carpellate and staminate regions. However, there are many
species of plants in which the two types of flowers are inter-
mingled throughout the entire inflorescence. In the cocoanut,
Cocos nucifera L., the flowers are monosporangiate and
monecious. The diphorism of the flowers, which are situated
on the branches of a large inflorescence, is very great. (Figs.
8 and 9). Both flowers have distinct vestiges of the opposite
set of organs. The dimorphism extends out to the perianth.
The sepals of the staminate flower are small and short, while
those of the carpellate flower are large and cover the entire
bud. The petals of the staminate flower are narrow while those
of the carpellate flower are broad. Figures 8 and 9 are sketches
of unopened buds drawn to scale and show plainly how the
dimorphic condition is expressed in the tissues far beyond the
sporangia. It is interesting to note that the vestigial carpels of
the staminate flower are only slightly united, while those of the
carpellate flower are completely syncarpous. The inhibitory
staminate condition does not permit a complete union and a
more primitive condition, apocarpy, is expressed. In the
carpellate flower the stamen vestiges are small; however, one
occasionally finds flowers in which one or more stamens are
rather prominent. The writer discovered one case in which a
vestigial stamen had developed a nearly perfect anther and the
other five vestiges were larger than usual.

Feb., 1918] Sexual Dimorphism 113

A further advance in the extent of the dimorphism is shown
by those plants in which the distinctive staminate and car-
pellate flowers are confined to definite parts of the same inflor-
escence. A good example of this condition is the wild rice,
Zizania aquatica L. This is a monecious species with the
flowers in a large panicle. The upper part of the inflorescence
is carpellate and the lower staminate. The change in condition
usually extends transversely across the inflorescence axis and
its branches. The staminate spikelets have a vestigial gynecium
with three minute stigmas, while the carpellate spikelets have
six distinct vestigial stamens. Here we have a considerable
extent of tissue involving a large number of spikelets in the
same sexual state. The amount of the vegetative tissue involved
is much greater than in cases like the cocoanut. But the most
striking peculiarity of this inflorescence is in the central part.
Here the spikelets are bisporangiate having perfect stamens and
gynecia. (See diagrams, Figs. 10, 11, 12). On the transition
zone neither state is established and so no inhibition occurs.
The sexual state is brought about in the sporophylls as is usual
in bisporangiate plants in general as well as in the lower grasses.
Examples like Zizania show plainly that the determination of
staminate, carpellate, and bisporangiate flowers takes place in
the vegetative tissues by the establishment of a certain physio-
logical state and has nothing to do with gross or cellular mor-
phology. The fundamental morphological conditions, both
gross and microscopic, are the same throughout the entire
panicle.

Zizania aquatica has also a remarkable sex-limited char-
acter. The lemmas of the staminate spikelets are awnless while
those of the carpellate spikelets are long-awned. The bispor-
angiate spikelets in the neutral region have short awns. (Figs.
13, 14, 15). There is every gradation of length of awn in pass-
ing from the awnless lemmas of the staminate region to the
long-awned lemmas of the carpellate region. No matter
whether one or many awn factors are involved, the character of
the awn is due entirely to the latency or activity of the awn
factor or factors under the influence of the sexual state. Here
then we have plain evidence of the nature of sex limited char-
acters in plant sporophytes. Both awnless and awned lemmas
have a common sporophyte heredity, but this heredity expresses
itself in all degrees of latency and activity depending on the

114

The Ohio Journal of Science [Vol. XVIII, No. 4,

sexual state of the tissue involved. If we hold to the reasonable
hypothesis that the presence of a positive character is due to
the presence of a factor or group of factors, then the absence of
an awn on the staminate lemma is due to latency caused by
the presence of the male condition in the given cells.

The relative position of the staminate and carpellate flowers
in the inflorescence is reversed in various genera from what it
is in the wild rice. In the aroids the staminate flowers are

Fig. 16. Staminate inflorescence of Zea mays L.
Fig. 17. Carpellate inflorescence of Zea mays L.

usually above and the carpellate below. In the gama-grass,
Tripsacum dactyloides L., the same is true and the two regions
of the inflorescence show a very remarkable dimorphism. The
lower part, containing the carpellate spikelets is remarkably
modified. The dimorphism exists not only in the flowers and
glumes but extends into the stem. The modified carpellate
spikelets are enclosed in pockets, formed of highly modified
joints of the stem which develops cleavage planes, the joints
being separated at maturity, giving rise to very remarkable
fruits.

Feb., 19 IS]

Sexual Dimorphism

115

The next step in the evolutionary progression of sexual
dimorphism may be represented by Indian-corn, Zea mays L.
Here as is well known the typical plant has a terminal staminate
inflorescence, and one or more carpellate inflorescences devel-
oped from the side of the stem. (Figs. IG and 17). The differ-
ence between the two branches is remarkable although they
consist of the same morphological elements. The carpellate
branch shows by far the greater deviation from the general
vegetative morphology.' The main differences are as follows:

Staminate stem.

1. Normal internodcs.

2. Normal sheaths.

3. Leaf blade normal.

4. Inrtorescence of comparatively
primitive type, branched.

5. Axes normal.

6. Character of glumes, membranous

and elongated.

7. Staminate flower.

8. Little or no vestige of gynecium.

9. Normal stamens.

10. Microsporangia,

11. Microspores.

12. Ordinary color in floral axis and

glumes.

Carpellate stem.

1. Intemodes greatly shortened.

2. Sheaths changed to husks.

3. Leaf blade absent or vestigial.

4. Inflorescence modified, with loss of

branches.

5. Axis a cob.

(). Character of glumes, chartaceous
and broad.

7. Carpellate flower.

8. Remarkable development of style

and stigmas (Silk).

9. Minute or no vestiges of stamens.

10. Megasporangia.

11. Megaspores.

12. In some varieties, red or other color

in floral axis and glumes.

The writer recently received an interesting ear of popcorn
from Mr. L. E. Thatcher which shows a zonal arrangement of
the sexual condition. This ear is a normal side ear with the
lower part typically developed. In the middle is a complete
zone, about an inch long, of staminate spikelets, while the
outer part is typical ear structure again with normal grains.
In this case there is a successive reversal of the growing axis.
First the axis develops a cob with normal carpellate spikelets,
then changes suddenly to a staminate condition and finally
resumes its growth in the carpellate state.

The buffalo-grass, Bulbilis dactyloides (Nutt.) Raf. is a
perennial, monecious grass with stolons which root at the
nodes. According to Plank* and to Hitchcockf any given node
produces but one type of inflorescence, and each kind of stolon
is supposed to propagate its own kind. If this is true, the
buffalo-grass would be an interesting plant for experimentation.

*Plank, E. N. Buchloe dactyloides Engelm. not a Dioecious Grass. Bull.
Torr. Bot. Club. 19 : 303. 1892.

fHitchcock, A. S. Note on Buffalo Grass. Bot. Gaz. 20 : 464. 1895.

116

The Ohio Journal of Science [\o\. X\^III, No. 4,

The final step in the development of sexual dimorphism of
the sporophyte is attained in the diecious condition. But here
again some species are more definitely diecious. There are
very few strictly diecious plants. Diecious sporophytes are
isolated developments arising from parallel evolutions scattered
from one end of the AnthophA^te phylum to the other. In every
instance they are derived from bisporangiate ancestors. In
some cases as will be shown below the entire transition may
occur in a single genus.

Fig. IS. A mauirc, d\'ing, slaroinale plant of Cannabis saliva L., gruwil in the

winter.
Fig. 10. A blooming, carpellate plant of Cannabis sativa L. of the same age as

Fig. 18.

A good example of a diecious species is the common hemp,
Cannabis sativa L. The plants are staminate and carpellate
with other differences in the flowers besides the sporophylls.
The carpellate plants are more robust and much longer lived.
Figures 18 and 19 represent two plants of dwarf size raised in
the middle of winter in the greenhouse. The plants photo-
graphed were transplanted to pots when mature. These abnor-

Feb., 1918J Sexual Dimorphism 117

mal plants usually showed not more than five to seven leaf
nodes, while plants raised in poor soil during the summer
showed as high as twenty nodes. The staminate plant was
dying of old age when photographed, although less than four
months old. In plants like the hemp, the sexual dimorphism is
as great if not greater than in the ordinary mammals. In fact,
in many mammals it is much more difficult to recognize the
sexes by somatic characters.

Among these hemp plants there were intermediates as is
common in most diecious species. Staminate plants with more
or less carpellate expression were considerably longer lived and
more robust than those which were purely staminate. Not
only did typical staminate plants sometimes produce bispor-
angiate flowers with more or less normal gynecia but some
carpellate plants even 43roduced stamens. This in spite of the
fact that the plants were differentiated in their vegetative parts
as typically carpellate. In plants grown later, out of doors,
from the same seed no abnormalities were seen, but as the
plants were not examined when they first began to bloom they
may nevertheless have been present. The great abundance of
intermediates in the winter, greenhouse plants was probably
due to the abnormal environment, mainly a lack of light. The
point to be emphasized in this connection is that we have here
a diecious plant which shows sexual dimorphism even in its
remote vegetative parts, but numerous individuals which are
thus specialized have the ability to produce the opposite primary
sexual generation and sexual cells, without any manipulation
whatever being employed, except that they were grown in an
unusual environment. How extremely impossible it would be
in this case to claim that the specific sex characters were due
to sexual Mendelian units, one individual being homozygous
for sex and the other heterozygous. The whole behavior in
these diecious plants is essentially the same as the sexual dif-
ferentiation in the less extreme cases, traced above from Marsilea
to Indian-corn. The maleness and femaleness represent states
which inhibit to a greater or less degree the development of the
opposite organs the heredity of which is potentially present,
since both the staminate and carpellate plants do produce both
male and female gametophytes. Dieciousness is a differential
state, hereditary, of course, in the ordinary sense, which permits

118

The Ohio Journal of Science [Vol. XVIII, No. 4,

under ideal conditions only the one set of spores and organs to
be expressed, although the hereditary abilities for the other set
are all present. The morphological expression is brought about
in the same way as such a differential development takes place
in the tissues of bisporangiate sporophytes, either with
monecious or bisporangiate flowers.

Explanation of Figures 20-26.

Fig. 20. A staminate flower, with oerianth removed, from a carpellate tree of

Acer platanoides L.
Fig. 21. Part of the same flower, showing the vestigial gynecium.
Fig. 22. Staminate flower, wiili perianth removed, from a staminate tree of

Acer platanoides L.
Fig. 2-3. Part of the same flower, showing the vestigial gynecium.
Fig. 24. Fruiting carpellate flower, with perianth removed, from the same tree

as Fig. 22, showing normal young samara and vestigial stamens.
Fig. 25. Tip of a staminate flower of Rumex acetosella L., showing one of the six

stamen filaments and the small vestigial gynecium with three minute

vestigial stigmas.
Fig. 26. Gynecium from a carpellate flower of Rumex acetosella L., showing one

of the three, large, branched stigmas, the other two being removed.

I

Feb., 1918] Sexual Dimorphism 119

The genus Acer presents a very good series, passing from
Acer platanoides L. through a number of species to Acer
negundo L., the boxelder. So far as the writer's observations
go, the boxelder is strictly diecious. Acer platanoides is in a
much more primitive condition being but a short distance
removed from the normal bisporangiate type. However the
plants are staminate and carpellate. The writer has found
carpellate trees which developed a few typical staminate flowers
with vestigial gynecia and normally developed stamens, although
the trees were covered with young fruit from the normal car-
pellate type of flower. (Figs. 20 and 21). Staminate trees
were also discovered which were producing many carpellate
flowers with vestigial stamens (Figures 22-24). These
carpellate flowers developed normal fruit. Figure 24 rep-
resents a partially developed fruit. In both cases the per-
centage of the unexpected kind of flowers was small, but the
staminate trees produced a much larger per cent, of carpellate
flowers than the carpellate trees did of staminate flowers.

Rumex is another genus which begins with a partial
bisporangiate state and ends in a diecious condition. Rumex
crispus L. has bisporangiate flowers and carpellate flowers,
the latter containing prominent vestigial stamens. Rumex
acetosella L. is strictly diecious. The carpellate plant shows
no vestige of the stamens or else these are too small to be seen
without making special sections. But the staminate flower
shows a small vestigial gynecium with three minute stigmas.
(Fig. 25). The persistence of the three vestigial stigmas may
be accounted for by the fact that the normal stigmas of Rumex
are very large and branched. (Fig. 20). In some of the
intermediate species of Rumex, as for example Rumex altissimus
Wood, the vestigial stamens are rather prominent and the
vestigial gynecium is of fair size with three small slightly
branched stigmas.

A most striking example to illustrate the stages in the
evolutionary development of dieciousness is presented by the
genus Fraxinus. Fraxinus cuspidata Torr. has flowers with
corolla and calyx. They are bisporangiate and very fragrant.
Fraxinus quadrangulata Mx. has bisporangiate flowers, but
no perianth. The merest vestige of a calyx is present. Fraxinus
americana L. has a minute calyx and is strictly diecious. If
the antiquated and misleading classification of the flowering

120 The Ohio Journal of Science [Vol. XVIII, No. 4,

plants, still generally in vogue, could be eliminated, more
botanists might realize the fact that monecious and diecious
species are the extremes and commonly the culmination points
of numerous parallel series rather than the first stages of
Angiosperm evolution.

One more example will be given to illustrate the progressive
differentiation of larger areas of staminate and carpellate tissues
in an ascending phyletic series. In the Cyperacese the least
specialized genera have bisporangiate flowers, like the genus
Scirpus. Some of the species of this genus have a fairly well
developed vestigial perianth of six segments. There are no
special structures difficult to interpret. But in the genus,
Carex, the perianth is absent and there is present the peculiar
perigynium and other unusual structures. Carex is monecious;
usually with staminate and carpellate flowers apparently with-
out vestiges of the opposite organs. In Carex nardina Fries, the
spikelets are all alike containing carpellate flowers below and
staminate flowers above. In the highest sedges like Carex
lupulina Muhl. the monosporangiate flowers are on separate
spikelets, the staminate spikelets above, the carpellate spikelets
below. In Carex, therefore, we approach the condition present
in Indian corn.

What is the nature and cause of this progressive change in the
area of the tissue involved? In the first case mentioned above,
the differentiation takes place in the sporophylls on the same
floral axis; in the second case the floral axes produce each but
one kind of sporophylls, but the two kinds of flowers are on the
same inflorescence axis; in the third case an entire spikelet
or group of spikelets is similarly affected. The whole matter
can only be interpreted, if at all, by a study of the evolutionary
series of expressions in related species. Isolated studies do
not give a true picture; in fact do not present the problem.
But in no study could the phenomena of segregating or associ-
ating chromosomes be involved but the much more difficult
problem of changing hereditary expressions and states in a
common vegetative tissue.

As an example of the changes of morphological expression
that take place in a growing bud, an ordinary grass may be
considered, like Bromus secalinus L. The grasses are derived
from plants whose buds produced three spirals of leaves in the
vegetative parts as well as in the flower, the ancestral flower

Feb., 1918] Sexual Dimorphism 121

being a trimerous, pentacyclic structure. In the sedges the
leaves are still in three spirals, but in the grasses they are in
two; in other words, the leaves of the grasses are two-ranked.
During the vegetative growth of Bromus, therefore, its bud gives,
off alternately incepts of leaves on opposite sides of the stem.
These leaves are differentiated through the activity of hereditary
units into the form characteristic of the species. This two-
ranked arrangement is a culmination type of morphological
expression. As stated, the species evolved from ancestors
in which three spiral incepts were successfully organized instead
of two. Now the cells of the terminal bud actually contain the
hereditary ability to develop threes as will appear below.
But for some cause the ability is suppressed. The bud, using
a metaphor, dances a two-step instead of a waltz. When a
spikelet begins to develop, the two-ranked condition continues
and the two-ranked empty glumes are produced and a number
of flowers, also in two ranks. The flower bud on the spikelet
also produces two-ranked glumes, the flowering glumes, unless
the palet represents more than one leaf. But the first set of
floral organs proper, the lodicules are produced in a three spiral
arrangement. The third, or posterior one, however, is sup-
pressed in harmony with the bilateral nature of the evolved
plant. One step down in the evolutionary scale, in the bamboo
tribe, the third lodicule is present in the proper position as it
should be in a monocotyl flower. In some way the bilateral
nature does not act so intensely as in Bromus and the higher
grasses in general. The lodicules probably represent a corolla,
the calyx being suppressed. Next the flower bud of Bromus
not only develops three incepts in typical tripartite arrange-
ment, alternate with the lodicules, but these organs pass into
that strange state which determines the nature of stamens
and male gametophytes with male cells or sperms as the final
goal. Now all of this up to the formation of microsporocytes
is a matter of vegetative growth. The cells in the stamens have
received the same heredity as those which produced leaves,
glumes, or lodicules. The three stamens are the only structures
which show the typical ancestral morphology in the entire life
cycle of the grass under consideration. Next the bud fails to
perform and the second, expected set of three stamens does
not develop. In many bamboos all the six stamens are present,
as in any typical monocotyl that is not specialized too much.

122 The Ohio Journal of Science [Vol. XVIII, No. 4,

Finally the bud does another "three-step" forming a united
tricarpellary gynecium, but here again the third stigma on the
outside in the bilateral plane is suppressed. All three stigmas
are usually present in Arundinaria and other bamboos. At the
same time the incept of the gynecium has set up in its cells a
sexual state just the opposite from that of the andrecium
immediately below although so far as any one knows and so
far as the evidence goes these cells have received exactly the
same hereditary units as have those of the leaves, the glumes,
the lodicules, and the stamens. But in some way a new
physiological state has been established which causes the
heredity to give rise to a new morphological expression along
with femaleness.

At each whorl there is a difference in hereditary expression, a
change in the activity and . latency of numerous hereditary
factors. This change in activity and latency, the writer
believes is due to changes in physiological state of the tissues
involved, whether chemical or otherwise. The progressive
changes of morphological expression in a vegetative tissue do
not find their explanation in any Mendelian formulae. Mendelian
ratios, segregations and associations have no direct bearing
on the problem. Just as these profound changes in the veg-
etative development are brought about by some state in the
cells which influence the hereditary activity, so essentially
similar changes in physiological activity bring about sexual
expressions, causing the cells or tissues to give rise to male or
female morphological structures with their accompanying
sexual activities.

It may be that in some cases physiological states or
hereditary factors may arise in an allosome or special chromo-
some which may assist in retaining and intensifying a male
or female state already established, but so far as the writer
can see the animal kingdom presents the same problems of
changes in sexual states in common tissues as does the plant
kingdom. The lower animals are hermaphrodites and the
higher unisexual forms are after all only modified hermaphro-
dites. Sex-limited and sex-linked transmission can readily be
explained without postulating a sex-determining allosome,
with no more complexity of hypotheses than if such mendelizing
units are assumed. And it must always be kept in mind that
with the assumption of sex-determining chromosomes the greater

I

Feb., 1918] Sexual Dimorphism 123

part of sexual phenomena becomes unexplainable and con-
tradictory. The presence of allosome difference in certain
sexual individuals does not make it necessary for us to amend
the proposition that sexuality either male or female is a state
or condition and not a mendelian factor or set of factors.

For the convenience of those who wish to make a study
of our more common or interesting species, the following short
list is given as an index to the general condition usually present
in monecious and diecious plants:

Four genera to illustrate the relation of diecious to
bisporangiate species — Thalictrum, Rumex, Acer, Fraxinus.

Thalictrum clavatum DC. Flowers bisporangiate.
Thalictrum dasycarpum Fisch. & Lall. All gradations of bisporangiate
to staminate and carpellate flowers on the same plant.

Thalictrum dioicum L. Plants diecious.

Rumex crispus L. With bisporangiate flowers and carpellate
flowers containing prominent vestigial stamens on the same plant. The
stigmas are much branched.

Rumex altissimus Wood. Monecious. The carpellate flowers have
six vestigial stamens; the staminate flowers have a vestigial g}^necium
with three slightly branched stigmas.

Rumex acetocella L. Diecious. The staminate flower has a minute
vestigial gynecium with three, unbranched, vestigial stigmas; the
carpellate flower apparently has no vestigial stamens; the stigmas are
much branched.

Acer platanoides L. Imperfectly diecious. Some carpellate trees
have staminate and some staminate trees carpellate flowers; both kinds
of flowers have prominent vestiges of the opposite organs. The flowers
have large petals and a prominent nectar disk.

Acer saccharinum L. Diecious. Carpellate flowers with vestiges of
stamens; staminate flowers with vestigial g\mecia. Some plants
occasionally have bisporangiate flowers. The petals are absent.

Acer negundo L. vStrictly diecious. The flowers are much reduced
and have no vestiges of the opposite organs.

Fraxinus cuspidata Torr. Flowers bisporangiate with a calyx and
corolla; very fragrant.

Fraxinus americana L. Strictly diecious. Flowers small without
corolla but with a small calyx; no vestiges of the opposite sporophylls.

124 The Ohio Journal oj Science [Vol. XVIII, No. 4,

List of a few monecious and diecious species with notes on
the condition of the reciprocal organs in the two types of flowers:

Sagittaria latifolia Willd. Monecious, occasionally diecious. The
staminate flower has prominent vestigial carpels, only slightly different
from the fertile carpels of the carpellate flower. The carpellate flower
has minute vestiges which, no doubt, represent stamens.

Sagittaria rigida Pursh. Monecious. The condition of the flowers is
the same as in S. latifolia.

Phoenix dactylifera L. The carpellate flower has six vestigial
stamens; the staminate flower has three vestigial carpels.

Cocos nucifera L. Monecious. The carpellate flower has six minute
vestigial stamens, one or more of which occasionally develop nearly
normal anthers; the staminate flower has three prominent, only slightly
united, vestigial carpels.

T3^ha latifolia L. Monecious. The flowers are much reduced and
neither type shows any vestiges of the opposite organs. More primitive
related genera, belonging to the PandanacccC, show vestigial structures.

Ariseema triphyllum (L.) Torr. More commonly diecious. Spadixes
occur which have only carpellate flowers; some have carpellate flowers
with a few imperfect staminate flowers at the top; some have normal
carpellate flowers below and normal staminate flowers above; some are
almost entirely staminate with two or three normal carpellate flowers
situated in about the middle of the spadix; and some are entirely
staminate.

Panicum virgatum L. Each spikelet contains a bisporangiate and a
staminate flower.

Zizania aquatica L. Monecious; with staminate spikelets below
and carpellate spikelets above, on the inflorescence; the carpellate
spikelet contains six vestigial stamens; the staminate spikelet contains
a vestigial g>mecium. In the transition tissue there are normal, bispor-
angiate spikelets. The awns are sex-limited, depending on the degree
of the carpellate condition.

Andropogon furcatus Muhl. In each pair of spikelets, the sessile
spikelet is bisporangiate; the stalked spikelet is staminate with a
vestigial g}mecium.

Tripsacum dactyloides L. Monecious; with the carpellate spikelets
below and the staminate above. The carpellate flower with three
vestigial stamens; the staminate flower with a vestigial gynecium.

Smilax hispida Muhl. Diecious. The carpellate flower has six
vestigial stamens; the staminate flower contains no vestige of the
gynecium.

Dioscorea villosa L. Diecious. The carpellate and staminate
flowers contain vestiges.

Thalyctnim dasycarpum Fisch. & Lall. On the same plant may be
found purely carpellate flowers, purely staminate flowers, carpellate
flowers with one stamen, staminate flowers with one carpel, and all
gradations between, some flowers having about half of each kind of
organs. Some species of Thalictrum are diecious.

Feb., 1918] Sexual Dimorphism 125

Menispermum canadense L. Diecious. The carpellate flower has
about twelve vestigial stamens; the staminate flower has no vestiges.

Sassafras sassafras (L.) Karst. Diecious. The staminate flower
has a vestij^ial carpel: the carpellate flower has six staminodes.

Zanthoxylum americanum Mill. Usually diecious. The staminate
flower has prominent vestigial carpels, the carpellate flower sometimes
has minute vestigial stamens.

Ptelia trifoliata L. Imperfectly monosporangiate and imprefectly
monecious. The staminate flower has a prominent imperfect gynecium;
the car])ellate flower has five vestigial stamens.

Ailanthus glandulosa Desf. Diecious. The carpellate flower has ten
prominent staminodes; the staminate flower has a prominent vestigial
gynecium of five carpels.

Napea dioica L. Diecious. The carpellate flower has a vestigial
stamen column; the staminate flower has no vestige.

Rumex acetosella L. Diecious. The staminate flower has a small
vestigial gynecium with three minute stigmas ; the carpellate flower has
no vestiges.

Aruncus aruncus (L.) Karst. Diecious. The carpellate flower has
vestigial stamens; the staminate flower has three vestigial carpels.

Gymnocladus dioica (L.) Koch. Diecious. The carpellate flower has
ten large staminodes; the staminate flower has a vestigial carpel.

Acer negundo L. Apparently strictly diecious with no vestiges of
the opposite organs. In the lower species of Maples the trees are
imperfectly diecious and the flowers have very prominent vestiges of
the opposite organs.

Morus rubra L. Diecious. Staminate flowers with a vestigial
gynecium; carpellate flowers without vestiges. Often there are
staminate and carpellate catkins on the same tree; sometimes there
are carpellate and staminate flowers on the same catkin; some-
times a carpellate catkin may contain a single staminate flower.

Cannabis sativa L. Diecious; but the plants are of various degrees;
some carpellate plants may have stamens and staminate plants may
have carpels.

Populus deltoides Marsh. Apparently strictly diecious, as also
some willows, but there are intermediate willows, occasionally.

Cucumis sativus L. Monecious. The carpellate flower has three
vestigial stamens; the staminate flower has a tricarpellate, vestigial
gynecium.

Diospyros virginiana L. Usually diecious. Carpellate flower with
vestigial stamens; staminate flower with a large vestigial gynecium.

Fraxinus americana L. Apparently strictly diecious. Various
species of Fraxinus present gradations from bisporangiate, conspicuous
flowers to diecious flowers with loss of perianth and nectar glands.

Ambrosia trifida L. Monecious. The staminate flower has a
vestigial gynecium; the carpellate flower shows no vestiges of stamens.

EUTETRAMORUS GLOBOSUS, A NEW GENUS AND

SPECIES OF ALGJE BELONGING TO THE

PROTOCOCCOIDEA (Family Coelastrida).

L. B. Walton.

While studying the plankton from "Mirror Lake," a small
pond on the campus of the State University at Columbus, Ohio,
exceedingly rich in phytoplankton at certain times of the year,
a form quite unique in structure was noted with much interest.
It consisted of 16 cells, each containing a chloroplast, the cells
being arranged in groups of fours and imbedded in an almost
invisible gelatinous matrix. The organism was non-motile with
no trace of flagella. The preparation was one taken from a

Eutetramorus globosus n. g., n. sp. (x2000.)

sterilized specimen bottle filled with water and floating alg^ —
mostly Cladophora — at the margin of the lake, Oct. 9, 1915,
the observation being recorded a few days later. An interval
of over two years has elapsed during which period various
samples of water from the lake have been studied without again
noting the species however.

The organism is referable to the Family Coelastridce {Coelas-
tracea) of the Protococcoidea* and constitutes a new genus quite
different from forms thus far known. The drawing (Fig. I)

*The endings of the Class and Family names are identical with those utilized
in a systematic review of the typically unicellular forms which will be published
shortly. They are an extension of those proposed by Poche (1911). Nomenclatural
methodology, particularly among the primitive plant-animal organisms is in a
somewhat chaotic condition.

Feb., 1918] Eutetramosru Globosus 127

is taken from a camera lueida sketch made at the time with a
Leitz Binocular, 2 mm. apochromatic objective.

Eutetramorus n. g.

Cells non-motile, light chlorophyl green; united into a col-
only of 16 cells arranged in groups of 4's within a gelatinous
like mucous covering; plane of each series of 4 cells perpen-
dicular to the square included by the groups.

Represented by a single species.

E. globosus n. sp.

Cells spherical, containing a chloroplast with central
pyrenoid, the 16 cells united in groups of 4's, the distance
between each group and the adjacent lateral group being
approximately one and one-half times the diameter of the
individual cell; reproduction unknown.

Diam. (single cell 5/x. (colony) 30^.

Distribution, Mirror Lake, Columbus, Ohio (U. S. A.).

Habitat, surface water at margin of lake.

The relationship of the form to Ccelastriim and the other
allied genera of the family seems clear, although the systematic
position must be a provisional one until the method of repro-
duction is known. In its organization it represents the lowest
form of the family where a definite colonial organization is
attained.

Kofoid (1914) has recently described a new genus and
species, Phytomorula regularis, from a reservoir at Berkeley,
CaHfornia, which is allied to Ccelastrum and is of unusual
interest in that it represents a 16 celled colonial form extremely
flattened, the cells being contiguous although not arranged in
the same plane. The species was extremely rare and he had
been unable, at the time of the presentation of the paper, to
obtain informa- tion as to its method of reproduction.

The family Ccelastridce now consists of five genera which may
be separated in accordance with the table given below. Three
of the genera have an extremely restricted distribution which,
however, may be the result of their comparatively rare occur-
rence. Eutetramorus is based on a single specimen obtained at
Columbus, Ohio. Phytomorula is described from a very few
specimens obtained in a reservoir at Berkeley, California.

128 The Ohio Journal of Science [Vol. XVIII, No. 4,

Burkillia is known only from Burma. The remaining two
genera, Coslastrum and Sorastrum are abundant and have a
wide distribution. The genus Hariotina based by Dangeard
(1889) on H. reticidatum is now included with Ccelastrum while
Selenosphcerium of Cohn (1879) is placed with Sorastrum.

TABLE OF GENERA.

A^ Cells comparatively smooth or at least not developing acute processes
or spike-like appendages; colonies with cells regularly arranged,
usually approximating the form of a sphere, which may be extremely
flattened.
B' Colonies formed of 16 cells; form not that of a true sphere.

C^ Cells arranged in groups of fours, the groups not con-
tiguous; colony not flattened 1. Gen. Eutetramorus

C^ Cells not arranged in groups of fours; contiguous; colony

flattened 2. Gen. Phytomorula

B^ Colonies formed of 2-32 cells; form approximately spherical.

3. Gen. Coelastrum
A'^ Cells developing acute processes or provided with acute or spike-like

appendage processes; colonies with cells not regularly arranged, not
approximating the form of a sphere.
B^ Cell walls gradually narrowed into an acute process.

4. Gen. Burkillia
B^ Cells provided with spike-like appendages or elongate processes.

5. Gen. Sorastrum

BIBLIOGRAPHY.

1915. Brunnthaler, J. Die vSiisswasser-Flora Deutschland, Oesterreichs, und der

Schweitz, Hft. 5. p. 1913, (G. Fischer, Jena.)
1914. Kofoid, C. A. Phytomorula regularis, A Symmetrical Protophyte related

to Coelastrum. Univ. of Calif. Pub., Botany, V. 6, No. 2, p. 35-40, PI 7.
1907. West, W. and West, G. S. Fresh-water Algae from Burma, including a few

from Bengal and Madras. Ann Roy. Bot. Gard. Calcutta, V. 6, Pt. 2.

1916. West G. S. Algae, p. 205 and 243 (Cambridge University Press, Cam-

bridge, England.

Kenyon College, Gambler, Ohio, December 5, 1917.

A STUDY OF CATTLE "TEMPERAMENT" AND ITS

MEASUREMENT.

By Arthur F. Pott,
State Normal School, Stevens Point, Wisconsin.

INTRODUCTION.*

The present phase of this problem has been evolved from an
attempt to correlate the so-called "dairj^ temperament" with
milk production. The term "dairy temperament" was first
promulgated by Ex-Gov. Hoard of Wisconsin in 1886, in a
lecture on "Nervous or Dairy Temperament in Cattle." He
called it a predisposing tendency in the animal to convert its
food either into milk or flesh." He called that temperament
pertaining to dairy cattle, "dairy temperament." A later
definition of the term was expressed by Prof. Haecker, of Minn. :
"An animal whose nervous system dominates the vital system,
has the inherited tendency to convert the nutriment in food
into milk." " Dairy temperament" today, is one of the strong-
est points of the dairy cow score card. The term and its defini-
tion are based on indications only, such as "eye full and
expressive," "clean face," "large nostrils," "long, light neck,"
"sharp withers," "prominent spinal column," etc.

The word "temperament" is perhaps a rather ambiguous
term to apply to cattle, but it was employed, no doubt, for
want of a better word. The usual definition of the word char-
acterizes it as a mental condition, or "special t^^pe of mental
constitution and development or mixture of characteristics,
supposed to have its basis in the bodily organism and to be
transmissible by inheritance,"! or again a "natural disposition."
Speaking of disposition, psychologically it is a "tendency left
behind by an experience, to give rise on suitable stimulation, to
a reaction which shows the influence of that experience, espe-
cially as applied to explain the phenomena of memory." The

*This paper embodies the essentials of a thesis submitted for the degree of
Master of Science in Agriculture, the work for which was carried out under the
direction of Professor C. S. Plumb, Head of the Animal Husbandry Department of
the Ohio State University.

The writer also wishes to express his appreciation to the following, for their
valuable suggestions and criticisms: Profs. Wm. M. Barrows, G. F. Arps and
A. P. Weiss, Ohio State University.

jFunk and Wagnalls, New Standard Dictionary, 1914.

129

130 The Ohio Journal of Science [Vol. XVIII, No. 4,

term "dair}' temperament," as it has been used, is based
entirely upon physical characteristics of the animal, and as
such is abstract and unusable. It is the purpose of this study
to attempt to measure "temperament" in cattle, and so place
it upon a quantitative basis.

Historical. — In the study of human psychology, many
experiments have been carried on by which various emotions,
and mental and nervous disturbances have been registered.
According to C. S. Stumpf — "our conscious states, without our
willing it — indeed, even in spite of us — are accompanied by
bodily changes, which very often can be detected only by the
use of extremely fine graphic methods." (10) These have been
based mainly, but not solely on respiration. Respiration in
animals is controlled almost entirely by the nervous system,
the respiratory center being located in the medulla oblangota.
Connected with it is the vagus center, which in turn receives
nerves from the lungs, heart and stomach. The respiratory
movements are controlled primarily by the nerves to the inter-
costal muscles and diaphragm. The nerves supplying these
muscles do not come from the respiratory center, but come
from the cells of the grey matter of the spinal cord. It is by
influencing the activities of these cells, that the respiratory
center controls the act of respiration. In its turn the respiratory
center is under control of the higher nerve centers of the brain.
Due to the afferent nerves from the viscera and sense organs, as
well as from the higher brain centers, respiration is influenced
by the heart beat, activities of the stomach, and internal
organs, as well as by external changes. The automatic activity
of the respiratory center is chiefly regulated by the amount
of acid in the blood and the temperature of the blood.
Therefore the respiratory movements are also regulated by the
metabolic activities of the animal. As the rate of the heart
beat accelerates, so, too, does the rate of respiration. Respira-
tion then is not a separate activity, but is in harmony with, and
closely allied to various other physical and mental activities.
Respiration has been found to be a good index to the nervous
reactiveness and "temperament" of an animal, and has formed
the chief basis of this study, being used as a means of measuring
"temperament" or nervous reactiveness. As Zoneff and
Neumann say: " Insoeben wurde ich bei gelegentlichen Ver-
suchen, auf den Unterschied aufmerksam, den das Athem mit

Feb., lUlSj

Cattle Tcmpcranioit

i;u

dem Brustkorb and die Zwcrclifellathmung beim Ausdruck der
Gefuhlc zcigcn. Ueberall, wo in unsern Versuchen der Athem
charackteristische Veranderungen aufweist, finden sich analoge
Erscheinungen im Puis." (12).

It can be shown by suitable curves recorded while the
animal is resting, that there is a fundamental rhythm in the
respiratory movements, which is peculiar to each individual.
Ordinarily this rhythm is obscured because external and
internal stimuli affect respiration to a very large extent. It
has been our experience, that those animals which were least

Fig. 1. Respiration tracings of 7H (above) and 2H (below) recorded while
feeding. At A, B and C are shown the deep expirations. This type of
respiration recalls the Cheyne-Stokes breathing sometimes seen in man,
and is typical of resting animals or those which have a stolid disposition.
The contrast between these tracings illustrates very well the difference
between the respiration of a stolid and nervous cow. Time intervals,
five seconds.

nervous and most easily handled, showed this fundamental
rhythm very often ; and further, that the fundamental rhythm
of these very stolid animals showed a very low variability when
compared with rhythms of more nervous ones, (see Figure 1).
As a result of considerable preliminary work and experience
with animals concerned, we have confined this study to the
measurement of the variability of the depth of breathing, shown
by four Holstein cows, which were tested as nearly as possible
under the same normal stable conditions.

132 The Ohio Journal of Science [Vol. XVIII, No. 4,

METHODS AND PROCEDURE.

An apparatus, similar to that used upon human beings, was
employed, with but a few minor modifications which adapted it
to use with cattle. This consisted of a pnuemograph, kymo-
graph, tambour, electric clock and signal magnet, which were
connected in the usual manner. The tambour registered the
respiratory activities upon a drum fitted with smoked paper,
w^hich revolved at a convenient rate. The speed was so timed
that the curves were easily read, and showed the minutest
variations. In all curves, the signal magnet registered five sec-
ond intervals below each curve. In every curve the up stroke
indicates expiration and the downstroke inspiration. These
pneumographic tracings, (see Figure 1), not only registered
respiratory movements, but also outward movements of the
animal's body, such as kicking, moving about, movements of
the head, switchings of the tail, etc. The pneumograph was
attached about the barrel of the animal, nearly over the dia-
phragm. In this position, all or nearly all of the movements
were recorded. As has been shown, every movement is con-
trolled by the nervous system and so has a direct bearing upon
the measurement of nervous activity. Thus a restless animal
showed a greater number of variations in its curve than a
more quiet one.

Four registered Holstein-Friesian cows were experimented
with — Seven H, Three H, Two H, and Five H. Three periods
of the day were selected for experimentation, which represented
three given conditions — before feeding, during feeding and
after feeding. Curves were made with the different animals
on different days under these conditions, over a period of three
months. After becoming familiar with the apparatus and
subject, each variation in the curve proved significant of some
definite activity. We were soon able to recognize kicks,
swallowing, switching of tail, etc., as recorded on the drum.
Our experience is in accord with that of Neumann, who says:
"Es zeigte sich bei diesen Versuchen, dass der Athem das
empfindlichste Reagens bei Gefiihlschwankungen bildete, und
das bei rightiger Behandling der pneumographischen Reg-
istierapparate die Athemveranderungen als sicherestes Kenn-
zeichen alle Veranderungen der Gefuhlslebens dienen konnen. "
(12.)

Feb., 1918] Cattle Temperament 133

RESULTS.

Treatment. — After sufficient data had been collected, (about
150 curves), a method of interpreting the pneumographic
tracings was used, whereby respiratory activities and the
corresponding variations were taken into account. The ampli-
tude of each inspiration and expiration was measured in milli-
meters, and these measurements were tabulated statistically,
and the data plotted in the form of a frequency polygon,
where the amplitudes of respirations were plotted against
their frequencies. The mean (M), standard deviation (S. D.),
coefficient of variability (Cv.), and their corresponding probable
errors, were calculated according to the usual formulae.

Many such frequency polygons were made of the different
subjects, under different conditions, and taken from different
days selected at random. After a sufficient number of such
polygons from each animal were plotted, the entire number of
polygons of each individual under a given condition, were
constructed into a composite curve or polygon, which latter
was taken as characteristic of that animal under the given
condition. The composite was constructed in the following
manner. The means of the individual polygons were super-
imposed upon each other, and then each polygon was plotted
about its own mean. Then, for example, where the various
lines of amplitude 4 crossed the different frequency lines, a
simple average was taken, and this average represented the
average frequency of amplitude 4, and was plotted as such
upon the composite. From the composite, another set of
figures was derived, and this taken as characteristic of the
given subject under the given condition. And so the mean,
standard deviation, and coefficient of variability of the com-
posite were taken as representative. This method of con-
structing composites was made necessary by the curious moving
of the mean up or dow^n (see below), and by variations in
pressure in the pneumograph and tambour due to changes in
tension and positions of the same.

134

The Ohio Journal of Science [Vol. XVIII, No. 4,

TABLE I.

Showing the means, M., standard deviations, S. D., and coefficients of variability,
Cv., of the respiration composites of the four cows under the experimental con-
ditions, before, during, and after feeding.

ANIMAL
XCMBER

BEFORE FEEDING

FEEDING

AFTER FEEDING

M. 6.4101

±

.067

M.

10.422 ±

.080

M.

5.349 ±

.077

Seven H

S.D. 2.025

±

.047

S.D.

2.581 ±

.057

S.D.

2.204 ±

.054

Cv. 31.59

=t

.813

Cv.

24.76 ±

.579

Cv.

41.20 ±

1.17

M. 6.698

±

.089

M.

9.122 =t

.084

M.

7.21 ±

.100

Three H

S.D. 2.570

=fc

.063

S. D

2.607 ±

.067

S.D.

3.042 =t

.070

Cv. 38.369

zt

1.06

Cv.

28.57 ±

.791

Cv.

42.19 ±

1.14

M. 6.732

=fc

.133

M.

9.971 ±

.132

M.

9.093 ±

.137

TwoH

S.D. 3.098

=fc

.094

S.D.

3.796 ±

.094

S.D.

3.458 =t

.097

Cv. 46.02

=fc

1.669

Cv.

38.07 ±

1.064

Cv.

38.02 ±

1.213

M. 8.227

±

.155

M.

11.97 ±

.174

M.

7.92 ±

.120

Five H

S.D. 4.154

±

.109

S.D.

5.162 ±

.123

S.D.

3.096 ±

.085

Cv. 50.49

±

1.67

Cv.

43.124 ±

1.20

Cv.

39.08 ±

1.22

Individual Variation. — For clearness and convenience, two
sets of comparisons will be made; first, variations within the
individual, and second, variations within the group. As
stated above, each subject was experimented upon under three
different conditions, before, during and after feeding. In order
to facilitate comparisons, all three polygons have been plotted
upon one sheet, i. e., those which pertain to the individual
animal; and at the same time the coefficients of these curves,
the mean, standard deviation and coefficient of variability
with their probable errors are shown in Table I.

The general appearance of the composites of all the individ-
uals would lead one to believe that the coefficient of variability
is the key to their interpretation. The coefficients do bear
a relation to the shape of the curve, but in their interpretation
they cannot be considered alone, but must be interpreted in
terms of the mean and standard of deviation, otherwise they are
misleading. For example, the after feeding polygons of Seven
H and Three H, (Fig. 3) are entirely different, and a single
glance would tell that the polygon of Three H is more variable
than that of Seven H, and yet their coefficients of variabihty
are practically the same, 42.19 and 41.20 respectively. But
considering the means and standard deviations, this is easily

Feb., 1918] Cattle Temperament 135

explained, that Three H is more variable than Seven H under
the same condition. The mean of Three H is 7.21 and that of
Seven H is 5.34, which shows that Three H had a large number
of respirations about 7 mm. in length, while Seven H had a
large number of respirations about 5 mm. in length. Their
standard deviations, 3.04 and 2.20 respectively, are more
significant, since they show that Three H deviated more from
its mean, than Seven H did from its mean. Now combining
the respective means and standard deviations of each individual,
it is seen that the mean and standard deviation of Three H
increased in about the same ratio, and hence the coefficients
of variability are practically alike, since by formula:

^ . ^ • 1 .,• standard deviation ^^^

— coefficient of variability = .100.

mean

But the fact remains that Three H is more variable than Seven
H, under the given condition, even though their coefficients of
variability do not show as marked difference as the polygons
themselves indicate.

Further, the fact that the mean and standard deviation of
Three H increased in the same ratio, is explained by the condi-
tion under which the tracings were made. This was not con-
stant. Above it was stated that the animals were subjected to
like conditions, but these conditions were not constant so far as
the time element is concerned. For sufficient reasons, as will be
explained later, the after feeding condition was not always
taken at a specific interval after the regular feeding time.
This, as will be seen, affects the means of the polygons, and in
turn the coefficient of variability. As the pneumographic trac-
ings of the after feeding condition were taken farther from or
nearer to the regular feeding time, the resulting polygons here
plotted moved farther from or nearer to the "o" ordinate, due
to the increasing or decreasing depths of breathing as the case
may be. The reason that the mean should vary directly with
the time of taking the after feeding tracings, is probably due to
the metabolism of the animal which also varies as the after
feeding condition draws near to or farther from the actual
feeding time. Consequently, the time element not being con-
stant, the means varied to such an extent as to make the
coefficients of variability misleading, and so the standard
deviation is of more consequence than the mean in interpreting

136

The Ohio Journal of Science [Vol. XVIII, No. 4,

fo

'" /A

K^ / \

iy 1 \

r" 1 \

if /fs \

// ^ \

'» - // ^ \

Si- 0 ' \ \

■^ / / \

^' ^ / \ \ Figure 2

yr vj / / \ \

I' 1 ' •■"■' \ \

f ^.y^-y"-^ ^ Vv ■■"".

iy/"'-'- / *-/1/n/>l'/uyes o) ficifiiraf-!i!i->'^ ._\^ v,^

^^'^ """"^-Ci^- ,^

^

Figure 4

\

I

Fig. 2. Curves showing respiration in the four cows while feeding: Seven^H;

— Three H; Two H; Five H. (All figures are drawn

to the same scale.) t ' IV*

Fig. 3. Curves showing respiration in the four cows after feeding, w l|H
Fig. 4. (Below). Curves showing respiration in four cows before feeding.

Feb., 1918]

Cattle Temperament

137

Figure 5

I ^ ,0 " /I i " 's /t n 't /ff jt " ii J

i 11 11- *> •" -"

/»' 2

i"

t

is-

'^ )

ij

/ /

': ^ /

*■•"

^/ /

io

/ ^J

/ ' .-'

r

.' /

Figure 6

K -• *

ij. /J z/ 'f__jt -i-.:^.'T A.'..

\ Figure 7

/■• f^ /r /

Fig. 5. Respiration curves of cow Five H under three conditions:

feeding; feeding, and after feeding.

Fig. 0. Respiration curves of cow Two H, under the three conditions.
Fig. 7. Respiration curves of cow Three H. under the three conditions.
Fig. 8. Respiration curves of cow Seven H, under the three conditions.

- - before

138 The Ohio Journal of Science [Vol. XVIII, No. 4,

the results. A further explanation of this will be given later, as
the specific cases come up for discussion. Finally, the appear-
ance of the composites themselves are clearly indicative of the
nervous reaction or "temperament of the animal.

The variations within each individual are beautifully
illustrated by the polygons themselves. Beginning with
Seven H, (Fig. 8), it will be seen that a marked difference is
shown between the feeding condition as compared with those of
before and after feeding. The composite of the former lies
considerably to the right of the other two, which explains the
fact that the mean has increased from 6.41 in the before feeding
condition to 10.42 in the feeding condition, or the length of
respirations increased, which indicates that feeding acted as a
stimulus to the animal. Comparing the standard deviations, it
will be seen that the feeding condition is the more variable.
The composite of after feeding is the most uniform, the mean
having gone down to 5.34 and the standard deviation to 2.20,
the latter figure is perhaps a little high because of a few" respira-
tions of extreme length. The results indicate that during
feeding, increased nervous activity or nervous reaction accom-
panied the feeding stimulus, and that before and after feeding
the animal was more quiet and not so nervous.

The composite of Three H, (Fig. 7), shows the same results,
but not as marked as with Seven H. The mean increased
during feeding, but the standard deviation remained nearly the
same as in the before feeding condition, while after feeding it
increased slightly as compared to the other two conditions. The
stimulus of feeding upon this animal did not show as marked
effects as upon Seven H, merely increasing the mean or ampli-
tudes of respiration, and so moving the polygon a little to the
right. Not much difference is displayed between the before
and after feeding conditions, both in appearance of the com-
posites and their respective means and standard deviations. As
a whole all three composites are very similar, which leaves the
impression that Three H, is not easily disturbed and that her
nervous make-up is of a stolid nature.

In the case of subject Two H, (Fig. 6), a more evident result
is evinced, namely, that the animal has a marked tendency to
be less active before feeding than after, which was not so
marked in Seven H and Three H. The mean 6.73, and the
standard deviation 3.09, in before feeding increased to 9.09 and

Feb., 191SJ Cattle Temperament 139

3.45 respectively in the after feeding condition. The condition
of feeding also showed an increase of the mean, 9.97, and the
standard deviation, 3.79, showing a marked reaction to the
stimulus of feeding, more so than was shown in Three H and
very nearly as great as in Seven H. Consequently, the nervous
activity of Two H increased during feeding.

Finally, studying the composite of Five H, (Fig. 5), we see
that a very nervous animal is indicated, which is readily shown
by the conformation of the three polygons. Many variations
exist and the polygons appear irregular and straggling. Here
again the feeding stimulus shows its effect in increasing the
nervous activity of the animal. The mean and standard
deviation, (8.22 and 4.15), in the before feeding condition,
increased to 11.97 and 5.16 respectively, in the feeding con-
dition. Comparing the before and after feeding conditions, the
mean S.22 and standard deviation 4.15 in the before feeding
condition decreased to 7.92 and 3.09 in the after feeding con-
dition, instead of increasing as was shown in the former subjects.
This may seem contradictory, but may be explained by the
fact that the pneumographic tracings of Five H in the after
feeding condition were taken too far away from the regular
feeding time. Consequently, the mean decreased instead of
increased. Had the after feeding condition been recorded
nearer the feeding condition, the mean probably would have
increased, and so established a like condition as in the first
three animals. Taking the after feeding record farther away
from the regular feeding time, up to a certain point, the nervous
activity decreased, and with it the mean, as has been before
explained. Because of the time element then the after feeding
condition in this case would tend to indicate less nervous
activity than in the before feeding condition.

In this connection an interesting fact might be brought
out regarding all the subjects studied. Looking at all the
polygons of the feeding condition, it will be noted they extend
to the right of the others and are wider and flatter. Then
in the after feeding condition the polygons move toward the
left and become higher and narrower. And in the before
feeding condition the polygons are a little to the right of the
after feeding condition and slightly wider and flatter than in
the after feeding condition, with the exception of Two H,
where the before feeding tracings were taken farther from

140 The Ohio Journal of Science [Vol. XVIII, No. 4,

the regular feeding time. Owing to inconveniences in obtaining
the subjects at definite times of the day, it was impossible
to work with them at stated intervals before and after feeding.
But had the penumographic curves been taken at definite
times and alike for each subject, the above fact would probably
have been illustrated more clearly, and would have shown that
each individual possessed a cycle of nervous activity, being
greatest at feeding, diminishing after feeding and continuing
to diminish until the before feeding condition was reached,
and then gradually increasing in nervous activity again until
the feeding condition had been reached, when it would be at
its height, and then this cycle would be repeated as the next
feeding time approached. Each animal of course would have
a definite point, which would be characteristic of that individual,
at which point the nervous activity would begin to increase or
decrease, depending on the nervous make-up of the animal in
question. The times at which these tracings were made are
different, but the attempt was made to choose approximately,
a definite time, in so far as existing conditions would permit.

A general comparison of all the composite polygons men-
tioned may also be made. It will be seen that each individual
has its own characteristic polygon, and that each is distinctly
different from that of any other. Those of Seven H appear
rather long and narrow. (It will be noted that all the polygons
are plotted to the same scale.) Those of Three H are shorter
and broader and more irregular, while the polygons of Two H
are still shorter and broader and rather flat, with about the
same amount of irregularities as those of Three H. Lastly,
those of Five H are the most irregular of the four and do not
show much symmetry, but are spread over the entire width of
the page. Thus each animal shows its individuality in its
respiration polygons, even under three different conditions, and
the peculiar conformations and variations are distinctly char-
acteristic of this one individual and not of any of the others.
All of the composites seem to possess a common feature, namely:
the shorter amplitudes show a higher frequency than those of
extreme length; in other words, the long respirations do not
occur as often as the short ones, which gives the polygons
an abrupt slope to the left and a more gradual slope toward
the right.

Feb., 11)18] Cattle Temperament 141

Variations Within the Group. — In making comparisons of
variations within the group of animals studied, the composites
of all the animals under a given condition were plotted on one
sheet, thus facilitating the interpretation of results and making
the comparisons more vivid. The following pages then repre-
sent the four animals under the same condition, i. e., before
during and after feeding. Studying the polygons of the before
feeding condition, (Fig. 4), it will be seen that each individual
is represented by a distinct composite different from the others.
The general conformation and appearance of each would lead
to the conclusion that Seven H is the most quiet and Five H
the most nervous, and Three H more nervous than Seven H and
less so than Two H. In the order of their nervous activity
then, they follow. Five H, Two H, Three H, and Seven H, as is
indicated by the height, symmetry and narrowness of each
polygon. This interpretation based on observation is entirely
in accordance with the actual results. The mean and standard
deviation of Seven H is G.41 and 2.02 respectively, which
increase to 6.69 and 2.57 in Three H, to 6.73 and 3.09 in Two H,
and to 8.22 and 4.15 in Five H. A curious fact is here brought
out that the coefficients of variability indicate the same order.
They follow, Seven H, 31.59, Three H, 38.36, Two H, 46.02,
and Five H, 50.49. This is because of the fact that the means
and standard deviations do not increase in the same ratio, and
the reason they do not increase in the same ratio, no doubt is
due to the fact that the before feeding tracings of all the subjects
were taken very nearly at the same time before the regular
feeding time. (See above.)

Turning to the polygons of the feeding condition, (Fig. 2),
the interesting fact heretofore mentioned is made more striking,
that the stimulus of feeding increases the nervous activity of
the animal. Comparing the shape of the polygons, the same
order of nervous activity is seen, namely — Five H, Two H,
Three H and Seven H. The same result is verified by the
respective means and standard deviations of the respective
animals. Here again the coefficients of variability read directly
and conform to the order of nervous activity given above —
beginning with 24.76 in Seven H and increasing to 43.12. It
will be noted that the coefficients of variability in the before
feeding conditions (Fig. 4), are as a whole relatively higher

142 The Ohio Journal of Science [Vol. XVIII, No. 4,

than those in the feeding condition. This fact might be explained
in that the means of the feeding condition are in turn greater
than those of the before feeding condition, and as a result
lower the coefficients of variability in the feeding condition.
The pneumographic tracings of the feeding condition were
taken under fairly uniform conditions, because here the time
element was practically the same, since the tracings were made
at the regular feeding time. Therefore these results are very
significant and indicative of the degree of nervous activity or
reactiveness of the four animals.

Lastly, taking up the after feeding polygons, (Fig. 3), we see
the results are somewhat different from those previously studied.
Taking into consideration the conformation and appearance of
the composites, the curves of Seven H indicate the least nervous
condition and next Three H, and Five H appears less active than
Two H. Turning to the actual results of the means and
standard deviations, the explanation for the difference will be
found. Seven H has a mean and standard deviation of 5.34 and
2.20, which are increased to 7.21 and 3.04 in Three H and to 9.09
and 3.45 in Two H, while in Five H the mean and standard
deviation decrease to 7.92 and 3.09 as compared to Two H.
This may be due again to the same time element, which was
explained above, when all the polygons of Five H were dis-
cussed. It is again brought out that the tracings of the after
feeding condition of this animal, were taken too far from the
regular feeding time, and so decreased the mean and standard of
deviation. While the coefficients of variability of the before
feeding and feeding conditions were in accordance with the
order of nervous activities of the animals, in the after feeding
condition they are not. They follow. Seven H, 41.20; Three H,
42.19; Two H, 38.02, and Five H, 39.08. Now here again the
time of taking the pneumographic tracings causes this result.
The after feeding polygons were taken in the morning, following
the morning feeding, but they were taken at various times and
not at a stated interval after feeding. This probably caused a
variation of results because of the changing nervous activities
of the animals, as the after feeding condition approached or
withdrew from the feeding time.

Feb., 1918] Cattle Temperament 143

DISCUSSION AND CONCLUSIONS.

The purpose of this study was primarily to try to establish a
method whereby the nervous activity of cattle, or so-called
"dairy temperament," might be measured. The results as
given would indicate that by means of the pneumographic
tracings the various nervous activities of cattle can be measured
and placed upon a quantitative basis. Under the three condi-
tions mentioned, the results point to the conclusions that
animal Five H was the most nervous, next Two H, then
Three H, and Seven H the least nervous. All the subjects
reacted similarly, differing only in the degree of intensity of
nervous reaction. And from the degree of intensity of nervous
reaction arose the conclusions as to which animal was the
more nervous.

Throughout the discussion, the term "temperament" and
"dairy temperament" have been avoided as much as possible,
and the term nervous acitivit}^ used instead. Nervous activity
or reactiveness as shown b}^ these results, is the response to
every-day stimuli through external and internal sensory con-
nections with the respiratory center. As has been mentioned
in the introduction, temperament, by definition is a mental
condition or development. Now "dairy temperament," today
is usually defined as a "predisposing tendency to convert food
into milk." We are not satisfied that this definition has any
quantitative basis. It is simply a deduction made from a
combination of physical characteristics. Our work takes no
cognizance of physical characteristics or outward indications, it
deals only with the actual reactiveness of the animal. Physical
characteristics as applied to "dairy temperament" have not
been standardized, since they mean different things to different
men, and what one man may call a prominent eye, another may
not consider as such. It would be better then to speak of
desirable physical characteristics, as dairy form, etc., alone, and
not involve them with "dairy temperament." Instead of using
this latter term, it might be better to speak of the degree of
nervous activity or reactiveness which an animal possess.

Whether this study, if continued, will produce results of
importance remains to be seen, but it is to be hoped that an
interest in this special line of experimentation may be developed
that will pave the way for further investigation. The conclu-

144 The Ohio Journal of Science [Vol. XVIII, No. 4,

sions drawn at best, can be but tentative, since but few animals
were studied. To come to definite conclusions further study
is necessary with more data. Only when several hundred
animals have been investigated, can the real value of this
method be determined. We would suggest that provisional
measures of "temperament " may be obtained from the standard
deviations of frequency polygons taken from several animals
which are studied at the same time under the same conditions
(preferably while feeding). Correlations between the "tem-
perament" or reactiveness of animals and their milk producing
abilities would, it seems to us, yield definite information as to
the value to be placed on the "temperament" or reactiveness of
dairy animals.

BIBLIOGRAPHY.

1. David Low — Domesticated Animals of the British Isles. MDCCLIII, p.-cvi.

2. E. L. and J. N. Sturtevant— The Dairy Cow, 1875, p. 85-92.

3. Guenon on Milch Cows — Orange Judd Co., 1888.

4. Ripley and Dance— Am. Cvc. D. Appleton Co., N. Y., 1876. Vol. XII.

5. Henry Davis— Philosophical Review, 1898, Vol. VII. pp. 162-180.

6. Hoard's Dairyman— 1907-1908, Vol. 38, p. 1038.

1912 Vol. 43, p. 588.

1904 Vol. 34, p. 85.

1905 Vol. 35, p. 1098.

Vol. 28, p. 776; p. 391; p. 690.

1915 Vol. 50, Nov. 5 issue.

1916 Vol. 51, March 24 issue.

7. Wisconsin Farmer's Institute Bulletin— 1887, Vol. I, p. 194; 1888, Vol. II, p.

175.

8. Sir John Lubbock — On the Senses, Instincts and Intelligence of Animals.

Int. fici. Series, Vol. LXIV.

9. Wesley Mills — The Nature and Development of Animal Intelligence. 1898,

McMillan Co., N. Y.

10. Oscar Pfungst— Trs. by Carl L. Rahn; Clever Hans, 1911. Henrv Holt &

Co., New York.

11. Chas. Darwin — The Expression of the Emotions in Man and Animals, 1905.

D. Appleton Co., N. Y.

12. Wm. Wundt— Philosophiche Studien. Vol. 18, pp. 1-16.

13. C. Lloyd Morgan — Introduction to Comparative Psychology. 1894. Chas.

Scribner & Sons, New York.

14. C. Lloyd Morgan — Can Animals Reason? The Spectator. Vol. 68, p. 683.

15. I. P. Pavlov — The vScientific Investigations of the Psj^chical Faculties or

Processes in the Higher Animals. Science N. S., Vol. 24, p. 613.

16. Mind.— Vol. XVT, p. 262.

Date of Publication, February 28, 1918.

THE

Ohio Journal of Science

PUBLISHED BY THE

Ohio State University Scientific Society
Volume XVIII MARCH, 1918 No. 5

TABLE OF CONTENTS

Blakk and Jackson — The Relative Intensity of the Harmonics of a Lecher

System (Experimental) 145

Blake — The Relative Intensity of Harmonics of alLecher System (Theoretical) 163

Sterki — Inland Mollusks 169

Drake — Two New Tingids from the West Indies (Hem -Heter) 174

THE RELATIVE INTENSITY OF THE HARMONICS OF
A LECHER SYSTEM. (EXPERIMENTAL).

By F. C. Blake and B. H. Jackson.

In their fourth paper* on the free vibrations of a Lecher
system, Blake and Sheard have shown how the tone intensity
depends very much upon the edge-on distance between the
plates, but they were unable on account of oscillator deteriora-
tion among other things to summarize this relation. We have
made a careful study of the various factors that help to deter-
mine the tone intensity and present the results of our study in
this paper.

I. Apparatus.

The apparatus used was that employed by Blake and Sheard
except that the brass plates used were 1 mm. thick instead of
4.5 mm. It is shown diagrammatically in Figure 1, where the
letters are the same as in Figure 1 of Blake and Sheard's paper
IV. This necessitated making eight small holes in pairs through
the plates around the circumference through which the strings
were passed and knotted. The diameter of each hole was about
0.6 mm. while that of the plates was 5.0 cm. The total area of
the holes was thus about 0.4 per cent of the area of the plate,

* Physical Review, N. S., IX, p. 177, 1917.

145

146 The Ohio Journal of Science [Vol. XVIII, No. 5,

hence the effect of these holes could be entirely neglected, par-
ticularly since the coupling was always very loose, viz., 11 cm.
On account of the tension of the strings and the thinness of the
plates, the plates were slightly bowed, that is convex toward
each other. This effect was never greater than S parts in 1100,
hence could be neglected.

c

tH (t

-c j; „■ J

TiecBitftr LjiJ Lacher Systtm |_j i] Oscillitor
Figure 1.

11. Variable Factors.

The following variable factors required study. First,
position of the bridge for each tone; second, influence of the
spark-gap length; third, length of the oscillator; fourth, edge-on
distance, y, between the plates; fifth, change of coupling, that is,
change of the face-on distance, .r, between plates; sixth, oscillator
deterioration, that is, irregularity due to deposit of soot upon
the metal rods of the spark-gap in oil; seventh, possible change
in the sensitivity of the thermocouple even though soldered.
While any one variable was being investigated, if another
variable suffered a change it constituted a source of error for
the time being.

Since the tension on the three circuits, oscillator. Lecher
and receiver circuits, was kept always uniform, small variations
of the lengths of the circuits could either be neglected or taken
into account as needed. Again as heretofore the receiver
circuit was always just half of the Lecher circuit. The fifth
variable above, .r, has been studied, but its importance demands
a special paper.

Our method of handling the first two variables was as follows :
With a given x and 7/, and a given oscillator length, having
set the spark-gap at about the position for maximum intensity
for a given tone, a bridge curve was taken across a given peak
forward and backward. Then the bridge being placed at the
peak of a curve, a spark-gap curve was taken. Without
stopping to plot the observations at the time, the correct
position of the bridge could be readily told to well within 0.5 mm.
by mere inspection of the observations.

March, 1918]

.1 Lecher System — Experimental

147

A sample of such curves is shown in Figure 2. For the
bridge at 150 the lower of the two curves of Figure 2 was taken,
then a bridge curve was taken and the reading of the maximum,
viz., 100, was so far above the maximum of Figure 2 (lower
curve) viz., 6G, that another spark-gap curve was taken, the
upper curve of Figure 2. These two curves illustrate very
clearly and forcibly the meaning of oscillator deterioration.
Evidently with so narrow a gap, viz., O.OOOK) inch, it is easy for
carbon soot from the decomposed oil to bridge, partially or
wholly, the gap and thus decrease the intensity materially.

^B*t iSo.o

£0 -

^ .at /OS.S

60

-S^-

■B-'t /93.4S'

9

V

/O o

Length o^ S/f^^f^-g^f "t 7>t distorts

Figure 2.

The matter would be easy to handle if the gap were occasionally
wholly bridged in this way, for observations under such con-
ditions could easily be detected, and hence eliminated. If,
however, in spite of the fact that we used only filtered oil the
gap was only partially bridged by the carbon soot there does
not appear any easy way to correct for this. When for the
sake of inspection the oscillator rods were removed, the ends
that constituted the spark-gap would be covered in places with
dark spots, due to extremely fine soot being deposited in an
extremely thin layer. This matter of oscillator deterioration
would not be serious provided one could always employ a
check receiver to advantage. This was indeed done in a con-
siderable part of the work here reported. But we shall show
later on in this paper that the profitable use of a check receiver
is very limited.

148

The Ohio Journal of Science [Vol. XVIII, No. 5,

y= ^s:^<:rtt. y= 69.U^

t5

/o»

\

it- o °

So

f

Aw

8c

6o -I \ 6o

4o o- 40 -

20

Zo

J I L

I . I I I I o I I I o I I I I I L? I 1 1 J 1 1 1 1 1

(/=8.o

CAtl.

JfAsf TtA^t 9fA^t-s'nl,dlt-
J/9t /eoi- />?'*'' /73+

i\- ioV- 8V- S

-I I /o

I ° I I ° I I I .1

o o y o 'i

(meaft)

'i

^a 9S AW /«*" 9a 9S- /o^ AST 90 ^O" /CO /of 90 9S 9o 96" So 9S' 9o 9S' 9o «*" »o #*" So *»«"

ZiCn^tA o^ Osci ll^tor-

Figure 3.

March, 191S] A Lecher System — Experimental 149

Neglecting momentarily the influence of oscillator deteriora-
tion, the only way in which the third and fourth variables could
be studied was to get for every new length of oscillator both
bridge and spark-gap curves for every peak ; all this for a given //.
And then for a new value of // we had to repeat all the above
operations. Table I shows the various values of y used.
It also shows the longest and shortest oscillator lengths, the
length of the Lecher wires and of the receiver. As heretofore
these lengths are the straight-away lengths, GH, LM, NT,
Figure I.

III. Summary Curves and Tables for Variable y.

Proceeding in the manner indicated above and plotting
all bridge and spark-gap curves we obtained summarizing
curves of intensity such as we have shown in Figure 3. They
were obtained by tabulating the peaks of the spark-gap curves
for different oscillator lengths.

It is plain from an inspection of Figure 3 that the higher
tones are proportionately greater the smaller the y is, a thing
Blake and Sheard found. For the curves for the values of y
shown in the figure up to and including 30.4 cm., the funda-
mental was taken with the bridge at 150. But for values of y
greater than 15 cm. the fundamental splits up into two peaks
as the curves of Figure 4 clearly show (see also Curve II,
Figure 5). Consequently in Figure 3 for y = 18.2 cm. and
30.4 cm., the relative intensity of the fundamental is too low.
A further study of Figure 3 shows also that as y increases the
length of oscillator for maximum resonance increases. Our
results on this matter are summarized in Curves III, Figure 5.
For harmonics above the third the intensity is so low for the
larger values of y that one is not able to examine how the
optimum oscillator length varies. For the fifths the optimum
oscillator length for y = l8 is unquestionably somewhat longer
than for t/ = 12, but the variation is proportionately less than
for the thirds and fundamental.

150

The Ohio Journal of Science [Vol. XVIII, No. 5,

IV. Various Corrections.

The sets of curves for Figure 3 for different values of y
cannot be directly compared with one another for several
reasons. First, those of 7/ = 8 cm. (together with the fourth
and sixth sets shown in Table I) were taken with a different
thermo-couple from the rest. Second, the different sets were
taken often several days apart and the oscillator deterioration
as well as possible change in sensitivity of the thermocouple
needed to be eliminated or corrected for. Third, the change
in room temperature, while not great, undoubtedly affected the
dielectric constant of the olive oil used in the spark-gap and
consequently the energy as read at the receiver.

y**»-^ y^^£z

0 9^.-^
L/13.7
"K 9Z.O

= 69.1

^ri d§e T^osi ti on

Figure 4.

In the light of our experiments there seems to be a distinct
deterioration in the sensitivity of our thermocouple even though
soldered. For with a pohshed spark-gap and other conditions
the same the intensity was often distinctly less for all the tones
of a set for a given y than when the same set was taken earlier.
Any figures we could give in illustration of this fact would not
be worth much, for manifestly the other factors, such as oscillator
deterioration and temperature change, are not readily differ-
entiated from the sensitivity changes of the thermocouple.
The fact that these three factors cannot be readily differentiated
one from another shows, too, the futility, for the object we
had in mind, of employing a check receiver. Moreover, while

March, 1918] A Lecher System — Experimental

151

for the bridge curves the check receiver helped to eUminate
oscillator variation, it proved wholly useless for spark-gap
curves. The check receiver was placed, after Blake and Sheard,
near the oscillator spark-gap, but in general the maximum for
the two receivers did not occur at nearly the same spark-gap
length, hence neither curve could be used to correct the other
except by an approximation method of some sort. Hence we
abandoned its further use. Accordingly, the only safe way to
make a direct comparison of the curves of Figure 3 was to get
some one tone, say the fundamental, to a common basis.

y

IS- 4S 65^
If tn c/rt.

^

i^^

-

o

a

jr ^

p^

—

o

, H

^^^

III.

fis

S 7 9 ^f

"ill

Figure 5.

Before describing the manner of reducing our corrections to
a common basis it is well to point out first, however, that the
optimum spark-gap length varies with the frequency-number
just as Blake and Sheard found (see their Figure 12). Curve
IV of Figure 5 summarizes roughly this relation as the mean
value for all the ys used. The optimum spark-gap length
for a given tone is not independent of y, however, as Curve V
of that figure shows for the fundamental. It is clear from
Curve IV, that the optimum spark-gap length decreases with

152

The Ohio Journal of Science [Vol. XVIII, No. 5,

the frequency numbers in general, reaches a minimum for the
third harmonic, and rises again for the fundamental. Curve
V shows that the optimum spark-gap length for the funda-
mental increases with increasing 7/ in a linear ratio. This is
probably true for the higher tones also, though the variation
is so little outside of the limits of experimental error that it
would be difficult to show this for the higher tones. Curves
IV and V show the absolute need for taking spark-gap curves if
the tone intensities are to be compared.

Figure 6.

To eliminate the various curves discussed above and to get
at a common basis for comparison of the various sets of curves
for the different values of y we proceeded as follows. Starting
with a given oscillator length GH, and a given total length GT
of the system. Figure 1, the value of y was changed from large
to small values by steps in as short a time as possible. For
each y we located the fundamental peak or peaks and then took
spark-gap curves. In this way Curve I of Figure 5 was obtained.
The curve is extrapolated to zero for y^b cm., for at this distance

March, 1918] A Lecher System — Experimental 153

the inner edges of the plates would touch. We then reduced
all the summarizing sets of Figure 3, together with all the other
sets not shown, to correspond to the standard for the fundamental
for 7/ = 18.2, viz., 142 (see Figure 3). In this way the curves
of Figure 6 were obtained. They certainly cannot be much in
error.

The curves for the ninth and eleventh harmonics represent an
average set of values since not every peak was investigated for
those tones for every change of oscillator. The other curves of
Figure 6 give the fundamental and the thirds, fifths and sevenths
on both sides of the fundamental. By taking the average of the
two thirds, the two fifths and the two sevenths, the damping is
eliminated and the results can be compared directly with the
fundamental.

V. The Question of Damping.

Before making the comparison it is well to ask two questions.
How is the intensity of a peak affected by the fact that it is
split in two? Is the damping sufficiently small that by taking
the arithmetical mean of any two corresponding tones equally
distant from the fundamental it can be eliminated? We
proceed to the answers to these questions.

For the results shown in this paper the coupling, x , was kept
constant at 11 cm. But we have shown before that for x = S
cm. and less and for small values of y the fundamental is
split up. Evidently, then, an increase of y acts like a decrease
of .r; it causes two periods for a given tone. In our arrange-
ment we detect these two possible periods (or wave lengths) by
means of a sliding bridge. Manifestly, when one period is
present, the other is absent, so far as the receiver is concerned.
In other words, all of the energy is in each of the two peaks of
the fundamental.

The second question, it would seem, can be answered in this
way. The fact that the peak on the oscillator side of the
fundamental is higher than the corresponding peak on the
receiver side of the fundamental makes it plausible to say that
if 7o is the intensity at the oscillator spark-gap say and / the
intensity at some point along the system the equation
/=7„e~" would hold where a is the logarithmic decrement.

154 The Ohio Journal of Science [Vol. XVIII, No. 5,

Whence for two different distances .ri and ,to we can say
7i = l^Q-""^ and /o = Le""'-^ whence

loge /1//2

a

.X-o-.Vi

Now for the third harmonic .r2 = ^^ and .ti = -^^ whence

4X3 1 ,. 2X3

.T2 — .Ti = X3. Similarly for the fifths and sevenths .vo — .Ti would
equal X5 and X- respectively. The ratio — ^ for the thirds is

= 1.099. For the fifths and sevenths this ratio is

101

^ = 1.044 and 114 = 1-017 respectively.
36.9 11.2

Using the figures for Xg given for 7/ = 8 cm. viz., X3=145,
X, = 86, X7 = 61, we get a to be 0.00065, 0.00050, 0.00028 respec-
tively. Thus the value of a apparently decreases for the higher
harmonics. It may fairly be questioned, however, whether
the question of damping can be treated as simply as here given.
Using the values of Xs corresponding to the optimum values of y,
which we shall call y^, viz., X3=150, Xs^^S? and X7 = 61, changes
the values of a but little. Now Blake and Sheard* have shown
that the electrostatic leakage, which they called "^ (z/) " was
greater for the lower tones than for the higher tones. This
would readily explain the larger values of a for the lower
tones.

The important thing at present, however, is that in any case
the value of a must be very small. Even though we measured
X, not from the oscillator spark-gap, but from the Lecher plates,
the value of a would be doubled only. There seems no good
justification for doing this, however.

For the third harmonic, using the value of a first given,
0.00065, and calculating the intensity / of the third at the
middle of the Lecher system we get 7=105.9. The arithmetic
mean of 111 and 101 is 106.0. This is the justification for
taking the arithmetic mean of the two tones either side of the
fundamental to get the tone intensity of any tone that can
fairly be compared with the fundamental. The question of
damping is thereby eliminated.

Physical Review, 1. c.

March, 1918] A Lecher System — Experimental 155

It seems very likely that the question of damping can be
handled thus simply for the cases here considered. Blake
and Sheard have shown that we are dealing here, when the
coupling is loose (.r=ll cm. is a very loose coupling), with the
free vibrations strictly and since our system is a strictly non-
radiant system the damping must be very small. It is perhaps
an open question whether the logarithmic decrement thus
measured was the decrement of the receiver merely or of the
total oscillating system. Likely the former, however, since
the oscillator and the distance y was purposely changed so as
to have the energy at the receiver the maximum possible for
each tone, with a given constant input of energy at the oscillator
spark-gap.

YI. Relation Between Tone-Intensity and Edge-on

Distance.

Thus the energy at the receiver is distributed among the
various tones as follows, the figures being read off directly
from the curves of Figure 6: fundamental 172, third 106, fifth
37.7, seventh 11.3, ninth 3.8, eleventh 1.8. Expressed in per
cent, the figures run 100%, 61.63%, 21.92%, 6.57%, 2.21%,
1.05%. With the more complete data given in this paper we
have thus been able to determine quantitatively the influence
of the edge-on distance between plates upon the tone-intensity.
Blake and Sheard were able to give a qualitative answer only.

If we plot the optimum distance between plates, t/s, against
the frequency- number, ^ve get Curve III of Figure 9. With
an average error of less than 1 per cent, and a maximum error
of 1.7 per cent, this curve gives the empirical relation

\ ~s~ loge 7/s = Constant.

We are not prepared at present to state whether such a
relation has an}^ important significance. Calculating the
distances t/s required to fit the formula we get the following
values: 36.8, 15.5, 11.1, 9.15, 8.00, 7.25. In the curves of
Figure 6 the short lines parallel to the y axis cutting across the
curves are shown at these points. Manifestly, within the
limits of error, they cut at the top of each maximum.

in

00

0.00
+0.02
—0.01
—0.05
—0.24
—0.48

0.00
—0.07
+0.05
+0.02
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+0.19

c: 00 r^ X O'^
o c: ci ca (M cj

d d d d d d

—0.53
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—0.12
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e] t^ ue -rf ei I
'— ei aD ic ei

^* o' o ^ ?■ J

1 I+++

1
1

1

C4

.H s^ |cr-'rfcec:cc

1^1 + M M

0.00
— 0..58
+0.88
+0.44
+3.28
+8.40

0.1)0
+0.92
+4.22
+6.21
+8.30
+ 11.1

—3.91
—1.51
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+33.0

—4.98
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+7.18
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—11.1

—3.33
+ 12.4
+42.1
+ 121.2

i

23

theor-
etical

14.60

t^

o
cc

e-1

12.24

12.00
11.79
11.50

OO

o

22

Is

theor-
etical

C^ Cvi C-l rj cc t^

cc -^^ cc CD *#_ CC

CD- CC d ic cc C'J

0-. CI —

l^ o =; cc "t t~

•* C^ CD cc CD --r

CD* CC d uc CC ci

97.24

24. .51)

11.11

6.18

3.84

2.62

97.15

24.74

11.31

6.31

3.97

98.25

25.23

11.65

6.54

4.15

98.10

25.33

11.66

6.59

4.22

M

theor-
etical

442.90
437.43
430.05
423.29
417.87
413.38

t^ c) o r^ cc o

c: -^ cr: cc CD CD

d »ic x ci r-^ cc

M- cc Cl Cl — —

-* -^ •* -^ ■* ^

cc S CD C-. cc cc t^ •* CI c: CC

d CD d -*- d CD- X- -^ d CC d
•^ cc cc CI ct — cc cc CI CI ^-

-rf ^ -rJH -rt* ^ M- -rj- tT -^ •* '^

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re — 5 ^. ^ 1 CD CO ei q co
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•* ce ce c^i ei ce re ce ei ei

^ ^ -TfH ^ TP -rj- Tf '^ -<t"rf

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O X c^

^ a> E

o ^ cc CJ — —

cc CD If: 'rj- t^ CD

-* -^ -^ ''J" cc —

oocjccic»c cc^frcC''*' — 00
tCcoocw— oc = — ^, oc o Tj;

cccccccc-^-^i^ "2— — — —

cccD'^'^o cexei'^ce 1
oco'-ob- coo — »oce
ei c^i CO -^ CO •— ei re -^ >je [

t^ O c: lO ■*
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d — cci^

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d d "rr r-^ c; — co

o

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s

experi-
mental

.96.32

23.40

10.32

5.66

3.64

2.82

t~^.f;-*>re= ^^occicc-- u^mujuc^ |^cjuc^uc

T». —. CD 00 -O cc C] -a- 00 C: CD cc — t^ =: QC — _ =i ^ ~; ^. •*.

cd' cc d ic cc ci t-: ^- d xc cc ci t-^ ■* — lo cc cc S H "-' "

c: CJ — o: o — ' c^i CI — C: CI —

98.10

25.. 53

11.11)

6.18

2.14

99.16

25.86

11.13

5.11

100.11

27.15

11.04

4.71

101.58
26.63
10.63

1— t
CO

00

experi-
mental

OV UC C = = ^ C — — =-;-• cc

c-J t^ d CC t^ — - d "^ d ci :y:' '^

rrcccCCl — — -^CCC^Cl — ^-

440.36
436.80
431.90
426.16
421.38
417.34

436.38
433.95
430.50
426.09
423.00

xxC"*— !if:ejo*cfM

ei ^ o ^ S 1 t- ^^ q CO CO

c: r^ »c — " CO' 1 -^ tc »c ce —
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437.01)
439.80
440.15
438.90

439.50
443.76
445.35
442.75

444.65
449.70
454.75

CO

od

5

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246.32

246.47

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ei

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250.11
251.58

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u

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37.40
37.58

37.67
37.60

^ X

ce CO

u»

t4

cs

46.34
46.43

46.46
46.41

46.82
46.70

47.00
46.59

47.40
47.05

47.98
46.94

S

t^

60.44
60.47

60.42
60.41

60.88
60.71

60.87
60.56

61.62
61.14

61.95
61.01

62.70

cqcc

CD CD
CC CC

CO
CD

•

s

«

(86.12)
86.00

86.01

85.82
85.78

(86.42)
86.36

86.13

86.10
85.84

(87.19)
87.00

86.60

(87.42)
87.00

86.44
88.03

— CO

rlicq

o

ci
X

5^

d

n

s

cc

145.84
145.81

145.04
145.14

145.60
145.48

144.65
144.82

145.86
146.04

145.14

145.54

146.60

4or^

COCJ

cc •^-

liOiO

o

coco

-^_C5

oici

oi

in

ll

442.90
442.90

440.07
440.07

440.36
440.36

436.38
438.72

43g.28
441.31

434.75
439.63

437.00

CO

in

CO

S

z

o
g

s

n

Z
<

11

(g4.60)
150.00
206.10

131.05

168.72

131.31
169.25

A

o

80.53
150.00
219.54

126.74

173.21)

79.90
103.00

173.35
196.75
220.36

126.50
173.50

102.65
126.34
173.70
197.44

126.17
174.15

CO

i^

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119.75

180.17

89.14
119.61

180.50

210.91

119.57
180.44

88.37
119.21
180.75
211.62

119.10
181.05

118.80
181.50

118.50
181.75

t-

to

30 o o c — _

cc t-^ d cc CO

cDO »oc; cc

107.08

192.91)

63.66
106.83

193.19

236.50

107.00
193.10

62.77
106.44
193.44
237.15

62.75
106,72

193.72
237.60

r- o

^ ei

it

105.71
194.78

104.75
195.70

<s

cc

77.05
1.50.00
222.89

77.51)
222.54

77.28
222.88

77.65
222.30

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1^ ci

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222.68

X Tj;

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t^Ci

'^ ce
CO ■<l«

oo
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70.74
230.18

m

7

150.00
150.00

150.09
150.00

CTS

ci

d

i

1 ^

O

ue

o

149.15

and

151.25

•»>

Oscil-
lator
cm.

103.7
to
90.1

102.2
to

88.2

102.4
to
90.0

110.0
to
86.4

102.1
to
89.0

106.3
to
88.3

102.1
to
89.4

100.6
to
89.1

105.4
to
94.0

in c

3s

M

Re-

:eiver

cm.

90.2
89.9

90.2

89.7
90.2

»je

ci
X

§

o
o

Oi

O
Ci

C4

Lecher

Wires

cm.

5S

X

cr.

180.44
179.41)

179.20
180.20

o

o'

X

X

o
oo

o

X

vH

'

7.15

o

00

9.0

10.0

:^

12.0

13.1

15.1

CI

00

o

CO

March, 1918] A Lecher System — Experimental 157

VII. How Does the Capacity of the Coxuexsers Vary
With the Edge-on Distance Between Plates
AND With the Plate Thickness?

Blake and Sheard found that for small values of y the value
of ho, K, that is, the end-capacity expressed in equivalent wire-
length, was practically constant, as calculated from the data for
the different harmonics. This was to be expected on account
of the small range of frequencies they worked with, viz., 1 to 9.
The range in this paper is but slightly greater, viz., 1 to 11,
hence we should expect to confirm their results in this regard.
This is indeed the case as Table I clearly shows. Columns 5
to 10 inclusive give the mean positions of all the peaks for all
the tones. The upper set of figures in columns 11 to 16 inclusive
give the experimental values of Xs, the wave-length correspond-
ing to the frequency number, s. In column 13 the figures in
parentheses are the observed values of the wave-lengths of the
fifths obtained by subtracting the two outer fifths from each

other and dividing by 2, e. g. 236.12—63.88^^^-^2. Blake and

Sheard observed that the outer internodal spaces for the fifths
were distinctly larger than for the inner internodal spaces.
Since they did not observe this for the sevenths and ninths
within the limits of experimental error they attributed this to
the presence of the small ebonite supports, Ri, Ro, R3, R4 in the
immediate vicinity of these outer peaks for the fifths. Our
results confirm theirs. In the calculations the outer fifths were
not used. We found, however, that the intensity of the outer
fifth near the oscillator is not as great as that of either of the
two inner fifths, whereas it should be somewhat greater. This
was true for all the values of y for which the outer fifths were
measured. Column 17 gives the position C, Figure 1, obtained
by adding 3.5H-1/2 (y — 2) to the plum-bob reading M as given
on the 3-meter stick. It is the reading of the back of the
Lecher plates next to the receiver. Column 19 is calculated in

158

The Ohio Journal of Science [Vol. XVIII, No. 5,

the manner indicated in Blake and Sheard's paper
Column 20 is obtained by means of the formula

Xs 27r

:q. v.).

Ko
K

tan

/2^

The first figure in each set in column IS is obtained by extra-
polation as given in Figure 7, full lines, and Figure 8, where the
points plotted are the remaining figures in column 18. Naturally
there is some degree of latitude in making this extrapolation.
For instance, one could so change Xi, in the sets mentioned
in the accompanying note, as to leave the first value in column
20 unchanged from the figure given in the table. We believe,
however, that the reader will agree from an inspection of the
curves of Figures 7 and 8 that the extrapolated values for Xi
given in the table are not greatly in error.

Column 23 was obtained in the following way. The first
set of column 20 is a decreasing set, the second an increasing
set but in each case the first four figures are fairly constant.
Using the first four figures of each set and allowing for the slight
decrease and increase respectively the first two figures were
taken as given in column 23. With these two given, the remain-
ing figures in that column were obtained by graphical methods
by assuming that Ko/ k varied with y according to the relation

1

loge

d+Vd^—b^

where d is equal to ^ and b is the radius of the plates,

viz., 2.0 cm. Thus all the figures in column 23 except the
first two may be said to be extrapolated from them. Columns
21 and 22 are similar to columns IS and 19 respect-

* It is to be noted that in column 5 the fundamental maximum is not exactly
at 150.00. For values of y less than 12 cm. this is to be explained by slight
inecjualities in the lengths of the wires, e. g., BM, HK, etc., leading to the plates
together with slight inequalities in the couplings at the four sets of plates. For
values of y = lo cm. say and above, at which the fundamental tends to split in two
the maximum is always at a length slightly greater than loO as the dotted line of
Curve II, Figure 5 shows. In column 19 the value of Is given for the fundamental
is given as if the peak were exactly at 150, i. e., l = p-150. If the value of the
position of the fundamental given in column 5 were used U would be slightly dif-
ferent from the figure given for it in column 19, being in general slightly less than
such figure. This would tend to raise slightly the value of the first figure in column
20 for sets 3 and 6 to 10 inclusive and to lower it in set 5.

March, 1918] .1 Lecher System — Experiniental

159

ively using the figures of column 2o instead of column 20.
In column 21 the first figure in each of the first three sets
is arbitrarily taken equal to the first figure in column 18,
the first figure in column 22 being necessarily equal to
that in column 19. The justification for this may be seen
in Figure 7 where columns 18 and 21 are plotted against
the frequency number for the first five sets, column 21
being represented b}^ the dotted lines. For 7/ = 8 the agree-

'i-foltne far y=0^n<i9rtn.

^O line ^or tj=^ ^/.Si/irr.

■^^O itfte /ory-/a.Or/rr.-

— ^■fOytrlfs/Uti »

160

The Ohio Journal of Science [Vol. XVIII, No. 5,

ment between the two columns is seen to be very good and
fairly good for r/ = 7.15, the variation being in the opposite
direction. For 7/ = 9 the variation is greater and in the same
direction as for z/ = 8 cm. For 7/ = 10 and 11 cm. it is plain that
the theoretical and experimental curves can no longer be
extrapolated to the same value. Plainly the agreement gets
worse as y increases, due to the lack of constancy in column 20
for a given set. In column 24 is given the percent, of error
in column 20 as against the value given in column 23.

4SS,

The lower set of figures in columns 11 to 16 inclusive is
obtained by calculating Xs from the equation given above,
using the values for Ko/ k given in column 23. Since the equation
is transcendental the method of approximation has to be used.
Multiplying this As (theoretical) by *s' gives column 21. Com-
paring the two sets of figures in columns 11 to 16 inclusive we
get the per cents of error given in column 25.

March, lUlS] A Lecher System — Experimental

161

Remembering that the thing that is experimentally deter-
mined is Xs the percents of error in column 25 are remarkabl}^
small. x\veraging for all the sets we may say approximately
that one per cent, variation in Xs with its consequent variation in
/s according to the formula

= />— I 15

L = p

O + Cf— a)

where .v is the frequency number 1, 3, 5, 7, etc., and a the
corresponding natural number 1, 2, 3, 4, etc., changes the
value of Ko/ K for values of s^\, 3, 5, 7, 9, 11, respectively
8, 11, 15, 22, 32, 44 per cent. This serves again to emphasize
the importance of an accurate measurement of Xg.

^' ^ ^

S

7 9

//

1

\ \

1 1

1

30

A

^

\

20

- \

/6

_ 0 °N. 2E-

/o

—

_ij

/4

-\^^

"0 — -

^0— .

~- 0.

JA

1 1

1 1

1

-o

1

wV

r 9 f3

y

/7 21

XS

29

Figure 9.

We have gone into the method of these calculations with
some detail because of the dependence placed upon the figures
in the theoretical paper by one of us that follows this paper.

Two things stand out from this discussion of Table I.
The value of Ko/k is very constant for small values of y\ and a
small error in the wave length determination makes a large
error in the constancy of Ko//c, the errors being larger propor-
tionately for the smaller wave-lengths. The first point was
made by Blake and Sheard and we thus confirm it. For the
first three sets of the table the average error in X^ is only 0.13 per
cent.

162 The Ohio Journal of Science [Vol. XVIII, No. 5,

Blake and Sheard showed that the inconstancy of Ko/ k
for larger values of y was due to the effect of the phase changes
introduced by the factor which they called "0 (t/) " due to the
fact that a portion of the wires was at right angles to their main
lengths and perhaps to the fact that at points like B and K,
Figure 1, there is a change of diameter of the wire where it
enters the small rod to which the plate is attached. It is
conceivable that the reflection coefficient at such points is not
necessarily negligible for the higher harmonics whereas it might
be for the lower tones. The greater y is the larger the phase
changes for the higher harmonics in comparison with the lower
harmonics. These phase changes act in the same direction as
the phase change y due to the end capacities, hence the apparent
Ko/ K would be larger for the higher tones.

If we plot the values of Ko/k given in column 23 against y we
get Curve I of Figure 9. For the sake of comparison the values
Blake and Sheard obtained for plates of the same diameter,
but of thickness 4.5 mm. instead of 1 mm. are shown in Curve II.
Thus, the extra capacity of the thicker plates for the coupling
11 cm. is about 10 per cent.

Summary.

We have determined experimentally the relation between
the tone intensity and the distance between the plates edge-on
and have found the optimum distance z/s for a tone of a given

frequency number .j to fulfill the empirical relation z/s = Ce — ^ s
A theory to account for the tone-intensity as a function of the
phase change y and the wave-length X is given in the next
paper.

We are indebted to Mr. B. F. Salisbury for assistance in
taking the observations and to the University for further
financial assistance.

Physical Laboratory, Ohio State University.

THE RELATIVE INTENSITY OF HARMONICS OF
A LECHER SYSTEM. (THEORETICAL).

By F. C. Blakk.

In the previous paper Blake and Jackson have shown the
dependence of tone-intensity upon the edge-on distance between
plates. They have found that the optimum condition for each
tone is such that, for a given in-put of energy at the oscillator,
the energy is distributed among the various tones as follows,
the fundamental being taken as 100%: third, 61.63%, fifth
21.92%, seventh 6.57%, ninth 2.21%o. eleventh 1.05%. In
this paper a theory is given to account for these results.

Figures 1, 2 and .3

The theory makes use of the Kirchhoff- Abraham generaliza-
tion of Kelvin's formula for the discharge of a condenser. As
given in Abraham's "Theorie der Elektrizitat," Vol. II, 2nd
edition, we consider electric waves coming from the negative X
direction (Figure 1) along two parallel wires that end in a con-
denser. Our experimental arrangement was necessarily dif-
ferent from that, but if we fix our attention upon the receiver
alone and remember that the optimum oscillator-length is
different for each tone it is obvious that, so far as the total
energy on the receiver is concerned, our arrangement (Figure 2)
is identical with Figure 1.

163

1G4 The Ohio Journal of Science [Vol. XVIII, No. 5,

Later in this paper we shall discuss the conditions under
which the practical case, Figure 2, may be said to conform to
the ideal case, Figure 3. For the moment we shall call them
identical.

Abraham's expression for the potential difference, 0 per
unit length of the wires (Figure 1) is

27rX \ o *
T ) cos ZTT 7lt

where A is a constant, k the capacity per unit length, X the
wave-length, 7 the phase change due to the end-capacity and n
the frequency. We consider the total energy on the receiver
that surges through the thermo-couple as made up of two parts,
that on the wires themselves and that on the condenser. Sup-
pose at a certain moment the condenser plates are charged to
a maximum value. Due to the distributed capacity of the wires
there is also at that moment a charge on them. A moment
later these charges discharge through the thermocouple thus
recharging the plates and wires with electricity of the opposite
sign. During the half period of the galvanometer, viz., 1.4 sec,
millions of vibrations surge through the thermocouple. The
galvanometer needle moves off until the loss of energy by heat
conduction and radiation from the thermocouple equals the
input of energy. The scale usually moves off in a very vigorous
fashion showing that the losses of energy are not appreciable
till near the end of the half period of the galvanometer. We
shall assume the rate of loss of heat energy by radiation from
the junction to be independent of the frequency of the tone
surging through it, and that Newton's law of cooling holds
with respect to the surroundings.

In order to calculate the energy that surges through the
thermocouple we must get the root-mean-square value of the
potential as it is distributed both as to space and time. If we

substitute — for t in the above expression we can say that

the total energy on the wires is given by the expression

n 0

2 2 \4 27r/

u

27ra;L ,
cos^ I dx

1

March, 191S] A Lecher System — Theoretical 105

The distance from the thermocouple to the back of the
4 ::;7r

receiver plates is ^.

4. Vir

Hence the total capacity of the wires is k( ^ —- j

The energy on the plates is represented by
1^..., 1 \2A y\ V.s

The condenser is at the point x — o. In the above expression

•> J

for 0 put x = o and we have </> = "' cos 7 per unit length.

K

The equivalent wire-length of the condenser is ^ .

But the plates are not charged all the while, hence V^ in the
expression for the energy on the plates must be taken

V- = — - J—~^ cos^ 7 • — 1 cos-j/ tdt.

K- ■iir- T -'

Now — ( cos2j;tdt= cos-xdx = —.

tJo vtJ o vt

But the plates are charged n times a second, and since
v = 2irn and 7/r = l, multiplication by n gives ~ = — ^

where no= frequency of the fundamental tone and s the
frequency number.

Integrating the above expression for the energy on the wires
we get as the expression for the total energy E,

+ 3 sin 7 cos'' 7 (l-fsin-7) — 5 sin5 7COS7 + -I sin 7 cos 7 > I .

Now the constant A depends upon the manner of setting
the receiver into vibration. Our case is similar to the acoustical
case treated by Lord Rayleigh.* The disturbing force which

varies as cos '^^-^ is not applied at a single point, but is dis-
\

tributed over the distance ^ . The disturbing force over
one-half of this distance concerns itself with the Lecher system.

* Theory of Sound. Vol. I, p. 189.

166 The Ohio Journal of Science [Vol. XVIII, No. 5,

That over the other half has to do with the receiver. Accordingly
we can take
rt2y\

A = 4"

2-

cos — ox = — ^ sm 7 cos 7 and A- = — ^ .

X 2x 4:Tr

t' 0

4 ' n

Call ° ° = C. Then our expression for E becomes

E = C X* sin-

7COS^7[-^7^^cos-^7^''+|('-,''-7)|(^-7)(2cos27+l)

+3 sin 7 COS''' 7 (1 + sin- 7) —5 sin^ 7 cos 7+4 sin 7 cos 7. > I .

Now Abraham's theory for the transmission of electric
waves along a pair of parallel wires does not take into account
the necessary bending of the wires leading up to the plates.
For obvious reasons the ideal case shown in Figure 3 cannot be
realized unless one has a rather large distance between the
Lecher wires, which means a long bridge and larger phase-
changes due to the bridge length. Blake and Sheard have
shown that there are two factors, which they called 6 (y) and
0(7/) controlling the relation between the edge-on distance
between the plates and the tone intensity. The former rep-
resents the electrostatic leakage to plates of the same circuit,
the latter the phase-changes due to that portion of the wires at
right angles to their main length. They have shown that these
two factors act in opposite directions and the preceding paper
confirms this. Abraham's theory does not consider either of
these factors. We have m.ade no attempt to determine the
nature of either of these functions of y, (though the exper-
imental data of Blake and Jackson are probably sufificient to
determine both). They have determined, however, the opti-
mum value of y for each frequency, that is, the value of y at
which the tone intensity is a maximum. Obviously, this
optimum value of y is that value at which these two factors
nullify each other.

If then we compare the tone-intensities for the various
optimum values of y they should agree with the simple theory
of Abraham that neglects the two factors mentioned above. In
making the theoretical calculations, however, some circum-
spection must be used. For instance, theory and experiment
should not be expected to agree except at or near those values

March, H)1S|

A Lecher System— Theoretical

!()■;

of 7/ where Ko/ k is perceptibly constant, at least so long as the
forms of d{ij) and 0(7/) remain unknown.

Now in the work of Blake and Jackson the per cents of error
were least for // = 8, as column 25 of their Table I shows as also
does their Figure 7. We have carried through the calculation of
E for the following values of jj: 7.15, 8.0, 9.0, 11.1 cm., and
have collected the results into Table I.

Table I.

y

Xs

Ko/k

7

EX10-«

Arbitrary

E

Cm.

Cm.

Cm.

Degre

es Min.

Units (C=l)

Percent.

442.90

11

42.1

3663.2

100.00

145.81

32

10.6

2143.6

58.. 52

7.15

86.01

14.60

46

50.8

692.4

18.90

60.47

56

36.5

209.0

5.70

46.43

63

9.3

71.47

1.95

37.58

67

43.3

28.77

0.78

442.90

14.60

100.00

145.84

14.64

58.56

7.15

86.00
60.44
46.34
37.40

14.58
14.42
13.71
11.61

18.89
5.76
2.12
1.13

440.07

11

4.1

3107.3

100.00

145.14

30

40.3

1921.0

61.82

8.0

85.78

13.70

45

6.0

683.2

21.99

60.41

54

56.3

220.6

7.10

46.41

61

40.1

76.64

2.47

37.60

66

24,3

32.33

1.04

441.00

10

37.8

2838.8

100.00 -

145.62

29

36.5

1823.5

64.24 .

9.0

86.19

13.17

43

50.0

691.5

24.36

60.74

53

43.3

234.5

8.26

46.71

60

33.4

86.20

3.04

37.86

65

24.9

36.07

1.27

440. 50

13.06

100.00

145.60

13.15

05.56

9.0

86.36
60.88
46.82
37.94

13.58
13.84
14.21
14.48

25.32
8.23
2.24
1.12

440.28

9

39.2

2235.6

100.00

145.68

27

12.5

1551.5

69.41

11.1

86.45

11.92

40

54.3

677.8

30.32

61.07

50

48.4

257.5

11.52

47.01

57

53.1

98.38

4,40

439.28

11.63

100.00

145.86

12.08

74.60

11.1

87.00
61.62

13.12
14.54

34.09
10.97

47.40

15.33

3.64

168

The Ohio Journal of Science [Vol. XVIII, No. 5,

The agreement between theory and experiment is remark-
ably good for 7/ = 8.0 cm., or for the mean values for 7/ = 7.15,
8.0 and 9.0 cm.*, as shown in Table II. Expressed in per cent
the largest error is nearly eleven per cent, but when, as here,
relative intensities are compared, it seems far more reasonable to
express the error in terms of the fundamental intensity. When
this is done the maximum error is less than one-half per cent.
In the experimental work, although we tried to read to fractions
of a division for small galvanometer throws, nevertheless such
things as the wandering of the zero during a reading, slight
unsteadiness of the zero particularly on windy days, together
with slight errors of experiment, mentioned above but unelim-
inated, served to make the experimental error as high as five
per cent, at least for the tones beyond the fifth.

Table II.

y

ENERGY IX PER CENT

cm.

S=l

3

5

7

9

11

7.15

100

58.52

18,90

5.70

1.95

0.78

8.0

100

61.82

21,99

7.10

2.47

1,04

9.0

100

64.24

24.36

8.26

3.04

1.27

Mean

100

61.53

21.75

7,02

2.49

1.03

Experimental

100

61.63

21.92

6.57

2.21

1.05

Expressed in per cent of the fundamental intensity the error
is thus small and we can accordingly say that a satisfactory
theory has been worked out to explain the observed curves of
Blake and Jackson's Figure 6.

* The value of Kq/k used in Table II is not exactly the value given in Table I
of Blake and Jackson's paper. The calculation was made taking ko/k = 13.17 cm.,
instead of 13.03 cm. This can affect our results but very little, for a change of
1 per cent in Ko/k with its consequent change in X affects the relative intensity
of the various tones from one to three per cent at most. The variation is not
always in the same direction for the different tones, however.

Physical Laboratory, Ohio State University.

INLAND MOLLUSKS
Directions for Collecting and Preparing.

By V. Sterkf.

Our land and fresh-water mollusca are very interesting
objects of study, not only for their anatomy, physiology, life
history, habits and classification, but also for their geographical
and local distribution and their variation.

In Ohio, we have over three hundred species, with many
varieties and local forms, but from a large part of the state the
fauna is still insufficiently known. Good work can be done
anywhere, toward an approximately complete survey. Besides,
in working up the ecology of various places, the mollusks
deserve consideration as well as any other group of animals.

LAND SNAILS.

These are found living in woods, copses, along old fences
and railway embankments, swamps and marshes (especially
their borders), banks of rivers and creeks above high water
mark. They are generally more common on hillsides than on
low ground and rocky slopes, especially of limestone, are often
wonderfully rich. During the day, in fair weather, they are
generally in their retreats, coming out at night, in the mornings
and evenings and often on damp, dark days and after rains.
Occasionally they ascend trees and other plants. As a
rule, they will be found under and about old logs and stumps
and in their cavities, under loose bark, pieces of wood, stone
slabs and bricks, in crevices of rocks, old stone heaps, etc.
Many are on the ground, under dead leaves and among thick
plant growth. Some Succinea are frequently found along
ditches, ponds and lakes, and often on reeds and rushes standing
in water.

During winter, they hibernate, either in the same places,
or buried in soft earth, muck, etc., often assembled in large
numbers.

A very handy tool for collecting is a small, strong hoe,
especially the "Ferris hoe." The blade of a garden hoe is
trimmed to a point in the middle and thus made heart-shaped,

1 69

17U The Ohio Journal of Science [Vol. XVIII, No. 5,

and sharpened, and the handle is sawed off to about three feet.
For collecting under dead leaves, etc., a small rake is helpful and
large stretches can be raked over in a short time.

The snails, as they are gathered, are put into a box or can,
with some moss or leaves at the bottom. It is a good plan
to have the box covered with canvas or soft leather, or still
better, with a thin rubber sheet such as a piece of an old tire,
with a slit in it. The snails can be put in easily and are pre-
vented from crawling or falling out, and there is no bother in
removing and putting on a lid. If the box is fastened to the
belt, both hands are left free for work.

Snails with very thin, fragile shells, e. g. Succinea, should
be kept separate, lest they be broken. The smallest should
be put into a wide mouthed bottle or vial or better in a wooden
pill box.

For a longer trip, in diversified country, several receptacles
should be provided. The specimens from a hillside should not
be mixed with those from lowlands, or from a swamp, etc.
To each collection a label should be added at once, noting the
locality, its nature, etc., and the date.

Naked or shelless snails or slugs should not be overlooked,
especially species of Philomyciis, which are from half an inch
to several inches long when extended. They are mostly found
under the loose bark of decaying logs or stumps. Large Limax
may be found in cellars, under board walks, in nurseries, etc.

In greenhouses and nurseries, especially old, established
ones, a number of snails and slugs can be found that have been
introduced with plants, etc., chiefly from Europe.

Collect large numbers of specimens wherever possible.
It is desirable to secure good, living snails, but even dead shells
should be taken if they appear in any way different from living
ones; they may represent different species, or forms, and
"a dead shell is better than none at all" for faunistic purposes.

SMALL SNAILS.

More than one half of the number of species of land snails
are small to minute, some measuring only one to two milli-
meters. These in part live at the same places with the larger
ones, but are frequently found also among mosses, lichens,
fine debris, etc. Large numbers of small Pupidae are occasion-
ally seen crawling on old stumps. They may be taken up

March, 1918J Inland Molliisks 171

singly, e. g. on the point of a knife, and put into a vial or small
box, but in general this is a tedious task and the results are not
in proportion to the time spent. The better way is to collect
them wholesale, so to speak. With a small brush, the under
side of pieces of wood, stones, old bricks, etc., the fine dirt
under loose bark or decaying logs and stumps is brushed down
into a pan, or on a piece of paper. Fine debris and dirt around
old trees and stumps, under brush heaps, etc., are scooped up
and the whole material is taken along.

Dead leaves, debris, moss from the ground, wood, rocks,
etc., are gathered on a large piece of paper, or preferably
muslin, (I use one two yards square), gently worked with the
hands, shaken well and gradually removed. A sieve is very
helpful in separating finer particles and dirt along with the small
snails and this material is carefully gathered. The leaves,
moss, etc., if wet, may be spread out on the sheet and left for
partial drying while other collecting is done; or if it is not far
from home or camp, they may be taken along in a large sack
or basket and taken care of there.

Some minute snails, like Vallonia and Vertigo often live in
lawns of cities and towns, sometimes by the thousands. If not
secured otherwise, they may be trapped on old boards, bricks,
etc., which are improved for this purpose if smeared with meat
or grease.

PREPARING.

The larger snails are taken care of first and their soft parts
must be extracted from the shells. A number of them are
put in a tea strainer, or a piece of muslin, and immersed in
nearly boiling water for about fifteen to thirty seconds, according
to size, so that the columellar muscles are detached from the
shells. Then the curved point of a long pin is stuck into the
fleshy part, and the animal extracted, slowly and carefully,
at the same time twisting the body or the shell. A safety pin,
straightened out more or less, is convenient, making a handle
at the same time. After some experience, one will succeed
in removing the soft parts entire and will also know how long
to scald for loosening the animal without "cooking" too
much. If a part is broken ofif and left in the shell, the latter
is filled w4th water and shaken vigorously, with the aperture
closed, and as a rule the remnant can be washed out. Then

172 The Ohio Journal of Science [Vol. XVIII, No. 5,

the shells are wiped clean, the inside with a wad of cotton or
sponge, and left to dry. Shells clothed with hairs, for example,
Polygyra palliata and P. inflecta, should not come into contact
with cotton. Freshwater snails are treated in the same manner.
The opercula of the operculate species should be kept with their
shells, but not gummed in the apertures.

Of the soft parts extracted, at least some may be preserved
in alcohol for future anatomical examination and they should
be labelled carefully. Especially specimens of doubtful identity
must be marked so that they are referable to their shells.
Alcohol of about 25% is used first, for a few days, then changed
to 40 or 50% and after a week or two to 60% or 70%.

Specimens especially wanted for anatomical examination
are best drowned in water (preferably boiled and cooled) in
corked bottles placed upside down or in sealed jars. In this
way, they will extend more or less and the parts will be more in
their natural position. Slugs wanted for the collection may be
treated the same w^ay before being put in alcohol.

Small and minute snails are left to themselves for days or
weeks, in wooden or pasteboard boxes, to give them time to
retire deep into their shells, when they may be dried in gentle
heat; medium sized ones may be scalded before drying. They
should not be dried in heat while fresh and this applies especially
to those which have lamellae and "teeth" in the apertures, or
deeper inside, as FupidcB, Strobilops, many ZonitidcB and the
small PoIygyrcE, for the animals would emerge and obstruct
those parts when dry, and thus make examination difficult or
impossible.

SIFTINGS AND BRUSHINGS.

These when dry may be kept with their labels in boxes,
paper bags or muslin sacks to be worked up at some convenient
time. They should be gently but thoroughly rubbed in the
hands in order to detach specimens clinging to leaves, debris,
etc. Then, work is made much easier if finer and coarser
materials are separated with a sieve or strainer, or with two of
different meshes, or by shaking to and fro on a piece of strong
paper. Portion after portion is spread out on the table and the
snails are picked out with a fine pair of pincers ; very small and
fragile shells are better scooped up or lifted with a moist camel's
hair brush. It is preferable to have a number of small boxes,

March, 1<)18] Inland MoUnsks 173

trays or dishes at hand, to separate the species at once, or at
least the genera or groups. This will save a good deal of
subsequent work and time.

After the specimens from a place are worked up, those of
each species, if identified, are placed in a box, tray or vial.
The latter may be corked if the specimens are perfectly dry,
but otherwise should be closed with a wad of cotton or sponge.

DRIFT.

Drift accumulated along rivers, creeks, brooks, ravines, etc.,
after freshets and floods, in quantities from a handful or less
to a wagon load or more, almost always contains moUusk shells,
sometimes in large numbers and of many species. Generally
they are dead shells, but these will be of interest for faunistics
and often as "pointers" indicating the occurrence of a species
or form which had not been found before. Fine drift is gathered
up carefully and from heaps of coarse material it may be sepa-
rated by sifting.

FOSSILS.

Fossil shells of land and fresh-water mollusks should be
carefully collected wherever there is a chance for doing so.
If the identity of the stratum cannot be ascertained, at least the
locality should be exactly noted. Fossils have been collected
at several places in the state, but not exhaustively, and there
are probably others.

IDENTIFICATION.

This is often difficult for the beginner, especially with small
and minute specimens. The best he can do is to send them to
an expert or specialist, who wdll separate, name and return
them. Send them all; often only a few are sent for naming
and the balance kept, on the supposition that they are all
identical, w^hile tw^o or several species may be mixed.

Sif tings and brushings may be sent as they are, "dirt and
all." Larger snails may be sent, separated or mixed up,
with some moss or dead leaves to prevent jarring. Many a
collector along other lines, an entomologist or botanist, may
have chances to collect specimens and to gather brushings and
siftings, w^hich, by the way, may yield insects or other organ-
isms of interest to himself. They will be of value in working
up our fauna and make additions to the state collection.

TWO NEW TINGIDS FROM THE WEST INDIES

(HEM.-HETER.)

By Carl J. Drake,
New York State College of Forestry.

Leptodictya bambusae sp. new.

Body elongate, narrow, very flat. Antennae long, slender;
first segment almost twice as long as the second; third segment
very long, slender, nearly four times as long as the first and
second conjoined; fourth segment broken off. Head armed
with five, long, slender, semi-erect spines, the anterior ones
extending slightly beyond the apices of the first antennal
segments. Eyes rather large, globular, the facets large. Rostrum
extending between the intermediate coxae. Pronotum finely
punctured, with the sides narrowed anteriorly, with three low
carinae; paranota formed as in L. tabida H.-S., with two rows
or areolae when viewed from above; the outer margin straight.
Legs rather slender. Wings extending slightly beyond the tip of
the abdomen. Elytra extending to far beyond the tip of the
abdomen, the outer margins much less rounded and straighter
than in L. tabida H.-S. ; costal area widest near the middle,
with three to four irregular rows of areolae, the row of areolae
along the distal three-fourths of outer margins a little larger
and more regular than the others; subcostal area narrow,
biseriate, the areolae small; discoidal area finely reticulated,
extending a little beyond the middle of the elytra, with five
rows of areolae at the widest part; sutural area broad, the
areolae becoming larger towards the apex. Hood a little longer
than broad, rather finely reticulate, covering the base of the
head. Length, 2.65 mm.; wddth, 1 mm.

Color: Areolae transparent, slightly irridescent, the nervures
yellowish white. Thorax beneath reddish-brown, the abdomen
testaceous. Legs testaceous, the tarsi darker. Head, eyes, and
a small portion of the pronotum just back of the hood black.
Bucculae and rostral laminae whitish. Antennae whitish, the
third segment slightly embrowned towards the apex.

Two specimens, taken on the leaves of bamboo, Bambusa
vulgaris Schard., at May agues, Porto Rico, July 21, 1914, by
Dr. R. H. Von Zwalenburg. It belongs to the group of Lep-
todictya which have the outer margins of the paranota distinctly

174

March, 1918] Tingids from the West Indies 175

straight as in L. tabida H.-S. and L. plana Heid. and not
rounded as in L. simnlans Heid. This species is probably most
closely allied to L. tabida H.-S., the sugar-cane Tingid, but
easily separated from it by the shorter spines on the head, the
much less rounded outer margins of the elytra, and the adven-
titious nervure in the discoidal area is wanting. Type in the
author's collection.

Leptostyla mcelfreshi sp. new.

Antennae very long, slender, reaching beyond the apex of
the abdomen; first segment slightly swollen, straight, about
four times as long as the second, the second segment short;
third segment very long, slightly curved, slender, cylindrical,
about four times the length of the first ; fourth segment sparsely
pilose, rather slender, equal to the first in length. Bucculas
contiguous in front, broad, rather finely reticulate. Rostrum
extending between the intermediate coxee. Pronotum finely
reticulate, tricarinate; median carina enormously elevated,
somewhat triangular, a little higher than the hood behind the
middle, quite widely reticulated, contiguous with the median
carina of the hood near the base; lateral carince composed of a
single series of large areolae, widely separated from the base of
the hood, parallel with the median carina. Hood rather large,
widely reticulated, obliquely conical with the sides somewhat
flattened and the apex back of the middle. Paranota very
enormously expanded, not very broad at the base and becoming
narrower outwardly, widely and irregularly reticulate, the
areolae becoming smaller towards the base, quite strongly
reflexed, the anterior and posterior margins slightly reflexed
and somewhat undulate, trough-shaped. Legs rather slender
and long. Head short, with five long slender spines, the frontal
spines directed forward, approximating at the tip and appearing
almost like a single spine, shortest; median spine semi-erect, a
little shorter than the first antennal segments; the two lateral
spines (one on each side) arising beneath the hood, extending
forward and strongly curved upward, slightly shorter than the
median spine. Wings not quite as long as the abdomen. Elytra
extending to far beyond the apex of the abdomen, diverging
posteriorly, distinctly constricted near the middle, narrowed at
the base and gradually widening, rounded at the tip; costal
area irregularly reticulate, narrowed at the base and apex, with

17G The Ohio Journal of Science [Vol. XVIII, No. 5,

five rows of areolae near the middle or widest part; subcostal
area narrow, finely reticulate, with three rows of areolse; dis-
coidal area rather small and short, finely reticulate, with four
rows of cells at the widest part, extending a little beyond the
apex of the posterior triangular process of the pronotum;
sutural area irregularly reticulate. Length, 3.6 mm. ; width
(elytra at constriction) 1.8 mm.

Color: Body beneath brownish. Legs testaceous, the tips of
tarsi darker. Rostrum testaceous, the apex infuscate. Antennas
testaceous, the basal segment partially embrowned and the
distal seven-eighths of apical segment dark fuscous. Bucculae
and rostral laminae whitish. Areolae mostly semi-transparent.
Nervures yellowish- white, the marginal nervure of paranota, a
few nervelets in discoidal and costal areas, a large spot on the
median carina, and an oblique fascia (formed by the three
nervures and nervelets connecting them) arising a little beyond
the discoidal area and extending to the apex of each elytron
brown. In this oblique fascia the middle nervure is lowered
and forms a shallow furrow.

Two male specimens, taken at Port au Prince, Haiti, during
the month of January. This species is most closely allied to
L. vesiculosa Champion and L. tumida Champion, but readily
separated from these species by the smaller hood, the longer
spines on the head and the enormously developed paranota.
Type in the author's collection. I have named the species in
honor of the late Frank M. McElfresh, a great student, collector
and worker in this group of insects. Some of the records pub-
lished by Osborn and Drake in the Ohio Journal of Science
(Vol. XVII) were taken from this collection. Although Mr.
McElfresh had excellent knowledge of this group of insects and
was preparing to monograph the Tingidae of North America he
left no notes or manuscripts and had published no papers on
the Tingids.

Syracuse, N. Y.

The Editor regrets the delay in the appearance of the
March issue of the Journal of Science. It was due to the loss
of a large part of the proof in transmission.

Date of Publication, April 25, 1918.

THE

Ohio Journal of Science

PUBLISHED BY THE

Ohio State University Scientific Society

Volume XVIII APRIL, 1918 No. 6

TABLE OF CONTENTS

Fenton — The Parasites of Leaf-Hoppers — Part 1 177

Oberholser— The Common Ravens of North America 213

DeLong — The Occurence of a Probable Gynandromorph in the Homoptera. . 226

DeLong— A New Species of Cicadellidse from Wisconsin 228

BuRRiLL AND Smith— A Preliminary List of the Ants of Wisconsin 229

THE PARASITES OF LEAF-HOPPERS.

With Special Reference to the Biology of the Anteoninae.

F. A. Fenton.

PART I.

CONTENTS. PAGE

I . Introduction 177

II. Historical 179

III. Systematic Position 181

IV. Methods 181

V. Distribution 181

Host List 182

VI. Relations with other parasites 191

VII. Biology and Life History 192

VIII. Description of Stages 203

IX. Internal Anatomy of the Larva 211

INTRODUCTION.

The study of the parasites of the leaf-hoppers presents an
interesting problem, not only from the practical standpoint, but
from the biological as well. The economic status of the leaf-
hoppers has been well estabhshed, and as a group they are now
considered as including some of the more important insect
pests. Therefore, their control by parasitism is significant as a
natural check, and their parasites must be considered as dis-

177

178 The Ohio Journal of Science [Vol. XVIII, No. 6,

tinctly beneficial. This fact has been utilized in actual practice
in control measures against the sugar-cane leaf-hoppers in
Hawaii with marked success.

From the biological viewpoint, the study of the interrelations
of these parasites with their host has a direct bearing upon the
interesting problem of insect parasitism in general. Thus the
specialization of the dryinid, and the morphological changes it
brings about in its host, the hopper, may and probably does have
parallels in the relations between other groups of parasitic
insects. with their hosts.

These Homoptera are subject to parasitism in all their
stages; namely, egg, nymphal, and adult. In North America
three orders of insects are known to contribute their quota of
the parasites of this group — the Hymenoptera, Diptera, and
Strepsiptera; and in Australia and Hawaii, certain of the
Lepidoptera are considered to be parasitic, though this relation
is not definitely established.

The eggs are known to be parasitized by species of the
PlatygastridcB, MymaridcB, Encyrtidce, Eiilophidce, and Tricho-
grammidcB (Perkins, 1905-1906). The nymphs and adults are
parasitized by the Anteonince, Pipunculidce, Strepsiptera, Epipy-
ropidcE and Encyrtidce (Perkins, 1905) and Chalcididce (Misra,
1917).

The relative importance of these parasites as natural checks
varies probably in different localities. However, in Ohio, at
least in the northern half, the Anteonince are the most important.
Strepsiptera have been recorded as being relatively numerous
in regions further south, and their rather obscure habits and
small size possibly may account for an underestimation of
their importance in this locality. Frequently their host survives
and thus it seems that their attack is not always fatal. Certainly,
however, they form an important link in the chain of natural
enemies.

The following paper is confined to an account of the Anteo-
nince. Material was more plentifully obtained for their study,
and their greater importance warranted a more extended
investigation. Most of the parasites used in the life history
studies were bred from hoppers taken near Sandusky and
Columbus, Ohio.

April, U)18] Parasites of Leaf-IIoppers 179

ACKNOWLEDGMENTS.

The writer is greatly indebted to Professor Herbert Osborn,
at whose suggestion the work was started, for his continued aid
throughout the entire study. He also wishes to thank Mr.
D. M. DeLong for the collection and identification of para-
sitized CicadellidcB from Tennessee, Mr. F. H. Lathrop for
the collection of material from Oregon, and Mr. S. H. Rohwer,
of the U. S. National Museum, for the use of type material in
the identification of species.

HISTORICAL.

Latreille (1805) described Lestodryinus {Dryinus) Jormicarius
which is the earliest record of description of a representative
of this group. Jurine (1807) described a species which now
forms the type of the genus Anteon. Ljungh (1810) described
Gonatopus formicarius as the first apterous form and Dalman
(1818) described Aphelopiis melaleuciis. Each of these species
are typical of the four tribes into which the Anteonince are now
divided.

The first host record was by Curtis in 1836, when he reported
Aph'rodes craticula to be parasitized in different stages by these
parasites. Edward Perris in 1857 thought that Platygonatopus
{Gonatopus) pedestris (Dalm.) was a hyperparasite of Eiiscelis
Athysaniis maritimiis, believing that the larval sac was the
primary parasite and distinct from the dryinid. G. von
Frauenfeld (1865) mentioned Platygonatopus {Gonatopus)
pedestris (Dalm.) in connection with (Typhlocyba). Since
Erythroneura {Typhlocyba) is parasitized only by the genus
Aphelopus in this family, it is probable that the host record is
incorrect.

Lichtenstein (1874) reported rearing a species, Gonatopus
ptinorum from the heetle Ptinus fur hnt Kieffer (1914) considers
this species as Cephalonomia xambeni Giard of the sub-family
BethylincB. From 15 to 20 larvae were found in the cocoon of
Ptinus, each spinning up an individual cocoon after issuing from
the host (Kieffer '14). Dale (1878) mistook a probable dryinid
larva for an Acarid and named it Homopterophagus dorsettensis.
It was reared from a black, "mustard seed-like structure,
noticed on the sides of diverse Ilomoptera, near the insertion
place of the wings." According to Kirkaldy this was the larva
of Gonatopus.

180 The Ohio Joimial of Scie?ice [Vol. XVIII, No. 6,

Mik (1882) described and worked out several important
stages in the life history of Gonatopiis pilosus Thorns, showing
the true nature of the parasite to the host and disproving
the hyperparasite theory. Giard (1889) noticed and described
the dryinid sacs attached to Erythroneura hypocastani in France
and remarked upon the sac-like structure believing it to be a
"true animal gall produced on an arthropod by another arthro-
pod." In a later article in this same year he succeeded in
rearing the adult parasite which proved to be Aphelopus
melaleuciis Dalm. He noticed the similarity of his own observa-
tions with regard to the hosts and life history with those of
Perris (1857) and Mik (1882) and concluded that as a group
the AnteonincB were generally parasitic on the {JassidcE) Cicadel-
lidcE. Giard also noticed certain secondary sexual modifications
on the host caused by the parasite and that the hoppers were
castrated.

Swezey (1903) reared both sexes of two dryinids from leaf-
hoppers and these were identified by Ashmead. He was thus
the first one to associate the males of the more highly specialized
apterous females and his true interpretation of the so-called
genus Labeo in connection with these marks an important step
in the knowledge of these insects.

Perkins (1905) gave a detailed account of the biology and
life history of the AnteonincB, together with descriptions of a
large number of genera and species new to science, he having
further divided the old genus Gonatopiis into a number of dis-
tinct genera based on more exact structures than the general
body form. He also gave an extended host list and the next
year supplemented this with new additions from Arizona.

Kieffer (1907) summarized the knowledge of the group to
that year in a work that was primarily systematic and later
(1914) monographed the group under the Bethylidce. Perkins
(1912) in reviewing Kieffer' s first paper added a number of new
species, publishing four excellent plates figuring a number of
typical AnteonincE.

Keilin and Thompson (1915) worked out some very impor-
tant points in the biology of Aphelopus melaleuciis Dalm.,
especially with regard to the very early stages and the pro-
duction and development of the peculiar hypertrophied tissue
in the host. In December of this same year and in 1916
Kornhauser worked out the biology of Aphelopus thelice

April, 1918] Parasites of LcaJ-IIoppers 181

Gahan, Mss. giving the first authentic record of a dryinid
being parasitic on a membracid and also being the first one to
observe polyembryony in this group. His work is concerned
primarily with the effects of parasitism on the host.

Misra (1917) in working upon the Indian sugarcane leaf-
hopper, Pyrin a aberrans Kirby, gave a number of pages to
the discussion of the biology of the two dryinid parasites of
this group.

SYSTEMATIC POSITION.

The Anteonince {DryinidcE) are now generally considered to
be a sub-family of the BethylidcE, the latter being included in
the super-family Proctotrupoidea of the Hymenoptera. A
further discussion of the systematic relationships of these
insects will be taken up in Part II of this paper.

METHODS.

All the parasites mentioned in the life history studies of
this paper were bred, it being very difficult to capture adults
in the field. All the hoppers were kept alive and the parasitized
ones isolated in separate shell vials provided with a layer of
damp soil and a cotton plug. Fresh leaves were added every
day until the host had been killed by the parasite. When
the latter had issued the host shell was preserved dry in gelatine
capsules. If the hopper died before the parasite matured, it
was preserved in 70 per cent, alcohol. The material for the
morphological studies was sectioned, or dissected in cedar oil,
being transferred into the latter directly from 70 per cent,
alcohol.

DISTRIBUTION.

Geological. — Kieffer reports the genera Dryinus and Clielogy-
nus from the lower Oligocene and Brues a new genus Dryinince
from this same stratum.

Geographical. — The AyiteonincB are found in all parts of the
world and will doubtless be found wherever their hosts are
abundant. According to Kieffer up to 1914 there have been
396 certain species described and 28 doubtful species. Of
these, 200 have been described from the Palearctic region, 98
from the Nearctic, 65 Australian, 32 Neotropical, 16 Oriental,
and 8 so far from the Ethiopian regions. To the list of countries
Alaska may be added, one species of Deltocephaliis being quite

182

The Ohio Journal of Science [Vol. XVIII, No. 6,

extensively parasitized. It is probable that they are cosmo-
politan in their distribution as a group.

Host Records. — As previously stated, the Anteonince confine
their attacks to the Fulgoridce, Cicadellidce and Membracidce so
far as is known at the present, with but one record in the last
named family.*

The following list summarizes the host records in so far as
have been ascertained to date. The names and systematic
position of the hosts have been arranged chiefly after Van
Duzee's latest catalogue, (1917).

HOST LIST.

MeMB RACIER.

Host Species

Stage of
Host

Parasite Species

Authority

Location

Thelia

bimaculata

nymphs

Aphelopus
thelias

Kornhauser

Evanston, 111.

ClCADELLID^.

Sub-family Bythoscopince.

Agallia

sanguinolenta

5th instar

?

F. A. F.

Marietta, Ohio.
H. 0.

Ipo
sp.

nymph and
adult

Parenteon

myrmecophilus

Perkins

Queensland,
Australia.

Eurinoscopus

sp.

nymph

Chelogynus
leiosomus

Perkins

Queensland,
Australia.

Eurinoscopus
sp.

nymph

Chelogynus
nitidus

Perkins

Queensland,
Australia.

Eurinoscopus
sp.

nymph

Chelogynus
dinudiatus

Perkins

Queensland,

Australia.

Eurinoscopus
sp.

nymph and
adult

Chelogynus
destructor

Perkins

Queensland,
Australia.

Idiocerus
sp.

nymph

Chelogynus
cognatus

Perkins

Queensland,
Australia.

Idiocerus
pallidus

nymph and
adult

?

F. A. F.

Corvallis, Oregon
F. H. L.

? Macropsis

nymph

Chelogynus
cognatus

Perkins

Queensland,
Australia.

I

*Ratzeburg reported rearing Aphelopus albipes (Ratzeb.) from pupae of
Bombyx pudibiinda, and Surface, Phorbas longicornis (Brues) from the cocoon of a
microlepidopteron. Both of these records are doubtful and in the latter case it is
probable that the cocoon of the dryinid itself was mistaken for the lepidopterous
cocoon.

April, 191S]

Parasites of Leaf-Hoppers

183

Sub-family Jassma.

Host Species

Stage of
Host

Parasite Species

Authority

Location

Parabolocratus
flavidus

adult

>

F. A. F.

Clarksville, Tenn.
D. M. D.

Scaphoideus

immistus

Adult and
nymph

?

F. A. F.

Venice, Ohio.

Tartessus
syrtides

?

Chalcogonatopus
gigas

Perkins

Queensland,
Australia.

Euleimomos
sp.

adult

Chelogynus
parvulus

Perkins

Queensland,
Australia.

Euleimomos
sp.

adult

Chelogynus
coriaccus

Perkins

Queensland,
Australia.

Euleimomos
sp.

adult

Pachygonatopus
melanicus

Perkins

Queensland,

Australia.

Euleimomos
sp.

?

Gonatopus
vitiensis

Perkins

Fiji.

Platymetopius
pyrops

adult

?

F. A. F.

Clarksville. Tenn.
D. M. D.

Platymetopius
arutus

nymph

?

F. A. F.

Columbus, Ohio.

Phrynophyes
sp.

?

Chalcogonatopus
decoratus

Perkins

Queensland,
Australia.

Deltocephalus
sp.?

nymph and
adult

Digonatopus
australias

Perkins

Queensland,
Australia.

Deltocephalus
sp.?

?

Gonatopus
dubiosus

Perkins

Queensland,
Australia.

Deltocephalus
sp.?

nymph

Chelogynus
pallidicomis

Perkins

Queensland,
Australia.

Deltocephalus
sp.?

nymph

Chalcogonatopus
optabilis

Perkins

Queensland,
Australia.

Deltocephalus
sp.?

>

Gonatopus
ombrodes

Perkins

Columbus, Ohio.

Deltocephalus
sp.?

nymph

Gonatopus

vitiensis

Perkins

Queensland,
Australia.

Deltocephalus
sayi

adult

Gonatopus
contortulus

F. A. F.

Castalia, Ohio.

Deltocephalus
sayi

adult

Gonatopus n. sp.

F. A. F.

Columbus, Ohio.

Deltocephalus
sayi

nymph

?

F. A. F.

Columbus, Ohio.

184

The Ohio Journal of Science [Vol. XVIII, No. 6,

Sub-family Jassintz — Continued.

Host Species

Stage of
Host

Parasite Species

Authority

Location

Deltocephalus
weedi

adult

?

F. A. F.

Clarksville, Term.
D. M. D.

Deltocephalus
infumatus

adult

?

F..A. F.

Corvallis, Oregon.
F. H. L.

Deltocephalus
inimicus

adult

Gonatopus
obscurrisimus

F. A. F.

Bay View and
Columbus, Ohio.

Deltocephalus
inimicus

nymph

Gonatopus
erythrodes

F. A. F.

Castalia, Ohio.

Deltocephalus
abdominalis

adult

?

F. A. F.

Katmai, Alaska.
J. S. H.

Deltocephalus
affinis

adult

?

F. A. F.

Bay View and
Columbus, Ohio.

Deltocephalus
oculatus

nymph

7

F. A. F.

Castalia, Ohio.

Deltocephalus
oculatus

adult

?

F. A. F.

Castalia, Ohio.

Deltocephalus
balli

adult

?

F. A. F.

Columbus, Ohio.

Deltocephalus
xanthoneuras ?

nymphs

Gonatopus

pilosus

Mik

Vienna, Austria.

Deltocephalus
silvestris

adult

?

F. A. F.

Mt. Katahdin,
Maine. H. 0.

Deltocephalus
imputans

adult

?

F. A. F.

Lexington, Tenn.
D. M. D.

Lonatura
catalina

adult

?

F. A. F.

Clarksville, Tenn.
D. M. D.

Nephotettix
sp.

nymph

Chalcogonatopus
optabilis

Perkins

Queensland,
Australia.

Nephotettic
sp.

?

Gonatopus
dubiosus

Perkins

Queensland,
Australia .

Nephotettix
plebius

adult

Gonatopus
pulcherrimus

Perkins

Queensland,
Australia.

Euscelis spp.

adult

Pachygonatopus
melanias

Perkins

Queensland,
Australia.

Euscelis
sp.

?

Digonatopus
koebelei

Perkins

New South Wales,
Australia.

Euscelis
sp.

?

Gonatopus
pallidiceps

Perkins

California.

April, 1918]

Parasites of Leaf -Hoppers

185

Sub-family JassincB — Continued.

Host Species

Stage of
Host

Parasite Species

Authority

Location

Euscelis
exitiosus

adult

?

F. A. F.

Wellington, Kas.
E. O., G. K.

Euscelia
humidus

adult

>

F. A. F.

Houlton, Bog,
Maine. H. 0.

Euscelis
curtisii

adult

Gonatopus
brunnescens

F. A. F.

Columbus, Ohio.

Euscelis
maritimus

nymph

Platygonatopus

pedestris

Dalman

Sweden.

Eutettix
sp.

nymph and
" adult

Chelogynus
nigricornis

Perkins

Queensland,
Australia.

Phlepsius
sp.

>

Chalcogonatopus
pseudochromus

Perkins

Columbus, Ohio.

Phlepsius
irroratus

adult

?

F. A. F.

Covington, Tenn.
D. M. D.

Phlepsius
truncatus

adult

?

F. A. F.

Clarksville, Tenn.
D. M. D.

Phlepsius
apertus

adult

?

F. A. F.

Houlton, Maine.
H. 0.

Thamnotettix
flavotinctus

adult

?

F. A. F.

Clarksville, Tenn.
D. M. D.

Thamnotettix
crumb i

adult

?

F. A. F.

Clarksville, Tenn.
D. M. D.

Thamnotettix
crumbi

adult

?

F. A. F.

Columbus, Ohio.

Chlorotettis
unicolor

nvmph and
"adult

Chelogynus n. sp.

F. A. F.

Castalia, Ohio,
Corvallis, Ore.
F. H. L.

Chlorotettix
nacreosus

adult

V

F. A. F.

Clarksville, Tenn.
D. M. D.

Chlorotettix
galbanatus

adult

?

F. A. F.

Clarksville, Tenn.
D. M. D.

Chlorotettix
necopina

adult

'?

F. A. F.

Clarksville, Tenn.
D. M. D.

Cicadula
. lepida

adult

?

F. A. F.

Clarksville, Tenn.
D. M. D.

Cicadula
sexnotata

adult

?

F. A. F.

Columbus, Ohio.

186

The Ohio Journal of Science [Vol. XVIII, No. 6,

Sub-family Typhlocyhince.

Host Species

Stage of
Host

Parasite Species

Authority

Location

Dikraneura
sp.

adult

Aphelopus
arizonicus

Perkins

Nogales,
Arizona.

Dikraneura
fieberi

adult

Aphelopus n. sp.

F. A. F.

Columbus, Ohio.

Empoasca
smargdula

adult

Aphelopus n. sp.

F. A. F.

Marietta, Ohio.
H. 0.

Empoasca
mali

adult

?

F. A. F.

Columbus, Ohio.

Empoasca
fiavescens

adult

?

F. A. F.

Columbus, Ohio.

Empoa
commissuralis

adult

?

F. A. F.

Mary's Peak,
Ore., F. H. L.

Erythroneura
sp.

?

Aphelopus
microleucus

Perkins

Nogales,
Arizona.

Erythroneura
sp.

?

Aphelopus
pulcherzimus

Perkins

Nogales,
Arizona.

Erythroneura
tricincta

adult

Aphelopus sp.

F. A. F.

Kelley's Island,
Ohio.

Erhtyroneura
comes

adult

Aphelopus n. sp.

F. A. F.

Columbus, Ohio.

Erythroneura
obliqua

adult

Aphlopus sp.

F. A. F.

Marietta, Ohio.
H. 0.

Erythroneura
vulnerata

adult

Aphelopus sp.

F. A. F.

Columbus,

Rockbridge, 0.

Erythroneura
douglasi

adult

Aphelopus
melaleucus

Girard

France.

Erythroneura
hippocastani

adult

Aphelopus
melaleucus

Girard

France.

Genus or Species Unknown or Record Doubtful.

Jassid
sp.

nymph

Jassid
sp.

Jassid
sp.

Large green
jassid on oak

Medium sized
jassid on willow

Jassid
sp.

Chelogynus
lusus

nymph

nymph

nymph

Chalcogonatopus
rapax

Digonatopus
plebius

Deinodryinus
paradoxus

Perkins

Perkins

Perkins

Perkins

Anteon

arizonensis

Chelogynus
funestus

Perkins

Tucson,
Arizona.

Nogales,
Arizona.

Java.

Nogales,
Arizona.

Nogales,
Arizona.

Perkins

Tucson,
Arizona.

April, 1918]

Parasites of Leaf-IIoppers

187

Family Fulgorid.e.
Sub-family Dictyophorince.

Host Species

Stage of
Host

Parasite Species

Authority

Location

Thanatodictya
sp.

nymph

Paradryinus
javanus

Perkins

Java.

Hasta
hastata

nymph

Paradryinus
Icptias

Perkins

Queensland,
Australia.

Sub-family Cixiince.

Cixius
contaminatus

Lestodryinus
formicarius

KiefTer

Europe.

Sub-family Tropiduchina.

Vanua
vitiensis

adult

Pseudogonatopus
kiefTeri

Perkins

Fi

Ji-

Sub-family Issince.

Brucomorpha
oculatus

nymph and
adult

Phorbas
mirabilis

Perkins

Columbus,

Sandusky, Ohio.

Brucomorpha
sp.

adult

Phorbas
arizonica

Perkins

Nogales,
Arizona.

^Sub-family AcanaloniincE.

Acanalonia
bivittata

nymph

Hesperodryinus
amphiscepae

Perkins

Nogales,
Arizona.

Acanalonia
bivittata

nymph

Perodryinus
amoenus

Perkins

Nogales,
Arizona.

Acanalonia | ?
immaculata

Apterodryinus
torvus

Perkins

Arizona.

Sub-family FlatincB.

Ormenis
prunosa

adult

Psilodryinus
arizonicus

Perkins

Nogales,
Arizona.

Ormenis
septentrionalis

5

Psilodryinus
arizonicus

Perkins

Nogales,
Arizona.

Ormenis adult
septentrionalisi

Psilodryinus
ormenidis

Swezey

U. S.

Misthanophan-
tia sonorana

nymph

Hesperodryinus
arizonicus

Perkins

Nogales,
Arizona.

188

The Ohio Jourfial of Science [Vol. XVIII, No. 6,

Sub-family FlatincB — Continued.

Host Species

Stage of
Host

Parasite Species

Authority

Location

Misthanophan-
tia sonorana

nymph

Apterodryinus
torvus

Perkins

Arizona.

Privesa
aphrophoroides

?

Neodryinus
raptor

Perkins

Queensland,
Australia.

Privesa
aphrophoroides

nymph

Paradryinus
venator

Perkins

Queensland,
Australia.

Scolypopa
sp.

?

Neodryinus
raptor

Perkins

Queensland,
Australia.

Gaetulia
sp.

?

Neodryinus
raptor

Perkins

Queensland,
Australia.

Siphanta
sp.

?

Neodryinus
raptor

Perkins

Queensland,
Australia.

Siphanta
sp.

nymph

Thaumatodryinus
koebelei

Perkins

Queensland,
Australia.

Siphanta
sp.

nymph

Paradryinus
venator

Perkins

Queensland,
Australia.

Siphanta
sp.

nymph

Paradryinus
threnodes

Perkins

Queensland,

Australia.

Siphanta
sp.

nymph

Paradryinus
koebelei

Perkins

Queensland,
Australia.

Siphanta
sp.

nymph

Paradryinus
varipes

Perkins

Queensland,
Australia.

Sephena
sp.

?

Neodryinus
raptor

Perkins

Queensland,

Australia.

Colgar
sp.

nymph

Paradryinus
leptias

Perkins

Queensland,
Australia.

Colgar
sp.

nymph

Paradryinus
threnodes

Perkins

Queensland,
Australia.

Colgar
sp.

nymph

Neodryinus
nelsom

Perkins

Queensland,
Australia.

Colgar
peracutus

nymph

Neodryinus
koebelei

Perkins

Queensland,
Australia.

Aphanophantia
sp.

?

Neodryinus
raptor

Perkins

Queensland,
Australia.

Massila
sp.

?

Neodryinus
raptor

Perkins

Queensland,
Australia.

Masilla
sp.

nymph

Paradryinus
venator

Perkins

Queensland,

Australia.

Massila
sp.

nymph

Neodryinus
koebelei

Perkins

Queensland,
Australia.

April, 1918]

Parasites of Leaf-IIoppers

189

Sub-family Delphacina.

Host Species

Stage of
Host

Parasite Species

Authority

Location

Stenocranus

?

Haplogonatopus
vitiensis

Perkins

Fiji.

Stcnocranus
dorsalis

nymph

Pseudogonatopus
stenocrani

Perkins

Ohio.

Perkinsiella
saccharida

nymph and
adult

Pseudogonatopus
saccharatorum

Perkins

Queensland,

Australia.

Perkinsiella
saccharida

nymph

Echthrodelphax
fairchildii

Perkins

Hawaii.

Hadeodelphax

nymph

Echthrodelphax
nigricollis

Perkins

Queensland,
Australia.

Hadeodelphax

nymph

Haplogonatopus
moestus

Perkins

Queensland,
Australia.

Aloha
ipomeas

nymph

Echthrodelphax
fairchildii

Perkins

Hawaii.

Pyrilla
sp.

nymph

Lestodryinus
pyrillae

Misra

India.

Pyrilla
aberrans

nymph

Lestodryinus
pyrillae

Misra

India.

Pyrilla
aberrans

nymph

Chlorodryinus
pallidus

Misra

India.

Pyrilla
perpusilla

nymph

Chlorodryinus
pallidus

Misra

India.

Pyrilla
pusana

nymph Chlorodryinus
pallidus

Misra

India.

Pissonotes
dorsalis

adult ?

1

F. A. F.

Orono, Maine.
H. 0.

Bostaera nymph Agonatopoides
nasuta synchromous

Perkins

Xogales,
Arizona.

Stobaera
sp.

nymph

Pseudogonatopus
arizonicus

Perkins

Xogales,
Arizona.

Stobaera
sp.

nymph

Pseudogonatopus
autoxenobius

Perkins

Nogales, Arizona.

Liburnia
sp.

?

Pseudogonatopus
americanus

Perkins

Columbus, Ohio.

Liburnia
sp.

~j

Pseudogonatopus
opacus

Perkins

Queensland,
Australia.

Liburnia
sp.

nymph and
adult

Pseudogonatopus
dichromous

Perkins

Queensland,
Australia.

190

The Ohio Journal of Science [Vol. XVIII, No. 6,

Sub-family Delphacince — Continued.

Host Species

^ HSt°^ Parasite Species

1

Authority

Location

Liburnia
sp.

nymph

Pseudogonatopus
perkinsi

Perkins

Hawaii.

Liburnia
sp.

nymph and
adult

Pseudogonatopus
juncetorum

Perkins

Queensland,
Australia.

Liburnia
sp.

nymph and
adult

Pseudogonatopus

palustris

Perkins

Queensland,
Australia.

Liburnia
sp.

?

Pseudogonatopus
kurandae

Perkins

Queensland,
Australia.

Liburnia
sp.

?

Paragonatopus
nigricans

Perkins

Queensland,
Australia.

Liburnia ?
sp.

Haplogonatopus
brevicornis

Perkins

Queensland,
Australia.

Liburnia
sp.

?

Haplogonatopus
apicalis

Perkins

Queensland,
Australia.

Liburnia
campestris

nymph and
adult

Haplogonatopus
americanus

Sweezey,
Perkins

Columbus, Ohio.

Liburnia
lutulenta

nymph and
adult

Haplogonatopus
americanus

Sweezey
Perkins

Columbus, Ohio.

Liburnia
verwandter

nymph

Echthrodelphax
nigricoUis

Perkins

Queensland,
Australia.

Liburnia
verwandter

nymph

Echthrodelphax
bifasciatus

Perkins

Queensland,
Australia.

Dicranotropis
sp.

?

Pseudogonatopus
nudus

Perkins

Java.

Genus and Species Unidentified.

Aphanophantia

nymph

Paradryinus
koebelei

Perkins

Queensland,
Australia.

Siphanta

>

nymph

Chloridryinus
pallidus

Perkins

Queensland,
Australia.

Fulgorid
species

?

Antonella
robusta

Dodd

Queensland,
Australia.

Fulgorid
species

?

Antonella
bicolor

Dodd

Queensland,
Australia.

April, 1918]

Parasites of Leaf-IIoppers

191

Twenty-two genera, including forty-nine known species and
five unidentified ones out of three of the six sub-families in the
Cicadellidcc are known to be parasitized by the dryinids. Most
of the host records are in the sub-family Jassince chiefly grass
living forms. In the Fnlgorida:, twenty-six genera, including
twenty-two species and four unidentified species are known to
be subject to parasitism by these insects. In this family most
of the host records are in the sub-family Delphacince. In all
fulgorids the nymphal stages are the most susceptible to attack,
the only adults parasitized being the weaker and less active
forms. It is likeh' that nearly every species in these two families
is subject to attack by one or more species.

CICAOIDAC ICICAPtULIOAll IFULGOHIOAtl CCRCOPIOAt It'EKBHACIDAil

lULOPfflDAt PLATYQASTRIDAE MYMARIDAI TBICHOORAMIIIDAt lAWTCOMIWAtl PIPUHCUtlDAi STREPSIPTCRA tPtPYMPtOAE'

OTHER PARASITIC
NYMINOPTIR

eORROOCNTIA LIPmOPTtRA

COLtOPTERA

OITTHOPTERA

DIPTERA

HY«EMOPTE»A
NON PARASITIC

Diagram 1.
Interrelations of Anteonince with other parasites.

The above diagram represents graphically the inter-relations
of the Anteonince and their hosts with other related parasites
of these hosts. It is seen at once how small a part they play
as a group in comparison with the other parasites in maintaining
nature's balance. They are but one link in a very complex
series. The leaf hoppers are parasitized also by the Eulophidce,
PlatygastridcE, Mymaridce, Trichogrammidce , Chalcididce, and
Encyrtidce of the Hymenopiera, by the Pipunculidce of the
Diptera, by the Epipyropidce of the Lepidoptera, and by the

192 The Ohio Journal of Science [Vol. XVIII, No. 6,

Strepsiptera. In addition to parasitizing the Ilomoptem, the
AnteonincB are themselves parasitized by the Encyrtidce and
EulophidcE.

Aside from parasitizing the Homoptera most of the groups
mentioned attack such other insect groups as the Hemiptera,
Coleoptera, Diptera, Orthoptera and Hymenoptera. In com-
parison with the other groups it is noted that the AnteonincB
and Pipuncnlidce are narrowly restricted in their range of
hosts.

BIOLOGY AND LIFE HISTORY.

Adult. — The adult dryinids are found living in grasses,
herbage and trees, in fact wherever their hosts, the leaf- and
tree-hoppers, are found. The females are much more active,
continually running and flying about in search of their prey.
In species where the female is wingless, the habits of the two
sexes are very different, the males generally being found resting
on the leaves and stems, or flying about in search of the females.
In describing the habits of Lestodryinus (Dryinus) pyrillce
Kieffer, Misra (1917) says: "The adult females are very
active and may be seen actively flying about the leaves, occasion-
ally settling down, especially near the nymphs, and then flying
away. The adults are prominent mostly during the hottest
part of the day. " Later in the sam.e paper, speaking of Chloro-
dryimus paUidus Perk., he says: "It is the female which is
mostly in evidence in the fields when the leaf-hopper is present
in numbers on the canes. The male is rarely seen about the
infested fields. "

Not all winged dryinids are as active as Misra has observed,
for Perkins (1905) says: "The prey is sought on foot, for while
most of the Anteonince {Dryinidce) are most active and rapid
runners, they are but poor performers on the wing. In most
of the winged forms, these organs are unduly short, and in
Echthrodclphax serve hardly more than to transport it from
one cane plant to another as occasion demands." There seems
to be no correlation between the presence of wings in this
family and the habit of attacking arboreal hoppers. Perkins
(1905) noticed that in Australia and generally in America, the
apterous forms of the Gonatopus type preyed upon hoppers that
fed on grasses and low herbage. "On the other hand," he
says, "the Hawaiian apterous species are essentially arboreal."

April, 1918] Parasites oj Leaf -Hoppers 193

Mimicry. — Along with the loss of wings in the female, there
has been developed the remarkable resemblance to ants in
many forms. This has been adaptive and due entirely to
the mode of life of the female. Not only have the wings been
lost, but the legs have been modified for running, and the
forelegs especially for grasping the prey. Thus the thorax has
become greatly modified and more closely fused with the
propodeum. In the more highly specialized species, many of the
thoracic sutures have been obliterated and segmentation of
the thorax is indistinct. The latter is divided into two nodes
by the narrow mesonotum, and may even be divided into three
nodes. This division of the thorax and the pedunculate form of
the first abdominal segment are adaptations to the stinging
habit of the female, enabling her to bend the entire body in an
arc while stinging the prey.

Many species have evidently taken advantage of this ant-
like form and are frequently found associated with ants, some-
times being found in the nests of the latter. This association
in the case of Platygonatop^is pedestris (Dalm.) was observed
by Haliday in 1834, when he says: "The first time I met with
this species, it was in company with some Myrmicce (not M.
rubra), under a stone, in a chalky lane. * * * In this
island, its haunts are on the sand-hills of the coast, among
which Formica emarginata swarms. The Dryinus, which is
not dissimilar in form and color, moves among them distin-
guished by its hitching gait, produced by the enormous length
of the tarsi and trochanters of its forelegs; it can run pretty
fast, however. * * * j witnessed an occurrence which
shows that it is not always quite amicable. Four ants were
bearing off one of the Dryinii quite alive and vigorous, though
not able to struggle much in their grip; my approach disturbing
them, three scampered off, but the fourth, more determined,
held on; the Dryinus however, as soon as she got fair play,
showed fight, and though her small jaws seemed ill calculated
to match those of the ant, the battle was maintained without
any visible disadvantage. * * * " Perkins (1905) says
that '' Paranteon no doubt has special habits; for the sluggish
hopper that it affects, forms flocks of greater or less extent, and
excreting much honey dew, is invariably covered with swarms
of ants, usually a moderate-sized species of Formicidce. The
Paranteon bears some slight resemblance to the ant in appear-

194 The Ohio Journal of Scie?ice [Vol. XVIII, No. 6,

ance, and this resemblance is enhanced by its actions. If a
number of parasites be bred together in a glass vial, they m.ay
be seen gathered in pairs, standing sub-erect on their four hind
legs, face to face, stroking one another, licking each other's
mouth, soliciting food. Now the ants that attend the hoppers
themselves behave in a similar fashion, and it is most probable
that they actually feed the Paranteon, which unless it were on
friendly terms with the ants, could never approach the hoppers
to lay its eggs in these. * * * Further, it would be interest-
ing to note what happens to the full-grown larva of the parasite,
which must emerge from its sac amongst a swarm of carnivorous
ants that are always prepared to kill and carry ofi any weaker
insect that falls in their way. Most probably not only are the
larvae not harmed by the ants, but they may be carried down by
them to their underground nest, and pupate therein. "

There are several records of dryinids being collected in ants'
nests. The following species have been recorded as associated
with ants: Dicondyhis distindits (Kief.), England, in colony
of Myrmica; Dicondyhis striatus (Kief.) Austria and England,
in nest of Formica fusca; and Gonatopus Myrmecophilus (Kief.),
Luxemburg and England, with Formica ruhibarhis.

Apterous species are often collected by turning over stones,
and are very rarely reported as being taken in the net, although
this may be due to their close resemblance to ants.

Both sexes feed readily on water sweetened with sugar.
Perkins (1905) noticed individuals of Echthrodelphax feeding
on the honey-dew secreted by their host, a certain fulgorid.
They may attack and devour their host, for the same author
says that "under unnatural conditions, such as in the confine-
ment of a small jar or glass tube, and probably under pressure of
hunger, the hoppers are frequently killed outright, and to some
extent devoured." In the act of oviposition, the female
invariably works her mandibles on the cuticle of the hopper,
and sometimes produces a wound which in some cases so weakens
the hopper as to cause its death. Again no sign of a wound is
noticed after oviposition.

The longevity of the adults varies with the sexes, the female
invariably being much the longer lived. In fact, the males
live often but a few^ days, often only for a day. Frequently,
according to Perkins (1905), the m.ale dies after copulation.
The female may live as long as seventeen days in captivity,
and undoubtedly much longer under natural conditions.

April, 11)18] Parasites of LeaJ-IIoppers 195

In the species studied, the act of pairing was unobserved.
In spite of the fact that with Gonatopus erythrodes (Perkins) and
Ilaplogonatopiis americanus (Perk.) males and females were
placed together for observation under as natural conditions as
possible, neither sex paid any attention to the other. The
females were constantly running around among and over the
grass-blades as if searching for their prey. On the other hand,
the males were as a rule inactive, resting upon the sides of the
cage or on the grass leaves. Perkins (1905) concerning Echthro-
delphax, states that "pairing of the sexes is of short duration,
and after copulation the male frequently never moves again,
and in general dies very quickly. "

Parthenogenesis occurs probably to a considerable extent
in this sub-family. This has been proven in captivity, and
undoubtedly occurs to some extent under normal outside con-
ditions. The great disproportion of sexes in some species,
and the fact that with others males have not been found, seems
to bear out that statement. In commenting on Pseudo gonatopus
Perkins (1905) says that but one in forty adults was a male.
Males have not been either captured or bred from species of
Eugonatopus or Agonatopiis. Adults from unfertilized eggs of
Gonatopus contortiilus Patton were all females, and were normal
and active in every way.

Polyembryony has been recorded for this group in the case
of Aphelopus thelicE Gahan ms. parasitic on Thelia bimaciilata.

The method of attack and oviposition is very similar for
nearly all species of the Anteonince. Upon nearing the prey the
female comes to a standstill, throwing her antennae backward
and swaying from side to side before making her jump. She
then throws herself upon the hopper, which may escape by a
quick jump. The chelate claws of the forelegs are thrust out
at the same time, and the prey is seized in this manner. Once
the hopper is captured, the female grasps it with all six legs,
and placing her body sidewise across that of the struggling host,
curls the tip of the abdomen under and thrusts the sting in
the ventral side of the thorax in the case with Gonatopus
erythrodes (Perk.) and Ilaplogonatopiis americanus (Perk.)
as observed by the writer, and with Echthrodelphax as noted by
Perkins (1905). With Gonatopus contortulus Patton, however,
the host is large and robust and winged, so that the method of
attack is varied. Here the prey is stalked from behind, and

196 The Ohio Journal of Science [Vol. XVIII, No. 6,

while clinging to it with the front pair of legs, the sting of the
dryinid is thrust forward beneath her body and into the ventral
side of the abdomen of the jassid. If the latter jumps, both fall
to the ground ; and in this case the process of stinging is similar
to the above-mentioned species. With other species the
sting is thrust in different places on the host, according to various
observers. While the hopper is struggling, the parasite may
be seen trying to locate a vulnerable spot in its host's armor.
This always seems to be in the thin membrane separating
adjacent segments. Once stung the hopper is paralyzed, and
it is then dragged to a convenient place for oviposition. In
several instances the parasite dragged the stunned jassid around
the vial for some time before stopping to oviposit. In doing
this the mandibles are used, the wasp backing around, dragging
the host after it. In many cases the host was oviposited in
immediately. In the case of opaque nymphs, before the
ovipositor is inserted, the sting can be seen working this way
and that with extreme rapidity, just beneath the cuticle.
It is finally withdrawn, and the ovipositor thrust in deeply into
the body of the nymph, and the egg laid. The entire operation
extends from one to often two or three minutes.

In Gonatopus contortidus Patton the egg may be laid on the
outside of the cuticle, although it is more often thrust beneath
it. In a Gonatopus species parasitic on Deltocephalus immicus
nymphs, the egg is thrust down between the segments, the tip
just protruding outside.

Perkins (1905) remarks that "occasionally after capture, the
prey is released without being stung, and that it is probable
that hoppers so released have already been stung by an earlier
captor." The writer is inclined to doubt this as the real
explanation, since more than one larval sac is often found on
one host. Once a nymph was found bearing a nearly mature
larval sac and an egg just protruding externally between the
segments. As there was only enough food present in this nymph
for the maturing of but one parasite, the fate of the youngest
dryinid is obvious. It is evident that it is just a matter of
chance whether the captured nymph has been previously
parasitized or not, and that when the female does not oviposit
the cause must be something other than that of previous
oviposition.

April, 1918] Parasites of LeaJ-IIop per s 197

In some species there is just enough food present in the hopper
to feed but one dryinid, while others may be seen bearing from
two to as many as eight parasites. In fact, with Brucomorpha
ocidatiis, two or three parasites are more often found attached
than one, and these may all mature if the host is a large, robust
individual. Generally the parasites on a hopper are of one
species, but may be of two different species. One sex seems
as readily attacked as the other. Some species confine their
attacks entirely to nymphs, others to adults, and others to
both nearly mature nymphs and adults. Once a nymph is
parasitized it never becomes adult, in so far as observed by the
writer. Keilin and Thompson (1915) state that the Typhlo-
cybidce parasitized by Aphelopiis malaleuciis Dalm. are ovi-
posited in before becoming adult. With Erythroueura tricincta,
parasitized also by a species of Aphelopiis, adults have been
kept in captivity as long as a week before a sac appeared
externally. It would seem, then, that with this species the
egg is laid after the host becomes winged. Nymphs parasitized
by Gonatopiis and its allies never become adults, no matter
if they are in their last instar. Gonatopiis erythrodes (Perk.)
attacks nymphs in the second, third and fourth instars, but
prefers nymphs of the fourth instar. The place selected for
oviposition with a given species of dryinid is generally restricted
not only to either the abdomen or thorax, but also to certain
segments of these, or in certain positions on these segments, as,
for instance, with Haplogoiiatopiis americanus Perk. Its sacs
are found invariably protruding dorsally between the last two
or three abdominal segments. Very often a species of dryinid
confines its attacks to one species of hopper. Others may
parasitize closely-related species; and still others may parasitize
species in different genera.

After oviposition the dryinid drops the hopper and runs
away. The latter recovers rapidly, and generally within five
minutes recovery is complete and feeding is resumed. The
number of eggs a female may lay is probably considerable,
though not as large as with many hymenopterous parasites.
In captivity the largest number of eggs obtained was thirteen,
and the greatest number in a single day was six. These figures
are unquestionably much too low for normal conditions. The
females are ready for oviposition as soon as they have fed after
emergence, so that the oviposition period must extend through-

198 The Ohio Journal of Science [Vol. XVIII, No. 6,

out the entire period of adult life, probably a month in nature.
Basing the above number of eggs per day on this period of
oviposition, the maximum number of eggs that can be laid is
one hundred and eighty.

Egg. — Since the egg is very minute and often is thrust down
deeply into the body, it is practically impossible to locate it
either by dissection or by sectioning the host. For this reason,
the date of hatching is still in doubt, and the incubation period
can only be estim_ated from the time the egg is laid to the
appearance of the sac outside the body-wall of the hopper.
In Gonatopus erythrodes (Perk.) a blackish discoloration precedes
the appearance of the sac by several days, and is noticed five
days after oviposition. Since in this case the discoloration
is due to the black sac beneath the cuticle, the egg must have
hatched within two or three days after being laid.

Larva. — The earliest stages of the dryinid larva have been
described by Keilin and Thompson (1915) for Aphelopus
malaleucus Dalman parasitic on Erythroneura (Typhlocyba)
hippocastani. They report finding an embryonic stage within
the tissues of the host, and do not consider it as the first
larval stage. The exact relation of this "embryo" to
the first stage larva is not known. It is possible that
this may be the true first instar, which may be modi-
fied and different from the following ones. In case the
egg is internal, just previous to emerging from the host, the
larva is seen to be curved into the form of a U, the apex of which
appears first between the segments. This is the second larval
stage, and it is covered almost entirely and protected by the
exuvium of the first instar. The appearance of the sac exter-
nally on the host takes place in from five to seven days after
oviposition. During this time the egg has hatched, the
"embryonic" stage has been passed, and a molt has taken
place.

The second molt occurs in from five to ten days after the
emergence of the sac, and is indicated by the rupturing of the
first exuvium along a median dorsal line. Between the gaping
halves of this split, the second exuvium is gradually pushed
outward by the growth of the larva within, now in its third
instar. Accompanying this split there is often a change in the
coloration of the sac, as from dark green to black.

April, IDIS] Parasites of LeaJ-IIoppers 199

As growth occurs, the second exuvium becomes spHt open
in the same manner as did the first, a fact coincident with a
third larval molt, and the third exuvium is pushed outward
by the larva, now in its fourth instar, exactly as before. The
fourth larval stage is not comparable to the other four. It is
characterized by the development of the m,andibles, beneath
which are formed those of the fifth stage. There is no complete
fourth molt, so far as is revealed by dissections, other than a
shedding of the head capsule. It is possible that this instar
is entirely eliminated in some of the more highly specialized
species. The larval sac has become of considerable size by now,
and is plainly visible on nymphs and brachypterous forms.
In winged species, the sac pushes one wing and elytron up out
of place. The host becomes very evidently affected by now,
being sluggish and easily caught. In from eighteen to nineteen
days after the appearance of the sac the fourth partial molt
occurs, and the larva enters upon its fifth and last larval instar.
Up until this time it has not touched any vital part of the host.
The mouth-parts of all the preceding instars are soft and slightly
chitinized, while in this last stage, sharp, heavily chitinized
mandibles are developed and the tissues of the host are
devoured. Growth is now very rapid, and soon the larva
breaks through the sac, first being seen as a narrow white band
between the slightly gaping halves of the exuviae. Peristaltic-
like movements take place spasmodically in the maggot, w^hich
increases visibly in size. In the case of Gonatopus erythrodes
(Perk.) parasitic on Deltocephalus inimicus nymphs, the last
stages are easily studied as the host is not very heavily chitinized
and opaque yellow in color. The dark jaws of the parasite can
be seen working in the interior of the host's abdomen first.
The entire abdomen is emptied of its contents before the parasite
directs its attention to the thorax. At this time, an hour after
the last molt, the hopper is still alive and responds to stimulus,
but it now dies very quickly as soon as the thoracic viscera are
attacked. The feeding seems to be a combination sucking, in
which the cone-like labium functions and the cutting by the
sharp mandibles. The head is attacked after the thorax, and
soon there is nothing left of the hopper but a mere shell. The
contents of the host seems to be partially liquified at the final
stages. About two hours elapse between the final molt and the
consumption of the host.

200

The Ohio Journal of Science [Vol. XVIII, No. 6,

Coincident with the growth of the parasite is the formation
and development of the sac. (Figs. 1-7). When the larva
hatches it is generally entirely or almost within the body of
the host and surrounded by its tissues. As it grows it gradually
assumes a curved position, due either to unequal growth or
pressure against the host's tissues. It also pushes its way along
the path previously made by the ovipositor toward the external
point of insertion of the latter. If the egg is laid internally,
it undergoes its first m.olt still within the body of the host, as

Figure 1.

1-6, side view of abdomen of Deltocephalus inimicus nymphs, showing stages in
parasitism by Gonatopiis sp.; 1, egg; 2, second instar; 3, third instar; 4, fourth
instar; 5, fifth instar; 6, empty sac; 7, dorsal view of abdominal segments of Delto-
cephalus sayi adult 'showing empty sac of Gonatopus sp.; 1st, 2nd. and 3rd. Ex.,
first, second and third exuviae; Lv., larva; Vp., ventral larval process.

noted; and instead of entirely breaking through the exuvium
dorsally in the normal manner, it splits the latter ventrally and
uses it as a buffer in forcing its way outward. It becomes
more and more curved in position, and finally breaks through the
cuticle of the hopper still with the protection of the first exuvium,
which surrounds it as a sort of hood. At no external point,
then, is the larva exposed, but with its head and caudal end still
in the tissues of the host, it feeds and the first exuvium becomes
fused with the cuticula of the hopper by a series of chitinous
hooks or other means. At the second molt, the second exuvium

April, 1918] Parasites of Leaf- Hoppers 201

is split ventrally as was the first, and it is pushed outward by the
growth of the third instar within, breaking through the first
exuvium which is firmly attached to the host. This process
is repeated in the third molt, the second exuvium becoming
split open by the third. The exuviae become chitinized and
darken in color in some species. In Phorpas mirahilis (Perk.)
parasitic on Briicomorpha ocidatiis, this darkening and hardening
of the exuvium begins in the region of the spiracles, and at first
there is on each segment an oval black spot around each.
These later become confluent and it seems that apparently
after the skin is shed the air has something to do with the
hardening of the exuvium. The exuvium just shed also adheres
very closely to the growing larva which at first breathes through
the spiracles of this cast skin. Tracheae can be seen in direct
connection between the first pair of spiracles, even in the late
phasis of this instar. Just before the fourth molt, then, there
is present the fourth instar parasite larva, protected dorsally
by the third exuvium and laterally by the first and second.
During the late phases of the second instar, the parasite pushes
itself completely outside of the host, remaining attached by the
chitinous ring of the sac only, and with its head and posterior
extremity just within the body of the hopper. Later on the
head alone remains within the body of the host.

With the final consumption of the host, the maggot with-
draws its head from within the shell of the latter, which it
quits, and goes off in search of a place to spin its cocoon. In
some species the dead shell of the host is jerked abruptly away
by the maggot (Misra, 1917). The larva is quite active, and
crawls around on its back similar to a Cotina grub by peculiar
peristaltic like movements, originating at the caudal end as a
constriction which progresses anteriorly. Sometimes as long as
a day is spent before the cocoon is started, and again it is begun
almost immediately, even beneath the dead host (Swezey, 1903).
The cocoons are spun either beneath the soil or above it on
leaves or other objects. Sometimes there is variation in
individuals in regard to the location of the cocoon. The first
generation cocoons of Gonatopus erythrodes (Perk.) were spun
below the soil, but some of these of the second were spun
above the soil on the sides of the cages. The cocoon is com-
pleted in from one to two days, although larvae that are to pass
the winter may be seen lining their cocoons for days after they
are apparently finished.

202 The Ohio Journal of Science [Vol. XVIII, No. 6,

Pupa. — Directly after completing the cocoon, the larva
becomes quiescent, and within one or two days assumes a
straight, rigid shape, becoming a prepupa. After five to seven
days pupation occurs, the larval skin being pushed down into
the bottom of the cocoon as a tiny crumpled mass. The
time spent in the pupal stage varies greatly — from three to
five weeks with different species. At the end of this time, the
adult issues, but does not as a rule emerge from the cocoon
until two or three days later. It then chews its way out at or
near one end, and is ready for active life immediately. The
total life cycle varies from a year to as low as forty-three days.

SEASONAL HISTORY.

Hibernation. — All species of the dryinids studied by the
writer have so far passed the winter as larvae within the cocoon ;
and this is probably true of the great majority of them. A few
got as far as the prepupal condition, but all larvse collected in
September and late August remained as such. There seems to
be no difference in the structure of the winter and summer
cocoons.

Generations. — The number of generations a year depends
upon the species of host attacked, and of course upon the
climate. Species attacking a host that has two generations a
year— as with Gonatopus erythrodes (Perk.) parasitic on Delto-
cephalus inimicus — have the same number of generations.
Others have but one. Two generations a year seems to be the
maximum in this latitude, though farther south there are
probably more. When a species attacks but one stage of its
host, it must spend most of the year in the cocoon, issuing only
when the right stage of its host is present. In one case, that of
Chelogyniis chlorotettixi n. sp. parasitic on Chlorotettix unicolor,
the nymphs of which are found only in June in the Lake Erie
region, spins its cocoon in early July and remains as an active
larva within its cocoon throughout the remainder of the summer
until the next spring, when it pupates and emerges in June in
time to attack the nymphs of its host. One cocoon of this
species was carefully split open in September, and an active
white larva revealed. Later it was noticed that this larva
had spun over the slit in its cocoon.

In some cases, where there are two generations a year,
part of the larvse of the first will pupate and the rest will remain

April, 1918] Parasites of Leaf-Hoppers 203

as larvae until the following spring. A similar instance was
noticed by Perkins (1905) in the case of estivating larvas.
He says: "The number that remain as larvae, when others
emerge after the usual period, may amount to not less than
twenty-five per cent. " From two thousand cocoons of (Dryinus)
sent to Perkins in Hawaii from North America and arriving
there in November, two males emerged a few weeks after
arrival, and one male a month later. After being removed to a
cooler and damper place, both sexes issued in a short time.
This retarding of development is characteristic not only in
localities where winters are long and severe, but also where a
hot, dry period is experienced. It is a phenomenon that has
been noticed with other insects, and is a means probably of
insuring a continuance of the species in case the immediately
following generation is not successful and perishes for want of
food.

DESCRIPTION OF STAGES.

Egg. — The egg of the Gonatopits species studied (Figs. 8 and
9) is either oval or kidney-shaped, and varies in color from light
yellow to dark grey or dark brown. In length it averages from
.15-.211 mm. and in width from .08-. 095 mm. There is no
sculpturing on the chorion of any studied and in the case of
those laid within the body of the host, no surface structures.
In one egg, that of a Gonatopus species on Deltocephaliis inimicus
nymphs, a circular ridge, hardily visible, was noticed at one
pole, and a black excrescence about half way down on one side.
When laid on, or partly exposed on, the cuticula of the host
it may be of the same color and almost invisible, as in Gonatopus
contort uhis Patton, or its color may be sharply contrasted with
that of the host.

Larva : First instar. — The earliest stage found was that of
a larva within its egg shell ready to hatch (Fig. 10). It was
structureless, with no visible appendages or segments. A small
dark brown spot present at one extremity might serve for the
purpose of breaking open the egg chorion, which is comparatively
hard and thick.

Second instar. — The second instar of Aphelopus comesi n. sp.
was studied, (Fig. 11). The specimen observed measured
.54 mm. in length and .23 mm. at its greatest breadth near
the head region.

204

The Ohio Jounmt of Science [Vol. XyiTlj No; 6\

The posterior third of the body is flexed downward and
forwards, the anal extremity pointing cephalad. Two lobed
processes projecting dorsally from the cephalic region are the

I

(
I

Figure 2. ' ' ' •

8, egg of Gonatopus sp.; 9, egg of Gonatopus erythrodes (Perk.); 10, first instar of
Gonatopus erythrodes (Perk.); 11, second instar of Aphelopus sp.; 12, third instar
of Gonatopus contortidiis Patton; 13, side view of mouth parts of fourth instar of
Anteon sp.; 14, dorsal view of fourth instar of Aphelopus sp.; 15, fifth instar of
Gonatopus contortulus dissected from sac; 16, dorsal view of mature larva of Gona-
topus sp.; 17, dor.sal view of head of Gonatopus sp., mature larva; 18, ventral view
of head of Aphelopus sp., mature larva; 19, ventral view of male Ha pi agonal opus
americanus pupa; 20, side view of same; 21, cavity in body wall oi Erythronenrd
comes produced by Aphelopus sp.; 22, dorsal view of head of Aphelopus sp., fourtl-^
instar showing relation to larval sac, and chitinous ring. 1st, 2nd., and 3rd. Ex.,
first, second and third exuviae; Vp., ventral larval process; md. int., mid intestine;
sg., supra-esophagial ganglion; hi., head lobe; phe., pharynx-esophagus; . vg.,
ventral nerve chain; p., proctodeum; h., head; spin., spinneret; ch., chitinous
head process; e., eye; Ibr., labrum; md., mandible; mxp., maxillary palp; lab.,
labium; a., antenna; Ibp., labial palp; sp., spiracle; ment., mentum; sm., subr
mentum; ph., pharynx; mdr., developing mandibles.

April, 1918] Parasites of Leaf- Hoppers 205

only mouth ])arts. A large mid-intestine beginning just back of
the head extends throughout the greater length of the body
to near the anal end. The pharynx-esophagus is a very narrow
tube, distinguished below the supra-esophagial ganglia. The
hind-intestine is contiguous with the mid-intestine, but there
is no connection between the two. In Aphelopus melaleucus
Dalm., according to Keilin and Thompson, the salivary glands
are very long and have a winding course. They were not seen in
the specimen studied by the writer. Twelve ventral ganglia
are easily distinguishable, the two anterior of which are doubtless
thoracic and cephalic. The second is connected with a more or
less elongated ganglion located just ventral to the first or sub-
esophagial ganglion. The above mentioned authors report
finding fourteen ganglia in the ventral nerve chain of Aphelopus
nielaleucus Dalm. instead of twelve, which is the ordinary
number in other Hymenopterous larvae. Nine pairs of spiracles
are present in this stage.

Third ijistar. — In this stage the larva assumes a more marked
curved position, being bent ventrally in the form of a U, the
cephalic and caudal ends being approximate, (Fig. 12). The
head and anterior third of the body lie buried in the tissues of
the host, the rest of the larva protruding externally. It is
completely enclosed in the second exuvium, except the head
region. The first exuvium lies midway across each side of the
body. In some species the ventral portion of the body immedi-
ately back of the head is prolonged into a peculiar appendage,
which is enclosed by a cone-shaped membrane, the cuticle of
the second exuvium. This is turned outwardly and posteriorly
and its tip lies just beneath the chitin of the host at the juncture
of two metameres, and is often visible externally in the case of
light colored nymphs as a dark brown process extending under
the cuticula from the point of attachment of the parasite.
Often there is no corresponding larval tissue within this and
in this case a small opening is noticed at the tip. It may serve
to tap the outside air and thus keep a supply under the second
exuvium for the enclosed larva. There is no connection between
the spiracles of the second exuvium and the larva unless the
thoracic pair retain this as they do in the next stages. The
mouth parts are similar to those of the preceding stage except
that they are larger. In Gonatopiis contortiihis Patton they appear
as two oval, fleshy lobes projecting prominently from the head

^06 The Ohio Journal of Science [Vol. XVIII, No. 6,

region. At their bases the head is constricted visibly and at
this constriction there are four triangular chitinous processes.
These lobes may represent the mouth parts alone and may also
be considered as the rudiments of the head. They are very
firmly embedded in the tissue and are often detached and
overlooked. Eleven distinct ganglia of large size comprise
the ventral nerve chain, the first two being the largest. Beneath
the second exuvium the cast tracheae and their branches can
be seen. Nine spiracles are present.

Fourth instar. — The exact status of this stage is not yet
determined. It is indicated only by the characteristic mouth
parts, (Figs. 13 and 14), and the exuvium has riot been noticed
if indeed there is more than a molting of the head parts. It
is obvious that the change from the peculiar soft lobed mouth
parts of the third instar to the heavy chitinized mandibles of
the last instar is too abrupt and that there must be an inter-
mediate stage probably having been mostly suppressed with the
specialization of the group. The mandibles in this stage are
distinct curved processes with broad rounded and notched
tips, thinly chitinized. A simple truncate labrum is present.
Shortly after this stage has been reached the fifth instar mandi-
bles develop back in the head, the point of each projecting into
•the base of the fourth instar mandibles. This instar is not
represented by any of the three exuviae composing the larval
sac.

Fifth instar. — Shortly before the appearance of the larva
from the larval sac the mouth parts of the last instar are com-
pletely developed, and larvae dissected out of sacs showing the
three larval exuviae present the same type of mouth parts as
the mature free-living larva, (Fig. 15). Though fully developed
late in the preceding stage the cutting mandibles do not function
until this instar. The mouth parts are represented by the
rounded labrum, the dark brown, curved, sharply pointed
mandibles, the maxillge, and labium. Just dorsal to, and at
the base of, the mandibles are the irregular pigmented eye
5pots. Nine spiracles are present, the first pair being of large
size and brown color and situated back of the head on the
second thoracic segment. All the other spiracles are indistinct.
The body still retains the U shape, though now the anal extrem-
ity reaches only as far as the first or second thoracic segments.
If the mature larva is hairy as in the case of Aphelopus species.

April, 1918] Parasites of Leaf-Hoppers 207

the hairs can be seen resting back along the body, those of the
first segment pointing cephalad, those of the rest caudad. In
some species the peculiar cone-like structure is still present
in this stage. This is evidently peculiar to only those species
ovipositing in the abdomen of the host and it is characteristic
of at least four stages since in nearly mature larvae a series of
four, all telescoping more or less within each other, can be
seen. It has been found in parasites ovipositing in the abdomen
of Chlorotettix iinicolor nymphs, Deltocephalus inimicus nymphs
and adults, and Deltocephalus sayi adults.

While in the sac the head of the dryinid larva is always
ventral and the caudal end of the body is curved upwards and
anteriorly lying dorsal to it. The spiracles of all the exuviae
are open, but are not connected in any way with those of any
of the larval spiracles. However, a trachea can be seen running
from the first spiracle of the third exuvium to the corresponding
spiracle of the larva itself, so that it is possible some air may be
taken in by the parasite in this way. The fifth instar is enclosed
laterally by the three exuviae and dorsally by the third only.

Mature Larva. — The mature larvae vary in size, those studied
measuring from 1.5 mm. to 4 mm. in length. They are blunt
and wider posteriorly and more pointed anteriorly (Fig. 16).
They are variously colored, agreeing in many cases with the
color of their hosts. They may be various shades of green and
even light pink, but as a rule they are white. Aphelopus larvae
are quite pubescent, but those of the more highly specialized
types are almost without hairs. The head is distinctly separated
from the body and is light brown or grey in color. Thirteen
segments are visible not including the head. They are more or
less concealed by the folds in the skin. On the head (Figs.
17 and 18) dorsally the two black pigmented eyes, the rudi-
mentary antennae and the short arcuate labrum are visible.
The former may be but slightly pigmented and very indistinct,
even in a species that normally has pigmented eyes. The
labrum is covered with short spines and appears as a fleshy
projection from the side. The curved heavy mandibles are
very prominent. They are one-jointed and articulated from
a chitinized basal piece. In Neodryinus and Paradryinus the
cutting edge is crenulate or denticulate (Perkins, 1905), or it
may be notched in Gonatopus (Mik, 1880). The maxillae are
less distinct. Laterally they are represented as short truncate

208 The Ohio Journal of Science [Vol. XVIII, No. 6,

fleshy lobes from which project the short one-jointed maxillary
palpi. The latter bear at their ends two minute papillae, each
with a spine at the tip. The labium is large and very prominent.
From the side it is seen as a fleshy elongated process with a
distinct apical half cut off from a broader basal piece. The basal
piece or sub-mentum of the labium extends nearly back to the
posterior margin of the head. To it is attached the mentum
which extends just beyond the tips of the mandibles and bears
at its tip a semi-circular transparent organ. A narrow tri-
angular piece is attached about half way back on the ventral
side of the mentum and projects to the tip of the labium.
It is the spinneret and at either side of this the small one-jointed
labial palpi are attached.

The pair of thoracic spiracles are nearly concealed under
a fold in the integument and the others are very small and
usually not visible. All the abdominal segments except the
last two bear a pair of spiracles, there being ten pairs in all.

Prepupa. — After spinning the cocoon the larva soon becomes
quiescent and shrinks down into the lower half of it. Within
a few days it becomes straight, rigid, extending the length of
the cocoon, and constricted near the middle. After assuming
this position it soon pupates.

Pupa. — The pupa (Figs. 19 and 20) is white, all the parts of
the adult insect being plainly distinguished. If the adult is
winged the wing pads are large and extend the length of the
thorax to the first abdominal segment. Seven abdominal
segments are visible. Pigmentation begins in the compound
eyes and gradually spreads over the entire body the pupa
becoming black before the adult issues.

Larval Sac. — As previously stated the laryal sac is merely
the cast skins of the larva itself modified for protective purposes
by being more or less chitinized, and firmly attached to each
other and to the host. The means of anchoring the sac to the
host varies in different genera. In Aphelopiis this attachment
consists of a sort of chitinous ring and two hooks, (Figs. 21 and
22). One of the latter is smaller, sharply pointed, and strongly
chitinized, the other is larger, blunt and but thinly chitinized.
The chitinous ring serves to keep the cuticula of the host open,
thus affording a place for the head and mandibles of the
parasite.

April, 1918]

Parasites of Leaf-IIoppers

209

Figure 3.

23-25, dorsal, ventral and side views of Erythroneura comes parasitized by
Aphelopus sp.; 26, ventral view of Agallia sanguinolenta parasitized by Anteon sp.;
27, dorsal view of Dellocephalus inimicus nj'mph parasitized by Gonatopus eryth-
rodes; 28, ventral view of Chlorotettix unicolor nymph parasitized by Chelogynus
sp.; 29, side view of Liburnia campestris adult parasitized by Haplogonatopiis
americanus; 30, dorsal view of Brucomorpha oculatus parasitized by Phorbas
mirabilis.

210 The Ohio Journal of Science [Vol. XVIII, No. 6,

The sac itself is of various shapes, sizes and colors. Those
of AphelopHS (Figs. 23, 24 and 25) are elongate oval, and
those of Chelogynus are rounded and much compressed. Between
these two types there are all sorts of gradations. Black is the
predominating color and there are variations from it to brown
and yellow. Many are green or blue-green, and others are
banded transversely with alternate stripes of light and dark
brown. In the latter case the colors are arranged segmentally.
When the skin that is to form part of the sac is molted it is
light colored and not pigmented and modified. As it becomes
exposed to the air it assumes the black or fuscous color of the
sac. In Phorbas the exuvium is pale yellow when first molted,
but soon each spiracle is surrounded by a black spot. As
previously noted, these enlarge and become confluent, the
exuvium finally becoming jet black in color. Invariably the
first exuvium is smooth and shining, the second often crinkled
or sculptured on its outer margin, and the third is generally
very distinctly rugosely sculptured by raised zig-zag lines or is
punctate. The sac is located in diverse positions, according
to the genus of the parasite. In Echthrodelphax, Paradryintis,
Thaumatodryinus and Neodryinus it is found on the thorax
beneath the wing. In Phorbas (Fig. 26), from various places
on the abdomen. In most of the Anteon species (Fig. 30), on
the ventral side under the hind legs, in Paranteon on the
abdomen, ventral as well as dorsal, seldom on the sternum or
propleura, in Pseudogonatopus on the dorsal and lateral sides of
the abdomen, in P. stenocrani Perk., upright between both
wings (Perkins, 1905). In Gonatopus (Fig. 27) it is on the side
of the abdomen, in Haplogoyiatopus (Fig. 28), on the dorsal
posterior part of the abdomen, in Chelogynus, (Fig. 29), on the
side of the thorax, and in Aphelopiis on the sides of the abdomen
between the first few segments. It always projects between
two segments. Often the spiracles of the different exuviae
composing the sac can be seen if it is light colored.

Cocoon. — The cocoon is either spun below the ground as is
the case with Aphelopus, Chelogynus, or Phorbas, or above on
some convenient object, as with Ha plo gonatopus, or some species
in Gonatopus. That of Aphelopus, (Fig. 31), is small, oval,
and white, of Chelogynus, (Fig. 33), larger, oval, plastered
over with sand grains or soil particles, and that of Bocchus,
(Fig. 32), tough, thick, brown and fibrous, with soil particles

April, U)1SJ

Parasites of Lcaf-IIoppers

211

mixed all through it. Many spun above the ground as in some
species in Gonatopus, (Fig. 34), are white, shining, and very
like a spider's cocoon. According to Perkins (1905) the cocoons
of Paradyinus are often adorned with bits of leaf tissue, that of
Ncodryiniis roofed over with the ruptured larval sac which is
removed from the hopper and attached. The cocoon of Pro-
dryinus ormcuidis (Ashm.) is spun beneath the dead hopper
(Swezey, lUOo). The cocoon is always composed of two parts,
and inner more compact structure, within a more loosely woven
part.

Figure 4.
31, cocoon of AphelopHS sp., 32, cocoon of Phorhas mirabilis; 33, cocoon of
Chelogynus sp.; 34, cocoon of Haplogonatopus americanus.

INTERNAL ANATOMY OF THE LARVA.

Internally the mature larva presents the following char-
acteristics. The mouth leads into a narrow pharynx-
esophagus the limits of either of the latter not being definable.
This opens into a very large, thick-walled mid-intestine, which
is blind at its posterior end, not being connected with the hind-

212 The Ohio Journal of Science [Vol. XVIII, No. 6,

intestine, which, however, is present at this stage. The
pharynx-esophagus is long, extending back to the hind thoracic
segment. It is very narrow with a small lumen for the greater
part of its length, but broadens out gradually at first, then
abruptly, into the mid-intestine. The latter occupies the
greater part of the body cavity and becomes greatly distended
at maturity, food being taken in much more rapidly than it
can be digested, so that the entire mid-intestine acts as a food
reservoir.

A set of powerful muscles is attached to the dorsal part of
the pharynx-esophagus and a smaller set below it. These act
in such a way that their contraction forces this structure open,
thus forming a sucking organ. Aside from these the only other
striped muscles in the larva of note are those controlling the
mandibles. The cells of the pharynx-esophagus are small.
Those of the anterior fourth of the mid-intestine are large,
columnar, and digestive in function, in contrast to those of the
greater part, which are elongate and flattened.

The salivary glands are very large, appearing as two sets of
long convoluted tubes, one on each side of the mid-intestine,
extending about three-fourths the way down it in the mature
larva. Each set is composed of two branches ending anteriorly
in a short lateral tube. These two lateral branches open into
a common duct which extends into the spinneret. They serve
primarily as silk glands in the spinning of the cocoon. The
cells composing the main duct are columnar, thus differing
distinctly from those of the rest of the glands. Undoubtedly
the true salivary function is principally confined to this section
of the glands.

There is no caudal vesicle on the larva as observed for
Apanteles (Tower, 1916), and no malpighian tubes have been
observed in any of the stages. Whether the waste products
are absorbed and discharged upon pupation, or whether they
are thrown off into the host is not known. The ten pairs of
spiracles lead into short lateral tracheae which open into the
two longitudinal trunks, the tracheal system in the mature larva
being very similar to the general type found. As in the third
instar the nervous chain consists of eleven ganglia.

THE COMMON RAVENS OF NORTH AMERICA.

By Harry C. Obekholser.

The subspecies of the common raven, Corviis cor ax Linnaeus,
are among the most difficult birds of the family Corvidae.
The differences characterizing them are almost wholly those of
size and proportion; and because great individual variation
complicates the case, these are largely average distinctions and
require series of specimens for proper elucidation. In the Old
World some 16 or 17 forms are at present recognized,^ but in
North America currently only two, Corvus corax principalis
of northern North America, with which the birds of the eastern
United States are considered identical, and Corvus corax sinuatus
of the western United States and Mexico. Another, however,
has been recently described by Doctors Rothschild and Hartert
as Corvus corax clarionensis,- from Clarion Island, in the Revilla-
gigedo group, western Mexico, and it is now proposed to
separate the bird from the eastern United States as a fourth
race. By the recognition of these two additional subspecies,
the measurements, characters, and distribution of the two
original forms are in need of considerable readjustment, and
they are, therefore, included in the discussion that follows.

The material used in the present study consists of about
400 specimens. This represents the collections of the United
States National Museum, including the Biological Survey;
the Academy of Natural Sciences of Philadelphia; the Museum
of Comparative Zoology at Cambridge, Massachusetts; the
American Museum of Natural History; the Brooklyn Museum
of Arts and Sciences; Dr. Louis B. Bishop, Dr. Jonathan Dwight,
Dr. L. C. Sanford, Dr. Witmer Stone; Messrs. William Brew^ster,
John E. Thayer, A. C. Bent, and E. A. and O. Bangs. To the
authorities of these museums and to the individuals here
mentioned the writer is indebted for placing their specimens at
his disposal.

^For an excellent account of the Old World forms of Corvus corax, cf. Hartert,
Vogel palaarkt. Fauna, Heft I, November, 1903, pp. 2-8; Heft VI, June, 1910,
pp. XIII-XIV.

^Novit. Zool., IX. Xo. 2. July 25, 1902, p. 381.

213

214 The Ohio Journal of Science [Vol. XVIII, No. 6,

All the measurements in this paper are in millimeters, and
have been taken as described in the author's paper on Butorides
virescens}

CoRVUS CORAX PRINCIPALIS Ridgway.

C[orviis\. corax principalis Ridgway, Man. North Amer.
Birds, 1887, p. 361 ("Northern North America, from Greenland
to Alaska, south to British Columbia, Canada, New Brunswick,
etc.").

Corviis corax var. littoralis Holboell, in Kroyer's Tidskrift,
IV, 1843, p. 390 (Greenland; Labrador) (nee Corvus littoralis
Brehm).

Chars, subsp. — Size largest of the North American races.

Measurements.^ — Male:^ wing, 426-457 (average, 446.5) mm.;
tail, 241-259 (250); exposed culmen, 67-76 (71.3); height of
bill at nostrils, 26-30 (27.7); tarsus, 66-71 (69); middle toe
without claw, 45.5-54 (49).

Female:* wing, 413-441 (average, 426) mm.; tail, 228.5-246.5
(250) ; exposed culmen, 68-72 (70.4) ; height of bill at nostrils,
25-27.5 (26.1); tarsus, 64-70 (66.5); middle toe without claw,
43.5-48 (45.5).

Type locality. — Saint Michael, Alaska.

Geographic distribution. — Northern North America. Breeds
North to northern Greenland, Grant Land, Melville Island,
Banks Land, Herschell Island, and the northern coast of
Alaska; west to the western coast of Alaska, the Aleutian Islands,
and Vancouver Island, British Columbia; south in the Pacific
coast region to Quinault, western Washington, and in the
interior to northern Mackenzie and northern Quebec (Ungava) ;
and east to northeastern Quebec (Ungava), and Greenland.

Remarks. — This North American race is distinguished from
Corvus corax corax Linneaus of northern Europe by its relatively

iProc. U. S. Nat. Mus., XLII, 1912, p. 533.

^In part taken by Mr. Robert Ridgway; but the measurements of exposed
culmen in the Bulletin of the United States National Museum, No. 50, part III,
1904, p. 259, are really those of the total culmen, given by mistake as exposed
culmen; and there is evidently also some mistake in the height of the bill at nostrils,
as this appears to be too large. Both these dimensions have, therefore, been
remeasured for the present use.

'Six specimens, from Alaska.

^Eight specimens, from x^laska.

April, 1918] The Ravens of North America 215

shorter and higher bill, less lustrous plumage, and less developed
and more purplish-hued lanceolate feathers of the throat;
and from Corviis corax behringianus Dybowski by the long
third primary, which equals or is longer than the fifth, instead
of being decidedly shorter. As has already been intimated,
the characters which distinguish this from the other North
American forms are wholly of size and proportions. The
greatest differentiation occurs in Alaska, whence came the type
of Corvus corax principalis; hence we have here used only
Alaskan specimens in the comparison of racial characters.
Birds from Greenland and northern Ungava are somewhat
smaller than those from Alaska, but are most satisfactorily
referred to this form. Examples from the western coast of
British Columbia are also intermediate between the present
race and Corvus corax siniiatus; those from western Wash-
ington are still smaller, and, indeed, almost half-way between
Corvus corax principalis and Corvus corax clarionensis; but
birds from both these regions are better referred to the present
form than to either of the others. This applies, however, only
to the coast region, since the interior birds are decidedly nearer
the southern races, as elsewhere more fully explained. By
the segregation of the birds from the eastern United States
and by the extension of the range of Corvus corax sinuatus into
the middle portion of Canada, the range of Corvus corax prin-
cipalis becomes limited to the extreme northern parts of North
America, excepting, as above stated, on the Pacific Coast,
where it reaches southward in a relatively narrow coastal
strip as far as the State of Washington.

Corvus corax europhilus, subsp. nov.

Chars, subsp. — Similar to Corvus corax principalis, but
smaller, with a relatively larger bill.

Description.— Ty^^e, adult male, No. 260039, U. S. Nat.
Mus., Biological Survey collection; Ardell, Alabama, April 4,
1915; L. J. Goldman; original number, 211. Entire plumage
glossy black, the secondaries and inner primaries somewhat
brownish, the head, back, rump, upper tail-coverts, and breast
with a slightly bluish sheen, the throat and wings with purplish
reflections; bill and feet black.

216

The Ohio Journal of Science [Vol. XVIII, No. 6,

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218 The Ohio Journal of Science [Vol. XVIII, No. 6,

Measurements. — Male:^ wing, 417-435 (average, 427) mm.:
tail, 230-256 (242.7); exposed culmen, 64-76 (70.5); height of
bill at nostrils, 25.5-28.4 (26.6) ; tarsus, 64.5-74 (69.8) ; middle
toe without claw, 44-51 (46.7).

Female:- wing, 399-422 (average, 411.5) mm.; tail, 227-244
(238.8) ; exposed culmen, 64.5-67.5 (66.3) ; height of bill at
nostrils, 24-25 (24.5) ; tarsus, 64-70 (66) ; middle toe without
claw, 38.5-43 (41.1).

Type locality. — Ardell, Cullman County, north central
Alabama.

Geographic distribution. — Eastern United States and south-
eastern Canada. Breeds north to Lance au Loup, southern
Labrador; central Quebec; and Lake Abitibi, central Ontario;
west to Marshall County, Minnesota; and Copeland, western
Arkansas; south to Copeland and Newport, northern Arkansas;
Nashville, central Tennessee; Ardell, north central Alabama;
Chattanooga, southeastern Tennessee; Young Harris and
Toccoa, northeastern Georgia; and Mount Pinnacle and Caesar's
Head, northwestern South Carolina; east to northwestern
South Carolina; Tryon and Grandfather Mountain, western
North Carolina; Cobbs Island, eastern Virginia; Tuckerton,
eastern New Jersey; Commack Hill, Long Island, New York;^
South Manchester, central Connecticut;- Tyngsboro, eastern
Massachusetts;^ Bar Harbor, southeastern Maine; Grand
Manan Island, southwestern New Brunswick; Eastport, Nova
Scotia; and St. Johns, eastern Newfoundland.

Remarks. — The status of the ravens inhabiting the eastern
United States has long been doubtful. The fairly good series
of specimens now available shows that the birds from this
region are apparently best treated as a separate subspecies.
They are, to be sure, to a certain extent intermediate between
Corviis corax principalis and Corvus corax sinuatus; and,
furthermore, from either of these two races are separated by
only average characters, but the same thing is true of all the
other races. In the measurement averages here given, only
specimens from Pennsylvania and New Jersey southward to
Alabama are included, and these together are considered
typical. Examples from New York, New England, central

'Seven specimens, from Alabama, Georgia, North Carolina, Virginia, Penn-
sylvania, and New Jersey.

-Four specimens, from North Carolina, New Jerse3^ and Pennsylvania.
^Not breeding.

April, 1918] The Ravens of North America 219

Ontario, Nova Scotia, New Brunswick, Newfoundland, and
even the southern Labrador coast are intermediate between
the bird from the southern Allegheny Mountains and Corvus
corax principalis, but seem to be decidedly nearer the southern
form. Those from Michigan and Minnesota are in like manner
intermediate, but verge also somewhat toward Corvus corax
siuuatus, though they average nearer Corvus corax europhilus.

There is apparently no name available for this race, as
Corvus carnivoriis Baird,^ adopted from Bartram, is clearly a
synonym of Corvus corax sinuatus, as explained under that
race. 2

Detailed measurements of specimens are given in the
table on pages 216-217.

Corvus corax sinuatus Wagler.

C[orvus]. sinuatus Wagler, Isis, 1829, p. 748 (Mexico) (ex
Lichtenstein MS.).

Corvus corax sinatus Ridgway, Ornith. 111., I, 1889, p. 331
(err. typ. pro Corvus sinuatus Wagler).

Corvus Cacalotl Wagler, Isis, 1831, p. 527 (Mexico).

Corvus catototl Bonaparte, Geog. & Comp. List Birds
Eur. and N. Amer., 1838, p. 28 (southern parts of North
America.)

Corvus major Wurtumburg, Erste Reise in Nordl. Amer.,
1835, p. 294 (Nebraska) (nomen nudum).

Corvus major Ridgway, Bull. U. S. Nat. Mus., No. 50,
III, Dec. 31, 1904, p. 263 (nom. nov. pro Corvus sinuatus
Wagler [in synonymy]).

Corvus nobilis Gould, Proc. Zool. Soc. Lond., V, 1837,
(Dec. 5, 1837) p. 79 (Mexico).

C[orvus]. splendens Bonaparte, Proc. Zool. Soc. Lond., 1837
(June 14, 1838), p. 115 (nomen nudum, but probably a lapsus
calami for Corvus nobilis Gould).

C[orvus]. lugubris Agassiz, Proc. Bost. Soc. Nat. Hist., II,
1846, p. 188 (nomen nudum).

Corvus carnivorus Baird, Rep. Explor. & Surv. R. R. Pac,
IX, 1858, p. 560 (Fort Randall, South Dakota).

Corvus lugubris Baird, Rep. Explor. & Surv. R. R. Pac,
IX, 1858, p. 560 (in synonymy), p. 561 (in text) (nom. nov. pro
Corvus carnivorus Baird).

iRep. Explor. & Surv. R. R. Pac, IX, 1858. p. 560.
2C/. postea, p. 221.

220 The Ohio Journal of Science [Vol. XVIII, No. 6,

Chars, siibsp. — Similar to Corviis corax europhilus, but wing
and middle toe shorter, and bill decidedly smaller.

Measurements.^ — Male:- wing, 415.5-459.5 (average, 434)
mm.; tail, 223-254 (239); exposed culmen, 64-70 (66.8); height
of bill at nostrils, 23-26 (24.4) ; tarsus, 68-73 (71.5) ; middle toe
without claw, 41-56 (46.5).

Female :3 wing, 419-432 (average, 422) mm.; tail, 236-252
(246.5); exposed culmen, 63-71 (66.3); height of bill at nostrils,
24-25 (24.5) ; tarsus, 65-73.5 (70) ; middle toe without claw,
44.5-47 (46.5).

Type locality. — Orizaba, Vera Cruz, Mexico.^

Geographic distribution. — Middle western Canada, middle
western United States, and Mexico, to Honduras. Breeds
north to Slave River and Fort Simpson, southwestern Mac-
kenzie; west to Sicamous, Shuswap, and Okanagan, central
southern British Columbia; northeastern Washington (proba-
bly); Fort Sherman, Idaho; western Wyoming; western
Colorado; Fort Wingate, western New Mexico; San Luis
Mountains, Mexican Boundary Line, southwestern New Mex-
ico; San Pedro River, at the Mexican Boundary Line, south-
eastern Arizona; Quitovaquita, southwestern Arizona; Guay-
smas, western Sonora; Mazatlan, western Sinaloa; and Tepic;
south to Tehuantepec, Oaxaca; Guatemala; and northern
Honduras; east to northern Honduras; Vera Cruz, Mexico;
San Antonio, central Texas; the Canadian River, central
Oklahoma; Fort Riley, eastern Kansas; Fort Randall, central
southern South Dakota; and Ramsey County, central northern
North Dakota.

Remarks. — Compared with Corvus corax principalis, this
race is smaller, with a particularly small bill, which is relatively
as well as actually more slender. As will be noticed in the
geographic distribution above given, the range of this form has
been restricted in southwestern North America by the recogni-
tion of Corvus corax clarionensis as a bird of the mainland
as well as the islands off the southwestern coast; and extended
by the inclusion of a large area in middle Canada, the birds
inhabiting which are much nearer this southern race than to

^Cf. footnote on p. 214, binder Corvus corax principalis.
^Nine specimens, from Guatemala, central and southern Mexico.
^Five specimens from Guatemala, central and southern Mexico.
^Here for the first time definitely fixed.

April, 191S] rhc Ravens of North America 221

Corviis corax principalis. Birds from central southern British
Columbia are clearly the present form; in fact, specimens
examined are practically typical, as indeed also are the few
seen from southwestern Mackenzie. There is not yet sufficient
material available to determine the northeastern limits of the
range of Corvus corax siniiatus in Manitoba. Although no
specimens have been examined from the northeastern part of
the State of Washington, the raven inhabiting that region
belongs probably to the present race.

This western raven was originally described by Wagler' from
a specimen taken in Mexico. In view of the multiplication of
races in North America, and since the specimen on which
Wagler' s name was based probably came from eastern Mexico,
we hereby designate Orizaba, Vera Cruz, Mexico, as the type
locality. Concerning none of the other synonyms above cited
in the synonymy of Corvus corax sinuatus is there any question,
excepting perhaps Corvus carnivorus Baird.- This is a com-
posite name, adopted from Bartram, whose specific names, of
being non-binomial, are unacceptable; and the status of course,
this one m^ust be determined, therefore, solely by the use made
of it by Baird, since he first gave it nomenclatural status.
His account'* includes all four of the North American races
here recognized, all of which he considers as belonging to a
single form, and which he called the "common North American
raven." The name Corvus carnivorus is certainly unavailable
for the bird from the eastern United States, since Baird's text
was evidently all written before he had seen a specimen of
that form, and his characters were undoubtedly drawn entirely
from western birds. The only specimen of the eastern raven
that he was able to see at all was a specimen from New Jersey, and
notice of that he subsequently inserted in a footnote.^ It
is also indicated by his text that neither can the name be
legitimately used for Corvus corax principalis Ridgway, a view
evidently taken by Mr. Ridgway in describing the latter sub-
species. Since most of Baird's specimens are referable to the
bird now called Corvus corax sinuatus, which is the commonest
and best known North American form, it seems most logical

^Corvtis sinuatus Wagler, Isis. 1829, p. 74S.

2Rep. Explor. &• Surv. R. R. Pac, IX, 1858, p. 560.

^Loc. cit.

*Loc. cit., p. 561.

222

The Ohio Journal of Science [Vol. XVIII, No. 6,

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The Ravens of North America

223

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224 The Ohio Journal of Science [Vol. XVIII, No. 6,

to restrict Baird's Corvus camivonis to this race rather than to
Corvus corax clarionensis from the southwest, specimens of
which formed a part of the material available to Professor Baird.
Since the specimen best agreeing with his description, and the
one with which he made his chief comparisons is No. 5186,
U. S. Nat. Mus., an adult male, from Fort Randall, South
Dakota, taken, October 18, 1856, it seems proper to consider
this the type, and Fort Randall, South Dakota, therefore, the
type locality. The name, of course, thus becomes a synonym
of Corvus corax sinuatus Wagler.

Corvus corax clarionensis Rothschild and Hartert.

Corvus corax clarionensis Rothschild and Hartert, Novit.
Zool., IX, No. 2, July 25, 1902, p. 381 (Clarion Island, Revil-
lagigedo Islands, Mexico).

Chars, subsp. — Resembling Corvus corax sinuatus, but wing,
tail, and other parts smaller, particularly the bill.

Measurements. — Male:^ wing, 365-421 (average, 398) mm.;
tail, 202-231 (219.2); exposed culmen, 63-67 (65.2); height of
bill at nostrils, 24-25 (24.4) ; tarsus, 64.5-73 (68.4) ; middle toe
without claw, 42-46.5 (44.1).

Female:- wing, 370-412 (average, 389.5); tail, 207-235
(218.8); exposed culmen, 57-68 (61.7); height of bill at nostrils,
21.5-24.5 (22.7); tarsus, 63-70 (66.6); middle toe without claw,
39-47.5 (42.2).

Type locality. — Clarion Island, Revillagigedo Islands, Colima,
Mexico.

Geographic distribution. — Southwestern United States and
extreme northwestern Mexico. Breeds north to Umatilla,
central northern Oregon; west to Narrows, central northern
Oregon; Fort Klamath, central southern Oregon; Humboldt
Bay, northwestern California; the Santa Barbara Islands,
southwestern California; and Cerros Island, western Lower
California; south to the Revillagigedo Islands, Colima, Mexico;
east to Porto Bolandra, southeastern Lower California; the
eastern coast of Lower California; Tucson and Apache, south-

'Six specimens, from the Revillagigedo Islands, Lower California, and the
Santa Barbara Islands, California.

-Sixteen specimens, from the Revillagigedo Islands, Lower California, and the
Santa Barbara Islands, California.

April, 1918] The Ravens of North America 225

eastern Arizona; San Rafael River, central Utah; Wells, north-
eastern Nevada; and Umatilla, central northern Oregon.

Remarks. — This race is so much smaller than Corviis corax
principalis or Corviis corax etirophilus, particularly so far as the
bill is concerned, that no further comparison is necessary. It
was originally described by Doctors Rothschild and Hartert^
from a specimen taken on Clarion Island in the Revillagigedo
group, and has been recently recorded by Mr. Ridgway- from
the Santa Barbara Islands, California. It is undoubtedly
worthy of recognition as a subspecies, but it has, however,
much more of a geographic range than hitherto supposed. Birds
from Lower California certainly belong to the same form ; while
those from California, Nevada, all but the coast region of
Oregon, as well as most of Arizona and Utah, are so very little
larger, though somewhat verging toward Corvus corax sinuatus,
that they are certainly referable to Corvus corax clarionensis .
No specimens from the coast region of Oregon have been
examined, and they also may belong under the present race.

Measurements of specimens, including some of those from
the mainland of the western United States and Lower California,
are given in the table on pages 222 and 223.

iNovit. Zool., IX, No. 2, July 25, 1902, p. 381.
2Bull. U. S. Nat. Mus., No. 50, part III, 1904, p. 265.

THE OCCURENCE OF A PROBABLE GYNANDROMORPH

IN THE HOMOPTERA.

DwiGHT M. DeLoxg.

Gynandomorphism occurs almost exclusively in insects and
has been found in different and quite diverse orders, including
Lepidoptera, Hymenoptera and Diptera, where it is commonly
found, and more rarely in Coleoptera. This condition has
never been reported, however, for a member of the Homoptera
although previously noticed no doubt by workers in this group.
Prof. J. G. Sanders has observed this condition in a single
specimen of Deltocephahis sayi.

This curious phenomenon may be manifest in two different
ways, either by ah anterior-posterior arrangement of both male
and female structures, or by a lateral arrangement. The latter
type is by far the most prevalent and very often a butterfly
or moth is found with one wing of male and the opposite one
of female coloring; also flies, ants and bees will often display
the lateral type, but it is very rare that anterior-posterior
gynandomorphism is found.

The specimen at hand is Chlorotettix unicolor Fh. which was
kindly sent to me with some other Cicadellidce by Frank H.
Lathrop and was collected at Corvallis, Oregon. The species is
of a uniform green color and the structural characters of typical
examples of the two sexes are just alike except the genitalia, so
the occurrence of both male and female characters in this
specimen is determined entirely by these genital structures,
which are quite different and distinct in the two sexes.

By a study of this specimen and a comparison of the geni-
talia with those of the typical male and female, some very
striking similarities and resemblances to parts of the genitalia
of each sex is noticed and a combination of typical characters
of both sexes is seen instead of a set of different or freakish
characters. I will attempt to point out these similarities with
the aid of the accompanying diagrams which show the genitalia
of the typical male and female, also the specimen having the
combination of characters.

If this is a gynandromorph which seems to be the case,
it is of the anterior-posterior type, the anterior structures being
female and the posterior male.

The last ventral segment of the abdomen is undoubtedly
a female segment and contains the median notch on the posterior

226

April, 1018] Gynandromorph in the Ilomoptera

227

margin which is characteristic of that sex. The segment in
this case is much shorter than the typical female, but resembles
it very much, while it bears no resemblance whatsoever to the
last ventral segment or the valve of the male. The male valve
in fact seems to be entirely missing and replaced by this female
segment. The plates, however, are well developed and typical
of the male* of the species. Although slightly larger in size,
the shape, structure and position are the same and the outer
margins are similarly armed with spines.

The pygofers resemble the male of unicolor more than the
female structures. The opening with keeled sides at the post-
erior end resembles- very closely the male, but the pygofers are
not so strongly inflated at the middle. There is no trace,
however, of an ovipositor which would be a conspicuous part
of the genitalia in case it were a female and should extend from
the base to the tip of the pygofers.

In the specimen in question a segment is found to lie just
beneath the last ventral segment, and the edge of which
protrudes slightly at the posterior end. This structure I am
not able to homologize with a similar one in the typical geni-
talia of either sex; but it is the only one not accounted for,
and which does not resemble in a very marked way some part
of one of the other specimens.

From this comparison it is seen that there is an entire absence
of male structures anteriorly, replaced apparently by typical
female structures and a condition just the reverse of this for the
posterior portion, so apparently is a good example of the anterior-
posterior type of this phenomenon. This is not commonly
seen and since it occurs in the Homoptera for which group it has
never been reported, perhaps deserves mention here.

A NEW SPECIES OF CICADELLID^ FROM WISCONSIN.

DwiGHT M. DeLong.

Deltocephalus marginatus n. sp.

Closely resembling D. bilineatus, but with longer head and a
black transverse band across margin of vertex between the
eyes. Length 4 mm.

Vertex produced, twice as long at middle as next the eyes,
width between the eyes equalling the length. Antennae two and
one-half times as long as vertex. Pronotum as long as vertex,
produced and very strongly curved on anterior margin. Elytra
quite long and slightly flaring; middle antiapical cell very long,
constricted along central half.

Color: Creamy white with testaceous and dark markings.
Face uniform creamy white, unmarked. Vertex with black,
longitudinal, median, impressed line on middle half; two pale
testaceous bands arise about one-third the distance from
apex and extend across pronotum to middle of apex, where
they end in two black spots
on the disc. Each of these is
bordered interiorly on the vertex
by a fine black line, broken into
spots, which converge at the

D mirjina-tyj.

apex; also partially bordered

exteriorly by a small black

dash. A black transverse band

extends across the margin of

vertex and recurves onto vertex

at either end along the eyes. Pronotum with four longitudinal

testaceous bands. Veins of elytra white; the clavus and claval

veins along suture broadly margined with testaceous. Outer

margins of inner and two outer apical cells and adjoining costal

cell, fuscous. Venter pale yellow, female segment with a large

round black spot either side of notch.

Genitalia : Female last ventral segment longer than preced-
ing, with posterior margin broadly notched, sides of notch
somewhat convexly rounded.

Described from two specimens from Amery, Wisconsin,
collected August 13, 1916. In the Wisconsin list these were
considered as D. bilineatus but with additional study it seems
that they are quite distinct in the points mentioned. It may
prove to be only a variety of that species, but I believe deserves
specific rank.

2 2S

A PRELIMINARY LIST OF THE ANTS OF WISCONSIN.*

A. C. Bi:rrili,,

Entomologist of The University of Idaho, Moscow, Idaho,

and
M. R. Smith,

Of The Truck Crop Insect Investigations, U. S. Department of
Agriculture, Baton Rouge, La.

The ants listed in this paper represent collections made by
Dr. W. M. Wheeler, of Harvard University, Mr. C. E. Brown,
of the University of Wisconsin, the writers and others. To
Mr. T. E. B. Pope, Curator of The Milwaukee City Museum, the
authors are indebted for a list of the species of ants in the
museum, and to the other gentlemen mentioned, the authors are
indebted for the use of their records.

The collector's initials, instead of his full name, are used for
brevity.

Subfamily Camponotinae.

Campouotus lierculaneus L. subsp. pennsylvaniciis De Geer.

A. C. B.; Lake Puckaway, Wis., September 1, 1908.
Campouotus hercidaneus L. subsp. ligniperda Latr. var. nove-
boracensis Fitch.

A. C. B.; Lake Puckaway, Wisconsin, September 1, 1908.
Camponotus castaneiis Latr. subsp. americaniis Mayr.

A. C. B.; Coons Valley, Wisconsin, September 21, 1911.
Camponotus hercidaneus L. var. whymperi Forel.

W. M. W. ; White Fish Bay, near Milwaukee, Wisconsin.
Camponotus Jallax Nyl. var. nearcticus Emery.

W. M. W.; Wisconsin.
Lasius niger L. var. americanus Emery.

A. C. B.; Milwaukee, Wisconsin, August 20, 1908.
Lasius niger L. var. neoniger Emery.

A. C. B.; Janesville, Wisconsin, July 26, 1911.
Lasius umbratus Nyl. subsp. mixtus Nyl. var. aphidicola Walsh.

C. E. B.; Milwaukee, Wisconsin, August 24, 1901.
Lasius claviger Roger.

C. E. B.; Madison, Wisconsin, July 12, 1902.

*Published by permission of the Secretary of the United States Department of
Agriculture.

2 2q

230 The Ohio Journal of Science [Vol. XVIII, No. 6,

Lasius interjectus Mayr.

C. E. B.; Milwaukee, Wisconsin, August 12, 1901.
Lasius latipes Walsh. i

A. C. B.; Lake Puckaway, Wisconsin, September 1, 1908. I

Prenolepis imparis Say.

M. R. S. ; Madison, Wisconsin, October, 1917.
Brachymyrmex heeri Forel subsp. depilis Emery.

A. C. B.; Vernon County, Wisconsin, September 22, 1912.
Formica fusca L. var. subsericea Say.

A. C. B.; Ozuakee, Wisconsin, September 13, 1913.
Formica cinerea Mayr. var. neocinerea Wheeler.

M. R. S.; Madison, Wisconsin, October, 1917.
Formica neogagates neogagates Emery.

C. E. B.; Milwaukee County, Wisconsin, May 13, 1901.
Formica pallide-fuha Latr. subsp. schaiifussi Mayr.

C. E. B.; Milwaukee, Wisconsin.
Formica pallide-fulva Latr. subsp. schaufussi Mayr. var. incerta
Emery.

C. E. B.; Milwaukee, Wisconsin, September 19, 1901.
Formica pallide-fulva Latr. subsp. nitidiventris Emery.

A. C. B.; Corliss, Wisconsin, July 19, 1911.
Formica pallide-fulva Latr. subsp. nitidiventris Emery, var.
fusca ta Emery.

A. C. B.; Corliss, Wisconsin, July 19, 1911.
Formica pregandei Emery.

C. E. B.; Beaver Lake, Wisconsin, June 20, 1901.
Formica rufa L. subsp. obscuripes Forel.

C. E. B.; Milwaukee County, Wisconsin, July, 1902.
Formica rufa L. subsp. aggerans Wheeler.

M. R. S.; Madison, Wisconsin, October, 1917.
Formica rufa L. subsp. aggerans Wheeler, var. melanotica Emery.

M. R. S.; Madison, Wisconsin, October, 1917.
Formica truncicola Nyl. var. integroides Emery.

C. E. B.; Milwaukee, Wisconsin, June 21, 1901.
Formica sanguinea Latr. subsp. rubicunda Emery. |

A. C. B.; Mayville, Wisconsin, July 25, 1909.
Formica sanguinea Latr. subsp. integra Emery.

A. C. B.; Milwaukee, Wisconsin, August 10, 1908.
Formica sanguinea Latr. subsp. subnuda Emery.

A. C. B.; Vernon County, Wisconsin, September 21, 1912.

April, 1918] The Ants of Wisconsin 231

Formica nlkei Emery.

A. C. B.; Cedarsburg, Wisconsin, July 22, 1910.

Formica dakotensis Emery var. specidaris Emery.
H. Muckerman; Prairie du Chien, Wisconsin.

Subfamily Myrmicinae.

Monomoriiim minimum Buckley.

A. C. B.; Juneau County, Wisconsin, July 24, 1908.

Monomoriiim pliaraonis L.

A. C. B.; East Milwaukee, Wisconsin, September 7, 1908.

Aphaenogaster tennesseensis Mayr.

A. C. B.; Lake Puckaway, Wisconsin, August 31, 1908.

Aphaenogaster fiilva Roger, subsp. aquia Buckley.

A. C. B.; Marquette, Wisconsin, September 1, 1909.

Aphaenogaster fulva Roger, subsp. aqiiia Buckley, var. picea
Emery.
A. C. B.; Marquette, .Wisconsin, September, 1909.

Stenamma brevicorne Mayr.

C. E. B.; Milwaukee County, Wisconsin, May 11, 1911.

Myrmica rubra L., subsp. brevinodis Emery.

C. E. B.; Milwaukee County, Wisconsin, June 16, 1900.

Myrmica scabrinodis Nyl.

C. E. B.; Milwaukee, Wisconsin, June 21, 1901.

Myrmica scabrinodis Nyl. var. sabuleti Meinert.

C. E. B.; Milwaukee County, Wisconsin, June 24, 1901.

Myrmica scabrinodis Nyl. subsp. schenkii Emery.

C. E. B.; Milwaukee County, Wisconsin, August 30, 1901.

Myrmica scabrinodis Nyl. subsp. schenkii Emery var. emeryana
Forel.

A. C. B.; Juneau County, Wisconsin, July 24, 1908.
Leptothorax acervorum Mayr. subsp. canadensis Provancher.

C. E. B.; Milwaukee County, Wisconsin, June 20, 1901.
Tetramorium guineense Fabr.

C. E. B.; Milwaukee, Wisconsin, August 21, 1908.

Solenopsis molesta Say.

A. C. B.; Fort Atkinson, Wisconsin, August 1, 1912.

Cremastogaster lineolata Say.

A. C. B.; Madison, Wisconsin, May 18, 1912.

232 The Ohio Journal of Science [Vol. XVIII, No. 6,

Subfamily Dolichoderinae.

Dolichoderus {Ilypoclinea Mayr.) taschenbergi Mayr. var.

A. C. B.; Juneau County, Wisconsin, July 24, 1908.
Tapinoma sessile Say.

A. C. B.; Madison, Wisconsin, May 18, 1912.

Subfamily Ponerinae.

Ponera coarctata Latr. subsp. peiuisyhanicus Buckley.

A. C. B.; Baraboo, Wisconsin, August 2, 1911.
Stigmatomma pallipes Haldeman.

A. C. B.; Osceola, Wisconsin, August 27, 1912.

OHIO ACADEMY OF SCIENCE.

Annual Meeting.

At a meeting of the Executive Committee, held in Columbus,
on December 8th, it was decided to hold the next Annual
Meeting of the Academy in Columbus, May 30 and 31, 1918.
The usual detailed notice will be issued later.

Saturday, June 1, will be available for field trips for such of
the sections as may desire to hold them, the late date being
unusually adapted to this purpose.

A more prominent place on the program will be given to the
exhibition of specimens, microscopic preparations, drawings,
models, scientific apparatus, etc., than at previous meetings. It
is expected that definite hours will be assigned for such dem-
onstrations, when the exhibitors will be present to give any
desired explanations.

Edward L. Rice, Secretary.

Date of Publication, May 6, 1918.

THE

Ohio Journal of Science

PUBLISHED BY THE

Ohio State University Scientific Society

Volume XVIII MAY, 1918 No. 7

TABLE OF CONTENTS

DkLong— Additional Records of Tennessee Cicadellidse (Hemiptera-

Homoptera) 233

Fenton — The Parasites of Leaf-Hoppers— Part II 243

Lyman and Bowers— The Digestibihty of Soy Bean Meal by Man 279

ADDITIONAL RECORDS OF TENNESSEE CICADELLID^.

(Hemiptera-Homoptera.)

DwiGHT M. DeLong.

Two years ago the first attempt was made at listing the
CicadeUid^e (Jassoidea) found in Tennessee. At that time I
published a paper* containing records for some 212 species and
varieties collected in various parts of the state, most of which
were the records of the previous summer.

In view of the fact that the proof was not seen by the author
after the manuscript was sent to press, a large number of errors,
mostly typographical, occurred in this bulletin, which I will
not attempt to correct here. In working over the material,
however, I find that a few species have been incorrectly cited
and I wish to change them at this time.

The species referred to as Spangbergiella vulnerata Uhl.
should be Spangbergiella mexicana Bak. Also Deltocephalus
signatifrofis V. D. should be D. sonorus Ball.

The specimens described as Typhlocyba nigridorsum DeL.
are no doubt extreme variations of E. vulnerata Fh. and with a
good series of specimens, I would now place this form under
vulnerata and very close to variety nigra Gill.

* Tenn. State Board of Entomology, Bull. 17,. 1916.

233

234 The Ohio Journal of Science [Vol. XVIII, No. 7,

In spite of the fact that Mr. Van Duzee has placed
Typhlocyba morgani DeL. as a variety under Empoa querci Fh.,
I am still certain that it is a good species and has no resemblance
to querci. In fact it belongs in the Genus Erythroneura and I
am sure Mr. Van Duzee has not seen specimens of this species.
At the present time I have a good series and find them to be
quite constant in their characters as described.

During the past two seasons additional material has been
collected and the following pages include records for these as
well as specimens unidentified at the time of publication of the
previous paper. These are new records for the state, five of
which are treated as new species and one as a new variety.
The total now brings the list for Tennessee to 241 species and
varieties.

Mr. Crumb has very kindly given to me his records of
species not listed previously and I have mentioned his name
as collector in each case. Specimens not otherwise designated
were collected by the author.

Macropsis occidentalis (V. D.).

Two specimens referred to this species were taken at Clarks-
ville, Tenn., July 5 and 17, 1915.

Macropsis tristis (V. D.).

Four specimens collected July 8 were swept from wild
plum by Mr. Crumb at Clarksville, Tenn.

Draeculacephala angulifera (Walk.).

Collected at Hixon during July, August and September,
1915, Mr. Crumb.

Gypona sp.

The species referred to is a new one in manuscript at present
and described by Gibson. I will not give the name here as I do
not wish to make this a manuscript species. One specimen
was collected at Clarksville, July 15.

Gypona scarlatina Fh.

Six specimens from Clarksville, collected June 21 to July 15,
five of them by Mr. Crumb.

May, 1918]

Tennessee Cicadellidce

235

Gypona spadix n. sp.

In general appearance resembling scarlatina, but shorter and more
robust with reddish-brown color and distinct genitalia. Length, 9 9 mm.
cT S mm.

Vertex decidedly produced, twice as long on middle as next the eye,
width between the eyes slightly more than twice the length. Ocelli
almost equidistant between eyes and middle line of vertex, and closer
to anterior than to posterior margin. Pronotum twice as long as
vertex and twice as wide as long. Elytra rather short and broad.

Fig. 1. Gypona spadix n. sp. ,

a, lateral view of head; b, dorsal view of head and pronotum; c,

female

genitalia; d, male genitalia.

Color: Face, eyes, lora3 and genae black cr dark fuscous, sometimes
narrowly margined with pale fuscous. Vertex, pronotum and scutellum
brownish yellow, lightly flecked with bright red. Ocelli red, a pair of
round spots behind these on the posterior margin of the vertex, brown.
Posterior portion of pronotum and some irregular blotches on anterior
half, darker. Elytra dvill and rather dark chestnut brown; veins
darker, dark bordered. In the female the outer veins are reddish in
color. Cross veins between first and second sectors, these between
apical and antiapical cells and the ends of claval veins along sutural
line, black. Veins of apical cells rather heavily bordered with fuscous.
Beneath yellow, bordered with fuscous, legs dirty yellow to fuscous.

236- The OJiio Journal of Science [Vol. XVIII, No. 7,

Genitalia: Female last ventral segment almost one-half longer
than penultimate. Posterior margin with narrowly rounded and prom-
inent lateral lobes ; broadly emarginate from these to a rather broad
central lobe which is distinctly sinuate at the middle. Ovipositor dark
in color. Male valve more than twice as long as preceding segment,
gradually narrowed either side from base to roundingly produced apex.
Plates longer than valve, half as broad as long. Outer margin undu-
lated, inner margins overlapping toward apex and convexly curved to
outer apical margins.

Described from one female and two males from Clarksville,
Tenn., collected August 7th, and a single female from Ulster
County, N. Y. Kindly loaned by Dr. Herbert Osborn. Type,
male specimen in author's collection.

This is different from any of the species described, so far as
I can determine from descriptions and authentically identified
specimens of most of the nearctic forms.

Aligia modesta (O. & B.).

Four specimens were taken during July, 1917, at Clarksville.

Scaphoideus carinatus Osb.

Three specimens from Clarksville, July 26, 28 and 29.

Scaphoideus cinerosus Osb.

Two specimens, Clarksville, July 13 and 16.

Scaphoideus opalinus Osb.

Five specimens swept from grasses in pasture land at
Clarksville, July 13 and 23.

D Otocephalus alboneura n. sp.

Resembling D. vinnulus Crumb in size and general appearance but
with more conically produced head, distinctive coloring and genitalia.
Length, 2 mm.

Vertex conically produced, twice as long at middle as next the eye,
and slightly longer than width between the eyes. Pronotum longer than
vertex, twice as wide as long, rather strongly produced on anterior
margin. Elytra scarcely longer than abdomen, with reticulations on
the clavus and middle antiapical cell divided.

Color: Face fuscous, marked with pale arcs and fading to dirty
yellow above. Vertex dirty yellow with dark median impressed line on
posterior two-thirds, a pair of spots at the apex, two either side along
margin between these and the eyes, the inner one lying close to the eye,
dark fuscous. A transverse fuscous band interrupted at the middle
crosses the disc of vertex between anterior margin of eyes and termi-
nates in a spot next either eye. Two oblique dashes either side of

May, 1918]

Tennessee CicadelUdce

237

median impressed line extend from this band to posterior marj^in, being
close to and parallel with inner marj^ins of eyes. Pronottnn dark
fuscous, outer margins, anterior median spot and five longitudinal
vittse, pale. Scutellt:m dark, a pale spot either side on outer margin half
way to apex. Elytra dark fuscous, almost black, ner\aires pale, reflexed
costal veins and inner apical cell broadly pale. -Beneath black, keel of
ovii)ositor and spots on pygofers pale. Legs dark set with white spines
which are black at base. '

Genitalia: Female last ventral segment concealed except a round-
ingly produced lobe at cither side of preceding segm.ent and extending
beyond its apex; penviltimate segment scarcely produced beyond pre-
ceding segment at the lateral margins but strongly convexly rounded at
middle to three times its length. The apex slightly notched either side
of middle so as to produce three distinct lobes. , ,

I . / I

,/ t

Fig. 2. Dellocephalus alboneiira n. sp.
a, dorsal view of head and pronotum; b, female genitalia.

Described from a single female specimen collected July 6,
1917, while sweeping short grasses on a rather steep hillside in
an open woodland at the State Experiment Farm, Clarksville,
Tenn. This specimen does not agree with any of the other
described species and apparently is distinct. Type in author's
collection.

Dellocephalus vanduzei G. & B.

One specimen from Clarksville, July 13, 1917.

Deltocephalus sonorus Ball.

Several specimens were taken during July and August at
Clarksville and TuUahoma.

Euscelis striolus (Fall.).

Very abundant on grasses during July and August at
Clarksville.

238 The Ohio Journal of Science [Vol. XVIII, No. 7,

Phlepsius carolinus Lthr.

This species was recently described from a single female
collected in South Carolina. During the past season I have
been able to collect great numbers of both sexes in Tennessee
and am describing the male genitalia here.

Male valve almost as long as last ventral seginent, gradually and
evenly sloping from lateral margins to obtusely angled apex. Plates
almost as broad as valve at base and three times its length, very slightly
concavely narrowed to rather broad, blunt and well rounded apices;
outer margins set with about six large dark spines and very densely
clotlTjCd with fine white hairs at basal lateral margins.

Phlepsius altus O. & B.

Abundant on short vegetation in open fields during August.

Phlepsius slossoni Ball.

Two specimens were collected by Mr. Crumb, October 8
and November 2 at Clarksville.

Dorydiella floridana Bak.

Two specimens from Clarksville, August 2, IV) 15. and
September 3, 1914, were swept from tall grasses by Mr. Crumb.

Thamnotettix morsei Osb.

One specimen from Clarksville, October 11, 191G, by Mr.
Crumb, and one collected at Elkmont, Tenn., September 27,
by Mr. W. B. Cartright.

Thamnotettix proprius n. sp.

Resembling shermani and the other members of this group in general
appearance, but with head narrower and more strongly produced, wing
venation, coloration and genitalia distinct. Length, 4.5 mm.

Vertex angularly produced, as long as width between the eyes, twice
as long at middle as next the eyes, slightly shorter than pronotum which
is almost twice as wide as long. Elytra with the antiapical cells, espe-
cially the iniddle one and the cell just anterior to it, short. Two cross
veins are found between the first and second sectors. Antennae four
and one-half times as long as vertex.

Color: Creamy white to light fuscous. Face with a heavy band
just below margin of vertex, a pair of arcs beneath this, a spot next
cither eye, and one below each antenna, black. Vertex with a black
band composed of three narrowly connected triangular spots either side
of apex, interrupted on the middle line. A large ochraceous blotch
either side of middle ari.ses behind this and extends to pronotum which
is crossed with five longitudinal white vittae. Basal angles of scutellum

May, 1918]

Tenjiessee Cicadellidcs

239

dark. Veins of elytra light, broadly margined with fuscous. Outer
apical cell, a spot at apex of costal cell, one behind first cross vein,
margin of clavus along suture, a spot at middle of outer clavus and
antiapical cells broadly margined with dark fuscous. Venter dirty
yellow, segments black at sides; male valve with a spot at apex and
one either side at base, also a spot at base and outer margins of apical
halves of plates, dark.

Fig. 3. Thamnotettix proprius n. sp.
a, dorsal view of head and pronotum; b, male genitalia; c, elytron.

Genitalia: Male valve almost as long as preceding segment, emar-
ginate either side of roundingly produced apex; plates narrow, more
than twice as long as valve, gradually narrowed to long, acutely pointed
tips and very thickly set with yellow and black hairs. Pygofers broad,
seen at either side of plates.

Described from two male specimens sw^ept from canebrake
August 8 and September 1, 1917. They agree in the
characters mentioned above and are distinct from the other
species of the genus. Type in private collection of author.

240 The Ohio Journal of Science [\o\. XVIII, No. 7,

Chlorotettix sp.

The species referred to here has been in manuscript for
almost a year and has been submitted for publication. I will
not give the name here as I do not wish to make it a manuscript
species.

Alebra albostriella var fulveola (H. S.).

Several specimens were collected during June and July at
Covington, Colliersville and Clarksville, Tenn.

Alebra albostriella var. rubrafrons n. var.

Resembling var. fulveola in form and size but with darker color and
face and vertex bright red. Length, 4 mm.

In stn.ictural characters this form is closely related to albostriella.
The head is not produced and broadly rounded. The comparative
width between the eyes is slightly less and the posterior margin of the
vertex is more emarginate, almost angtdarly excavated in some spec-
imens. The wing venation is also different.

Color: Deep orange yellow, darker than in fulveola. Clypeus, frons,
face and vertex densely and almost uniformly bright, cardinal red;
ocelli and rudiments of the arcs on the front pale. Scutellum with basal
angles light, the remainder often flecked with red and the apical margin
always red. Elytra with apical portion smoky sub-hyaline.

Male pygofers more inflated than in fulveola.

These characters are constant and distinct in the five male
specimens at hand, which were collected at Clarksville, July 11,
1917, while sweeping small shrubs in an open woodland at the
Experiment Farm. The ranking of this form will depend entirely
upon the possibility for variation in the wing venation and how
much emphasis should be placed upon this character, for the
venation here is different from albostriella and its other varieties.

Alebra bicincta n. sp.

Resembling albostriella in form and size but with two brown cross
bands extending across elytra. Length, 4 mm.

Head broadly rounded, not produced, almost parallel margined
with anterior margin of pronotum. Venation of wings slightly different
from albostriella.

Color: Milky white tinged with yellow. Face, disc of pronotum,
basal angles of scutellum, and sutural margins of the wing washed Avith
yellow. Two rather broad transverse bands extending across elytra,
dark brown or fuscous. The anterior of these is just behind the apex of
scutellum and extends obliquely to the costal margin; the posterior one
covers the entire apical portion to a little anterior of the cross veins.
Beneath pale yellow, ovipositor dark fuscous.

May, 1918]

Tennessee CicadellidcE

241

Genitalia: Last ventral sej^ment of female rather strongly and
broadly roimdini^h' produced. Pygofers large, inflated.

Described from two female specimens taken at Clarksville,
Tenn., August 13 and 31. The latter specimen was taken
from sassafras while collecting at night with a flash-light.
Type in private collection of author.

Fig. 4 — a, b, Alebra hicincta n. sp. a, elytron; b, dorsal view of head and

pronotum.
c, d, Alebra eburnea n. sp. c, dorsal view of head and pronotum;

d, elytron,
e, elytron of Alebra alboslriella var. rubrafrons n. var.

Alebra eburnea n. sp.

Resembling alboslriella somewhat in general appearance but smaller
with very narrow head and distinct wing venation. Length, 3.2 mm.

242 The Ohio Journal of Science [Vol. XVIII, No. 7,

Vertex slightly produced, rather broadly rounded. Width between
the eyes one-half greater than length. Pronotum twice as wide as long
and decidedly wider than head. Elytra distinctly longer than abdomen.

Color: Ivory white; face, vertex, pronotum and scutellum, creamy
white, the latter with two somewhat darker lines extending across
middle from pronotum to apex. Elytra milky white, opaque from base
to beyond cross veins, tips smoky. Beneath creamy white, last ventral
segment of female with a dark line one-half the way to base from middle
of apical portion. Ovipositor yellow.

Genitalia: Female last ventral segment strongly rounded and
slightly emarginate at middle.

Described from a single female from Covington, Tenn.,
swept from wild-grape June 18, 1915. By a comparison with
the species of the genus this seems so distinct in size and form
that I describe it here. Type in author's collection.

Alebra fumida Gill.

A single specimen collected July 6 at Clarksville has been
referred to this species. The specimen at hand is shorter and
more robust than typical individuals of fumida which are from
its type locality. It also has a different wing venation, but
because of its marked resemblance in general appearance, I
hesitate to describe it as a different form until more material is
available for study.

Dikraneura cruentata Gill.

Four specimens were collected during July. All of these
lack color markings except the black lines on the cross veins.
The specimens are decidedly smaller and have a different
character of the wing from that pictured by Gillette, but no
doubt belong here.

Erythroneura hartii Gill.

Abundant on apple during July and August.

Erythroneura rubroscuta Gill.

One specimen from Clarksville, June 30, 1915.

Erythroneura comes var. maculata Gill.

Abundant on sycamore during August and September.

THE PARASITES OF LEAF-HOPPERS.
With Special Reference to Anteoninae.

F. A. Fenton.

PART II.

TABLE OF CONTENTS.

Phylogeny 243

Taxonomy 247

Life Histories of Species 248

Systemmatic 258

PHYLOGENY.

The relationship of the Anteonince to the Ilomoptera is shown
in the accompanying phylogenetic tree. It is seen that they
are parasitic on the Membracidce, Fulgoridce, and Cicadellidce —
three rather closely-related families. Up to the present time
there are no records of any dryinid being parasitic on a Cercopid.
If this holds true, it may be said that the spittle-forming habit

FULQORIDAE CERCOPIDAE MEMBRACIDAE CICAPELLIDAE

COCCIDAE APHIDIDAE

of this group may protect the nymphal stages of these insects,
and that the adults are usually too large and active for the more
highly specialized AnteouincB. There is but one record of
dryinid parasitism among the Membrac-idce^that of a poly-
embryonic form Aphelopus thelice Gahan mss. parasitic on
Thelia bimaculata^ (Kornhauser, 19 15-19 16).

In the Cicadellidce, as noted, three of the six sub-families are
at present known to be subject to parasitism by Anteonince, as
indicated in the above phylogenetic tree of the family. Fifteen
genera in the Jassince are attacked, which are chiefly grass-

* According to Professor Kornhauser, the adult parasite was separately
identified by both Mr. Brues and Mr. Gahan as belonging to the Anteonince.

243

244

The Ohio Journal of Science [Vol. XVIII, No. 7,

■ ' • 1 ■. ' ■ ' ( ! '
• I : . ( I ( I • ■. I !

TYPHPLOCYBINAE

JASSINAE

BYTHOSCOPINAE

PAROHNAE

6YP0NINAE

CICADELUNAE

inhabiting forms, and but eight genera in the Bythoscopince,
Cicadellince and TyphlocybincB. It is interesting to note that
the only genus parasitizing the Typhlocyhince is Aphelopiis,
which is the most primitive and generalized genus in this
sub-family. This same genus is the one found parasitizing
Membracid nymphs. Anteonince parasitizing genera in the
other sub-families noted are all higher and more specializd
types.

DELPHACINAE

FLATINAE

ISSINAE

May, 1918]

Parasites of Leaf-IIoppers

245

By far the greatest number of host records today in the
family Fidgoridce are in the sub-families DelphacincD and FlatincB,
the highest specialized groups of this family. Ten genera in
each of these are paratitized, while but one or two genera are
parasitized in the Issince, AcanaloniincB, Tropidiichince, CixiincB,
and Dictyophorince. No species in the others are at present known
to be dryinized. Only the highest specialized genera in the
AnteonifKB parasitize this group, thus suggesting that as a
group they were first parasitic on the Cicadellidce, later becom-
ing adapted for parasitizing the FiilgoridcB. Most of the species
are subject to parasitism in their immature stages, the adults
apparently being free, except in the less active, smaller, and
short-winged forms as Libiirnia.

Apterous species with both arms of chela
lamellate, hosts Fulgoridae, sexual dimorphism
marked. Pseudogonatopus type. , '•

Winged species with both arms of
chela lamellate, hosts Fulgoridfe, sex-_
ual dimorphism marked. Echthrodel-
phax type.

Winged species with one arm of
chela lamellate, hosts Cicadellidae, >
sexual dimorphism, Chelothelius type.

Winged species, chela in some spe-
cies only partly extensile, hosts ^
Cicadellidae, sexual dimorphism but
not extreme. Chelogynus type.

Winged species, no chela, hosts
Cicadellidae and Membracidae, no>
sexual dimorphism. Aphelopus type.

Apterous species with one arm of
chela lamellaite, hosts Cicadellidae,
sexual dimorphism marked. Gonato-
pus type.

Winged or partly apterous species,
lateral arm of chela serrate, median
arm bare, hosts Fulgoridae, sexual
dimorphism. Phorbas type.

Winged species, chela fairly special-
ized and freely extensile, hosts Cica-
'dellidae, sexual dimorphism. Deino-
dryinus type.

Winged species, chela in some
/species not extensile, hosts Cicadel-
^ lidae. Anteon type.

Primitive Dryinid

Ohio Journal of Science.

Vol. XVIII, Plate I.

F. A. Fentoit.

May, IDIS] Parasites of Leaf -Hoppers 247

TAXONOMY.

The relationships of the AnteonincB in the Hymowptera is a
much -debated question, but it is now generally concluded that
they are a sub-family of the BelhylidcB in the super-family
Proctotrupoidea. The larval sac is a structure which at once
distinguishes all A^iteonince, and at the same time shows their
possible affinity with such other forms that present this peculiar
characteristic. Of the three other sub-families that Kieffer
(1914) places with them under the BethylidcE, little is known of
the life history of the EmboIemincB and the Sclerogibbince. How-
ever, with the sub-family BethylincB, the genus Harpagocryptus
Perkins shows its relationship in the similar larval sac found
on the Orthopteron Trigonidium, which resembles that of the
genus Aphelopus. Perkins (1912) says: "It is interesting to
note that the elongate larval sac of Harpagocryptus and Aphel-
opus, so different from those of the other Dryinidce ( Bethylidce) ,
is correlated with the possession of simple front tarsi in the
females." In working out the life cycle of Rhopalosoma poeyi
Cresson, parasitic on the jumping tree-cricket, Orocharis
saltator Uhl., Hood (1913) brought out the interesting fact that
here there is also a larval sac very similar to that of the Anteon-
incE. In commenting on Mr. Hood's paper in the same meeting,
Mr. Rohwer said: "One is confronted with the remarkable
resemblance between the larva of Rhopalosoma and some of the
dryinids. Perhaps the Rhopalosomidce and the {Dryinidce)
Anteonince had a common origin, as the larvae would indicate,
and the adults have specialized along different lines, though
retaining certain characters in common." Ashmead placed this
genus in a new family, which he included in the super-family
Vespoidea; and Rohwer believes that this is the more nearly
correct view. However distinct the adults of Rhopalosoma may
be from the Anteonince, the fact that larval sac is a feature in
common with both, and the development of this structure on
two widely- separated insect groups — the Homoptera and
Orthoptera — indicates at once an adaptation to the jumping
habit of these insects.

The development and specialization of the chelate front
tarsus in the female may be taken as a criterion showing the
evolution of the group. The chelate tarsus is an adaptation of
the parasite to the structure and habit of the host. Along with

248 The Ohio Journal of Science [Vol. XVIII, No. 7,

the specialization of this structure, such other modifications as
the loss of wings and shortening of the antennae in the female —
all leading to sexual dimorphism — can be correlated. Thus,
as is shown in the above phylogenetic tree, we start with
Aphelopus as the most primitive and generalized group in which
there is no sexual dimorphism. We note specialization pro-
ceeding along two different lines : one a loss of wings, probably-
due to the general activity and abundance of the host; the
other a development of the chelate tarsus as an aid to catching
and holding the active prey. With the latter again specializa-
tion has taken place with regard to the kind of host attacked.
Perkins (1905) has observed that Anteonince parasitizing jassids
have the lateral claw of the chela without lamellae, and generally
curved, (Plate I, Fig. 7). Those confining their attacks to the
Fidgoridce having the lateral claw of the chela with lamellae,
generally nearly straight and with a slight notch near the tip,
(Plate I, Fig. 8). So characteristic is this structure of the chela
that the host family can be ascertained at once by examining
the female.

Specialization has taken place primarily and more markedly
in the female, the males of even those species having the most
highly specialized females being very similar to the males of
the most generalized genus. This is due to the greater need for
activity on the part of the female in the hunting and capturing
of the active hosts. In fact, males of some of the more highly
specialized genera are still unknown and these species are
possibly entirely parthenogenetic.

LIFE HISTORIES OF SPECIES.

Gonatopus erythrodes (Perkins).

July 3, 1917, a large number of Deltocephalus inimicus
nymphs bearing dryinid larval sacs were collected in a run-down
meadow near Castalia, Ohio. About thirty per cent of the
nymphs of this species were parasitized. They were placed
individually in shell vials for separate study. This field was
revisited several times on successive days, but after July 15 the
parasites had disappeared, the remaining nymphs having
matured. July 9 the first parasite killed its host, and July 17
the last one of the series of sixteen individuals collected had
entered the ground to pupate.

May, 1918]

Parasites of Leaf-IIoppers

249

Adults issued between Agust 3 and 11, the time spent in the
cocoon averaging about twenty-five days. One female issued
August G and was allowed to oviposit August 17, without
having been fertilized by a male. This female pounced upon
the first nymph and oviposited in it, and within five minutes
repeated the operation upon another. She oviposited in
thirteen nymphs from August 17 to 21 inclusive. Each nymph
after being parasitized, was placed in a separate vial. After
two or three ovipositions, the parasite rested, paying no atten-
tion to other nymphs. The greatest number of eggs laid in a
single day was six, although this number is probably too low
for natural conditions. Nymphs in the third and fourth instars
are parasitized, the fourth instar being the most susceptible.

The egg measures .2 mm. by .12 mm. and is oval, pale yellow
in color, without any sculpturing on the chorion.

Two of the nymphs did not develop parasites, although they
were paralyzed and the ovipositor inserted. Instead, they
matured and were perfectly normal in every way, so far as could
be observed. This indicates that the sting itself will not pro-
duce castration nor stop development, but that the larva itself
must produce some condition that brings about these changes.
Six nearly-developed eggs were found in the ovaries of this
female, thus proving that the eggs are not developed all at one
time but instead are gradually matured, and that the life of the
adult and extent the oviposition period are relatively long.
This female lived seventeen days and died only because of
abnormal conditions. The total life of this individual is summed
up below.

Date

Issued

Date

First
Egg Laid

Date

Last
Egg Laid

Total

Oviposition

Period

Total

Number
of Eggs

Average
per
Day

Died

Total
Life

Aug. 6

Aug. 17

Aug. 21

5 days

12

2+

Aug. 27

17 days

This female issued from a sac located on the abdomen of the
host, and invariably oviposited in this same position on the
nymphs. In this species the greatest number of sacs are found
between the fourth and fifth abdominal segments. They are
found as frequently attached to the right side of the body as to
the left. The place of attachment is shown below.

250

The Ohio Journal of Science [Vol. XVIII, No. 7,

Seprnents. .. ...

1-2

2-3

3-4

4-5

5-6

&-7

Right

Left

0
0

0
0

3
2

3
3

1
2

0
0

Five days after oviposition, a dark discoloration is noticed
on the parasitized nymph near the region where the ovipositor
is inserted. This is due to the presence of the developing par-
asite within the tissues, surrounded by its first exuvium, the
dull gray color of which shows through the opaque yellow body-
wall of the host. Thus it seems likely that the egg hatches
very shortly after being laid, and that the first instar is rel-
atively of short duration. Two days later, a week after ovipo-
sition, a tiny gray vesicle protrudes between the segments of
this nymph. This is the second larval instar enclosed within the
first exuvium. At this phase, the entire outlines of the parasite
can be seen within the body of the jassid, the anterior end
reaching as far anteriorly as the posterior margin of the pre-
ceding segment.

The second molt occurs in from seven to ten days after the
appearance of the sac, and the third three to four days later.
The larva enters the fifth instar and kills the host eighteen or
nineteen days after the sac becomes external. Within a few
hours the host is devoured, and the parasite leaves it. The
nymphs are killed in from twenty-four to twenty-five days
after oviposition.

The larval sac is about 1 mm. in width, oval, compressed
and dull black in color, being composed of the typical three
exuviae, the first being smooth and shining, the second dull and
roughened on the outer half, and the third being distinctly
rugosely sculptured.

Immediately after emerging the white, grub-like larva seeks
the soil, beneath which it spins a compact oval, white cocoon
covered with soil particles. Some of the larvae in the second
generation spin their cocoons above the soil, attached to some
convenient object, a fact not observed in those of the first or
summer generation. The outer cocoon is completed in a day,
but longer time is required for the completion of the inner. The
larvae of the last generation can be seen spinning within the
cocoon for days after it is apparently completed. The length of

May, 1918]

Parasites of Leaf -Hoppers

251

time spent in the cocoon in the summer is from twenty to
thirty-one days, averaging twenty-five and a half days. In
one case an adult was noticed within the cocoon ready to issue
seventeen days after it was spun. This issued two days later.
Of ten cocoons spun during the summer, seven contained larvae
that pupated soon, the rest remaining as larvae over winter.
The life cycle is summarized in the following table :

Maximum

Minimum

Number of
Specimens

Time between oviposition and appearance
of sac

7 days

19 days
31 days

5 days

IS days
20 days

5

Time between appearance of sac and death
of host

4

Time spent in cocoon..

6

Total

57 days

43 days

There are two generations a year, the adults for the first
issuing during June, and those for the second in early August.
Winter is passed in the larval stage in the cocoon. This species
is parthenogenetic.

Gonatopus contortulus Patton.

This species parasitizes Deltocephaliis sayi adults and has
been reared from no other host. The oval, dark grey egg is
always laid between the anterior abdominal segments, generally
being thrust entirely beneath the cuticle but occasionally being
attached externally in the suture between the segments. In the
majority of cases it is laid between the second and third abdom-
inal segments, either on the right or left side, but it may also
be laid between the first and second or third and fourth. The
female used in the experiments was unfertilized, having been
bred from a parasitized hopper taken in the field. She issued
August 17 and died August 30, thus living thirteen days. A
total of 17 eggs was laid in a period of nine days as the table
below indicates:

Date — August

Total

18

19

20

21

22

23

24

25

26

9 days

No. of
Eggs..

2

0*

2

3

0*

4

2

1

3

17 eggs

* No hoppers were given to the female August 19 and 22.

252 The Ohio Journal of Science [Vol. XVIII, No. 7,

The normal longevity and number of eggs a female of this
species can lay under natural conditions is probably greater
than the above records indicate. However it can be seen that
the adult life of the female extends throughout several weeks
and that the oviposition period is prolonged. Females dissected
show but few eggs matured in the ovaries and this fact, together
with the presence of all stages of the parasite in the field at a
given time, seem to prove this. The female generally made no
attempts at oviposition after two or three hoppers had been
parasitized. At least twenty minutes and generally over an
hour elapsed before the second attempt.

An adult Deltocephalus oculatus was placed in the cage to
determine whether the parasite was confined to sayi as its host.
The hopper was seized and partly eaten without any attempt
at oviposition.

The first larval stage is passed within the tissues of the host
and when the egg is laid externally the young larva must pen-
etrate into the host of its own accord. Shortly after the first
molt the larva begins to push its way out between the segments
through which the egg is laid. It molts for the first time and
enters the second instar four days after oviposition. Nine
days later larvae are found to be in the third instar which is
characterized by the two large head lobes. Within twenty-five
days after the egg is laid the larva enters into the fifth stage.
It issues from the sac and kills the host in from 26 to 31 days
after the latter has been parasitized. The mature larva is 4 mm.
in length, white, with no visible hairs.

The larval sac is oval, and shining black in color, the third
exuvium being distinctly rugosely sculptured. The oval, white
cocoon is spun above the ground on the grass blades and is
4}/^ mm. long and 3^ mm. wide. The pupal period extends
from 22 to 24 days, the adult remaining in the cocoon for two
or three days before issuing, through an irregular hole near
one end.

The species parthenogenetic, unfertilized females producing
females. There are two generations a year, the winter being
passed in the larval stage within the cocoon. The following
table summarizes the life history experiments. The pupal stage
was passed through in the green house and is included in the
table because the length of time under artificial conditions is
dentical with that of normal summer conditions.

May, H)1S]

Parasites of Leaf-IIoppers

253

254

The Ohio Journal of Science [V^ol. XVIII, No. 7,

Haplogonatopus americanus Perkins.

Nearly mature nymphs and adults of Liburnia campestris
and L. Lutulenta are found bearing the larval sacs of Haplo-
gonatopus americanus throughout the latter part of July and
early August in the vicinity of Sandusky, Ohio, and in June and
August about Columbus, Ohio. L. campestris is by far the
more heavily parasitized species of the two, possibly due to its
greater abundance. The larval sac of this parasite is always
dorsal in position and attached to the posterior region of the
host's abdomen. The greater number of sacs are found pro-
truding between the 5th and 6th segments, but there is some
variation as the following table shows :

Segments

1-2

2-3

3-4

4-5

5-6

&-7

7-8

Dorsal median

Dorsal right

0
0
0

0
0
0

0

1
0

2
2
3

6

7
7

0
0

1

0
0

Dorsal left . . ...

0

Unlike the other two species studied the females do not
oviposite readily in captivity and so but few data were obtained
in the oviposition experiments. At the time the adults issued
nearly all the Liburnias had matured and there is a possibility
of this species living until a second generation of nymphs is
produced. This is not likely since this species parasitizes adults.

The process of oviposition was observed in a few instances
and is similar to that already described for Gonatopiis erythrodes
(Perkins), except that the tip of the ovipositor is curled upward
and thrust in the dorsal side of the abdomen. Parasitized hop-
pers taken in the field bearing sacs containing larvae in the
second instar are killed by the mature larva within eight to
eleven days.

When the larval sac becomes visible on the outside of the
body of the host, it is dull grey or brown in color but when fully
developed it is brown or dark yellow. It is then 1 mm. in
length and oval in shape. The first exuvium is smooth and
shining and is always darker in color than the other two. These
comprise the greater part of the sac and are dull and roughened.
The spiracles are dark brown and distinctly visible along the
median line of the different exuviae.

After leaving the host the white elongate larva almost imme-
diately starts spinning the cocoon on some convenient object
above the ground. The cocoon is completed in two days and is

May, 1918]

Parasites of Leaf-Hoppers

255

elongate oval and white in color being 3 mm. long and 1 1-3 mm.
wide. Soon after the cocoon is completed, generally within
three days, the larva pupates. The pupa is typical, being 2^/2
mm. in length and .9 mm. wide. In from nine to eleven days
during the summer and in twenty-six days during the fall, the
adult issues through a hole at or near one end of the cocoon.
The entire time spent in the cocoon varies from fourteen to
seventeen days during the summer and from twenty-four to thirty-
five days during the fall. Males are as numerous as females.

There are two generations a year in the latitude of northern
Ohio and three in the vicinity of Columbus, larvae of the third
hibernating within the cocoons.

T
Life Cycle of Ha

\BLE II.

plogonatopus americamu

',

Exp.
No.

-

Date

Cocoon

Spun

Date
Pupa-
ted

Date
Issued

Length
Pupal
Stage

Time
Spent in
Cocoon

Sex
of
Par-
asite

Stage

of
Host

Sex

of
host

Host Species

22a

July 20

Aug. 6

17 days

cf

nymph

?

Libumia nymph

23a

July 18

Aug. 2

15 days

d^

nymph

?

Libumia nymph

23b

July 18

July 25

Aug. 3

10 days

16 days

9

adult

o^

Libumia camp-
estris winged

23d

July 20

July 26

Aug. 3

9 days

14 days

d'

nymph

?

Libumia nymph

24a July 20

Aug. 5

16 days

&

nymph

?

Libumia nymph

24b : July 21 ' Aug. 6

16 days

16 days

9

nymph

?

Libumia nymph

24c

July 21

Aug. 6

16 days

&

nymph

p

Libumia nymph

24e

July 20

Aug. 5

16 days

9

nymph

?

Liburnia nymph

25d

July 22

July 27

Aug. 6

11 days

15 days

d'

nymph

?

Libumia nymph

25e

July 20

Aug. 5

16 days

9

nymph

?

Liburnia nypmh

25f

July 22

July 27 Aug. 6

11 days

15 days

9

nymph

■?

Libutnia nymph

25g

July 20

July 26 Aug. 5

11 days

16 days

9

nymph

?

Libumia nymph

25h

July 22

July 27

Aug. 6

11 days

15 days

cf

nymph

?

Libumia nymph

35b

July 27

Aug. 7

11 days

9

adult

cf

Libumia lutu-
lenta winged

39a Aug. 2

Aug. 26

24 days

9

nymph

9

Libumia nymph

42 1 Aug. 5

1
1

Aug. 11

pupa p

reserved

&

adult

?

Libumia lutu-
lenta

51a

Aug. 26

Sep. 4

Sep. 20

26 days

35 days

nymph

?

Libumia numph

256 The Ohio Journal of Science [Vol. XVIII, No. 7,

Chelogynus osborni n. sp.

This species parasitizes both nymphs and adults of Chloro-
tettix unicolor Fitch, infested individuals of which are found
from late June to middle July in the vicinity of Sandusky, Ohio.
The sac is always located on the thorax either to the right or
left side between the meso- and metathorax in the suture just
below the middle coxa. When it first appears it is very small
and blue green in color, being almost invisible against the green
of the host. After five days it assumes the normal black color
and when fully developed it is nearly circular, laterally strongly
compressed, and dull black in color. The third exuvium is
strongly rugosely sculptured. Nine days after the appearance
of the sac the parasite maggot issues and kills the hopper. It
is relatively large, green, and bears few scattered hairs. The
male parasite is usually much smaller and does not completely
devour the host. The cocoon is spun in the soil and is 3.5-4.5
mm. in length, broadly oval, white, with sand or soil particles
plastered over the outside. There is but one generation a year
in this region, the larvae hibernating and pupating the following
spring.

Phorbas mirabilis (Perkins).

This species attacks both nymphs and adults of Bruco-
morpha oculatus Newm., the latter being more often parasitized.
From one to three sacs may be attached to one individual and
all may mature even though they are of different sizes. The
sacs are always found protruding between various segments on
the hopper's abdomen. In one specimen three sacs were found
attached between the second and third, third and fourth, and
fourth and fifth abdominal segments respectively. The fully
developed sacs vary from one to two mm. in diameter. They
are shining black in color, the outer exuvium being distinctly
rugosely sculptured. The larva is of a pale purple color when
it first matures. The cocoon is spun in the soil and is large
varying from 2 to 4 mm. in length and 1 mm. in width. It is
very tough and composed of white silk, intermixed with soil, so
that it is of a dull brown color. The adult escapes through a
large, ragged hole cut through at one end. There are two
generations a year, the first occurring in July and the second in
September.

May, 1918]

Parasites of Leaf -Hoppers

257

Aphelopus dikraneuri n. sp.

Aphelopus dikraneuri n. sp. parastizes adults of Dikraneura
fieberi (Low), which is found to be quite extensively infested
with this parasite in the vicinity of Columbus in July and
September. The sac is found attached to the anterior region of
the abdomen either to the right of left side, the position of
attachment varying somewhat as the following table shows:

Abdomen.

Segments

1-2

2-3

3-4

4-5

5-6

6-7

Right

Left

1
1

7
8

1
3

0
0

0
0

0
0

The sac is opaque yellow in color, elongate oval, and 1 mm.
in length. Because of its small size, the first exuvium is usually
overlooked, but it is smooth and shining, while the other two
are dull and finely punctate. The second exuvium is often
brown in contrast to the other two. Under reflected light the
empty sac is irridescent.

The mature larva is white, elongate, with numerous hairs.
The cocoon is spun just below the surface of the soil. It is
small, white and oval, generally being coated over with soil.
There are two generations a year.

The above description and life history is true of all species
of Aphelopus parasitizing Dikraneura, Empoasca, or Erythro-
neura. It is particularly similar to that of Aphelopus come si
n. sp. parasitizing Erythroneura comes Say. This species was
found to be very abundant in one grape vineyard in Columbus
during the latter part of October, 1917. As high as 80% of the
grape leaf hoppers were parasitized. Adults of A. comesi issued
in May the following year.

258 The Ohio Journal of Science [Vol. XVIII, No. 7,

SYSTEMMATIC.

The following classification is based upon Kieffer's Mono-
graph of the Bethylidae (1914).

Subfamily Anteonin^.

Dryinidae (.part.), Haliday, Ent. Mag., v. 4, p. 411, 1837.
Dryininae, Haliday, Hym. Syn., p. 3, 1839.
Dryinoidae, Forster, Hym. Stud., v. 2, p. 94, 1856.
Dryinini, C. G. Thomson, Ofv. Ak. Forh.. v. 17, p. 175, 1860.
Anteonidae, Kicffcr, Bull. Soc. Metz, v. 27, p. 108, 1911.
Anteoninae, Kieffer, Das Tierreich, 41 L, 1914.

Female: Body sometimes lengthened or very elongate, sometimes
somewhat compact. The length varies between 1.5-10 mm. Head
viewed dorsally transverse, almost square or rounded. Mouth on the
anterior end of the head. Mandible three or four-dentate, in one
species stated to be two-dentate. Antennae ten- jointed originating close
behind the clypcus, slender, filiform, or distally slightly and clavate.
Eyes very large and prominent (Lestodryinini and Gonatopodini) or
moderately large in the others. Ocelli three, mostly forming a triangle,
in apterous forms often lacking. Wings and tegulae often entirely
lacking, seldom reduced. Fore wings with pterostigma, two or three
closed basal cells (subcostal, median, and submedian), one distal and
generally anteriorly open radial cell. Anal vein often distinct. Traces
of a cubital as well as two discoidal cells, namely, one distal median
cell and one distal submedian cell; in the AphcJopini and in two other
species only costa and radius are developed. Hind wings lobed. Legs
slender, femur in form of a reversed club, tibia only slightly thickened,
spur of fore tibia with a transparent lamella, extending for its length,
abruptly ending before the tip, the spur thus appearing bi-lobed, claws
of the four hind legs and in the male in the fore legs generally with one
broad proximal or divided tooth.

In all females, except in the tribe Aphelopini the fore legs are mod-
ified. The coxa is excessively elongated, often more than half as long as
the femur, the trochanter is a long, often stalked proximally and some-
what curved joint, which is often five times as long as the corresponding
joint on the other legs, femur proximally strongly club-shaped, tibia
thicker and shorter than the others, the tarsus ending in an almost
bare chela, which is generally thrown backward, hdng close to the
tarsus, ventrally or dorsally, and generally reaching to the proximal end
of the third, seldom the fifth, the fourth or the second joints; the fore
legs are therefore called "Raubfusze," "Pedes raptorii," the third tarsal
joint, often also the second, present a proximally, oblique or perpen-
dicular ]3rojccting process, from which long, stout, bristles project, and
lie against the distal end of the chela, while the fourth joint, for the
same purpose, on the whole ventral side appears more or less convex
or flattened; also the third and fourth joints on both sides bear single,
very long, stout bristles, which geiierally provide the chela with a
support. The medial chela arm, which generally lies against the under-

May, 1918] Parasites of Leaf-Hoppers 259

side of the former, is composed of the fifth tarsal joint, which proximally
is more or less elongated, only in some Antconini is the fifth joint aljout
normal, without elongation; on the ventral side, that is, on the side
lying against the lateral chela arm, the median chela ann bears nmnerous
rows of hyaline diversely shaped lamella and bristles. The lateral
chela ami is slender, pointed, mostly saber-shaped, ventrally with or
without rows of lamellae and l)ristles, the medial chela arm usually so
rests that both ends cross; there is presented a strong, lengthened
claw; the other claw in contrast is reduced and wrapped around by
the lobes of the more or less strongly lengthened empodium. If the
fifth tarsal joint is much shorter than the proximal j^rocess, then both
chela anns are movable, whereby the morphological proximal end of
the fifth tarsal joint becomes apparently the distal end; in the contrary
case only the lateral chela joint is movable. Abdomen slightly depressed
from above, seldom latcralh' depressed together, second segment
somewhat bell-shaped, the following gradually shorter and smaller.

Male: Generally the male is much smaller than the female, mostly
only half as long, the eyes are pubescent and almost half spherical, while
in the female they arc bare and oval. The pronotum of the male is not
\'isible from above, the hind angles of the pro thorax reach the tegulae
always in the male, while in the female they often do not, the parapsidal
furrows can be seen very plainly often extending across the mesonotum
in the male, while in the female they may be lacking. In Deinodryinus
the pterostigma is broad in the male and only moderately broad in the
female, in two other genera the veins of the basal cells are obliterated
in the male, while they appear well developed in the female, the legs
of the male are not long and slender as in the female, but short and
quite thick, the fore tarsus lacks the chela.

Key to Tribes of the Anteonina.

FEMALES.

1. Thorax divided by a deep constriction into two nodes, apterous, fore-tarsi

with chela Tribe Gonatopodiiii

Thorax not so divided, winged 2

2. Pterostigma small, lanceolate, female with chela Trihe Lestodryini

Pterostigma broad 3

3. Wings with two basal cells, female with chela Tribe Anteonini

Wings without basal cells, female without chela Tribe Aphelopini

M.\LES.

1 . Pterostigma broad 2

Pterostigma lanceolate 3

2. Wings with two basal cells Anteonini

Wings without basal cells Aphelopini

3. Vertex angulate, head triangular in profile, ocelli generally widely sep-

arated Gonatopodini

Vertex rounded, head not triangulate in profile, ocelli not widely separated,

Lestodryinini

Tribe Lestodryinini.

Dryininae, Kieffer, Gen. Ins. fasc. 54 p. 3, 1907.
Lestodryinini, Kieffer, Das Tierreich, 4.16, 1914.

Female: Head mostly transverse; eyes long, strongly projecting;
mandibles three or four dentate; antenna? .slender. Pronotum mostly

260 The Ohio Journal of Science [Vol. XVIII, No. t,

not attaining the tegulai. Wings present, with two or three closed
basal cells, and a generally distally open radial cell; pterostigma small,
lanceolate or linear-lanceolate; in two or three siDecies the wings are
rudimentary or shortened; fore legs strongly lengthened, coxa and the
generally stalked and curved trochanter long, femur proximally strongly
thickened, tibia distally thickened, first and fourth tarsal joints long,
second, third and fifth short, the fifth always much shorter than its
proximal elongation, forming with it the median chela arm, this always
with rows of lamellae, lateral chela arm often with rows of lamella or
spines. The four hind legs more slender than the fore legs, coxa and
trochanter much shorter, femur and tibia longer, first basally strongly
thickened, the last distally slightly thickened; abdomen weakly
depressed from above.

Male: Head transverse, vertex convex, ocelli situated close together
in a triangle, seldom in an arc, eyes oblong oval, pubescent. Antennae
filiform, pubescent. Prothorax not visible from above, the hind angles
attaining the tegulag on the sides, mesonotum with or without parapsidal
furrows. Wings with two basal cells; radius curved or angulate, the
distal part generally as long or longer than the proximal, somewhat
shorter in Phorbas. Pterostigma linear-lanceolate or small-lanceolate.

Genus Phorbas Ashmead.

Phorbas Ashmead, Bull. U. S. Mus., v, 45, p. 90 (cf), 1893.
Bocchus Ashmead, Bull. U. S. Mus., v. 45, p. 91 (9 ), 1893.
Eukoebeleia Perkins, Rep. Exp. Sta. Hawaii, Ent. v. 1, p. 59, 1905.

Female: Head large, broad, vertex slightly convex, the sides con-
verging behind the eyes; occiput very slightly concave, eyes large, oval;
ocelli close together in a triangle, antennee subfiliform, slightly thickened
toward the tip, scape longer than first flagellas joint. Maxillary palpi
six-jointed, labial palpi three- jointed; mandibles three-dentate. Thorax
normal, pronotum large, narrower than mesothorax. The hind angles
attaining the tegula-, parapsidal furrows distinct, .somewhat converging,
propodeum distinctly truncate. Wings present or rudimentary; if
developed with lanceolate stigma, two basal cells, and an open marginal
cell, the radius long and curved. Fore legs of medium length, fourth
tarsal joint very long, but shorter than first; median chela ann not
widened and provided with lamellae but \^entrally with continuous
raised process; lateral arm ventrally dentate. Abdomen globose,
distinctly petiolate, the petiole slender, cylindrical.

Male: Head broadly transverse, wider than thorax, vertex convex,
eyes oblong oval, pubescent; longer than thorax. Maxillary palpi five-
jointed, labial palpi three-jointed, mandibles three-dentate. Thorax
with sparse short white hairs; mesonotum with two distinct parapdisal
furrows. Wings hyaline, covered with fine white hairs and very shortly
ciliated. Venation pale, pterostigma lanceolate, two basal cells and an
open marginal cell, radial vein pale and curved, distally distinctly longer
than proximal]}'. Legs short. Abdomen subpetiolate, oval.

May, 191S| Parasites of Leaf -IIof)p>crs 201

Note. — It has been suspected that Phorhas Ashmead was
the male of Bacchus Ashmead and that Eukoebeleia Perkins was
synonomous with the latter. The difference mentioned is that
Bacchus has a four-jointed maxillary palpus while that of
Eukoebeleia is five-jointed. Examination of Ashmead's type
specimen of Bacchus showed that it had a five-jointed maxillary
palp, and that the male of Eukoebeleia was synonymous with
Phorbas. In Kieffer's key to Bocchus (1914) it is mentioned as
having bare eyes. There is no note of this is Ashmead's descrip-
tion and those of his type were slightly hairy. Since the females
of Phorbas have not been described it is impossible to include
such species in the following key:

Key to Species.

1. Wings rudimentary 3. mirabilis

Wings developed 2

2. Scape as long as the third antennal joint, pronotum as long as the mesonotum..3
Scape shorter than the third antennal joint, pronotum longer than the

mesonotum 4

3. Prothorax yellow, head and mesonotum tinged with brown. . A. flavicol lis
Prothorax black and most of body black 3. arizonica

4. Third antennal joint almost twice as long as the first, abdomen reddish

brown 2. alriceps

Third antennal joint but slightly longer than the first, abdomen black,

4. schafferl

1. P. flavicollis (Ashmead).

Bull. U. S. Mus., V. 45, p. 91 (9 ). 1893.

Female: Black, except as noted, wings hyaline, with fuscous band
across marginal cell, two-thirds width of wing. Length, 3 mm.

Marquette, Mich. Collected.

2. P. atriceps (Brues).

Bocchus atriceps Brues, Canad. Ent.. v. 36, p. 118 (9), 1904, Kieffer, Das

Tierreich 41 L p. 45, 1914.
Chelogynus atriceps Brues, Bull. Wis. Soc. ser. 2 v. 3, p. 184, 1905.

Female: Reddish brown, head black above antenna. Wings
hyaline with fuscous band as in flavicollis. Length, 5 mm.

Moshola, New York. Collected.

3. P. mirabilis (Perkins). Plate I, Figs. 2 and 4.

Eukoebeleia mirabilis Perkins, Rep. Exp. Sta. Hawaii, Ent. v. 1, p. 59, ( 9 ) 1905.

Female: Dull red to black, in black specimens only posterior region
of vertex dorsal part of prothorax and mesonotum dull red. Antennae
cxcejjt two or three basal joints black; mandibles and legs yellowish-
brown or testaceous, abdominal pedicel pale. Head and thorax densely
minutely punctate, thorax and sides of head distinctly pilose. Pro-
podcum rugose-areolatc dorsally, posteriorly rugose with median area
defined by raised lines and this bordered by a similar narrower area on
each side. Wings rudimentary, pointed, the fore pair reaching beyond

262 The Ohio Journal of Science [Vol. XVIII, No. 7,

posterior margin of thorax. Abdomen smooth, polished, posterior seg-
ments with scattered stout hairs. Length, 2.5-3 mm.

Male: Black, pubescent, tibise and tarsi of fore legs fuscous, tarsi
of middle and hind legs, except fifth joint which is fuscous, testaceous or
pale yellow. Head distinctly more than twice as wide as long, posterior
margin arcuate, occiput concave. Ocelli in a triangle, the anterior one
its diameter distant from the lateral ones which are much nearer to
each other than to the eyes. Eyes oblong, oval, pubescent. Antennae
of medium length, pubescent, scape twice the length of three, this twice
longer than two, very slightly longer than four, four to nine subequal,
nine somewhat smaller than four, ten slightly longer than nine and
pointed; clypeus arcuate, mandibles black except the brown teeth,
maxillary palpi dark brown, long, extending beyond the posterior
margin of the head, with three joints visible beyond the geniculation.
Head and thorax densely punctate. Mesonotum broad, wider than
long, with distinct parapsidal furrows which slightly converge and
extend to the posterior margin of the mesonotum being widely sep-
arated there; scutellum much smaller than mesonotum being nearly
square and only slightly wider than long; metanotum somewhat shorter
than scutellum, posteriorly rounded; propodeum rugose-areolated,
with median area defined by raised lines, wings hyaline, posterior wings
ciliate, clothed with very fine white hairs. Venation pale, subcostal
vein brown, radius curved, not reaching the wing margin, the distal
part shorter than the proximal, pterostigma, lanceolate; brown, often
white medianly. Legs of medium length. Abdomen polished black,
with scattered stout hairs laterally and posteriorly. Length, 1.5-2 mm.

Described from two specimens. Columbus, Sandusky,
Ohio. Bred from Briicomorpha oculatus adults.

4. P. schaeflferi (Bmes).

Chelogynus schcefferi Brues, Bull. Wis. Soc, ser. 2 v. o, p. 101 (9 ), 1907.
Bacchus schcpfferi, Kieffer, Das Tierreich, 41 L. p. 45, 1914.

Female: Black; base of antennae, mandibles and clypeus yellowish;
prothorax, four anterior legs; coxae and basal parts of femora of hind
pair bright ferruginous. Frons and vertex rugulose, prothorax, meso-
notum and scutellum, smooth and polished, with scattered punctures;
scutellum with transverse crenulate impressed line at its base. Pro-
podeum rugulose, the posterior face nearly smooth above. Abdomen
shining black. Body conspicuously whitish pubescent except flagellum
of antennse. Length, 5 mm.

Huachuca Mountains, Arizona. Collected.

P. schaefferi Var. a. (Brues).

Chelogynus schcefferi Brues, Bull. Wis. Soc ser. 2 v. 5, p. 102 (9 ), 1907.

Female: Colored like P. atriccps with entire thorax reddish, but
structurally similar to above.

Brownsville, Texas. Collected.

May, H) 18] Parasites of Leaj -Hoppers 263

6. P. arizonica (Perkins).

Eukwbelia arizonica Perkins, Rep. Exp. Sia. Hawaii, Ent. v. 2, Hull. 4, p. 44
( 9 ), lfl07.

Female: Reseml^les mirabilis. h\x\. \vinj.;s well dcvelo])efl, and raised
lines on j^roiKideum are more distinel. Ix'n<:[lh, '.\ mm.

Nogales, Arizona. Bred from Bruconiorpha.

Tribe Gonatopodini.

Gonatopodinae, Kieffer, Andre. Spec. Hym. Eur., v. 0. p. 499, 1906.
Gonatopodini, Kieffer, Das Tierreieh, 41 L, 1914.

Feinale: Head transverse or almost transverse, much broader than
the thorax, dorsalh' mostly concave, seldom flat or sli«[htly convex,
posteriorly seldom rounded; the sloping surface of the posterior head
concave. Eyes very large, almost occupying the entire side of the head,
projecting outward, bare, posteriorly strongly diverging. Ocelli small
and approximate, often indistinct or lacking. Maxillary palpus two to
six jointed, labial j^alpus two or three jointed. Mandible slightly
convex, three or four toothed. Antennae originating close behind the
ch'peus, slender yet less so than in Lestodryinini, seldom reaching the
hind end of the thorax, third joint the longest, the following gradually
becoming shorter until the penultimate, the tenth somewhat longer
than the preceding. Thorax strongly lengthened and of striking form,
namely, divided by a generally pedunculate construction in a fore and a
hind node, in one species asserted to be divided by two constrictions
into three nodes. The fore node is composed of the pro thorax and
carries ventrally the fore coxae; it is generally divided through a trans-
verse furrow into two parts, of which the anterior is generally the
shorter. The hind node bears ventrally the middle and the hind coxae,
and shows generally anteriorly a transverse suture, which continues
itself laterally obliqueh' towards the posterior part and ends between
the iniddle and the hind coxae, this node is composed of the metathorax,
the hind part of the mesothorax and the propodeum. The fore part of
the mesothorax becomes fonned through the nearly pedunculate con-
striction, which is joined to the fore node, without forming a jjart of
the same. Mesothorax and scutellum not visible, the latter often
recognizable as a trace on the anterior part of the hind node. In Gyn-
ochelys the mesonotum and scutellum are distinct though quite small.
Tegula and wings quite lacking, except in Gynochelys where the tegula
is present and the wings replaced by a barely visible, scale-like process.
Fore legs because of the divergent form of the thorax very widely sep-
arated from the middle legs, coxa and trochanter greatly lengthened,
the latter generalh' proximally more slender than distally, also club-
shaped femur and tibia shorter and thicker than those of the other legs,
the femm- proximally greatly thickened, the tibia distally and less
strongly thickened, meta-tarsus and fourth joint of the tarsus long, the
three other joints short, in Cryprtogonatopus the fourth joint is also
short, median chela arm ventrallv with rows of lamellae; lateral chela

264 The Ohio Journal of Science [Vol. XVIII, No. 7,

arm often with lamellae, sometimes with rows of teeth, sometimes
unarmed; middle and hind legs long and slender, femur in form of a
reversed club, tibia straight and very thin, distally hardly thicker,
claws simple. Abdomen slightly flatly compressed, in circumference
mostly shortly oval, petiole short, second segment longer than third,
this longer than any of the following.

Male: Head transverse, triangular viewed from the side, vertex
angulate, ocelli not situated close together but generally widely sep-
arated. Eyes rounded, pubescent. Occiput deeply concave. Pro-
thorax either not visible from above or barely so, the hind angles
attaining the tegula? on the sides; mesonotum with parapsidal furrows.
Wings with two basal cells, radius curved or somewhat angulate, ptero-
stigma lanceolate. The males differ from those of the Lestodryinini in
the triangular head, angulate vertex, rounded eyes, and the arrangement
of the ocelli.

Genus Haplogonatopus Perkins.
Haplogonatopus Perkins, Rep. Exp. Sta. Hawaii, Ent. v. 1, p. 39, 1905.

Female: Vertex distinctly and fairly deeply concave, mandible four-
dentate; maxillary palpus two-jointed, third antennal joint the longest
twice as long as the following one. Pronotum not divided by a trans-
verse impression into two parts. Fore trochanter same length as coxa,
fore femur twice as long as trochanter basally much swollen; chela
reaching to the base of the third tarsal joint; median arm with two
rows of lamcllffi which are large, pointed and set far apart, and with a
row of long spines, distally distinctly curved with a cluster of lamellae
at distal end; lateral arm nearly straight but distinctly curved and
notched at its distal end, medianly with a row of lamellae. First and
fourth fore tarsal joints the longest, the first being somewhat the longer,
the second and third much shorter, the second being the shortest.

Male: Head transverse, not wider than thorax, vertex angulate,
occiput deeply concave; eyes large, rounded, finely pubescent; ocelli
in triangle, the fore ocellus situated below vertex in a concavity twice its
diameter in front of the lateral ocelli. These situated nearer to the
eyes than to each other and directly on the hind margin of the head.
Antennas filiform, pubescent, scape one-third longer than two, three to
six joints the longest, two less than half as long as three and somewhat
swollen, seven to ten subequal, seven slightly shorter than three. Max-
illary palpi not visible ventrally. Mandibles three dentate. Thorax
pubescent, prothorax barely visible from above the hind angles attaining
the tegulse, mesonotum with two distinct posteriorly converging parap-
sidal furrows which are approximate on the posterior margin. Wings
hyaline, finely haired and ciliated; venation distinct though pale, two
basal cells and a nearly closed marginal cell, pterostigma narrow, radius
distinct, long and curved almost extending to the wing margin. Legs
long, especially the hind legs. Abdomen subpetiolate, oval. Five
species of which one is North American.

May, 1918] Parasites of Leaf-Hoppers 265

H. americanus Perkins.

Gonaiopus bicolor Ashmead, Swezey, The Ohio Naturalist, v. Ill, p. 447-8

(9), 1903.
Labeo lon^itdrsiis Ashmead, Swezey, The Ohio Naturalist, v. Ill, p. 447-8

(c^), 1903.
Haplogonatopus americanus Perkins, Rep. Exp. Sta. Hawaii, Ent. v. I, )). 40

(9), 1905.

Female: Fcnaiginous to testaceous in color, abdomen black to
brownish. Antenna black, apical joint and two or three basal ones
pale. Head nearly smooth, pronotum minutely punctured, propodeum
dull with fine rugulosity. Length, 2.5 mm.

Male: Black, mandibles white, teeth brown, maxillary palpi white.
Scutellum smooth and shining, metathorax posteriorly rounded, smooth
and shining at top, but with lateral diagonal impressions separating
punctate areas. Length, 2 mm.

Columbus, Sandusky, Ohio. Bred from Liburnia campestris
and L. lutidenta nymphs and adults.

Genus Gonatopus Ljungh.

Gonatopus, Ljungh, Beitr. Naturk, v. 2, p. 161, 1810, Kieffer, Andre Spec. Hym.

Eur., V. 9, p. 487 nota., 1906, Perkins, Rep. Exp. vSta. Hawaii, Ent. Bull. 11,

p. 13, (non Perkins, 1905), 1912.
Dryninus, Dalman, Analecter ent., p. 14, 1823.
Neogonatopus Perkins, Rep. Exp. Sta. Hawaii, Ent. v. 1, p. 42, 1905.

Female: Head dorsally more or less slightly concave, maxillary
palpus four-jointed, labial palpus two-jointed. Mesonotum and scu-
tellum not visible. Fore tarsal joint, one and four long, the three others
short; median chela arm distally bent inwards, ventrally with rows
of lamellaj; lateral chela arm without lamellae or teeth, at the most
with a tooth before the distal end, often ventrally with rows of spines.

Male: Differs from Haplogonatopus chiefly in configuration of the
head as follows: Concavity on the frons much smaller and vertex less
angulate. There may be exceptions to this, however, as for instance, the
male of Gonatopus erythrodes (Perk.). The maxillary palpus is four-
jointed in comparison to that of Haplogonatopus which is two-jointed.

Key to Species.

1. The entire thorax smooth and shining 2

Thorax at least in part dull and transversely rugulose or punctate 3

2. Hind thoracic node distinctly hairy, head brown 16. brimnescens

Hind thoracic node microscopically finely hairy, head yellow.. 17. pallidiceps
Thorax not hairy 7. decepiens . . . .

3. The whole thorax finely and thickly transversely rugulose 4

Thorax at the most only in the smallest part transversely rugulose 5

4. Thoracic spiracles situated on distinct elevations 8. conlorlulus

Thoracic spiracles not situated on elevations 9. punctalus

5. Hind thoracic node with distinct erect hairs 6

Hind thoracic node without hairs 12

6. Thorax entirely ferruginous 7

At least the hind node of the thorax dark 8

7. Lateral chela arm with a tooth before the distal end 11. mimiis

Lateral chela arm without tooth before distal end 15. mimoides

266 The Ohio Journal of Science [Vol. XVIII, No. 7,

8. Thorax black or dark brown 9

Thorax red, metathorax black 14. erythrodes

9. Head ferruginous 10

Head black 11

10. Pronotum with transverse impression 2. affinis

Pronotum without transverse impression 1. peculiaris

11. Head deeply concave 12. obscurissimus

Head slightly concave 1.3. inimicis

12. Head and thorax yellow, mandible 4-dentate 4. bicolor

Head in greatest part and thorax black or dark brown 13

13. Head finely hairy, mandible 3-dentate » o. californicus

Head not hairy 14

14. Abdomen flecked with reddish brown 6. flavifrons

Abdomen black 15

15. Head finely rugulose, behind black 3. cyphonotus

Head very finely punctate, behind yellow or red 10. ombrodes

1. G. peculiaris Brues.

Trans. Amer. Ent. Soc. v. 29, p. 125 (9 ), 1903.

Female: Polished black, except greater part of head, sutures of
legs and base of antennae, which are reddish or yellow. Pronotum
smooth, shining, lacking the transverse impression. Length, 2.75-3 mm.

Austin, Texas. Collected.

2. G. aflSnis n. sp.

Female: Differs from above chiefly in possessing the emargination
on the pronotum. Dark brown to black, scape of antennas white,
mandibles and frons below antennas testaceous, thoracic constriction,
central area of hind node just before spiracles, and extreme posterior
portion of propodeum yellow to testaceous. Legs ferruginous. Pro-
notum shining with minute punctures, thoracic constriction reticulately
sculptured, hind thoracic node minutely reticulately rugose anteriorly
and posteriorly, but smooth in middle. Abdomen polished, both abdo-
men and thorax with short scattered hairs. Length, 3.5 mm.

Male: Black, pubescent, vertex angulate, antennae?* head triangular
w^hen viewed from side : vertex minutely and densely punctate ; occiput
deeply emarginate and excavated. Three ocelli visible from above, the
anterior being below the vertex and twice its diameter distant from the
lateral ones which are nearer to the eyes than to each other. Eyes
nearly spherical, pubescent. Antennae of medium length, pubescent.
Mandibles white, teeth brown; maxillary palpi white, two joints
visible. Prothorax scarcely visible from above attaining the tegulas on
the sides; mesonotum broad, with white pubescence very finely retic-
ulately punctate; parapsidal ftirrows distinct, converging and meeting
posterior margin of mesothorax but a small distance apart; scutellum
much smaller than mesonotum, smooth and shining; propodeum
minutely reticulately punctate. Wings hyaline, ciliated and clothed
with fine hairs. Venation pale, pterostigma short and lanceolate. Two
basal cells present. Radius pale, curved, the distal part much longer
than the proximal, running almost to the margin of the wing, forming a
nearly closed cell. Legs long, alternately marked with yellow and

*

Specimen imperfectly emerged.

May, 1918] Parasites of Leaf-IIoppers 267

brown. Coxae and most of middle and hind femora brown, front femora
lij^ht at ends and vcntrally but brown otherwise, middle and hind tibiae
brown but yellow at distal and proximal ends, fore tibia) yellow with
brown band across middle, tarsi yellow, fifth tarsal joint brown. Abdo-
men fuscous, depressed dorso-ventrally, with fine whitish pubescence.
Length, 2 mm.

Described from one specimen, Bay View, Ohio, bred from
Deltocephalus ajjinis. Type deposited in Entomological Museum
at Ohio State University.

3. G. cyphonotus Bradlev.

Canad. Ent. v. 38, p. 380 (9 ), 1906.

Female: Black, except scape, pedicel, face, mandibles, trochanters,
all coxa) beneath, posterior and middle coxae in middle, and anterior
tibiae and tarsi above, lemon yellow, rest of legs reddish yellow. Thorax
above and abdomen smooth and polished, side of thorax thoracic con-
striction above, head and coxae finely roughened; hump of thorax
without transverse impression. Differs from G. contortulus in that
entire thorax lacks fine transverse striation and that thoracic construc-
tion less marked. Length, 2.5 mm.

British Columbia. Collected.

4. G. bicolor Ashmead.

C. ashmeadi Kieffer, Andre. Spec. Hym. Eur. v. 9, p. 108, 1904.
G. bicolor Ashmead, Bull. U. S. Mus. v. 45, p. 85 (9), 1893, Kieffer, Das
Tierreich, 41 L. p. 73, 1914.

Female: Yellow or reddish yellow; 4-5 basal antennal joints yellow,
rest brown. Middle and posterior knees and tips of posterior tibiae
black; abdomen piceous black. Mandibles four-dentate. Propodeum
smooth, polished, posterior face slightly transversely acictdated. Length,
3 mm.

Silma, Alabama; Texas. Collected.

5. G. californicus Ashmead.

Bull. U. vS. Mus. V. 45, p. 85, (9 ), 1893.

Female: Piceous brown, except apex of metathorax and the abdo-
men, which are more or less black. Antennae, tarsi slender part of
posterior femora, and middle posterior tibiae honey-yellow. Head
finely and closely punctate, pubescent, metathorax transversely rugulose.
Length, 3 mm.

California. Collected.

6. G. flavifrons Ashmead.

Bull. U. S. Mus. V. 45, p. 84, (9), 1893.

Female: Black, shining, with fine shagreened punctation. Occiput,
face, mandibles, palpi, antennae except three terminal joints which are
fuscous, and legs, yellow. Anterior femora above almost entirely black.

268 The Ohio Journal of Science [Vol. XVIII, No. 7,

tibia with black streak above, middle and posterior coxae and femora
basally more or less black above. Comes nearest to G. contortulus but
is larger, head broader, and differs in color. Length, 4.4 mm.

New York and Hull, Canada. Collected.

7. G. decipiens Provancher.

Addit. Hym. Quebec, p. 179 (9 ), 1897.

Female: Black, antennae testaceous; tibia testaceous, fore tarsus
brown, hind tarsi testaceous. Head large, flat, finely punctate, thorax
smooth and shining. Length, 2.4 mm.

Kap Rouge, Canada. Collected.

8. G. contortulus Patton.

Canad. Ent., v. 11, p. 65, (9), 1S97.

Female: Black; frons black to testaceous; scape and first flagella
joint fuscous to pale testaceous; pronotum with central black area
bordered by testaceous color laterally and on posterior margin. Anterior
coxae testaceous, brown at sides, anterior trochanter pale testaceous,
anterior femora, tibiae and basal two tarsal joints dark fuscous or dark
testaceous, rest of tarsus and chela testaceous. Middle and hind legs
fuscous to dark testaceous. Abdomen black or fuscous polished and
sparsley haired. Prothorax faintly reticulately sculptured, mesonotum
longitudinally rugose, propodeum finely reticulately sculptured ante-
riorly, posteriorly distinctly transversely rugose. Length, 3 mm.

Redescribed from four specimens. Sandusky, Ohio. Bred
from Deltocephalus sayi adults.

9. G. punctatus n. sp.

Female: Similar to G. contortulus except as follows: pronotum dis-
tinctly and finely punctate; propodeum entirely and distinctly punctate
and propodeum spiracles sessile. Thorax and propodeum without hairs.
A distinct lateral, oblique suture separates the meso-and meta-thorax.
Length, 3.5 mm.

Male: Black, pubescent; vertex angulate, head being triangular
when viewed from side; occiput deeply hollow^ed out. Ocelli in triangle,
all visible from above, the anterior not situated in a concavity. A very
slight concavity on frons below anterior ocellus. Lateral ocelli nearer
to the eyes than to each other. Eyes spherical, pubescent; antennae
nearly as long as the body and pubescent; two slightly shorter and
thinner than scape and swollen at middle, three to six and ten, five times
as long as wide, seven to nine successively becoming shorter, nine four
times as long as wide. Mandibles, except the brown teeth, and max-
illary palpi sordid ^^ellow; latter short wdth two visible joints. Pro-
thorax not visible from above, attaining the tegulae on the sides, meso-
notum finely sculptured. Parapsidal furrows converging for three-
fourths their distance towards the base then becoming parallel and end-
ing distinctly separated at the base; scutellum smooth and shining,
being slightly wider than long. Metanotum somewhat wrinkled and
somewhat shining, shorter than scutellum, posteriorly rounded;

May, 1918] Parasites of Lcaf-IIoppers 269

propodcuni roughened and denseh- and coarsely punelale, no area
separated by raised lines. Wings hyaline, ciliated, clothed with short
hairs. Venation distinct, pterostigma lanceolate. Radius curved prox-
imally distinctly longer than distally but not reaching ti]j, sub-discoidal
vein tincly distinct and extending clear to margin of wing, cubital vein
visible short distance from wing margin. Legs long, fore pair testaceous,
last two pairs fuscous. Abdomen same length as thorax, black, ])ilose,
de])ressed dorso-ventrally. Length, 2.4 mm.

Described from one male and three females. Columbus,
Ohio. Bred from Deltocephahis sayi adults. Types deposited
in Entomological Museum, Ohio State University. Paratypes
in writer's collection.

10. G. ombrodes (Perkins).

Neogonalopus ombrodes Perkins, Rep. Exp. Sta. Hawaii, Ent. v. 1, p. 42 (9),

1905.
Gonatopus ombrodes Perkins, Rep. Exp. Sta. Hawaii, Ent. Bull. II, p. 13, 1912.

Female: Black; vertex of head, and pronotum often piceous or
brown, face and occiput yellow or ferruginous; basal two or three
antennal joints pale; legs pale yellow brown or testaceous. Head and
pronotum minutely punctate, propodeum dull and very densely sculp-
tured. Abdomen smooth and glabrous. Length o-3.4 mm.

Columbus, Ohio. Bred from Deltocephalus sp.?

11. G. mimus (Perkins).

Neogonalopus mimus Perkins, Rep. Exp. Sta. Hawaii, Ent. v. 2, Bull. 4, p. 33

(9), 1907.
Gonatopus mimus Perkins, Rep. Exp. Sta. Hawaii, Ent. Bull. 11, p. 13, 1912.

Female: Ferruginous or testaceous, abdominal petiole blacky
abdomen often darker than the thorax or brown; first two or three
antennal joints testaceous, rest black. Head in front of ocelli smooth,
behind dull with minute sculpture, antennas short and stout. Chela
with fifth joint having lamella? not reaching to articular cavity but
replaced towards tip by bristles. Length, 2.5-3 mm.

Nogales, Arizona. Bred from jassid sp.

12. G. obscurissimus (Perkins).

Neogonalopus obscurissimus Perkins, Rep. Exp. Sta. Hawaii, Ent. v. 1, p. 43

(9). 1905.
Gonatopus obscurissimus Perkins, Rep. Exp. Sta. Hawaii, Ent. Bull. 11, p.

13, 1912.

Female: Black, pronotum, mesonotum, and some of leg joints dark
brown or piceous. Basal 2-3 antennal joints, tarsi and part of hind
tibiae and femora pale, yellow, or testaceous. Head and pronotum
shining, latter faintly sculptured; propodeum with minute dense surface
sculpture, posteriorly finely transversely rugose. Length, 3 mm.

Columbus, Ohio. Bred from Deltocephalus spp.

270 The Ohio Journal of Science [Vol. XVIII, No. 7,

13. G. inimicis n. sp.

Female: Nearest to G. obscurissimus but differing as follows: Head
slightly concave, general body color fuscous, head smooth, without hairs
inesothorax ferruginous, fuscous anteriorly, fineh' reticulately rugose,
propodeum fuscous, polished, smooth posteriorly with fine indistinct
punctation. Thoracic spiracles on elevations. Length, 2.75 mm.

Described from two specimens. Bay View and Columbus,
Ohio. Bred from Deltocephahis inimicus adults. Type deposited
in Entomological Museum, Ohio State University.

14. G. erythrodes (Perkins). Plate I, Figs. 1 and 5.

Neogonatopus erythrodes Perkins, Rep. Exp. Sta. Hawaii, Ent. v. 1, p. 43

(9), 1905.
Gonatopus erythrodes Perkins, Rep. Exp. Sta. Hawaii, Ent. Bull. 11, p. 13, 1912.

Female: Ferruginous; propodeum black, abdomen testaceous to
brown; antennae black, three basal joints pale; legs yellow-brown to
testaceous. Head and pronotum shining, latter with fine indefinable
punctation; propodeum dull, similar to G. onibrodes but posterior face
distinctly transversely rugose. Length, 3 mm.

Male: Black, pubescent, vertex angulate, head being triangular
when viewed from side; occiput deeply emarginate and excavated.
Anterior oceUus situated in a concavity on the frons and not visible
from above. Lateral ocelli nearer eyes than to each other. Eyes oval
pubescent; mandibles testaceous, teeth brown. Maxillary palpi sordid
white with two joints visible. Antennas pubescent, of medium length
being about same length as thorax, two shorter than one, three distinctly
longer than one being three times as long as wide, four-nine subequal
shorter than three, ten pointed and narrow, slightly longer than nine.
Head and prothorax densely punctate, latter not visible from above,
attaitiing the tegulae on the sides; mesonotum broad, shining, minutely
sculptured, parapsidal furrows converging being but narrowly separated
at the base; scutellum nearly square, polished, with transverse furrow
near the base; metanotum polished and half-length of scutellum;
propodeum rugose. Wings hyaline, ciliated, and clothed with fine hairs.
Venation pale, pterostigma elongate lanceolate. Two basal cells present.
Radius curved, the distal part longer than the proximal, nearly reaching
margin of wing. Legs long, fore legs testaceous, tip of femora and bases
of tibiae lighter color, last two pairs of legs fuscous; abdomen as long as
thorax, black, pilose dorso-vcntrally depressed. Length, 1.5 mm.

Described from one specimen. Sandusky, Ohio. Bred
from DeUocephalus inimicus nymphs. Type of male deposited
in Entomological Museum, Ohio State University.

15. G. mimoides (Perkins).

Neogonatopus mimoides Perkins, Rep. Exp. Sta. Hawaii, Ent. v. 2, Bull. 4,

p. 34, (9), 1907.
Gonatopus mimoides Perkins, Rep. Exp. Sta. Hawaii, Bull. 11, p. 13, 1912.

Female: Ferruginous or testaceous, petiole black, mandibles pale
yellow. Antennae black, three basal joints pale. Pronotum very

May, 1918] Parasites of Leaf-JIoppcrs 271

minutely punctate with sparse short hairs, thoracic constriction long;
propodciim closely and minutely punctured, i)osteriorly transversely
rugulosc. Abdomen sparsely pilose. Length. :> mm.

Nogales, Arizona. Collected.

16. G. brunnescens (Perkins).

Neogonatopus brunnescens Perkins, Rep. Exp. Sia. Hawaii. Ent. v. 1, p. 44,

(9), 1912.
Gonatopus brunnescens Perkins, Rep. Exp. Sta. Hawaii, Ent. Bull. 11, p. 13,

1012.
Female: Brown or yellow-brown, abdomen dark Ijrown to black,
or sordid testaceous; propodeum posteriorly dark brown or pitchy.
Basal two or three antennal joints and all legs pale colored. Head very
little concave on \'ertcx. Head and propodeum smooth, abdomen
pilose. Length, 2.r)-o mm.

Columbus, Ohio. Bred from Eiiscelis {Athysanus) curtisii.

17. G. pallidiceps (Perkins).

Xeogonatopus pallidiceps Perkins, Rep. Exp. Sta. Hawaii. Ent. v. 1, p. 45,

■(9). 190.5.
Gonatopus pallidiceps Perkins, Rep. Exp. Sta. Hawaii, Ent. Bull. 11, p. 13, 1912.

Female: Black; head, apical margin of pronotum, and the neck in
front of it, three basal antennal joints, and all legs pale, yellow, or
ferruginous. Head smooth, pronotum very finely punctured, pro-
podeum with minute surface sculpture, posteriorly very finely trans-
versely rugose. Abdomen deep black. Length, 2.5 mm.

Alameda, California. Collected.

Tribe Anteonini.

Anteoninae, Kieffer, Andre, vSpec. Hym. Eur., v. 9, p. .510, 1906.
Anteonini, Kieffer, Das Tierreich, 41 L, 1914.

Thorax not divided into two nodes by a constriction. Wings almost

always developed, only atrophied in Myslrophorns. Pronotum attaining

the t'egulae. Pterostigma broad, half ellipsoidal in shape, only small and

elongate in Deinodryinus; venation composed of costa, subscota, a

radius not reaching the fore wing margin, medial, submedial, a basal

vein originating at the distal end of the subcosta and a transversal vein,

other veins obliterated, only \4sible as transparent lines. Fore-legs not

strongly elongated and not excessively slender, coxa and trochanter

short, fere tarsus with a chela in the female, lateral chela arm always

without lamella:.

Genus Chelogynus Haliday.

Gonatopus (part), Dalman, Svenska Ak. Handle, p. SI, 1818.
Dryinus (part), Dalman, Analecta Ent., p. 9, 1823.
Chelogynus Haliday, Ent. Mag., v. 5, p. 518, 18.38.
Antaeon (part), T. A. Marshall, Cat. Brit. Hym., Oxyura, p. 7. 1873.
Anteon (part). Kieflfer. in Andre, Spee. Hym. Eur., v. 9, p. 130, 1905.
Neochelogynus and Prosanteon (part), Perkins, Rep. Exp. Sta. Hawaii, Ent. v. 1,
p. fi0. f)(i, 1905.

272 The Ohio Journal of Science [Vol. XVIII, No. 7,

Distinguished from Anteon by the fourth joint of the front tarsus,
which in the female is longer than the metatarsus, by the median chela
arm, which is provided ventrally with rows of lamellse and whose free
distal end is as long or longer than the basal. As in .-1 nteon the parapsidal
furrows are at least lacking behind, the eye is bare, the radius mostly
angled, the proximal part much longer than the distal.

Key to Chelogynus.

1. Wings hyaline 2

Wings with one or two dark transversal bands 6

2. Antennaj brown 3

Antennae partly red or testaceous, at least the distal end black 4

3. Pronotum longer than mesonotum and coarsely punctate 4. osborni

Pronotum as long as the mesonotum and very finely punctate. . .3. canadeniss

4. Only the end joint of the antennae black, the others testaceous 1. liisus

Not only the end antennal joint black 5

5. Coxas black for the greatest part, pronotum with rugulosities . . . .9. funestus
Coxae pale 2. melanacrias

6. Wings with one dark band, head and thorax red 5. atriventris

Wings with two dark bands 7

7. Entirely ferruginous 6. ferrugineus

Black 8

8. Frons with three longitudinal ridges, sides of face reddish brown. .7. grandis
Frons without longitudinal ridge, face black 8. henshawi

1. C. lusus Perkins.

Rep. Exp. vSta. Hawaii, Ent. v. 2, Bull. 4, p. 50, (9 ), 1907.

Female: Black, head and pronotum densely minutely shagreened,
mesonotum with fine surface sculpture, scutellum polished, propodeum
rugose, posteriorly dull, with very dense and fine rugulosity. Mandibles
white, antennae except apical joint concolorous and testaceous.
Length, 2 mm.

Tucson, Arizona. Bred from jassid nymph.

2. C. melanacrias Perkins.

Rep. Exp. Sta. Hawaii, Ent. v. 2, Bull. 4, p. 49, (9 ), 1907.

Female: Differs only from C. lusus in color and sculpture as follows:
antennae testaceous on basal joints, four apical joints black. Pronotum
somewhat transversely rugulose anteriorly. Length 2-2.5 mm.

Nogales, Arizona. Collected.

3. C. canadensis Ashmead.

Bull. U. S. Mus., V. 45, p. 93, (9 j, 1893.

Female: Black; head finely punctate, pronotum as long as meso-
notum very finely punctate, mesonotum and scutellum smooth, i)ro-
podeum rugose. Length, 2.5 mm.

Ottawa, Canada. Collected.

4. C. osborni n. sp. Plate I, Fig. 6.

Female: Black, antennas brow^n, scape testaceous, flagellum brown
except third flagellar joint which is dark brown, legs testaceous except
middle and hind coxae and trochanters which are fuscous, and distal end
of hind femora which is darkened. Mandibles testaceous except teeth

May, 11/18] Parasites of Lcaf-IIoppcrs 273

which are brown, maxillary palpi pale testaceous with three joints
visible. Ocelli in trianj^le, the anterior situated medianly on the vertex,
the lateral ones closer together than to eyes and three times their
diameter distant from the anterior. Eyes oval and Ixire. Antennae
somewhat elbowed, the scape long, the flagellum slightly clavate, three
two-thirds as long as one, more slender and twice as long as two, four
shorter than three but longer than five, six to nine subequal twice as
long as wide, ten slightly longer than nine and pointed. Vertex with few
scattered very fine punctures, pronotum with few large punctures,
longer than the smooth mesonotum. Parapsidal furrows distinct,
extending half-way length of mesonotum, con\'erging. Scutellum
smooth, over half as long as mesonotimi, with deep transverse fvirrow
across anterior margin and a row of punctures across base. Metanotum
half length of scutellum, shining with deep furrow across anterior
margin. Propodeum distinctly reticulately rugose with middle rugulose
area marked b>' raised lines. Median chela ami with two rows of
lamella} distally which converge into a single row medianly, tip spoon -
shaped with cluster of lamellae. Wings ciliated, clothed with fine hairs,
with two basal cells, radius straight, the distal part very short and
angled. Abdomen globose, shortly pedunculate, polished, black.
Entire body pubescent, face thickly so, vertex and dorsal part of thorax
with few scattered hairs, propodeum with numerous hairs, abdomen
with few indistinct hairs. Differs from C. canadensis chiefly in character
of antennae, and body sculpture. Length, 2.5 mm.

Described from, two specimens. Sandusky, Ohio. Bred
from Chlorotettix unicolor nymphs. Type deposited in Ento-
mological Museum, Ohio State University. Paratypes in
writer's collection.

5. C. atriventris (Cresson).

Dryinus atriventris Cresson, Trans. Amer. Ent. Soc, v. 4, p. 193, (9 ), 1872.

Female: Ferruginous, with pale pubescence. Face, mandibles,
and base of scape pale yellowish; tips of antennae black, propodeum
rugose, posterior face depressed and transversely aciculated. Length,
4.5 mm.

Texas. Collected.

6. C. (?) ferrugineus Brues.

Bull. Wis. Soc, V. 3, p. 183, (9 ), 1905.

Female: Ferruginous, except darker tips of antenna? and tarsi,
scape w^hite at base below^ mandibles yellow. Head rugose-punctate,
pro thorax finely punctured shining covered with short pubescence;
mesonotum shorter, polished, with few punctures and distinct parapsidal
furrows; propodeum finely reticulated; abdomen polished and
unpunctured. Length, 5 mm.

Texas. Collected.

274 The Ohio Journal of Science [Vol. XVIII, No. 7,

7. C. (?) grandis Brues.

Bull. Wis. Soc, V. 3, p. 184, (9), 1905.

Female: Black, legs black except four anterior tibiae and tarsi, base
of anterior trochanters, and tips of anterior femora. Sides of face below
and antenna except five apical joints, rufous, mandibles black, max-
illary palpi fuscous. Occiput shining, finely punctured, and sparsely
pubescent. Pro thorax strongly contracted; pronotum closely punc-
tured; mesonotum polished delicately punctate; scutellum shining;
propodeum finely rugulose. Length, 7 mm.

Riverside, Massachusetts.

8. C. (?) henshawi Ashmead.

Bull. U. S. Mus., V. 45, p. 93, (9), 1893.

Female: Black; antennae and legs pale rufous; the six terminal
antennal joints fuscous; clypeus and mandibles rufous. Head and
pro thorax finely rugose ; mesonotum and scutellum smooth ; propodeum
coarsely rugose. Wings hyaline with two transverse fuscous bands.
Length, 5 mm.

Milton, Mass. Collected.

9. C. funestus Perkins.

Rep. Exp. Sta. Hawaii, Ent. v. 2, Bull. 4, p. 48, (oo), 1907.

Female: Black; basal four antennal joints ferruginous; front
femora pale to black; mandibles yellow. Head and frons dull, minutely
rugulose; antennal flagellum subclavate, pronotum with extremely fine
transverse rugulosity; mesonotum with few shallow punctures; scu-
tellum smooth and shining; propodeum truncate, rugose, posteriorly
minutely rugulose without a definite central area. Length, 2-5 mm.

Male: Black; anterior tibiae and tarsi testaceous, middle and hind
tarsi pale or brown. Head sculptured as in female, antennae as long as
head and thorax, flagellum submoniliform, mesonotum with faint
siirface sculpture, scutellum polished; propodeum as in female.
Length, 2 mm.

Tuscon, Arizona. Bred from jassid nymphs found on small
bush.

Tribe Aphelopini.

Anteoninae (part), Kieffer, Andre Spec. Hym. Eur., v. 9, p. 495, 1906.
Aphelopinae, Perkins, Rep. Exp. Sta. Hawaii, Ent. Bull. No. 11, p. 7, 1912.
Aphelopini Kieffer, Das Tierreich, 41 L., p. 214, 1914.

Distinguished from the Anteonini as follows: Wings hairy and
ciliated, with costa, subcosta, radius and a broad truncate pterostigma;
the rest of the veins obliterated. Legs not thickened, fore legs without
chela in 9 •

May, 1918] Parasites of Leaf-IIoppers 275

Genus Aphelopus Dalman.

Gonatopus (part) Dalman. vSwenska A. k. Handl., p. 82, 1818.
Dryinus (Aphelopus) Dalman, Anatccta ent., p. 14, 1823.
Aphelopus Haliday, Ent. Mag., v. 1, p. 273, 1833.
Ceraphron (part), Ratzchurg. Ichneum, v. 2, p. 141, nr. 2, 1848.

Head transverse, anteriorly slightly narrowed, posteriorly deeply
arcuate. Eyes almost reaching the hind margin, oval, with a short,
scattered, indistinct pubescence. Ocelli forming a triangle, the lateral
ones nearer the hind edge than to the eye, farther apart than from the
eye. Mandible twice as long as broad, distally truncate and three-
dentate, the teeth three-angled, of equal length, the middle often with
a little tooth at its base. Maxillary palpus with fine long joints, labial
palpus two or three jointed. Antennae originating close or the clypeus
and farther apart than from the eyes, pubescent. Pronotum barely
visible from above; mesonotum convex, transverse, anteriorly rounded,
with two posteriorly converging parpasidal furrows extending only
part way down the mesonotum, seldom without such. Scutellum
posteriorly rounded, anteriorly with a transverse impression; propo-
deum large, at first horizontal, then gradually sloping; propleura
concave; mesopleura strongly convex, with an oblique longitudinal
furrow, separated from the metapleura by a ridge reaching from the
wHng bases to the hind edge of the middle coxae. Venation as above
given. Hind wings without distinct veins, with four frentila. Spur of
fore tibia hairy, narrowed on the whole ventral side to a base, hyaline
lamella, which is discontinued suddenly before the distal end of the
spur, this thus appearing two-lobed, all claws simple. Abdomen
strongly laterally compressed, as long or shorter than the thorax.

Key to Species of Aphelopus.

1. Thorax without white color 2

Thorax in part white, at least first two pairs of legs white or pale yellow. . 10

2. Radius very short one-third as long as the pterostigma, abdomen reddish

brown 1 . riifiventris

Radius at least as long as the pterostigma, abdomen black 3

3. Propodeum w'ith a smooth median field, head without red coloring 5

Propodeum without a medium field 4

4. Face, clypeus, mandible and gena; ferruginous 2. affinis

Face white to just above bases of antennas 9. comesi

Face black 3. dikraneuri

5. Mesonotum smooth and shining 5. americanus

Mesonotum dull and coriaceous 6

6. Head with mandibles above white 7. arizioncus ( 9 )

Head with white facial markings 7

7. Clypeus alone white 4. microleucus

More than clypeus white 8

8. White facial markings extending a little above the antennae. . . .8. viduus
White facial markings extending at least as far as the anterior ocellus,

along the eye margins 9

9. Mesonotum densely minutely shagreened 7. arironicus (cf)

Mesonotum coriaceous 6. varicornis ( 9 )

10. Only pronotum and prostemum white, the rest of the thorax black

10. albopictus

Mesonotum, except middle section, pleura, and sternum white

11. pulcherrimus

276 The Ohio Journal of Science [Vol. XVIII, No. 7,

1. A. rufiventris Ashmead.

Bull. U. S. Mus., V. 45, p. 100, (9 ). 1893.

Female: Head and thorax black, minutely punctate; abdomen
rufous; antennee and legs honey yellow. Antennas short, subclavate.
Propodeum coarsely rugose. Length, 2 mm.

Jacksonville, Florida. Collected.

2. A. affinis Ashmead.

Bull. U. S. Mus., V. 45, p. 102, (9 ). 1893.

Female: Black, face from frons ferruginous; antennee brown; legs
except posterior tibia; which are fuscous, honey yellow. Head and
thorax minutely punctate, propodeum rugulose. Length, 2.2 mm.

Canada. Collected.

3. A. dikraneuri n. sp. Plate I, Fig. 3.

Black; clypeus ferruginous, mandibles yellow, maxillary palpi
long, white, extending beyond the base of the head with four segments
visible. Legs fuscous to brown; fore legs may be testaceous, if so
coxae are fuscous. Antennae brown entirely. Ocelli in triangle, the
anterior located medianly on the vertex and farther from the lateral
ones than they are from the hind margin of the head, these nearer to
the eyes than to each other. Eyes oval, without hairs. Antennae fili-
form, two same length and width as one in male, both wider than
three; one somewhat longer than two in female; four-nine equal in
length three times longer than wide, ten distinctly longer than nine,
tapering at tip. Vertex and mesonotum minutely reticulately sculp-
tured. Parapsidal furrows distinct, converging and extending half way
the length of the mesonotum in male, shorter and not extending to
middle in female. Scutellum finely reticulately sculptured, metathorax
smooth and shining, propodeum reticulately rugose without median
smooth area. Radius slightly curved and part beyond usual point of
angelation obliterated. Abdomen compressed laterally. Head and
thorax pubescent, abdomen with numerous hairs on ventral half of
side and posteriorly. Length, 1.5 mm.

Columbus, Ohio. Bred from Dikraneura fieheri, adults.
Described from one male and three females. Type deposited in
Entomological Museum, Ohio State University. Paratypes in
writer's collection.

4. A. microleucus Perkins.

Rep. Exp. Sta. Hawaii, Ent. v. 2, Bull. 4, p. 55, (9 ), 1907.

Male: Black; clypeus and mandibles white, two basal antennal
joints dark brown, all coxae and trochanters pale yellowish white or
white, front legs testaceous; middle legs darker, and hind legs black.
Head and thorax minutely sculptured, propodeum reticulately rugose,
small smoother area on posterior face. Antennae filiform. Length, 2 mm.

Nogales, Arizona. Bred from Erythroneura sp.

May, 1918J Parasites of Leaf-Ifofypcrs 277

5. A. americanus Aslinicad.

Bull. U. S., Mus. V. 45, p. 100, (9), 1893.

Male: Black; antennae reaching to middle of abdomen; brown
except scape which is \'cllow. Legs honey yellow, jjosterior femora and
tibia fuscous or black. Finch' punctate, except mesonotum and scutel-
kmi which arc smooth with few indistinct scattered punctures.
Length, 1..') mm.

Jacksonville, Florida. Collected.

6. A. varicornis Bnies.

Bull. Wis. Soc, V. 4, p. 143, (9 ), 190G.

Female: Black; legs and first two antennal joints light yellow; face
below ocelli and maxillary palpi white. Head finely shagreened, shining;
mesonotum shagreened. Parapsidal furrows distinct extending two-
thirds the length of mesonotum. Propodeum rounded behind, areolated
and rugulose. Radius a little longer than stigma, faintly curved at tip.
Length, 1.75 mm.

Woods Hole, Massachusetts. Collected.

7. A. arizonicus Perkins.

Rep. Exp. Sta. Hawaii, Ent. v. 2, Bull. 4, p. 53. (d" 9 ), 1907.

Female: Black; basal two antennal joints yellow or yellow-brown,
legs testaceous, except hind femora and tibiae which are black or pitchy.
Head and mesonotum finely shagreened, antenna; sub-clavate, pro-
podeum finely reticulately rugose, posteriorly with smooth shining area.

Male: Similar. Two basal antennal joints darker brown, face
yellow below, legs entirely testaceous, antenna; more elongate.
Length, 2 mm.

Nogales,- Arizona. Bred from Dikraneura sp.

8. A. viduus Perkins.

Rep. Exp. Sta. Hawaii, Ent., v. 2, Bull. 4, p. 54, (9), 1907.

Male: Black, lower part of face and mandibles pale yellow or cream-
colored. Front and middle legs pale yellow, tarsi infuscate, hind
femora, tibiae, and tarsi, pitchy. Head and mesonotum densely
shagreened. Antennae long, slender and filiform. Propodeum reticulately
rugose, a postina median area present. Length, 2 mm.

Nogales, Arizona. Collected.

9. A. comesi n. sp.

Male: Black, face white to jitst above bases of antennae. Mandibles,
palpi, and short antennae brown, the basal two joints testaceous. Fore
legs, coxas and trochanters of middle and hind legs and femora of middle
legs white, tibia of middle leg darkened, femora and tibiae of hind legs
fuscous, tarsi of middle and hind legs sordid yellow. Abdomen brown.
Head finely coriaceous shining, clypeus truncate. Eyes oval with fine,
almost invisible pubescence. Antennae filifonn, three shorter than four.

278 The Ohio Journal of Science [Vol. XVIII, No. 7,

which is shorter than five, seven and eight almost equal, slender, longer
than four or five, more than twice as long as wide, ten long, slender.
Head dull, finely coriaceous, mesonotum coriaceous with indistinct
parapsidal furrows, converging and being obliterated slightly beyond the
middle and almost indistinguishable anteriorly. Propodeum without
median field and distinctl}^ rugosely sculptured. Wings hyaline, radius
pale, curved extending nearly to margin of wing. Abdomen with sparse
hairs, smooth. Length, 1.5 mm.

Described from one specimen. Columbus, Ohio. Bred from
Erythroneura comes, adult. Type deposited in Entomological
Museum, Ohio State University.

10. A. albopictus Ashmead.

Bull. U. S. Mus., V. 4.5, p. 101, (9 ), 1893.

Black, face below front ocellus, maxillary palpi, anterior and middle
legs, pronotum to tegvdae, and propectus white or yellow- white. Antennae
dark brown. Head finely punctulate, propodeum rugose. Length,
1.5 mm.

Washington, D. C, and Bladensburg, Md. Collected.

11. A. pulcherrimus Perkins.

Rep. Exp. Sta. Hawaii, Ent., v. 2, Bull. 4, p. 54, sex? 1907.

Sex?: Black, whole face white, scape, legs white or pale yellow,
mesonotum exteriorly to the parapsidal furrows, whole thorax beneath
and pleural portion, cream colored. Antennae except scape testaceous.
Head and thorax densely shagreened, posterior area of metanotum
minutely shagreened. Length 1.5 mm.

Nogales, Arizona. Bred from Erythroneura sp.?

THE DIGESTIBILITY OF SOY BEAN MEAL BY MAN.*

By j. F. LYiMAN and W. G. Bowkrs,

There always has been considerable interest, in connection
with human nutrition, in the seeds of the legumes. The common
white or navy bean has played a prominent part in the affairs
of the United States in supplying nourishment to soldiers in the
service and to laborers on the farm and in the factory. The
bean has been recommended as the "poor man's beefsteak,"
supplying as does meat, large quantities of proteins.

In China and Japan another legume, the soy bean, is exten-
sively used as human food. This is ordinarily prepared for the
table by processes not in general use in this country, usually
involving some kind of fermentation. These preparations supply
a very important part of the food requirement, especially of the
protein requirement, of the Chinese and Japanese, and have
been found to be well utilized in the body, in some cases 96 per
cent of the protein being absorbed.^

The soy bean has been cultivated rather extensively in the
United States in recent years, but it has been used almost
entirely as forage or for animal feed. Recently a soy bean
meal, a by-product in the manufacture of soy bean oil, has
been produced in considerable amounts in some sections of the
South. The composition of this material and of the whole beans
in comparison with common white beans suggests that we have
here valuable supplies of human food which should not be
neglected. The following table gives the average composition of
the different materials.

^White beans (dry) .

^Soy beans

Soy bean meal

Moisture
Percent

12.6

10. n

.3.7

1

Protein
Percent

Fat

Fiber

Percent Percent

22.5

38.29

46.66

1.8

14.89

5.1

4.4

4.64

5.5

N-free
Extract
Percent

Ash
Percent

55.2

26.64

32.3

3.5

5.54

5.75

Obviously, chemical analyses alone do not measure the
nutritive value of food. We must know also the digestibility of
the food, the relative quality of its proteins, carbohydrates,
fats and minerals, the extent to which accessory substances or
vitamines occur, and finally whether, for any reason, it exerts a
harmful influence on the body.

* From the Laboratory of Agricultural Chemistrj^ and Soils, The Ohio State
University, Columbus.

279

280 The Ohio Journal of Science [Vol. XVIII, No. 7,

Alendel and Fine'' found the protein in mush prepared from
soy beans to be from 74.5 to 87.6 per cent digestible by dogs
and 85.3 per cent digestible by men (uncorrected values). In
fact the protein of so^^ beans was only 2.6 per cent less digestible
than was the protein in a mixed diet of wheat, meat, eggs and
peanut butter. The protein of the common white bean, baked
in the usual way, was distinctly less digestible, viz., 77.9 per
cent, in the human subject. The experiments of Wait^ on the
digestibility of white beans agree with this result. Daniels and
Nichols,*^ from experiments on rats, have concluded that the
proteins of the soy bean are highly efficient in promoting the
growth of young animals, in fact, about as efficient as casein of
milk. On the other hand, the proteins of the white bean appear
to be decidedly poorer in quality or about as valuable as the
proteins of corn.^ Osborne and MendeP found in soy beans
abundant amounts of water-soluble accessory or vitamine, while
the fat-soluble vitamine, though not abundant, appears to
exceed the amount found in other seeds and grains commonly
used for human food.

The nutritive quality of the white bean is further lowered
by the presence of considerable amounts of carbohydrates,
e. g., hemicelluloses, which are indigestible in man, and which
ferment in the intestine with the production of large quantities
of gas, resulting in discomfort and even in injury to the intestine
because of the stretching effect of the gas." The carbohydrates
of the soy bean, while resembling those of the white bean, appear
to be considerably more digestible and less prone to intestinal
fermentation. Mendel and Fine^ in comparable experiments
with dogs, fed in one case soy bean flour and in the other a
preparation made from hulled white beans, and found five
times as much fiber and over three times as much hemicellulose
in the feces of the white bean fed animal. In the human subject
the weight of the air dry feces from the soy bean feeding was
increased only four per cent above the mixed diet period, while
on the white bean diet the increase in weight of air dry feces
was 68 per cent, indicating a good utilization of the non-
nitrogenous material in the first case and a poor utilization in
the second. In our own experiments the liberal use of soy bean
meal in the diet, 120 grams per day for a three-day period, did
not result in voluminous stools, on the contrary, both the protein
and carbohydrates (N-free extract) of the soy bean meal was
highly digestible, as will appear from the following experiment.

May, 1918.1

Digestibility of Soy Bean Meal

281

EXPERIMENT I.

The subject was a healthy man, aged 3<S, weight 150 pounds,
engaged in moderately active laboratory work each day. Since,
obviously, it is impossible to consume with relish an exclusive
diet of soy bean meal, a mixed diet was selected containing a
few articles of known digestibility in addition to soy bean meal,
the chief constituent. The feces from the three-day experi-
mental period were marked off with lampblack taken with the
appropriate meals. Soon after collection of the stools they were
moistened with alcohol containing a little sulphuric acid, dried
on the steam bath, brought to an air-dry condition by exposure
to the atmosphere of the laboratory and finely ground. Protein
was determined by the Kjeldahl method (N x 6.25) ; fat by the
method of Gephart and Csonka,'' nitrogen-free extract, fiber
and ash by the ordinary routine methods.^"

The soy bean meal used in this experiment had the following
composition :

Moisture 6.5 percent

Ash 5.7

Protein 44.1 "

Crude Fiber 5.9

Fat 3.3

N-free Extract 35.5 "

Two hundred grams of this meal were mixed with 400
grams of patent wheat fiour and 10 grams of sugar and made
into yeast-raised bread. This bread was of good texture and
highly palatable.

The diet for the three days was as follows, approximately
one-third being consumed each day.

Grams

Protein
Grams

Fat
Grams

N-free
Ext.
Grams

Calories

Soy bean meal

Patent wheat flour

Milk..

200
400
3000
200
200

88.2

53.2

103.1

6.6

6.0

12C.0

170.0

71
288
141

200

696
1420
2064

Butter

Cane sugar. .

1530
800

Total...

4000
1330

244.5
81.5

302.6
100.8

700
233

6510

Per Day

2170

282 The Ohio Journal of Science [Vol. XVIII, No. 7,

The feces from the three days weighed 248 grams moist, and
66.6 grams air-dry. They contained:

Percent Grams

Moisture 7.25 4.82

Ash 19.70 13.12

Protein 27.66 18.82

Crude fiber 8.46 5.63

Fat 22.01 14.65

N-free extract 15.22 10. 14

In addition to undigested food residues, feces always contain
nitrogenous metabolic products derived (1) from the digestive
fluids and (2) from mucous and epithelial cells from the walls of
the digestive tract. In order to determine the amount of these
metabolic products a basal ration consisting of milk, butter,
sugar, starch and agar, and designed to yield feces of about the
same bulk as in the soy bean period was eaten. The diet was
as follows:

3000 grams milk
275 grams butter
150 grams cane sugar

130 grams potato starch, boiled in water to make a pudding.
36 grams agar, eaten dry and raw.*

One-third of the above was consumed daily for a three-day
period. The resulting feces weighed 270 grams moist, and 55.6
grams air-dry, and contained:

Percent Grams

Moisture 5.36 2.98

Ash 19.50 10.85

Protein 11.90 6.60

Fat 31.20 17.34

Crude fiber 1.40 .77

N-free extract 30.60 17.03

The protein in the feces of this period represents metabolic
products from the digestive juices and intestinal walls as well
as some unabsorbed protein from the milk. Deducting this
from the protein eliminated in the feces during the soy bean
period we have 18.82 — 6.60=12.22 grams, which represents the
unabsorbed protein from the wheat flour and soy bean meal.
This gives a coefficient of digestibility for the total protein of
the bread of 91.3. Assuming that protein of patent wheat flour
is 88.6^^ per cent digestible the coefficient of digestibility of the
soy bean meal protein becomes 93. This is hardly fair, however,
since the coefficient 88.6 was obtained without making a cor-

* The agar contained 6.35 percent of ash and 15.75 percent of moisture.

May, 1918]

Digestibility of Soy Bean Meal

283

rection for metabolic products in the feces. We feel safe in
concluding, however, that soy bean meal prepared as in this
experiment is as digestible as regards its protein as is the protein
of fine wheat flour which stands supreme among the cereal
grains in this respect.

The carbohydrates of bread made from fine wheat flour are
97.7 per cent absorbed in man," milk sugar 99^'^ per cent and
cane sugar probably 100 per cent. Using these values the
nitrogen-free extract of the soy bean meal is found to have a
coefficient of digestibility of 96.9 per cent.

EXPERIMENT II.

In this experiment the soy bean meal was prepared as a

porridge by cooking five hours in a double boiler. This sample

of meal contained more grit than that used in Experiment I,

but not enough to increase the ash content greatly. It had

the following composition:

Percent

Moisture 4.17

Ash 5.80

Protein 49.31

Crude fiber 5 . 10

Fat 6.50

N-free extract 29. 12

The diet for the three-day period contained the following,
approximately one-third being eaten each day:

Grams

Protein
Grams

Milk

Butter

Cane sugar. . . .
vSoy bean meal.

Total

Per Day. .

2000
150
300
370

70

182

2750
917

252
84

Fat
Grams

N-free
Extract
Grams

80.
127.5

24'0

94

300
108

231.5

77.2

502
167

Calories

1376
1147
1200
1375

5098
1699

The fecal residue from this food weighed 224 grams moist
and 58.1 grams air dry and contained:

Percent Grams

Moisture 4.40 2 56

Ash 20.64 11.19

Protein 39.90 23.20

Crude fiber 7.00 4.06

Fat 18.96 11.00

N-free extract 9.00 5.22

284 The Ohio Journal of Science [Vol. XVIII, No. 7,

Assuming that 6.0G grams of protein represents the unab-
sorbed protein of the milk plus the metabolic protein from the
digestive fluids, etc., an assumption that can not be far from
the truth, then the unabsorbed protein from the soy bean meal
must be 23.20 — 6.60 = 16.60 grams. This gives a coefficient of
digestibility for the protein of the soy bean meal of 90.9. If
the carbohydrates of milk are 99 and cane sugar 100 per cent
absorbed then the coefficient of digestibility for the nitrogen-
free extract of the soy bean meal is 96. In this experiment, as in
Experiment I, we cannot calculate accurately the digestibility of
the fat but we were not particularly concerned with that phase
of the subject at this time. By adopting the proper experimental
procedure one might determine satisfactorily the digestibility
coefficient of soy bean fat, but for that purpose the amount of
the experimental fat should be largely increased while other fats
in the diet should be reduced to a minimum.

Summary: The protein of soy bean meal, prepared as a
bread by mixing with wheat flour or as a porridge, is highly
digestible by man. The digestibility factors obtained in the
two experiments were 91.3 and 90.9.

The carbohydrates of soy bean meal appear to be much more
highly digestible and less prone to intestinal fermentation than
are the carbohydrates of the common white bean. The digesti-
bility factors obtained in two experiments were 96.9 and 96.

The chemical composition, digestibility and palatability of
foods prepared from soy bean meal from which most of the oil
has been expressed, indicate that such a meal is an excellent
food for man.

References.

1. Oshima, Bui. 159 (1905), U. S. Dept. Agr. Office Exp. Stations.

2. Bui. 28, U. S. Dept. Agr. Office Expt. Stations.

3. Street and Bailey, J. Ind. Eng. Chem. 7, 853, (1915).

4. Mendel and Fine. J. Biol. Chem., 10, 447, (1914).

5. Bull. 187, U. S. Dept. Agr. Office Expt. Stations.

6. Daniels and Nichols, J. Biol. Chem., 32, 91, (1917).

7. McCollum, Simmonds and Pitz, J. Biol. Chem., 29, 521, (1917).

8. Osborne and Mendel, J. Biol. Chem., 32, 369, (1917).

9. Gephart and Csonka, J. Biol. Chem., 19, 521, (1914).

10. Bui. 107, U. S. Dept. Agr. Bureau Chem,

11. Report Maine Expt. Station, 1898.

12. Farmers Bui. 363, 22.

Date of Publication, June 30, 1918.

THE

Ohio Journal of Science

PUBLISHED BY THE

Ohio State Univkrsity Scientific Society

Volume XVIII JUNE. 1918 No. 8

TABLE OF CONTENTS

Fenton— The Parasites of Leaf-Hoppers— Part III 285

Barrows— A List of Ohio Spiders 297

HiNE— Description of Seven Species of Asihis (Family Asilickt) ;->19

Drake — North American Species of Teleonemia Occurring North of Mexico. 323

THE PARASITES OF LEAF-HOPPERS.

With Special Reference to Anteoninae.

F. A. Fenton.

PART III.

The Effect of Parasitism on the Host.

There has been comparatively little work done in regard to
the internal or cytological changes in the insect host brought
about by its parasite. Giard (1889) in working out the biology
of Aphelopus melaleucus parasitic on Erythroneura (Typhlocyba)
hippocastani, misinterpreted the real nature of the larval sac,
supposing that it was a true animal gall, "formed by a gradual
dilation of the hypodermis which secretes an abnormal cuticle
* * * " He proposed the name thylacies to those galls
produced in animals comparing the typhlocybid larval sac
with the tumors caused by Cuterebra on the skin of mammals.
He compared the genitalia of normal and parasitized E. hypo-
castani and E. doiiglasi. In the females of these two species the
ovipositor in parasitized individuals is greatly reduced and
functionless. In the males of E. douglasi very little change
is brought about in the penis which is comparatively simple, but
in E. hippocastani where this structure is a very complex eight-
branched organ it is reduced to six or even three branches, thus
greatly affecting the specific characters. Because of this
superficially it might be confused with E. rosce. Certain acces-
sory sex organs were also found to be affected by the dryinid.

285

286 The Ohio Journal of Science [Vol. XVIII, No. 8,

Marchal (1897) studied in detail the pathological conditions
that Trichacis remulus Walk, produced in its host Mayetiola
destructor, the Hessian fly. He observed that the Trichacis
larva is always in intimate connection with the nervous system
of the cecidomyid larva and he noted the remarkable cell
proliferations that are brought about by the parasite. Although
he did not find the early stages and therefore was unable to
state anything about their origin, he made some striking con-
clusions, saying that ' ' These groups of giant cells are evidently
destined to accumulate nutritive materials necessary for the
parasite. They are a kind of internal animal gall developed
by the presence of the Hymenopteron. "

Keilin and Thompson (1915) noticed the peculiar mass of
hypertrophied tissue formed within Erythroneura hippocastani
parasitized by Aphelopus melaleiiciis and traced its origin to
the hypodermal cells which are stimulated to abnormal growth
by the presence of the parasite. The early stages of the dryinid
were found to be enclosed by this tissue similar to that in the
Trichacis larva, shutting the parasite off from the viscera of
the host. This cyst was noticed to be surrounded by a mem-
brane which persisted after the parasite larva had assumed the
curved position and had broken through to the exterior of the
host. They believe that the parasite draws its nourishment
from its host through the cyst membrane until the fifth stage
is reached, and that the tissue is not a phagocytic cyst. They
compare the cyst to the placenta in animals or to a vegetable
gall and term the parasitism " placentaire'' or " gallicole. "

Kornhauser (1915-16) in studying the effects of Aphelopus
parasitism on Thelia bimaciilata observed marked changes in
the external characteristics of the sexes, especially in the size
and form of the male and a reduction of the external genitalia
of both sexes, stating that "the presence of parasites in the
male nymph brings about lower oxidation, storing of fat,
retarded rate of development, increased size; and with this
change in metabolism comes a change in some of the secondary
sexual characters. But changed metabolism is not powerful
enough to change the external genitalia, it merely reduces
them in size. "

In studying the effects of the parasitism of Aphelopus
comesi on Erythroneura comes the writer was able to confirm
many of Keilin and Thompson's observations and to add a
number of interesting points. The early phases were not studied

June, 1918] Parasites oj LeaJ-IIoppers 287

owing to a lack of material, but comparisons of the later stages
of parasitism in these hoppers with normal individuals were
made. When the dryinid is in its second instar and has become
partly external the hypertrophied tissue has reached its maxi-
mum size (Plate II, Fig. 2). It is seen as a large ovoid mass
of cells occupying most of the body cavity in the hind thoracic
and anterior abdominal regions, displacing and pushing back-
wards the large storage stomach of the host. It extends for
three somities as a rule, its anterior extremity lying between the
muscles of the third thoracic segment. It is in intimate con-
nection w4th the point of attachment of the parasite. In
cross section the thin membrane surrounding it can be seen,
except at that part nearest the parasite, w^here the cells of the
cyst are in contact with the mouth parts of the latter, (Plate
IV, Fig. 3).

In the earliest stages studied the cell walls and nuclei are
sharply defined and part of the tissue is composed of dividing
cells, (Plate IV, Fig. 2; Plate VI, Fig. 1). The cells are filled
with food globules and are vacuolated. Later division ceases
and the cells become more noticeably vacuolated, (Plate IV.,
Figs. 3 and 4). Late in the fourth instar of the parasite dis-
integration of the cells begins, (Fig. 5, Plate IV, Figs. 3 and 4).
The cell walls are broken down, the nuclei become disintegrated
and the protoplasm becomes very largely filled with round
vacuoles. Directly after the fourth molt of the parasite this
cyst is attacked first and devoured after which the host viscera
are devoured.

The cyst stains easily with Delafields haemotoxylin, being
thus sharply contrasted with the surrounding tissues which
take the eosin stain more readily. It reacts to stains similarly
and resembles fatty tissues.

The function of such a cyst is problemmatical. It is not
found in a great many other insect hosts of Hymenopterous
parasites and has not been observed in any other genus of the
Anteoniyia so far. It cannot be phagocytic because the mandi-
bles of the parasite are not developed or are functionless until
the last larval molt, and it is not absorbed. Except for cyto-
plasmic changes it remains unchanged until the fifth instar is
reached by the dryinid. It doubtless serves as a means for
absorbing, storing up, and then in turn giving up in a modified
manner food for the parasite that otherwise would have been
utilized by the host. This food is probably absorbed from the
blood of the hopper through the cyst membrane.

288

TJie Ohio Journal of Science [Vol. XVIII, No. 8,

This hypertrophied tissue is developed principally at the
expense of the gonads of the host, (Plate III, Figs 1 and 2).
These are lacking or almost so in parasitized individuals.
The digestive system is also modified to some extent in that the
storage stomach is forced backwards and displaced by the
abnormal tissue. The malpighian tubules are much smaller in
parasitized individuals. Normally these are large and much
swollen, in parasitized specimens they are hardly enlarged at

Text Fig. 5. Hypertrophied tissue in dryinized male Erythroneura comes

(high power).

all. When the parasite pushes apart the abdominal somites
and begins to grow outwards the cavity thus formed in the
body wall is enlarged until it becomes almost circular. Around
this cavity a layer of hypodermal cells is developed as an abortive
response by the host tissues to heal over the wound, (Plate IV,
Figs. 2 and 4).

No other host was studied in as much detail in regard to
the cytological changes as was the above. However it is certain
that in the case of Gonatopus contortuliis parasitic on Delto-
cephalus sayi, (Plate VI, Fig. 2), there is no such development
and from general dissections of all the other species studied in

June, 191S] Parasites of LeaJ-IIoppers 289

this paper no sueh tissues were found. Since many Cicadellid
species are attacked and parasitized by dryinids after they have
become sexually mature it is doubtful in these cases whether the
gonads would be completely disintegrated. In fact in para-
sitized DeltocepJialiis sayi they are present, but modified in
that the germ cells are not matured after a certain point is
reached in the parasites growth, so that if reproduction is
not entirely stopped it is greatly impaired.

In Erythronenra comes the hypertrophied tissue is probably
caused by a stimulation set up by the presence of the parasite
in the host's tissues or to some toxic substance secreted by it.
That the sting alone will not produce such a result is proved
by the fact that hoppers that have been stung but not oviposited
in develop and mature in a perfectly normal manner.

A comparison of the genitalia of both sexes in Erythroneiira
comes in normal and parasitized individuals showed no
differences.

CONCLUSIONS.

1. The larva of Aphelopiis species secretes some toxic
substance into the tissues of the host or otherwise stimulates
the abnormal development of the hypodermal tissues into a
hypertrophied cell mass.

2. This is developed at the expense of the gonads which
do not develop upon the maturity of the host.

3. This is surrounded by a membrane and functions as a
means of absorbing food from the body of the host for the
parasite and is itself not consumed until just before the death of
the host.

4. The genitalia of the host are not modified by the parasite.

5. A wall of cells is developed around the wound produced
in the body wall of the host.

G. The malpighian tubules of the host are underdeveloped.

7. Gonatopiis, Ilaplogouatopus, and Chelogynus species
that parasitize nymphs stop further development of the host
which does not become mature.

8. Adults parasitized by these genera, that have become
sexually mature, may still reproduce at first but there are
evidences that soon reproduction is greatly impaired or entirely
stopped.

9. The hypertrophied tissue has not been found produced
by any genus other than Aphelopiis.

290 The Ohio Journal of Science [Vol. XVIII, No. 8,

BIBLIOGRAPHY.

AsHMEAD, W. H. — Monograph of the North American Prototrvpidce. Bulletin

U. S. National Museum, 45, pp. 80-102. 1893.
Brues, C. T.

Descriptions of New Ant-Like and myrmecophilous Hymenoptera. Trans-
actions American Entomological Society, Vol. 29, p. 125, 1903.

Some New Species of Parasitic Hvmenoptera. Canadian Entomologist,

Vol. 36, pp. 117-119, 1904.

-Notes and Descriptions of North American Parasitic Hymenoptera.

Wisconsin Natural History Society, Vol. Ill, pp. 183-185, 1905.

-Notes and Descriptions of North American Parasitic Hymenoptera,

Number 11, Wisconsin Natural History Society, Vol. IV, p. 143, 1906.

-Notes and Descriptions of North American Parasitic Hymenoptera,

Number V, pp. 101-102, 1907.

-Some Notes on the Geological History of Parasitic Hymenoptera. Journal

New York Entomological vSociety, Vol. XVIII, No. 1, pp. 1-16, 1910.

-Drynida;, in Hymenoptera of Connecticut. Connecticut State Geological

and Natural History Survey. Bulletin 22, pp. 613-616, 1916.
Bradley, J. C. — Contributions to the Entomology of the Selkirk Mountains of

British Columbia. Canadian Entomologist, Vol. 38, p. 380, 1906.
GiARD, A. — Sur une galle produite chez le Typhlocyba rosce. par une larve d' Hymen-
optera. Comptes Rendus, Ac. Sc. Paris,' CIX, pp. 79-82, 1889.
Sur la castration parasitaire des Typhlocyba par une larve d'Hymenoplere

(Aphelopus malaleucus Dalm.), et par une larve de Diptere {Ateleneura spuria

Meig.). Comptes Rendus, Ac. Sc. Paris, CIX, pp. 708-710, 1889.
Haliday, a. H. — Notes on the Bethyli and on Dryinus pedestris. Entomologists

Magazine, Vol. 2, pp. 219-221, 1834.
Hood, J. D. — Notes on the Life History of Rhopalosoma poeyi Cresson. Proceedings

of the Entomological Society of Washington, Vol. XV, No. 4, pp. 145-148, 1913.
Keilin, D., and Thompson, W. R. — Sur les larves des Dipteres et Hymenopteres des

Tvphlocvbes. Comptes Rendus des Seances de la Societe de Biologv, T.

LXXVI'II, pp. 1-9, 1915.
KiEFFER, J. J. — Family Dryinidce. Genera Insectorum, Fasc. LIV, 1907.

Monograph of the Bethylidce. Das Tierreich, 41 Lieferung, pp. 1-22, 1914.

KoRNHAUSER, S. J. — Changes in Thelia himacidata (Fabricius) Induced by Insect

Parasites. Abstract Amer. Soc. of Zoologists, Columbus, Ohio, 1915.
Further Studies on Changes in Thelia bimaculata brought about by Insect

Parasites. Anatomical Record, Vol. 11, No. 6, pp. 538-540, 1916.
Marchal, p. — Les Cecidomyies des Cereales et leurs Parasites. Annales de la

Societe Entomologique de France, pp. 1-105, 1897.
MiK, J. — Zur Biologv von Gonatopus pilosus Thoms. Wiener Entomologische

Zeitung, 1, pp. 215-221, 1882.
MiSRA, C. S. — The Indian Sugar-cane Leaf-hopper Pyrilla aberrans Kirby.

Memoires of the Department of Agriculture in India, Vol. V, No. 11, 1917.
Patton, W. H. — Descriptions of Several New Proctotrupida and Chrysidce. Can-
adian Entomologist, Vol. 11, p. 65, 1879.
Perkins, R. C. L. — Leaf-hoppers and their Natural Enemies. (Part I, Dryinidce).

Report of Work of the Exp. Sta. of the Hawaiian Sugar Planters Association,

Bull. 1, part 1, 1905.

ibid. Bull. 1, part 10, pp. 483-494, 1906.

ibid. Bull. 4, pp. 1-55, 1907.

ibid. Bull. 11, pp. 5-17, 1912.

Vau Duzee, E. p. — Catalogue of the Hemiptera of America North of Mexico.

University of California, Technical Bull. Vol. 2, 1917.
Swezey, O. H. — Observations on Hvmenopterous Parasites of Certain Fulgoridce.

Ohio Naturalist, III, pp. 444-451, 1903.
Observations on the Life History of Liburnia campestris with Notes on a

Hymenopterous Parasite infesting it.' Bull. U. S. B. E. 46. pp. 42-46. 1904.

June, 1918] Parasites oj Leaf-IIoppers 291

EXPLANATION" OF PLATES.

Explanation ou Platu L

Fig. 1. GonatopHs crytlirodes, female.

Fig. 2. PItorbas mirahilis. male.

Fig. 3. Aphelopus dikraneuri, male.

Fig. 4. Phorlxis niirahilis, female.

Fig. 5. Gonalopus crythrodes. male.

Fig. 6. Chelogynus oshonii. female.

Fig. 7. Chela of Gouatopiis erythrodes.

Fig. 8. Chela of Ilaplogonatopus americanus.

Explanation of Plate II.

Fig. 1. Normal male Erythroneura comes.
Fig. 2. Parasitized Erythroneura comes.

Tes., testes; sto., storage stomach; hyp., hypertrophied tissue; par.,
parasite, fourth instar.

Explanation of Plate III.
Fig. 1. Parasitized male Erythroneura comes, longitudinal section.
Fig. 2. Normal male Erythroneura comes, longitudinal section.

par., parasite, fourth instar; hyp., hypertrophied tissue; tes., testes.

Explanation of Plate IV.

Fig. 1. Normal male Erythroneura comes, cross section through second abdominal

segment.
Fig. 2. Parasitized female Erythroneura comes, showing nature of hypertrophied

tissue in early phase, cross section through second abdominal segment.
Fig. 3. Parasitized male Erythroneura comes, showing nature of hypertrophied

tissue in late phase, cross section through point of attachment of

parasite on first abdominal segment.
Fig. 4. Rarasitized male Erythroneura comes, cross section of same individual as

Figure 3, through head of parasite on second abdominal segment.
Sto., storage stomach; tes., testes; hyp., layer of hypodermal cells

developed around the wound produced in the host by the parasite;

par., parasite; es.. esophagus; hyp., hypertrophied tissue; ch., heavy

layer of chitin at point of attachment of parasite.
Camera lucida drawings, ocular 7.5. obiective 1(3 mm., Spencer

Microscope.

Explanation of Plate V.

Fig. 1. Normal male Erythroneura co)ues, cross section through second abdoininal
segment.

Fig. 2. Parasitized male Erythroneura comes, cross section through metathorax,
showing extreme anterior part of parasitic cyst.

Fig. 3. Parasitized female Erythroneura comes, cross section through first abdomi-
nal segment just before attachment of parasite.

Fig. 4. Parasitized male Erythroneura comes, cross section, third abdominal
segment.
Sto., storage stomach; tes., testes; hyp., hypertrophied tissue; gon.,
rudiment of ovary; par., parasite.

Explanation of pl.\te VI.

Fig. 1. Parasitized male Erythroneura comes, cross section through second
abdominal segment at point of attachment of parasite.

Fig. 2. Parasitized male Deltocephalus sayi, longitudinal section through testis
showing atrophy of this structure and disintegration of surrounding
tissues. This individual was dying from the effects of parasitism
at the time of fixation, the parasite being in the last stage.
Es., esophagus; hyp., hj'pertrophied tissue; m., membrane stirrotuiding
cyst; par., parasite; tes., testis; int., intestine.

Ohio Jourxal of Science.

Vol. XVIII, Pl.\te II.

F. A . Fenloii

Ohio Journal ok Science.

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A LIST OF OHIO SPIDERS.*

By W. M. Barrows.

The following list of 255 species of spiders is the result of
five years of intermittent study of the spiders of the state.
The work during the last two years has been carried on as a
project of the Ohio Biological Survey.

I am much indebted to Mr. J. H. Emerton, of Boston, for his
kindness in identifying the specimens collected in 1913, 1914,
1915, 1916, and part of 1917, and for much other help and
advice. In the list, the specimens determined by Mr. Emerton
are marked by an (E) placed after the name. A few specimens
were determined by Mr. Nathan Banks. These are indicated by
a (B). I am also indebted to the following persons for small
collections made in various parts of the state: J. S. Houser,
Wooster, Ohio; P. R. Lowry, Wooster, Ohio; R. J. Sim, Jefferson,
Ohio; T. L. Guyton, Winterset, Guernsey Co., Ohio, and
Chester I. Bliss, Sandusky, Erie Co., Ohio.

As far as possible the list follows the arrangement of the
Catalogue of Nearctic Spiders by Nathan Banks, U. S. National
Museum Bulletin 72, except that in each family the genera are
here arranged alphabetically. Included in the list are the
records of specimens collected in Ohio by Wm. Holden. These
are recorded as notes by Emerton in a Collection of the Arachno-
logical Writings of N. M. Hentz, Boston, 1875. These records
are marked (W. H.). Two records of Attidse are taken from the
Peckham's Revision of the Attidee of North America, and are
indicated by a (P).

Several new species found in the collections will be described
in a later paper.

* Contributions from the Department cf Zoology and Entomology, Ohio State
University, No. o2.

297

298 The Ohio Journal of Science [Vol. XVIII, No. 8,

THERAPHOSID.^.

Atypus milberti Walckenaer (B).

9 Cedar Point, Ohio, August 15, 1913.
This specimen was taken from the stomach of a frog by Mr. C. J.
Drake.

Atypus niger Hentz.

d^ Marietta, Ohio. (W. H.)
Brachybothrium pacificum Simon (B).

o^ 9 Rockbridge, Ohio, September 7, 1915.
9 Bainbridge, Ohio, August 17, 1917.
This very interesting "Folding-door Spider" builds its burrows
along the bases of the moist overhanging cliffs. Its habits and life-
history will be taken up in a separate paper. Up to the present time
it appears to have been recorded only from the West Coast.

PHOLCID^.

Pholcus phalangoides Fuessly.
cf 9 Columbus, Ohio.
A cellar or house species. Specimens were taken in the cellar of
the Denton Brothers Packing Co. plant.

Spermophora meridionalis Hentz.
Marietta, Ohio. (W. H.)

DYSDERID^.

Ariadne bicolor (Hentz).

d" 9 Cedar Point, Ohio, June 19, 1913. Ohio. (W. H.)
Common all over the state, building its tubular web in cracks of
trees, buildings and rocks.
Dysdera interrita Hentz.

d" 9 Columbus, Ohio, June 13, 1913.
Common around Columbus, under boards and rocks.

DRASSID^.
Callilepis imbecilla Keyserling. (E).

d Cedar Point, Ohio, June 19, 1913.
9 Columbus, Ohio, June 11, 1914.

Cesonia bilineata (Hentz). (E).

9 Cedar Point, Ohio, August 18, 1914.
Drassus hiemalis Emerton.

o"^ 9 Buckeye Lake, Ohio, June 24, 1917.

Gnaphosa bicolor (Hentz).
cf 9 Ohio. (W. H.)
Herpyllus ecclesiasticus Hentz. (E).

9 Rockbridge, Ohio, April 11, 1914.
cT 9 Ohio. (W. H.)
Widely distributed. Lives under loose dry bark of dead or injured
trees. Probablv winters in the adult state.

June, 1*) IS] A List of Ohio Spiders 299

Prosthesima atra (Hentz). (E).

cf 9 Rockl)ri(lgc, Ohio, June 12, 1915.
cf Columbus, Ohio, Au<^ust, 1914.
9 Marietta, Ohio. (W. H.)

Prosthesima depressa (Emcrton).

cf 9 Buckeye Lake, Ohio, June 24, 1917.

CLUBIONID^.

Agroeca minuta Banks.

9 Flint Ravine, eleven miles north of Columbus, July 4, 1917.
Anyphaena rubra? Emcrton. (E).

VouHK, 9 Rockbridge, Ohio, Fall, 1912.
Castianeira bivittata Keyserlinj^. (E).

9 Rockbridge, Ohio, June 1914.
(^ 9 Ohio. (W. H.)
Castianeira crocata (Hentz). (E).

o" 9 Cedar Point, Ohio, Julv 4, 1914.

9 Zanesville, Ohio. (W.^H.)
A female was fotuid laying eggs at Columbus on November 10, 1917.
Late in the summer this is one of the commonest black or black and red
spiders on the ground in dry fields and pastures.
Castianeira pinnata (Emerton).

cf Columbus, Ohio, September 2, 1915.
Castianeira trilineata (Hentz).

9 Athens, Ohio. (W. H.)

Chiracanthium viride Emerton. (E).
(^ 9 Columbus, Ohio, July, 1916.

9 Cedar Point, Ohio, July 20, 1903.
Taken on shrubs and low vegetation in the woods which fringe
streams and in moist woods. Hibernates nearly full grown under
leaves and rubbish on the ground.

Clubiona crassipalpis Keyserhng. (E).

9 Columbus, Ohio, June 11, 1917.
cf 9 Rockbridge, Ohio, June IS, 1916.
Clubiona minutissima Petrtmkevitch. (E).

d' 9 Columbus, Ohio, June 23, 1916. Delaware, Ohio. Rock-
bridge, Ohio.

Clubiona mixta Emerton. (E).

9 Cedar Point, Ohio, Julv 5, 1914.

d^ Cedar Point, Ohio, August 31, 1914.
Clubiona ornata Emerton.

9 Wooster, Ohio, August-September, 1917.
Taken while sweeping in grassland.
Clubiona pallens Hentz. (E).

cT Rockbridge, Ohio, June IS, 1916.

9 Columbus, Ohio, July, 1916.

300 The Ohio Journal of Science [Vol. XVIII, No. 8,

Clubiona rubra Emerton. (E).

cT 9 Columbus, Ohio, October 20, 1917.
These specimens are simiilar to the figures given bv Emerton, Trans.
Comm. Acad., Vol. VIII, 1890. Plate V, Fig. 0. 'They were found
under logs and boards in a pasture.

Clubiona tibialis Emerton. (E).
c^ 9 Cedar Point, Ohio.

Gayenna (Anyphaena) calcarata (Emerton). (E).
Erie Co., Ohio, 1915.
cT 9 Columbus, Ohio, July, 191G.
d" 9 Delaware, Ohio, June 24, 1916.
It is possible that two species are included under this name as two
forms of males have been taken at the same time. These live in young
trees in rather dense woods.

Gayenna (Anyphaena) incerta Keyserling. (E).

9 Sugar Grove, Ohio, December 26, 1915.
Taken in sifting leaves under the snow.

Gayenna (Anyphaena) saltabunda (Hentz). (E).
9 Rockbridge, Ohio, July 1, 1916.

Micaria longipes Emerton. (E).

9 Columbus, Ohio, September 6, 1917.
cf Sugar Grove, Ohio, September 11, 1917.
Found running in pastures and dry upland prairie. The young
have been taken in sweeping tall grass.

Phrurolithus alarius (Hentz). (E).

9 Columbus, Ohio, June 12, 1916.
9 Zanesville, Ohio. (W. H.)
cf Rockbridge, Ohio.
This and the following species are usually taken in moist leaves or
rotting wood in deep woods.
Phrurolithus borealis Emerton. (E).
cf 9 Columbus, June 24, 1916.
Phrurolithus formica Banks. (E).

9 and young. Columbus, Ohio, November 10, 1917, in ants

nest,
cf Rockbridge, Ohio, October 4, 1914.
These spiders were found living with the ant Crematogaster lineolata.

Trachelas ruber KeyserHng. (B).

cf Rockbridge, Ohio, June, 1914.

9 Cedar Point, Ohio, August 19, 1914.

9 Columbus, Ohio, October 22, 1917.

AGELENIDyE.

Agelena naevia Walckenaer. (E).

This species occurs everywhere. The mature males and females
are found from June to October, the height of the mating season being
September. The females lay the eggs in sheltered places, preferably
under bark and usually remain near the eggs until they die some time
in the earlv winter.

June, 1018] A List of Ohio Spiders 301

Cicurina arcuata Keyscrling. (E).

cT 9 vSuji;ar Grove, Ohio, December 26, 1915.
Cicurina brevis (Emerton). (E).

cf Columbus, Ohio, November 10, H)17.
9 Rockbridge, Ohio, July 1, 1916.
This species winters in the adult condition quite often with ants.
Mr. R. J. Sim collected several pairs for me at Jefferson, Ohio, on Nov.
21, 1917. These were living peaceably with ants (Acanthamyops
latipes Walsh.). Excepting that they are rather pale, they appear
perfectly normal.
Cicurina pallida Keyserling. (E).

(^ 9 Sugar Grove, Ohio, December 26, 1915.
Coelotes calcaratus Keyserling. (E).

9 Rockbridge, Ohio, October 4, 1914.
Coelotes montanus Emerton. (E).

d' 9 Rockbridge, Ohio, April 11, 1914.
Coras medicinalis (Hentz). (E).

Young, Cedar Point, Ohio, August 6, 1910.
cf 9 Columbus, Ohio, November 10, 1917.
9 Sugar Grove, Ohio, December 26, 1915.
Hahnia agilis Keyserling. (E).

9 Columbus, Ohio, November 10, 1917.
cT Sugar Grove, Ohio, December 26, 1915.
Hahnia cinerea Emerton. (E).

9 Columbus, Ohio, June 12, 1916.
Sugar Grove, Ohio, July, 1915.
Tegenaria derhami (Scopoli). (E).

cf Columbus, Ohio, October 3, 1915.

DICTYNID^.

Amaurobius sylvestris Emerton. (E).

9 Columbus, Ohio, November 10, 1917.
This spider is found under logs, in crevices of bark and rocks all
all over the state.

Argenna obesa (?) Emerton. (E).

Trans. Conn. Acad. Vol. 16. p. 399.

d^ Rockbridge, Ohio, June 12, 1915.
Mr. Emerton comments on this as follows: "You have one little
Argenna like our coast species, but twice as large as any I have seen.
Here it lives under straw along the shore and around the salt marshes
in exactly the same situations as Lophocarenum arenarium at Sandusky"
Dict3ma frondea Emerton. (E).
9 Cedar Point, Ohio.

Dictyna longispina Emerton. (E).

cf 9 Delaware, Ohio, June 30, 1917.
9 Columbus, Ohio, May 24, 1916.
Rather common in woods near the Olentangy River where it occurs
in hollow stems and under bark during May and June.

302 The Ohio Journal of Science [Vol. XVI II, No. 8,

Dictjma minuta Emerton.

cf 9 Coltimbus, Ohio, June 13, 1917.

Dictjma muraria Emerton.

d^ 9 Wooster. Ohio.

cf 9 Buckeye Lake, Ohio, June 14, 1917.
Dictyna rubra Emerton. (E).

cf Columbus, Ohio, June 24, 1916.

Dictjma volupis Ke\serling.

(f 9 Colum.bus, Ohio, June 11, 1917.
A very common and abundant species. It winters nearly mature
and becomes mature early in the spring.

Titanoeca americana Emerton. (E).
9 Cedar Point, Ohio.

Titanoeca brunnea Emerton.

9 Delaware, Ohio, June 30, 1917.

THERIDIID^.

Ancylorrhanis hirsuta (Emerton). (E).

& Rockbridge, Ohio, July 1, 1916.

Argyrodes cancellatum (Hentz). (E).
o^ 9 Clifton. Ohio, July 14, 1917.
cf Rockbridge, Ohio, June 1914.
This pair was taken from the web of an Acrosoma spinea in rather
deep woods. Other specimens have been taken at Cedar Point and
Rockbridge.
Argyrodes trigonum (Hentz). (E).

Rockbridge, Ohio, October 4, 1914.
9 Cedar Point, Ohio, August, 1913.
cT 9 Sugar Grove, Ohio, July 15, 1915.
Very common in the webs of Frontinella communis in the Sugar
Grove region.
Asagena americana Emerton. (E).

9 Columbus, Ohio, June 13, 1917.
o^ Columbus, Ohio, June 10, 1917.

Ceratinella brunnea Emerton. (E).

9 Columbus, Ohio, June 9, 1916.
Ceratinella emertoni Cambridge. (E).

cf 9 Columbus, Ohio, May 24, 1916.
Ceratinella fissiceps (Cambridge).

9 Sugar Grove, Ohio, June 10, 1917.

Ceratinella minuta Emerton.

9 Flint Ra\-ine six miles north of Columbus, Ohio.

Ceratinopsis alternatus Emerton. (E).

d" 9 Rockbridge, Ohio, June 27, 1917.
Ceratinopsis interpres Cambridge. (E).

cf 9 Rockbridge, Ohio, June 12 and 18, 1915.
Common on the Mountain Laurel and Huckleberry bushes in the
dry upland woods of Hocking and Fairfield Counties.

I

June. 191,S] A List of Ohio Spiders 303

Ceratinopsis nigripalpis Emertt)n. (E).

cf 9 Rockbridge. Ohio, June IS, 191(i.
Sugar Grove. Ohio. July 4. 191.3.
Cornicularia pallida Emerton.

(f Sugar Orove, Ohio, June 10, 1917.

Crustulina guttata (Emerton). (E).

9 Columbus. Ohio, October. 1917.
Found under the edges of logs and stones in low ]jastures.
Enoplognatha marmorata (Emerton). (E).
9 Cedar Ponit. Ohio, July 4, 1914.
9 Columbus. Ohio.
Found in the usual places, under edges of logs, stones and boards
in the edges of woods near the river.

Erigone autumnalis Emerton. (E).

cT Columbus, Ohio. October 10, 1917.

Erigone dentigera Cambridge.

d" Delaware, Ohio, June 28, 1916.
Erigone longipalpis Emerton. (E).

d" 9 Sugar Grove, Ohio, April, 1913. *

Euryopis argentea Emerton. (E).

9 Sugar Grove, Ohio, July 15, 1915.

Euryopis funerea Emerton. (E).

d Rockbridge. Ohio. July 1, 1916.
9 Cedar Point, Ohio, August, 1913.
Very abundant in July on the lower branches of trees on the edges
of a swamp one mile south of Delaware.

Latrodectus mactans (Fabricius). (E).

a" 9 Rockbridge. Ohio, October 4, 1914.
cT N. Kenova, Ohio, August 24, 1914.
9 Ohio. (W. H.).
9 Marietta, Ohio. (W. H.)
Very common on the ground in the upland prairies of Hocking
County.
Lophocarenum arenarium Emerton. (E).

Trans. Conn. Acad. Arts and Sci., Vol. 16, p. 391.
9 Cedar Point.
This specimen was taken under mbbish at the waters edge on
Sandusky Bay, Sandusky, Ohio. It has been found by Mr. Emerton
on the salt marshes near Lynn, Mass., where it lives in quite similar
situations.

Lophocarenum (Diplocephalus) erigonoides Emerton. (E).

cf 9 Columbus, Ohio. October 20, 1917.

(f 9 Buckeye Lake, June 24, 1917.
Specimens have been taken on the ground, in the sweep-net, and
from fences while they were balooning.

Lophocarenum (Hypselistes) florens (Cambridge).
9 August-September, 1917, Wooster, Ohio.
Taken in sweeping in grassland.

304 The Ohio Journal of Science [Vol. XVIII, No. 8,

Pedanostethus riparius Keyserling.

cf 9 Columbus, Ohio, September 22, 1917.

cT 9 Rockbridge, Ohio, June 12, 1915.
Very common in the moist woods which fringe the Olentangy river
where it lives under rocks and boards.

Pedanostethus terrestris Emerton. (E).
Jour. N. Y. Ent. Soc. XXII, p. 2(32.
cf 9 Bainbridge, Ohio, August 17, 1917.
Found under stones and leaves in a very deep moist ravine.

Spintharus flavidus Hentz.

d^ Brinkhaven, Ohio, September 15, 1917.

Spiropalpus (Cornicularia) spiralis Emerton. (E).
cf Columbus, Ohio, June 23, 1916.

Steatoda borealis (Hentz). (E)

Cedar Point, Columbus, Rockbridge, Brinkhaven, Ohio.

(W. H.)
Besides living under bark and boards in most moist situations, this
species makes a very large tangled web about a foot from the ground
in the very thick growth of nettles and other plants in the low moist
woods along the Olentangy River. They hibernate as adults.

Teutana triangulosa (Walckenaer) .

d^ 9 Columbus, Ohio, October 1915.
d^ 9 Lancaster, Ohio. (W. H.)
Rather common in comers and behind cases in the Botany and
Zoology Building, Ohio State University.
Theridium differens Emerton. (E).

d" Rockbridge, Ohio, June IS, 1916.
cT 9 Delaware, Ohio, June 28, 1916.
d" Cedar Point, Ohio, August 1, 1913.
Theridium frondeum Hentz. (E).

9 Cedar Point, Ohio, July 4, 1914.
d" 9 Delaware, Ohio, June 28, 1916.

9 New Lexington, Ohio, (W. H ) \

Widely distributed. Usually found in tall grass and shrubs near
water.
Theridium kentuckyense Keyserling. (E).

d' 9 Rockbridge, Ohio, July IS, 1916.
Found on ferns and other vegetation in the extremely moist dark |

ravines of Hocking County. ■

Theridium murarium Emerton. (E).

9 Columbus, Ohio, June 9, 1916.
Cedar Point, Ohio.

Theridium rupicola Emerton. (E).

d" 9 Rockbridge, Ohio, June, 1914.
9 Sugar Grove, Ohio, July 15, 1915.

Theridium spirale Emerton. (E).

c^ Columbus, Ohio, August, 1914.

i

June, 1918] A List of Ohio Spiders 305

Theridium tepidariorum Koch. (E).

cf 9 Buckeye Lake, Ohio, June 24, 1917.

cf 9 Cedar "Point, Ohio, August G, 1910.

d^ 9 Ohio. (J. H. E.)
These spiders are commonly found in houses and other buildings.
On the cliffs at Clifton Gorge, Bainbridge, Rockbridge and Brinkhaven,
however, they appear to be living in the "wild" condition, not in any
way associated with man.

Theridula sphaerula (Hcntz). (E).

(f Bainbridge, Ohio, August 17, 1917.

Tmeticus (Gonglydium) flaveolens Emerton. (E).
cf Columbus, Ohio, November 10, 1917.
These were taken balooning.

Tmeticus longisetosus Emerton.

d" Columbus, Ohio, March 12, 1918.
Taken near the river under a sheet of tin.

Tmeticus probatus (Cambridge). (E).

o^ 9 Rockbridge, Ohio, July 27, 1917.

Tmeticus terrestris Emerton.

d" 9 Columbus, Ohio, March 12, 1918.

Tmeticus tridentatus Emerton.

cf Rockbridge, Ohio, September 30, 1917.

Tmeticus trilobatus Emerton.

d Columbus, Ohio, March 12, 1918.

Ulesanis americanus Emerton. (E).

c^ Rockbridge, Ohio, June IS, 1916.

LINYPHIID^.

Bathyphantes micaria Emerton. (E).

Bathyphantes concolor (Reuss). (E).

d Columbus, Ohio, Oct. 28, 1917.

Bathyphantes alboventris (Banks). (E).
cf 9 Columbus, Ohio, June 11, 1917.
Hibernates in the adult state.

Bathyphantes nigrina (Emerton). (E).

d Columbus, Ohio, October 20, 1917.
9 Columbus, Ohio, June 11, 1917.
Very common in moist situations near or on the ground under
rubbish or matted vegetation. Some of the males and many of the
females survive the winter.

Drapetisca socialis Emerton.

9 Rockbridge, Ohio, September 30, 1917.
Found running on the bark of beech trees growing with hemlocks
in the moist ravines in Fairfield and Hocking Counties. I have taken
specimens in the same kind of situations at Forest Hills, Mass.

30G The Ohio Journal of Science [Vol. XVIII, No. 8,

Frontinella (Linyphia) communis (Hentz). (E).
(f Sugar Grove, Ohio, July 1."), 1915.
These si^iders build tall rather striking dome webs, sometimes three
feet high in the pine woods west of Sugar Grove in the same situations
in which the Pink Lady's Slipper grows.

Lepthyphantes nebulosa (Sundervall) .

o" 9 Columbus, Ohio, October 28, 1917.
This pair was found cojHilating under a flat stone near the river.

Linyphia (Diplosytla) brevipes (Kevserling).
d" 9 Columbus, Ohio, March 12, 1918.
Found under tin and boards near the river.

Linyphia (Neriene) clathrata Sundervall. (E).
(f 9 Columbus, Ohio, June 13, 1917.
cf Rockbridge, Ohio, June 12, 1915.
9 Buckeye Lake, Ohio, June 24, 1917.
In the woodlot north of the University this species builds its webs
in the angles between the roots of the large trees.

Linyphia mandibulata Emerton. (E).
cf 9 Jefferson, Ohio.

cf Sugar Grove, Ohio, April, 1913.

Linyphia marginata Koch. (E).

cf 9 Rockbridge, Ohio, June, 1914.
d^ 9 Ohio. (W. H.)
Very common in the moist dense woods.

Linyphia phrygiana Koch. (E).

Young, Rockbridge, Ohio, May 27, 1916.
9 Loudonville, Ohio, June 6, 1915.
d" 9 Ohio. (W. H.)
Found only on the undersides of branches in very moist woods,
usually close to streams. This species here does not live in the varied
situations in which it is found in New England. Matures early in May.

Microneta cornupalpis (Cambridge).

cf Columbus, Ohio, November 10, 1917.

Nesticus paUidus Emerton. (E).

cf 9 Columbus, Ohio, June 24, 1917.
Found under sheets of tin and boards near the river.

Stemonyphantes bucculenta (Clerck).

cf 9 Cokmibus, Ohio, March 2(), 1918.

Tapinopa bilineata Banks.

d Rockbridge, Ohio, September 30, 1917.
d Columbus, Ohio, August, 1917 (immature).

MIMETID^.

Mimetus interf actor Hentz. (E).

cf Columbus, Ohio, August, 1917.

cf Sugar Grove, Ohio, July 15, 1915.
Adults and young have been taken several times from the tops of
weeds.

June, 1918] A List oj Ohio Spiders 307

TETRAGNATHID.T..

Eugnatha pallidula Banks.

d" 9 Buckeye Lake. Ohio, July 21, H)17.
Eugnatha straminea (Emerton). (E).

cf 9 Rockbrid^^e, Ohio, June, H)l().
cf Cedar Point, Ohio.
Eugnatha vermiformis (Emerton). (E).

Erie County, Ohio, 1913; vounj<, Buckeve Lake, Ohio,
Sept. 13, 1913.
Pachygantha tristriata Koch. (E).

d" 9 Columbus, Ohio, October 20, 1<»17.
d" Rockbridge, Ohio, Fall, 1912.
Found under logs in grassland and pastures. They hibernate as
adults.
Tetragantha extensa (Linnaeus). (E).

d" Delaware, Ohio. June 18, 1916.
d" 9 Cedar Point. Ohio, August 6, 19 lU.
Tetragantha grallator Hentz. (E).

d" 9 Columbus, Ohio, May 24, 1916.
Rockbridge. Ohio, July 1, 1916.
9 Buckeye Lake, Ohio, June 24, 1917.
cf 9 Cedar Point, Ohio. Ohio. (W. H.)
In central Ohio this species is usually found only near water, very
often on grass which overhangs the water. Near the shore of Lake
Erie, howe\'er, it lives in trees and on buildings often at a considerable
height.

Tetragnatha laboriosa Hentz. (E).
cf 9 Columbus, Ohio. June,
cf 9 Guernsey County. Ohio. June 16, 1916.
cT 9 Ohio. ("W. H.).
cT 9 Rockbridge, Ohio, May 27, 1916.
9 Buckeye Lake, Ohio. Jtme 24, 1917.
This is one of the commonest of the meadow and grassland forms.
Its webs are built high up in the tops of the tall grasses. It matures
early in June.

EPEIRID.E.

Acrosoma reduvianum McCook.

9 Bainbridge, Ohio. August 17. 1917.
9 Rockbridge, Ohio.
Apparently rather rare in Ohio. The webs are slung between trees
or branches four to six feet above the ground in open woods.

Acrosoma gracilis ( Walckenaer) . (E).

cf Rockbridge, Ohio, June 22, 1914.
9 Ohio. (W. H.).
Females abundant in the late summer in dry beech, pine and oak
woods. The webs are usually about breast high and connected to the
supports by very long tough guy-lines.

308 The Ohio Journal of Science [Vol. XVIII, No. 8,

Acrosoma spinea (Hentz). (E).

d" Rockbridge, Ohio, July 1, 1916.
cT 9 Columbus, Ohio, July 7, 1914.
9 Ohio. (W. H.)
On low weeds in open places and in open woods where there is plenty
of light. Wideh' distributed, but never very abundant.

Argiope aurantia Lucas. (E).

One of the most familiar and striking meadow spiders. Universally
distributed. The web is placed low in thick grass or weeds. The
nearly mature female feeds very largely on grasshoppers. The males
mature in July when both sexes are small and inconspicuous. After
mating the female grows to a very large size.

Argiope trifasciata (Forskal). (E).

cf Winterset, Ohio, September 17, 1915.
cf 9 Columbus, Ohio, August-September.
Ohio. (W. H.)
This is fully as common and widely spread as the preceeding, but
is less conspicuous. It lives in the same situations and matures some-
what later.

Cyclosa conica (Pallas). (E).

cf Loudon ville, Ohio, June 6, 1915.

cT Rockbridge, Ohio, May 27, 1916.
Taken in hemlock woods. The webs are usually slung between tree
trunks five or si.K feet above the ground.

Cyclosa turbinata McCook. (E).

c^ Rockbridge, Ohio, July 1, 1916.
9 Ohio. (W. H.)
This species has been found only below cliffs where it builds its
small orb web in weeds in the wettest situations. It seems to prefer
i:)laces where a fine mist falls on it or around it.

Epeira attestor (Petrunkevitch). (E).

9 Rockbridge, Ohio, July IS, 1916.

Epeira angulata Clerck. (E).

This species is closely related to cavatica and is found in similar
situations.

Epeira cavatica (Keyserling). (E).

cf Sugar Grove, Ohio, July 4, 1915.
Rockbridge, Ohio, October, 1913.
E. cavatica in Ohio seems to be limited to the faces of overhanging
cliffs. Mr. Emerton's comments on these specimens are interesting:
"It was first described from Kentucky caves, but here in New England
it is a northern species, living in great abundance on piazzas, in wagon
sheds, and on barns all over Maine, New Hampshire and Vermont,
coming south into northern Massachusetts, but not around Boston or
further south. Westward I have it from Spokane. "
Epeira domicilorum Hentz. (E).

9 Columbus, Ohio, September, 1916.
Rather common on porches and barns in Columbus.

June, 1U18J A List oj Ohio Spiders 309

Epeira displicata llcntz. (E).

d" Rockbridge, Ohio, May 27, 1910.

9 Loudonville, Ohio, June 6, 191.").

cf Marietta, Ohio. (W. H.)
Epeira ectypa Kcvserhng. (E).

9 Rockln-idge, Ohio, October 4, 1914.

Epeira insularis Hentz. (E).

cf Bainbridge, Ohio, August 17, 1917.
Cedar Point, Ohio.
Ohio. (W. H.)
cf 9 Columbus, Ohio, September IS, 1917.

9 Rockbridge, Ohio, June 1914, and d" vSeptember 30, 1917.
A very common woodland form. Found in low bushes in moist
woods though usually in rather light situations.

Epeira juniperi Emerton. (E).

d" 9 Rockbridge, Ohio, June 18, 1916.
Taken while beating branches of hemlock.

Epeira patagiata Clerck. (E).

cf Rockbridge, Ohio, June, 1914.
Erie County, Ohio, 1915.

Epeira placida Hentz. (E).
Epeira prompta Hentz. (E).

Rockbridge, Ohio, May 27, 1916.
c/' Buckeye Lake, Ohio, June 24, 1917.
cf Columbus, Ohio, August, 1914.
9 Cedar Point, Ohio.
Epeira sclopetaria (Clerck). (E). (B).

c^ 9 Cedar Point, Ohio, June 28, 1913.
Very abundant on buildings near Lake Erie.

Epeira strix Hentz. (E).

d" N. Kenova, Ohio, August 24, 1915.
o" Rockbridge, Ohio, July 27, 1917.
d' 9 Buckeye Lake, Ohio, June 24, 1917.

Epeira thaddeus Hentz. (E).

9 Rockbridge, Ohio. October 4, 1914.
9 Columbus, Ohio, August, 1914.

Epeira trifolium Hentz. (E).
Widely distributed.
\"ery common over the state. It seems to prefer tall weeds, par-
ticularlv those which grow on the river flood plains.
Epeira trivittata Keyserling. (E).

cf 9 Delaware, Ohio, June (J, 1916.
cf 9 Buckeve Lake, Ohio, June 24, 1917.
c^ 9 Cedar "Point, Ohio, July 21, _1910.
cf 9 Bainbridge, Ohio, August It, 1917.
This spider is almost always present in tall grass, weeds, reeds and
bushes.

310 The Ohio Journal of Science [Vol. XVIII, No. 8,

Epeira verrucosa Hentz. (E).

9 Bainbrid^e, Ohio, August 17, 1917.
9 Rockbridge, Ohio, August, 1915.
9 Columbus, Ohio, July, 1917.
cf 9 Ohio. (W. H.)
This striking species is rather common in the southern part of the
state. It builds its webs very high up between the trees in the thick
moist woods. The webs sometimes have guy-lines fifteen or twenty
feet long.

Epeira vulgaris Hentz.

9 Brinkhaven, Ohio, September 1."^, 1917.

Glyptocranium cornigerum (Hentz). (E).

9 Columbus, Ohio, November 1, 1915.

Leucauge hortorum (Hentz). (E).

c^ 9 Rockbridge, Ohio, June 19, 1914.
9 Cedar Point, Ohio, June 27, 1913.
This striking green and silver spider is rather strictly limited to
moist woodland. It is widely distributed in the state.
Mangora gibberosa (Hentz). (E).

cf 9 Bainbridge, Ohio, August 17, 1917.
cf 9 Columbus, Ohio, August, 1917.
cf 9 Guernsey Co., 1916.
Erie Co., 1915.
Very common in the tops of grass and grain.

Meta menardi (Latrielle) (E).

Immature d^ 9 Bainbridge, Ohio, August 17, 1917.
A cave species. Found in dark wet situations along cliffs, or in the
entrances of caves.
Metepeira labyrinthea Hentz. (E).

c^ 9 Cedar Point, Ohio, July, 1915.

9 Rockbridge, Ohio, October 4, 1914.
d^ 9 Columbus, Ohio.
Commonly found in the branches of dead trees in the edges of
woods all over the state.
Plectana stellata Walckenaer. (E).

9 Cedar Point, Ohio, August 11, 1910.
9 Guernsey Co., Ohio, 1910.
9 Columbus, Ohio, September, 1910.
9 Rushville, Ohio. (W. H.)

Singa keyserlingi McCook. (E).

9 Cedar Point, Ohio, June 27, 1913, and August 13, 1910.
This species makes a small orb web in the tops of the dune grass
(Andropogon). During the day it stays in the hollow stems of the
dead grass. This form is probably not the same as S. pratensis.
Theridiosoma gemmosum (L. Koch). (E).
cf 9 Sugar Grove, Ohio, July, 1915.
This form may always be found in the wet moss on the faces of
cliffs and in other wet situations in the deep woods.

June, 1918] A List oj Ohio Spiders 311

ULOBORIDyE.

Hyptiotes cavatus (Hcntz.) (E).

9 Rockbiidj^e, Ohio, October 4, H)14.
cf 9 Sugar Grove, Oliio, September 10, 11)17.
Found in dark woods and ravines. Its triangle web is usually
built in branches a few feet above the ground.
Uloborus plumipes Lucas (E).

o^ 9 Sugar Grove, Ohio, July 13, 1915.
cf 9 Columbus. Ohio, June 24, 1916.

THOMISID^.

Coriarachne versicolor Keyserling. (E).
cf 9 Columbus, Ohio, June 13, 1917.
9 Cedar Point, Ohio, July 24, 1918.
9 Buckeye Lake, Ohio, June 24, 1917.
Very abundant on or under the bark of trees.

Misumena celer (Hentz).

9 Marietta, Ohio. (W. H.)
Misumena vatia (Clerck) . (E).

cr^ Rockbridge, Ohio, Mav 27, 1916.

9 Cedar Point, Ohio, August, 1913.

Misumessus asperatus (Hentz). (E).

c^ 9 Columbus, Ohio, May 24, 191G.
Philodromus bidentatus Emerton. (E).

d" Rockbridge, Ohio, June IS, 191G.

d^ 9 Cedar Point, Ohio.

Philodromus canadensis Emerton.

Canadian Entomologist, 1917, p. 270.

cT Cedar Point, Sandusky, Ohio, June, 1915.
Emerton states that this species is common about Montreal and
Ottawa and westward to Lake Nipigon and Prince Albert.

Philodromus macrotarsus Emerton. (E).

Canadian Entomologist, August, 1917, p. 271.
cf Columbus, Ohio, September 1, 1917.

Philodromus minutus Banks.

(^ Buckeye Lake, Ohio, June 24, 1917.

Philodromus ornatus Banks. (E).

9 Cedar Point, Ohio, August, 1913.
Philodromus pictus Emerton. (E).

cT Loudon ville, Ohio, June G, 1915.
Philodromus vulgaris Hentz. (E).

9 Buckeve Lake, Ohio, June 24, 1917.

9 Ohio. ' (W. H.)
Runcinia aleatoria (Hentz). (E).

cT Guernsey Co., Ohio, 191G.

9 Columbus, Ohio, August 23, 1917.

312 The Ohio Journal of Science [Vol. XVIII, No. 8,

Synema parvula Hentz. (E).

Young, Cedar Point, Ohio.
This southern species occurs on the vegetation of the hot, dry-
sand dunes.

Thanatus coloradensis Keyserhng.

Cleveland, Ohio, December 1915.
One specimen was collected by Emerton in the railroad restaurant.
In a note in the Entomological News in 1917, he sums up situation in
regard to this spider in the east as follows: "So it appears that we
have here a western spider that takes readily to life about houses and is
spreading across the country. ' '

Thanatus lycosoides Emerton. (E).

cf Rockbridge, Ohio, October 4, 1914.
9 Columbus, Ohio, March 20, 1918.

Tibellus duttoni (Hentz). (E).

9 Cedar Point, Ohio, 1913.

Tibellus oblongus ( Walckenaer) .
9 Erie Co., Ohio, 1915.

Tmarus caudatus (Hentz). (E).

9 Rockbridge, Ohio, April 10, 1915.

Xysticus fratemus Banks. (B).

cf 9 Columbus, Ohio, June 12, 191G.
Rockbridge, Ohio, June 11, 1916.
This species lives on the ground among the leaves in woodland.

Xysticus gulosus Kevserling. (E).

(f 9 Columbus, Ohio, October 20, 1917.
9 Rockbridge, Ohio, June, 1914.
Very abundant in grassland. In the fall this species with stomach-
osus makes up a large part of the balooning individuals.

Xysticus limbatus Keyserling. (E).

9 Cedar Point, Ohio, August, 1913.

Xysticus nervosus Banks. (E).

d" 9 Columbus, Ohio, October 28, 1917.
cf 9 Rockbridge, Ohio, October, 1913.

Xysticus quadrilineatus Emerton. (E).

9 Rockbridge, Ohio, October, 1913.

Xysticus stomachosus Emerton. (E).

9 Buckeye Lake, Ohio, June 24, 1917.
9 Columbus, Ohio, June, 1917.
cf 9 Marietta, Ohio. '(W. H.)

Xysticus triguttatus Keyserling.

of 9 Guernsey County, Ohio, June IG, 191G.
cf Rockbridge, Ohio, June, 1914.
Common in grassland.

June, 1918] A List of Ohio Spiders 313

PISAURID^.

Dolomedes idoneus IMontj^omcry = vernalis Emerton. (E).
d" 9 RockbridK^e, Ohio, June 12, 1915.

9 Columbus, Ohio, June, 1917.
d" 9 Ohio. (W. H.)
Thi.s large spider is rather common along river courses where it
lives under loose bark and under logs. Where cliffs occur near streams
it becomes a distinct cliff species. Mating occurs early in June. The
males which are smaller than the females, die and the females grow to
be very large by midsummer.

Dolomedes sexpunctatus Hentz. (E).

cf 9 Columbus, Ohio, June 25, 1916.
cT 9 Malta, Ohio, August 12, 1915.
Common aroi:nd permanent ponds.

Dolomedes urinator Hentz. (E).

d" Columbus, Ohio, April 4, 1913.

9 Rockbridge, Ohio, September 18, 1915.

9 New Lexington, Ohio. (W. H.)
These are probably the largest spiders which occur in Ohio. Common
in the ravines of Hocking County, on logs overrunning streams.

Pisaura brevipes Emerton. (E).

Trans. Conn. Acad. Arts and Sci., Vol. 16, p. 400.
9 Sugar Grove, Ohio.
9 Ashtabula Co., Ohio.

Pisaurina undata (Hentz). (E).

9 Delaware, Ohio, June 24, 1916.

Young, Columbus, Ohio, June 10, 1914.
9 Rockbridge, Ohio, July 27, 1917.
Marietta, Ohio. (W.'H.)
The young during the summer are found on vegetation everywhere.
One specimen from a cave was nearly white, but was otherwise normal.

LYCOSID^.

Allocosa rugosa (Keyserling) . (E).

cf 9 Columbus, Ohio, June 12, 1917.
cf 9 Rockbridge, Ohio, June, 1917.
Running on the ground in warm, dry situations.

Lycosa avara Keyserling. (E).

d" 9 Rockbridge, Ohio, October, 1913.
Lycosa baltimoriana (Keyserling). (E).

9 Cedar Point, Ohio, August, 1913.

Lycosa carolinensis Walckenaer. (E).

d" 9 Columbus, Ohio, October 13, 1917.

cT 9 Ohio. (W. H.)
Probably the commonest burrowing spider in Ohio. It makes its
burrows in lawns, pastures, and the edges of fields.

314 The Ohio Journal of Science [Vol. XVIII, No. 8,

Lycosa (Trochosa) cinerea (Fabricius). (E).
d^ 9 Cedar Point, Ohio, July 21, 1910.

9 Marietta, Ohio. (W. H.)
This spider is at its optimum in the hot sands of the lake shore.
Here it burrows into the loose sand, sometimes to a depth of six inches
during the day. At night it emerges and hunts crickets and other
insects.
Lycosa communis Emerton. (E).

cf Buckeve Lake, Ohio, June 24, 1917.

c^ Columbus, Ohio, June 28, 1918.

9 Marietta, Ohio. (W. H.)
Lycosa frondicola Emerton. (E).

9 Rockbridge, Ohio, April 28, 1912.

9 Cedar Point, Ohio, July 21, 1910.

Lycosa kochii Kevserling. (E).

9 Delaware, Ohio, June IS, 1916.

9 Columbus, Ohio, November 10, 1917.

Lycosa lenta Hentz. (E).

9 Delaware, Ohio, June 30, 1917.
d^ 9 Columbus, Ohio.
Lycosa (Geolycosa) missouriensis (Banks). (E).

9 Cedar Point, Ohio, July 20, 1903.
One of the burrowing spiders which lives in sandy regions. Widely
distributed in Eastern U. S.
Lycosa nidicola Emerton. (E).

9 With eggs. Cedar Point, Ohio, July 12, 1903.
Widely distributed. Hibernates as adult. Common.
Lycosa pratensis (Emerton). (E).

9 Cedar Point, Ohio, August, 1913.
Lycosa punctulata Hentz. (E).

9 Rockbridge, Ohio, September, 1913.

cT Rushville, Ohio. (W. H.)

Lycosa scutulata Hentz. (E).

cT 9 Rockbridge, Ohio, September 30, 1917.
d^ 9 Ohio. (W. H.)

c^ N. Kenova, Ohio, August 24, 1915.
Very abundant in the dry upland prairies in the southern part of
the state.
Lycosa tigrina McCook. (E).

d" 9 Rockbridge, Ohio, September 30, 1917.

Pardosa albopatella Emerton. (E).

cf Rockbridge, Ohio, June 18, 1916.
d^ 9 Columbus, Ohio, June 11, 1917.
Abundant in open ground early in June.

Pardosa glacialis (Thorell). (E).

9 Buckeye Lake, Ohio, July 1, 1917.
A distinctly northern species. Found on the sphagnum bog in
the center of Bucke\'e Lake.

June, 1918] A List oj Ohio Spiders 315

Pardosa lapidicina l^nicrlon. (E).

cf 9 Columbus, Ohio, June-July, 1910, Sandusky, Ohio.
Found on hot stones and clay banks near streams all over the state.
Pardosa nigropalpis Emerton. (E).

d" 9 Columlms. Ohio, June 11, 1917.
d" 9 Rockbridge, Ohio, June, 1914.
Ver\- abundant in open ground in June.

Pardosa tachypoda Emerton.

9 Erie Count w Ohio, 1915.

Pirata minuta Emerton. (E).

cf Brinkhaven, Ohio, June 5, 191o.

9 Columbus, Ohio, wSeptember S, 1917.

Pirata montana Emerton. (E).

cf 9 Brinkhaven, Ohio, June (i, 19 Ki.

Pirata sylvestris Emerton. (E).

d" 9 Buckeye Lake, Ohio, July 21, 1917.

Schizogyna bilineata (Emerton).

9 Buckeye Lake, Ohio, June 24, 1917.

Schizogyna ocreata (Hentz). (E).

d" Marietta, Ohio. (W. H.)
cf 9 Columbus, Ohio, June 13, 1917.
cf 9 Rockbridge, Ohio, June, 1914.
Ver\" abundant in fields and open woods.

Schizogyna relucens (Montgomery). (E).

d^ Columbus, Ohio, October 1, 1915.
cf Sugar Grove, Ohio, July 4, 1915.

OXYOPID^.

Oxyopes salticus Hentz. (E).

d" 9 Rockbridge, Ohio, July 27, 1916.
9 Guernsey Co., Ohio, July 16, 1916.
This is a southern species and does not appear to occur north of
Guernsey Co. Found in the upland prairie vegetation.

ATTID.-E.
Attidops youngi (Peckham).

cf 9 Brinkhaven, Ohio, September 15, 1917.
9 Rockbridge, Ohio, September 10, 1917.
Several individuals were found crawling slowly on the face of cliffs
about two miles north of Brinkhaven. One female was taken under
hemlock bark at Rockbridge.
Dendryphantes capitatus (Hentz). (E).

d" 9 Columbus, Ohio, June 12, 1916.
of 9 Buckeye Lake, Ohio, June 24, 1917.
9 Cedar Point, Ohio.
9 Rockbridge, Ohio.
Very common on grass and weeds in fields and at the edge of woods.

316 The Ohio Journal of Science [Vol. XVIII, No. 8,

Dendiyphantes castaneus (Hentz). (E).

9 Buckeye Lake, Ohio, June 24, 1917.
Dendryphantes militaris (Hentz). (E).

d" Rockbridge, Ohio, July 1, 1916.
cf 9 Guernsey Co., Ohio, June 16, 1916.
(^ 9 Bucke^^e Lake, Ohio, vSeptember 13, 1913.
d^ Marietta, Ohio. (W. H.)

Fuentes lineata (C. Koch). (E).

9 Buckeye Lake, Ohio, June 24, 1917.
9 Rockbridge, Ohio, October 4, 1914.
Habrocestum parvulus (Banks). (E).

d" 9 Rockbridge, Ohio, Alay 27, and June IS, 1916.
Habrocestum pulex (Hentz). (E).

d" 9 Cedar Point, Ohio, August, 1913.
Columbus, Ohio, June 9, 1916.
Buckeye Lake, Ohio, June 24, 1917.
Brinkhaven, Ohio, June 6, 1915.

Homalattus cyaneus (Hentz). (E).

c^ Columbus, Ohio, July, 1916.

Hyctia pikei Peckham. (E).
(d 9 Cedar Point, Ohio.
LTsually taken on cat-tails.

Icius hartii Emerton. (E).

9 Buckeye Lake, Ohio, June 24, 1917.

Maevia vittata (Hentz). (E).

cT 9 Columbus, Ohio, May 23, 1916.
d" Rockbridge, Ohio, June 22, 1914.
Widely distributed. Usually found on the ground or on low-
vegetation.

Marpissa binus (Hentz).

d" 9 Gypsum, Ohio, August 25, 1914.

Marpissa undata (DeGeer). (E).

cf 9 Gypsum, Ohio, August 25, 1914.
d^ 9 Ohio. (W. H.)

9 Buckeye Lake, Ohio, June 24, 1917.
On fences and bark of trees.
Myrmarachne albocinctus (C. Koch). (E).
d" Rockbridge, Ohio, June 22, 1914.
Much like an ant in appearance though not usually associated
with ants.

Parnaenus (Phidippus) chryseus Peckham. (E).
cf 9 Sugar Grove, Ohio, July 4, 1915.
9 Columbus, Ohio.
The specimen taken at Columbus was wintering in a curled leaf on a
low branch. A southern species.

Pellenes agilis (Banks). (E).

d^ Cedar Point, Ohio, July 8, 1903.

June. 191SJ A List of Ohio Spiders 317

Pellenes borealis (Banks). (E).

d^ 9 Cedar Point, Ohio, 1913.

d^ Columbus, Ohio, May 24, 1910.

Pellenes coronatus (Hentz). (E).

cf Brinkhaven, Ohio, June 0, 1915.
It is interestini^ to note that this southern species is found in the
same region in wliich occurs the most northern stand in Ohio of the
southern pines, Pinus rigida and Pinus v^irginiana.

Pellenes hoyi Pcckham. (E).

d" 9 Delaware, Ohio, June 28, 191().

Phidippus audax (Hentz). (E).

Common all over the state. Winters half grown and matures in
Mav or June. The three spots on the abdomen of the young are
orange; these become white in the adult.

Phidippus insignarius (C Koch). (E).

cf 9 Rockbridge, Ohio, June 12, 191.3.
Phidippus mccooki (Peckham). (E).

cf 9 Guernsey Co., Ohio, September 2, 191().

Phidippus multiformis Emerton. (E).

W^ry common (j\-er the state. It is found in grassland and in open
woods. Matures in July and August.
Phidippus mystaceus (Hentz).

d" 9 Ohio. (W. H.)
Phidippus putnami (Peckham).

d" Columbus. Ohio, August 20, 1917.
Urbana, Ohio. (P.)
Phidippus whitman! Emerton. (E).

d' Columbus, Ohio, June 1, 1914.
cf Wooster, Ohio, June 0, 1915.

Phlegra leopardus (Hentz).

cT 9 Marietta, Ohio. (W. H.)

Salticus senicus (Clerck). (E).

Common on buildings and fences ever\'where. Matures early in
June.

Sittacus palustris (Peckham).

d" Flint, Ohio, (North of Columbus), Julv 4. 1917.

Synemosyna picata Hentz. (E).

cf Columbus, Ohio, June 24, 191(5.

Synemosyna formica Hentz. (E).

cT 9 Brinkliaven, Ohio, June (>, 1915.
d" Rockbridge, Ohio, July 1, 1916.

Synemosyna scorpiona Hentz. (E).

9 Buckeve Lake, Ohio, June 24, 1917.
Marietta, Ohio. (W. H.) Ohio. (P.)

318 The Ohio Journal of Science [Vol. XVIII, No. 8,

Thiodina puerpera (Hentz). (E).

cf 9 Rockbridge, Ohio, June 12, 1915, and June IS, 1916.
A distinctly southern form. Taken in beating trees.

Tutelina elegans (Hentz). (E).

d" 9 Delaware, Ohio, June 28, 191(3.
c^ 9 Cedar Point, Ohio, June 19, 1913.

Tutelina (Icius) formicaria (Emerton). (E).
cf Sugar Grove, Ohio, July 15, 1915.
9 Rockbridge, Ohio, September 30, 1917.
A very striking spider. One from the face of a cliff.

Wala mitrata (Hentz). (E).

cT Rockbridge, Ohio, July IS, 1916.

Wala palmarum (Hentz). (E).

d^ 9 Buckeye Lake, Ohio, September 13, 1913.
cf 9 Brinkhaven, Ohio, June 6, 1915.

Zygoballus bettini Peckham. (E).

d^ 9 Columbus, Ohio, Mav 24, 1916.

Zygoballus sexpunctatus (?) (Hentz). (E).
d" 9 1916.

a

(

I

DESCRIPTIONS OF SEVEN SPECIES OF ASILUS.

(Family Asilidae).

Jas. S. Hine.

This genus was considered in Volume II, pages 13()-172 of
the Annals of the Entomological Society of America, and
Banks added one additional species and gave a key to a section
of the genus in Psyche, Volume XXI, page 131. Since pub-
lishing on these robberfiies I have accumulated much material
and from this the following additional species are described at
this time.

Asilus gilvipes n. sp.

Large reddish species, legs all red with black bristles, posterior
margin of the scutellum with four bristles, wings hyaline, oviduct
conical, male forceps short. Total length 20 mm.

Face and front yellow pollinose, mystax composed mostly of yellow
bristles, but with a few black ones above, beard straw yellow, antennae
palpi and proboscis black; thorax yellow pollinose, bristles of the dorsum
black, wings hyaline, legs in all their parts red with black bristles;
abdomen yellow pollinose, middle parts of the segments darker, giving
a banded appearance, bristles mostly yellow; ovipositor short and
distinctly conical, male forceps about as long as the seventh abdominal
segment.

Holotype female from Saguache, Colorado, collected July 16,
1911, by A. K. Fisher, in the author's collection. An additional
male and female with the same data, and an injured male from
New Mexico belonging to the American Museum of Natural
History.

Asilus comosus n. sp.

A rather large robust species with a hairy body, large mystax and
brownish wings. Coxse and basal half or more of the femora black,
remainder of the legs red. Length of the female IS mm.

IMystax very large, long and black, reaching nearly to the base of the
antennae, beard pale yellowish, postorbital bristles numerous and black.
Thorax with numerous long black bristles dorsally, scutellum distinctly
hairv above and with several long black bristles on the posterior margin.
Wings plainly tinged with brown, nearly uniform colored all over,
legs black to' beyond the middle of the femora, otherwise red, with
black bristles and pale hair. Abdomen dark, hind borders of the

319

320 The Ohio Journal of Science [Vol. XVIII, No. 8,

segments dorsally pale, bristles and hair ]3ale; transverse rows of numer-
ous bristles before the incisures; female genitalia black, rather short,
compressed.

Holotype female collected by W. M. Wheeler in Monterey
County, California, July 5, 1896. Property of the American
Museum of Natural History. Two other females with the
same data.

This species is very distinct from all others known to me,
and for that reason I allow myself to describe it from the
female only. It seems to fall in the subgenus Antipalus, not
heretofore reported from America.

Asilus vescus n. sp.

Small black species with the sides of the thorax and posterior margins
of the abdominal segments gray pollinose. Male genitalia dark reddish
brown. Total length 10 mm.

Mystax black above, white beneath; antennas and proboscis black,
beard white, thorax with two black stripes above very narrowly sepa-
rated with a gray line, sides of the thorax gray pollinose, scutellum
with some very fine gray hairs dorsally and two quite prominent black
bristles on the posterior margin; wings hyaline, slightly gray on the
posterior margin and at the apex; legs with femora wholly black, tibiae
and tarsi dark reddish. Abdomen black, venter and posterior margins
of the segments dorsally gray pollinose. Male genitalia dark reddish
brown, but not contrasting strongly in color with the abdominal seg-
ments, cut out at the apex above and produced backward and inward
at the apex below, so that a small, but distinct, open space may be seen
enclosed from dorsal view with a lens. Female genitalia shining black.

Holotype male from Monterey County, California, collected
July 2, 189G, by W. M. Wheeler, property of the American
Museum of Natural History. Several other males and females
taken at the same time by the same collector.

Asilus delicatulus n. sp.

Body gray with pale bristles, wings hyaline, legs red, scutellum
distinctly hairy with a row of white bristles on the posterior margin.
Total length 1 1 mm.

This is a very distinct species and suggests at once its sand-
inhabiting habits. It appears to be related to A. leucopogon,
although the female ovipositor does not have a circuit of spines
at the tip as in that species. The male has the last two abdomi-
nal segments and the genitalia bright yellowish red and the
female has the ovipositor of the same color. The delicate

June, 1918] Seven Species of Asil us 321

appearance, the uniform gray color of the body, white bristles
everywhere and the row of numerous white bristles on the margin
of the scutellum are amply sufficient to identify the species.

Holotype male from Las Cruces, New Mexico, collected
by J. M. Aldrich June 1(), 1917 and deposited in his collection.
He procured two males and four females from the same place
on the same date.

Asilus formosus n. sp.

Body yellowish gray ijollinose all over, legs all red except each
femur has a large darker spot on its anterior surface, wings clear hyaline,
total length IS mm.

Mystax pale yellowish, antenna:, palpi and proboscis black, beard
white, "postorbital bristles white, hair and bristles of the thoracic dorsum
black, posterior margin of the scutellum with two black bristles, legs
red with black bristles, each femur with a darkened patch on its anterior
surface. Wings clear hyaline, bristles of the abdomen pale.

Holotype male from Clary County, Kansas, August 29,
1911, and another male from Stanton County, Kansas, July 29,
1911, both collected by F. X. Williams; in the author's collection.
One male from Garden City, Kansas, August, 1895, H. W.
Menke, collector.

Asilus citus n. sp.

Body dark, abdomen with alternate dark and gray bands, thorax
thinly gray pollinose, middle of the dorsum with two black stripes
very narrowly separated; femora, except base and apex, black; other-
wise legs red with black bristles; genitalia red. Total length 12 mm.

Mystax white with a few black bristles above, beard white, post-
orbital bristles largely white, a few black ones near the vertex. Bristles
of the thoracic dorsum black, scutellum with two black bristles on its
posterior margin, wings hyaline, apex and posterior margin faintly
gray.

Holotype male from Santa Rita Mountains, Arizona, taken
in July; F. H. Snow; in the author's collection. Fifteen males
and females from the same locality.

The banded abdomen and the dark red genitalia of both
sexes, together with the small size suggest this species.

Asilus compositus n. sp.

Body slender, dark brown in color, legs yellowish brown, antennal
bristles about twice as long as the third segment which bears it. This
species is related to gracilis, which has the long antennal bristle also.
Total length 17 mm.

322 The Ohio Journal of Science [Vol. XVIII, No. 8,

I

Mystax white, very small, leaving a long distance between it and
the base of the antennae, beard white and sparse, posterior orbital
bristles all ]3ale; thorax lightly gray pollinose with black bristles dorsally,
two black bristles on the posterior margin of the scutellum ; legs yellowish
brown, darker in places on the femora, bristles of the legs largely pale, i

but some black ones on the tibiae and tarsi; wing hyaline, feintly gray
on the hind margin and at the apex; abdomen dark with pale bristles.
Genitalia dark brown, shining in both sexes.

Holotype male from San Diego, California, June 30, 1913, 1

collected by E. P. Van Duzee; in the author's collection; a
female with the same data, a male from Los Angeles County,
California, collected by Coquillett, a male and female from
Claremont, collected by Baker, and a male and female from
Kerryville, Texas, collected by F. C. Pratt.

This species differs from A. gracilis in the much darker color
of the whole body. The legs are brown instead of yellow and
the mystax is white and not straw colored.

THE NORTH AMERICAN SPECIES OF TELEONEMIA
OCCURRING NORTH OF MEXICO.

Bv Caul J. Dkakk,
Xrw York State College of Forestry.

The genus Teleonemia of Costa belongs to that group in the
family Tingida? (^Hemiptera-Heteroptera) which have the
nervures or veins of the delicate lace-like structure much
thickened and the peculiar lacy appearance which is so char-
acteristic of the family is somewhat obscured. In the culmina-
tion of species the genus, no doubt, attains its maximum in
the warmer climates, both tropical and subtropical America
being represented by a number of closely allied forms; in fact
it seems to be the genus which is most characteristic of the
family Tingidae of tropical America. The most northern
localities represented by specimens before me are Utah, Kansas,
Missouri and both North and South Carolina. Stal (1873,
p. 131) enumerates eleven species, two from North America
and nine from South America. Two species are listed by
Uhler (1886, p. 22) for North America, elongata being a manu-
script name. Champion (1898a, p. 34) gives fourteen species
for Mexico and Central America, twelve being described as
new. Both Banks (1910, p. 57) and Van Duzee (1917a, p. 26
and 1917b,, p. 221) catalogue five species that have been taken
north of Mexico. The present paper includes eight North
American species that occur north of Mexico, schwarzi, consors
and barberi being described herein as new.

Material Examined: The author is very much indebted to
Professors Lovett, Ayers, Dean, Osborn, Ferris, Van Duzee,
Johannsen and Knight for the loan or presentation of specimens
of Teleonemia. Many thanks are also due to Mr. Edmund
Gibson who has very kindly permitted the writer to study
the determined species as well as the undetermined specimens in
the National Museum.

According to Champion (1898a, p. 34) and Van Duzee
('1917b, p. 221) the bibliography and synonomy of the genus
Teleonemia may be stated as follows:

323

324 The Ohio Journal of Science [Vol. XVIII, No. 8,

Genus Teleonemia Costa.

Type fimera Costa.

Costa, Annuario del Museo Zoologico della R. Universita di Napoli, II, p. 144,

1864.
Stal, Enumeratio Hemipterorum, III, pp. 122 and 131, 1873.
Champion, Biologia Centrali- Americana, Rhynchota. II, p. 34, 1898.
Champion, Transactions of the Entomological Society of London for 1898, p. 61.
Distant, Fauna of British India, Rhynchota, II, p. 142, 1903 (names /«» era Costa

type).
Van Duzee, Catalogue of the Hemiptera of America North of Mexico, p. 221, 1917.
Americia Stal, Enumeratio Hemipterorum, III, p. 131, 1873 (subgenus of

Tingis; type, albilatera Stal.
Amanrosterphus Stal, Enumeratio Hemipterorum, III, p. 131, 1873 (subgenus

of Teleonemia: haplotype, morio Stal.
*Lasiacantha Lcthierrv et Severin, Catalogue General des Hemipteras, III,
p. 18, 1894 (in part).

Narrow and elongate in shape. Head usually with five spines, the
spines arranged as in related genera; bucculae closed in front; antenni-
ferous tubercles somewhat prolonged, obtuse. Antennae rugulose,
distinctly or indistinctly pilose, contiguous at the base; first and second
segments short; third segment long, cylindrical or subcylindrical,
obliquely truncate at the tip; fourth segment moderately long, more or
less lanceolate or ovate. The length of the antennae varies in different
species and sometimes a little in the same species. Rostral groove
uninterrupted, the rostrum varying in length in different species.
Pronotum tricarinate; paranota narrow, uniseriate (except in schwarzi),
folded back against or nearly against the pronotum proper; in schwarzi
the paranota are almost wanting, barely more than carinate and with
only two or three exceedingly narrow areolae near the anterior margin.
Metastemal orifices distinct. Hood indistinct or entirely wanting,
but quite large in some Mexican and Central American species. Elytra
extending considerably beyond the apex of the abdomen; discoidal and
sutural areas co-elevated; costal and subcostal areas varying in width
and number of rows of areolae in different species. Wings present.

Key to the American Species of Teleonemia Occurrinx. North of Mexico.

1 — Subcostal area almost entirely biseriate 2.

— Subcostal area uniseriate throughout 6.

2 — Rostrum very long, extending beyond the apex of the rostral sulcus 4.

■ — Rostrum much shorter, extending to or very slightly beyond the meso-meta-

sternal suture 3.

3 — Antennas varying slightly in length, usually barely reaching the base of elytra,
the third segment less than twice the length of the fourth; paranota with

distinct areolse T. nigrina Champion

— Antennae barely reaching the base of the elytra, the third segment slightly
more than twice the length of the fourth; paranota almost wanting, scarcely

more than carinate T. schwarzi sp. new.

4 — Insects 4 mm. or more in length; antennee moderately stout, very slightly pilose;

lateral carinas diverging posteriorly 5.

— Smaller insects, about 3.8 mm. long; antennas rather slender, distinctly and
quite densely pilose; lateral carinas parallel T. consors sp. new.

* Name cited in error.

June, \\)\S\ Teleonemia North of Mexico 325

■i-Costal area uniseriate throughout or uniseriate to beyond the middle and

ime ularh biseriate for a short distance towards the apex; general color

St^:cous or brown.sh testaceous, the elytra -ottled^ w.th ..^scou^s ^or

-CosTalareauniseriate' throughout; general color dark brown or fuscous-brown;
thTrd antennal segment becoming slightly smaller distally and^about three

e-DlSoidal^rea^Jci^ ^HbcSj^costal area wUh" rather large areolae; antenn.
^'rSch,ng slightly beyond the base of the elytra, ^'-^^ly^pdos.^^^^^^ ^^^^_

-DiscoidaVarea glabrous'; costal area with small or very small areolae; antennae
reaching bcvond the base of the elytra, mdistmctly pi use . ■■■,■■■'

-GSa\^ot;whiS tc^aceous; sometimes sliglnly mottle., with bro»^;
costal area with larger areola:; anterior margra of the pronotum neariy tn-
angSflar the me<l,a„' earina distmetly elevated anter.orU. anu™a:^rafter
slender

Teleonemia nigrina Champion.

Teleonemia nigrina Champion, Biologia CentraU-Americana,
Rhvnchota, Hemiptera-Heteroptera, Vol. II, P- 41, Tab^III,
Figs. 13, 13a and 13b. Monanthia nigrina Uhler MS. (Texas
specimen in the British Museum).

Form moderately elongate, narrow; head with rather short, W^nt and
more or less prominent median spine. Antenna) rather short, shghtly
variable in length, usually barely reaching the base of the elytra the
third segment less than twice the length of the fourth. Rostrum extend-
inc^ to the meso- metastemal suture, the rostral groove abruptly and
gr?atlv widened on the metastemtmi. General color above grayish or
brownish grav, the elytra mottled with black, the body beneath black
or piceous. Pronotum rather finely punctate, subtruncate m front,
rather sharplv tricatinate, each carina wi^th a row of small areoke^
Elvtra moderately long, slightly constricted beyond the middle, costal
area uniserate, the areolae long, narrow and separated by strong trans-
verse nervures; subcostal area biseriate. In typical specimens the
terminal ventral segment in the female is armed with a long, stout^
obliquely projecting tubercle on each side. Length o-3.b mm., width
1-1.5 mm.

Mexico, Guatemala and Texas (Champion, 1898a, p. 41);
New Mexico (Uhler, 1904, p. 362) ; Texas (Barber, 190G, p. 281) ;
California (Van Duzee, 1914, p. 11). I have before me over
250 specimens of nigrina. representing a great number ot
different localities in Texas, New Mexico, Arizona and Cali-
fornia, also specimens from Kansas, Utah, Missouri, Arkansas,
Georgia, North Carolina and South Carolina.

326 The Ohio Journal of Science [Vol. XVIII, No. 8,

Van Duzee (1914, p. 11, and 1917c, p. 261) states that
nigrina is "a common species everywhere in Cahfornia" and
has taken it in vSan Diego County feeding upon Rhus laurina
and a leguminose plant allied to alfalfa. Specimens are at hand
that bear the food plant labels as follows : Sugar beets (Spreck-
les, California, Sept. 20, 1904, collected by E. S. G. Titus) ;
Adenostegia iCordylanthiis) filafoUa (Campa, California, July
29, 1917, by W. D. Pierce); Adenostegia pilosa (Permanente
Cr., Santa Clara County, California, August 12, 1917, by
G. F. Ferris, who states "It occurs in great numbers on both the
leaves and flowers of its host plants"); Sphaeralcea aiigustifolia
(Rio Grande, Brewster County, Texas, by Mitchell and Cush-
man); Verbena (Marion County, Arkansas, July 15, by Mr.
McElfresh, also Riley County, Kansas, September and October,
by Mr. J. B. Norton); Ilelenium tennifolium (Wolf Creek|
Texas, August 5, 1906, by Dr. F. C. Bishop).

I have seen specimens of nigrina Champion labeled " Teleo-
nemia elongata Uhler MS.," and this is probably the species
listed by Uhler (1886, p. 22) and Smith (1909, p. 149) and
Van Duzee (1917b, p. 222). The antenna? are slightly variable
in length, but the third segment is always less than twice the
length of the fourth. Van Duzee (1917c, p. 261) states, "Some
of the females taken in company with the typical form have
mere tubercles on the last ventral segment in place of the
prominent horn-hke processes figured by Champion." The
tubercles of genital segment vary in size in the same specimen
and sometimes they are entirely wanting.

Teleonemia schwarzi sp. new.

Moderately elongate, rather small; elytra testaceous, the ner\-ures
vanegated with brownish fuscous. Head black, armed with short slender
spines, the two frontal spines converging and the median spine very
short or almost entirely wanting. Eyes black. Pronotum coarsely
punctate, black or reddish black, the pubescence whitish, the surface
of the disc shining; tricarinate, each carina very low and thin, the
niedian carina with barely distinct areolae, the lateral caiina? slightly
diverging anteriorly and without distinct areolae; paranota almost
wantmg, barely carinate. In some specimens the pubescence on
the pronotum seems to be almost entirely rubbed off and disc is
shming and appears much like the dorsal surface of the bodv in the
genus Alveolingis O. & D. Rostral groove much widened" on the
metastemum, the rostrum extending to the meso- metastemal suture.
Body beneath black or reddish black, the. pubescence rather sparse and

June, lUlS] Teleonemia North of Mexico 327

whitish Rostral lamina^ ^•cllo^vish brown. Legs reddish black or
Teddsh brown, the distal portion of tibia, somewhat testaceous; apex
of tibia and tarsi blackish. Length, 2.(J4 mm.; width, .SO mm.

Antenna barelv reaching the base of the elytra, moderately strong,
densely and rather longlv pilose, dark iDrown; first and second scKm en ts
very short nearly equal in length; third segment a httle more than twice
The leu °[h of the fourth; fourth segment subequal to the Icnigth ci he
first aud second conioined. Elytra slightly narrowed beyond he
middle rounded at the apices; costal area narrow, umscnate, the
areoke long narrow and separated by strong transverse ncrxures;
subefstal a?ea almost entireh' biseriate. Wings a httle longer than the
abdomen.

Several specimens: San Diego (April, Coquillett) and
Palm Springs (January 18 and March 9, Hubbard) California;
Santa Rita Mountains, Arizona, collected by Dr. Schwarz.
The species is named in honor of Dr. E. A. Schwarz of the
National Museum, who has collected a great number of speci-
mens of Teleonemia and other Tingidce. Type m the Nationa
Museum, Washington, D. C. Pamtypes in the National
Museum, the California Academy of Science, Cornell Uni-
versity and the author's private collection.

Teleonemia consors sp. new.

Form similar to variegata Champion, but somewhat smaller. A^tenn^
rather slender, quite densely and distinctly pilose; first segment shghtl>
\ol7ev than the second; third segment a little more than three times the
len^h of the fourth. Pronotum nearly truncate m front, sparse y
pubescent, distinctly tricarinate, the lateral carina parallel; paranota
Ced back against the pronotum proper. Rostra g--e f ad^^^^^^
widening posteriorly, the rostrum extending beyond the first ^entra
^ture. General color fuscous or dark testaceous, the nervures partially
darkened, the areola more or less grayish; body beneath Wmh
fuscous or blackish, the pubescence sparse and grayish. Legs dark
fuscous or reddish fuscous with the tarsi darker. Rostral lamm^
yellowish brown. Elytra long, somewhat oval m shape; costal area
uniseriate, the areote rather large and mostly transparent ,subcotal
area biseriate; discoidal area large, widely reticulated. The el>tra
except sutural area, are sparsely pubescent Wings a little longer
than the abdomen. Length, 3.5-3.6 mm.; width, l.o2 mm.

Six specimens; Bonita, Post Cr. Can., Arizona, July 16,
1917 by Mr. H. H. Knight; Oracle, Arizona, July. Type m
author's collection. Paratypes in collection of H. H Knight,
Cornell University (Heidemann Collection) and author s col-
lection.

^^^ The Ohio Journal of Science [Vol. XVIII, No. S,

Somewhat allied to variegata Champion and pilicornis
Champion, but readily separated from the former by characters
given m the key and from later by its shape, much longer
rostrum, etc.

Teleonemia barberi sp. new.

Closely allied to variegata Champion, but readily separated
from It by the much darker color, the uniseriate costal area, the
much slender spines on the head and the shorter third anteknal
segment.

Elongate, rather broad, slightly smaller than variegata Champ
general color dark fuscous or dark brownish testaceous and slightly
mottled with blackish. Pronotum subtruncate in front, very coarsely
punctate, distinctly tricarins, the outer carinje divaricating posteriorly
the cannae without distinct areolae. Rostrum extending to the posterior
margin of the first ventral segment. Antennae stout, slightly pilose
rather long; third segment subcyhndrical, almost three times the
length of the fourth. Elytra similar in shape to variegata, the subcostal
area bisenate, the costal area rather broad and uniseriate. Length
3.9-4 mm.; width, 1.1-1.25 mm. '

Numerous specimens, collected on ' ' desert willow, ' ' Chilopsis
m the Huachuca Mts., Arizona, July 23, by Prof. H. G. Barber!
I have also a single specimen from Valentine, Texas, July 8,
1917, that was collected by Dr. J. Bequaert.

Type in the collection of H. G. Barber. Paratypes in the
collections of Cornell University, H. G. Barber and the author.

Teleonemia variegata Champion.

Teleonemia variegata Champion, Biologia Centrali- Ameri-
cana, Rhynchota, Hemiptera-Heteroptera, Vol. II, p. 42
Tab. Ill, Figs. 15 and 15a, 1898.

^'''^^^}^' -^^^^^l ^'■°'''^' ^^^^ ^^^S^s^ species in the genus occurring
north of Mexico. General color testaceous or brownish testaceous the
elytra mottled with fuscous or black; body beneath reddish brown or
tuscous. Head with moderately slender or quite stout spines the
spmes varying slightly in length in different specimens, the frontal
spme either porrect or adpressed against the head. Pronotum coarsely
punctate, tricannate, the carinas without distinct areola, the median
carina not raised antenorly; paranota turned back against the pronotum
proper. Antenna moderately stout, slighth- pilose, long, the third
segment from three to three and a half times the length of the fourth
Kostral groove gradually widening posteriorly, the rostrum extendina
to or a little beyond the second ventral suture. Elvtra long, somewhat

June, 1918] Teleonemia North of Mexico 329

ON-al in shape; eoslal area ])rominent, entirely uniseriate or uniseriate

to bcvond the middle and irreKmlarly biseriate for a short distance

towards the apex, the areoke lar-e. Legs sometimes marked or slightly

annulated with fuscous, the tarsi black; apical segment of the antenna

partly or entirely fuscous. Length, 4.1-4.5 mm.; width, l.S-l.SS mm.

Barber (1910, p. 38) first records this species north of

Mexico from specimens taken in the Huachuca Mts., Arizona.

Champion (1898a, p. 42) describes the species from specimens

taken in Mexico (Omilteme in Guerrero, 8,000 feet) and

Guatemala (Capetillo, 4,000 feet). I have examined specimens

from Prescott and Huachuca Mts. (Barber, collector), Arizona

and Cordoba, Mexico (Fred Knab, collector).

Teleonemia scrupulosa Stal.

Teleonemia scrupulosa Stal, Enumeratio Hemipterorum,
Band, III, p. 132, 1873. Teleonemia scrupulosa Champion,
Biologia c'entrali- Americana, Rhynchota, Vol. II, p. 40, Tab.
Ill, Figs. 12 and 12a, 1898.

Moderately elongate, rather narrow. General color dark gray or
brownish the elvtra with darker markings, the pubescence whitish.
Antennce'stout, moderately long, distinctly pilose, the third segment a
little less than three times the length of the fourth. Rostrum extendmg
to the meso- metastemal suture, the rostral groove gradually ^vldenlng
behind the anterior coxee. Pronotum with the canuce moderately
elevated and uniseriate, the paranota uniseriate, not qmte turned
back against the dorsal surface of the pronotum. Elytra constricted
back of the middle and widening towards the apices, with a transverse
fascia before the apex; costal area moderately broad, the areolae rather
lar-e and hyaline; discoidal area finely pubescent, the outer margin
curved or sometimes nearly straight ; subcostal area uniseriate. Length,
3.25-3.9 mm.; width, 1.1-1.3 mm.

Champion (1898a, p. 41) reports this species as the most
common Teleonemia occurring in Mexico and Central America.
Numerous records have been published for the West Indian
Islands and the tropical and subtropical regions of both North
and South America by Stal (1873, p. 132), Champion (1898a,
p 40) and Van Duzee (1907, p. 22). Barber records the species
for Florida (1914, p. 507) and Texas (190G, p. 281). Seventy-six
specimens are at hand, representing the following localities:
Grenada, Jamaica, Guatemala, Mexico, Hayti, Florida and
many records from Texas.

Scrupnlosa has been taken on mint (Prof. Avers) and poppy
mallow, CalUrhoc involucrata. (Mr. J. D. Mitchell) m Texas.

330 The Ohio Journal of Science [Vol. XVIII, No. 8,

As stated by Champion (1898a, p. 41) the shortly pilose antenna
and pubescent discoidal area are the principal characters of the
species.

Teleonemia sacchari Fabricius.

Acantkia sacchari Fabricius, Entomologia Systematica,
Tom. IV, p. 77, 1794 ("Habitat in Americae meridionalis
InsuHs"). Tingis sacchari Fabricius, Systema Rhyngotorum,
p. 126, 1803. Alonanthia sacchari Herrich-Schaffer, Die Wan-
zenartigen Insecten, p. 85, V, Tab. CLXXIII, Fig. 533, 1839.
Alonanthia sacchari Fieber, Entomologische Monographein,
p. 70, Tab. VI, Figs. 22-25, 1844.

Antennae slender, indistinctly pilose, brownish, the apical segment
sometimes partially or entirely dark brown or fuscous; third segment a
little less than three times the length of the fourth; fourth segment
slightly longer than the first and second taken together. Head reddish
brown, the median spine semi-erect and the dorsal spines a little longer
than in belfragei. General color brown , the elytra more or less variegated
with fuscous, the body beneath reddish brown. Pronotum sparsely
pubescent, brown or reddish brown, the carina; uniseriate, a little
thicker and not so highly elevated as in belfragei. Median carina
much less elevated anteriorly and the anterior margin of pronotum
more rounded than in belfragei. Legs pale brown, the tips of tibiae
and tarsi black. Elytra with the costal area narrow, the areola long
and very narrow; subcostal area biseriate; discoidal area glabrous.
Length, 3.7-3.85 mm.; width, 1-1.2 mm.

Brazil and Mexico (Fieber, 1844, p. 76, and Herrich-
Schaffer, 1839, p. 85); Cuba and Island of St. Bartholomew
(Stal, 1873, p. 132); Lower California? (Uhler, 1894a, p. 278);
Jamaica (Van Duzee, 1907, p. 22); Balthazar, Is. St. Vincent
(Uhler, 1894b, p. 202-203, in part); Biscayne Bay and Key
West, Florida (Barber, 1914, p. 507). Of this species I have
examined tw^enty-nine specimens, representing the following
localities: Island of St. Vincent, Grenada, Cuba, Jamaica
(Mandeville, Van Duzee coll.) and Florida (Key West, Jan. 30,
1869, L. Worth, Feb. 6, 1887, and Biscayne, Sept. 20, 1889).

The general color, length of the antennas and width of the
costal area is somewhat variable in this species. In a series of
specimens from Grenada the third segment of the antennae
varied .28 mm. in length. In most specimens the areolae in
the costal area are very narrow.

June. 1918] Tclconemia North of Mexico 331

Teleonemia belfragei Stal.

Teleonemia belfragei Stal, Enumeratio Hemiptcrorum, Band.
III. p. 132, 1.S73. Champion, Transactions of the Entomologi-
cal Society of London for 1898, p. 02, PI. Ill, Fig. 8.

General color whitish Icstaceotis; head, pnjnolum anteriorly and
body beneath slighth- tinged with ferruginous; elytra usually with a
few brown or fuscous s])ots on the nervurcs. Antenna? slender, indis-
tinetl>- pilose, whilish testaceous, the apical segment brown or fuscous;
first and second segments subequal; third segment nearly three times
the length of the fourth; fourth segment subequal to the length of the
first and second taken together. Pronotum sparsely pubescent,
tricarinate, each carina uniseriate and strongly elevated; median carina
elevated anteriorly, the anterior margin of the j^ronotum nearh- tri-
angular and projecting slightly o\'cr the base of the head. Head
sparsely pubescent, the median spine suberect. Legs mostly pale
testaceous, the tarsi black. Elytra with the costal area moderately
broad, uniseriate, the areola; hyaline, a little longer than broad, the
transverse nervurcs strong, brown and usually four or five of them black.
Rostrum extending slightly beyond the meso-metasternal suture.
Length. 3.4-3.55 mm.; width, 1.1-1.25 mm.

Texas (type locality, Stal, 1873, p. 132). Florida: Crescent
City, Sanford and Biscayne (Van Duzee, 1909, pp. 173-174);
Crescent City, Biscayne Bay and Lakewood (Barber, 1914, p.
507). I have examined seven specimens from Jacksonville
(Ashmead) and Crescent City, (Heidemann and Van Duzee)
Florida.

The slender antennae, the elevated median carina anteriorly,
lighter color, etc., readily separate this species from T. sacchari
Fabr. Numerous specimens taken at Gainesville, Fla., May
1918, have the elytra slightly mottled with brown. This is a
very common species in Southeastern United States and feeds
on the French mulberry, Callicarpa Americana in Florida.

332 The Ohio Journal of Science [Vol. XVIII, No. 8,

LITERATURE CITED.
Banks, Nathan.

1!I10. Catalogue of the Nearctic Hemiptera-Heteroptera.
Barber, Harry G.

1906. Hemiptera from Southwestern Texas. Mu.seum of the Brooklyn

Institute of Arts and Sciences. Science Bulletin, Vol. I, No. 9,

pp. 255-289.
1910. Some Hemiptera Heteroptera New to the Fauna of United States.

Journal of the New York Entomological Society, Vol. XVIII, pp.

34-39.
1914. Insects of Florida. II. Hemiptera. Bulletin of the American Museum

of Natural History, Vol. XXXIII, Art. XXXI, pp. 495-535.
Champion, George C.

lS9Sa. Biologia Centrali-Americana, Rhynchota, Hemiptera-Heteroptera,

Vol. II, pp. 1-48.
189Sb. Notes on American and other Tingidae, with descriptions of two New

Genera and four new Species. Transactions of the Entomological

Society of London, pp. 55-64, pis. II and III.
Distant, William L.

1903. Fauna of British India. Rhynchota, Vol. II.
Fabricius, Johann C.

1794. Entomologia Systematica, Tom. IV.

1803. vSystema Rhyngotorum.
Fieber, Franz Xavier.

1844. Entomologische Monographein.
Herrich-Schaffer, Gottlieb, A. W.

1839. Die Wanzenartigen Insecten, V.
Lethierry, L. et Severin, G.

1894. Catalogue General des Hemipteres, Tom. Ill, Heteropteres.
Smith, John B.

1910. Catalogue of the Insects found in New Jersey, 3rd Edition.
Stal, Carl.

1868. Hemiptera Fabriciana. ,

1873. Enumeratio Hemipterorum, Band. III.
Uhler, Philip Reese.

1886. Check List of the Hemiptera-Heteroptera of North America.

1894a. Hemiptera of Lower California. Proceedings of the California
Academy of Science, Vol. IV, pp. 223-295.

1894b. On the Hemiptera-Heteroptera of the Island of Grenada, West Indies.
Proceedings of the Entomological Society of London, pp. 167-224.

1904. List of the Hemiptera-Heteroptera of Las Vegas, Hot Springs, New

Mexico, Collected by Messrs E. A. Schwarz and Herbert Barber.
Proceedings of the United States National Museum, Vol. XXVII,
pp. 349-364.
Van Duzee, Edward P.

1907. Notes on Jamaican Hemiptera. Bulletin of the Buffalo Society of

Natural History, Vol. VIII, pp. 3-79.
1909. Observations on some Hemiptera taken in Florida in the spring of 1908.

Bulletin of the Buffalo Society of Natural History, Vol. IX, pp.

149-230.
1914. A Preliminary List of the Hemiptera of San Diego County, California.

Transactions of the San Diego Society of Natural History, Vol. II,

pp. 1-57.
1917a. Check List of the Hemiptera North of Mexico.
1917b. Catalogue of the Hemiptera of America North of Mexico.
1917c. Report upon a collection of Hemiptera made by Walter M. Giffard in

1916 and 1917, Chieflv in California. Proceedings of the California

Academy of Science, Vol. VII, pp. 249-318.
Walker, Francis.

1873. Catalogue of the Specimens of Heteropterous-Hemiptera in the col-
lection of the British Museum, Vol. VI.

AUTHOR INDEX- VOLUME XVIII.

PAGIC

Adkins 5S

Barrows 297

Blake 1 03

Blake and Jackson 145

Bowers. Lyman and 279

BuRRiLi. AND S.Mrrn 229

Claassen ()2

DeLong 22(i, 22N, 233

Drake 174, 323

Fenton 1 77, 243, 2So

FORMAN AND ReED 1

Hine 319

Jackson, Blake and 145

Leonard 33

Lyman and Bowers 279

Xapper 7

Oberholser S5, 213

Pott 129

Reed, Forman and 1

Rice 14

Schaffner 99, 101

Smith, Burrill and 229

Sterki 1 ()9

Walton (5. 12()

WiTHROW (15

333

SUBJECT INDEX TO VOLUME XVIII.

Acad, of Science, Ohio, 14.

Additional Tenn. Cicadellida;, 233.

Additions to Cat. Ohio Vascular Plants
99.

Algag, Eutetramorous Globosus, new
spec, and gen. of, 126.

America, Ravens of North, 213.

America, Relation War to Chem. in, 65.

American Species Teleonemia North of
Mex., 323.

Anteoninae, Parasites of, 177, 243, 285.

Ants of Wisconsin, 229.

Acjuatic Micro-organisms, Axial Rota-
tion of, 6.

Asilus, Description of Seven Species,
319.

Asteriae of Ohio, 33.

Axial Rotation of Acjuatic Micro-organ-
isms, 6.

Bean Meal by Man, Digestibility Sov,
279.

Biology of the Anteoninae, 177, 243, 285.

Catalog, Ohio Vase. Plants, Addit.
to, 99.

Cattle "Temperament" and Its Meas-
ure, 129.

Chemistry in Am., Relation War to, 65.

Cicadellidae from Wisconsin, 228.

Cicadellids, Records of Tenn., 233.

Common Ravens of N. Am., 213.

Concretionary Forms Greenfield Lime-
stone, 7.

Description Seven Species Asilus, 319.

Digestibility Soy Bean Meal bv Man,
279.

Dimorphism in Heterosporous Sporo-
phytes. Sexual, 101.

Dogs, Tumors in, 1.

Empidonax Trailii, Status of, 85.

Eutetramorous Globosus, New Gen.
and Spec. Algse, 126.

Expression Sex. Dimorphism in Hetero-
sporous Sporophytes, 101.

Greenfield Limestone, Concretionary
Forms, 7.

Gynandromorph in Homoptera, Occur-
rence of, 226.

Harmonics of Lecher Syst. (Exper.),
145; (Theor.), 163.

Heteroptera, Ohio, 58.

Heterosporous Sporophytes, Sex. Di-
morphism in, 101.

Homoptera, Occurrence Gynandro-
morph in, 226.

Inland MoUusks, 169.

Leaf-Hoppers, Parasites of: Pt. I, 177-

Pt. II, 243; Pt. Ill, 285.
Lecher Syst., Harmonics of (Exper.).
145; (Theor.), 163.

Lichens of Northern Ohio, 62.

Light on Status of Empidonax Trailii, 85.

Limestone, Concretionarv Forms
Greenfield, 7.

List of Ants of Wis., 229.

List of Ohio Spiders, 297.

Man, Digestibility Soy Bean Meal
by, 279.

Meal by Man, Digestibility Soy Bean,
279.

Measurement, Cattle "Temperament"
and Its, 129.

Micro-organisms, Axial Rotation of, 6.

Mollusks, Inland, 169.

New Light on Status Empidonax
Trailii, 85.

New Spec. Cicadellidag from Wis., 228.

North Am., Ravens of, 213.

North Am. Spec, of Teleonemia, 323.

Northern Ohio, Lichens of, 62.

Occurrence Gynandromorijh in Homop-
tera, 226.

Ohio Acad, of Sci., 14.

Ohio Asteria;, 33.

Ohio Heteroptera, 58.

Ohio, Lichens of Northern, 62.

Ohio Spiders; List of, 297.

Ohio Vase. Plants, Addit. to Cat., 99.

Parasites of Leaf-Hoppers: Pt. I, 177-
Pt. II, 243; Pt. Ill, 285.

Plants, Addition to Ohio Vascular, 99.

Preliminary List Ants of Wis., 229.

Ravens of North America, 213.

Records of Tenn. Cicadellidae, 233.

Relation War to Chemistry in Amer.,65.

Relative Intensity Harmonics of Lecher
Syst. (Exper.), 145; (Theor.), 163.

Sexual Dimorphism, Heterosporous Spo-
rophytes, 101.

Species, Description of Asilus, 319.

Species of Teleonemia, N. Amer., 323.

Spiders, List of Ohio, 297.

Sporophytes, Sex. Dimorphism in Hete-
rosporous, 101.

Study of Cattle "Temperament" and
Its Measure, 129.

Teleonemia, N. Amer. Species of, 323.

"Temperament" and Its Measure.
Study of Cattle, 129.

Tennessee Cicadellidae, 233.

Tingids from the West Indies, 174.

Two New Tingids from the West
Indies, 174.

Tumors in Dogs, 1.

Vascular Plants, Addit. to Cat. Ohio,99.

War to Chem. in Amer., Relation, 65.

West Indies, Tingids, 174.

Wisconsin, Ants of, 229.

Wisconsin. Cicadellidae from, 228.

334